✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY VOL. 61, No. 1, PP. 1-314 FEBRUARY 17, 2025 UNIVERSITY OF FLORIDA GAINESVILLE ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ The FLORIDA MUSEUM OF NATURAL HISTORY is Florida’s state museum of natural history, dedicated to understanding, preserving, and interpreting biological diversity and cultural heritage. The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is an on-line, open-access, peer-reviewed journal that publishes results of original research in zoology, botany, paleontology, archaeology, and museum science. Multi-author issues of related papers have been published together, and inquiries about putting together such issues are welcomed. Address all inquiries to the Editor of the Bulletin. The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online regis- tration system for the ICZN. The ZooBank Publication number for this issue is 55195832-AC66-4098- 96D8-B4E45788E56F. Michal Kowalewski, Editor for this issue Bulletin Committee Jonathan Bloch Jason Bourque Sarah Fazenbaker Alan Franck Robert Guralnick Michelle J. LeFebvre Verity Mathis Larry Page Jerald Pinson Roger W. Portell Pamela Soltis ISSN: 2373-9991 Copyright © 2025 by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, images and other media are for nonprofit, educational, and personal use of students, scholars, and the public. Any commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum. Publication Date: February 17, 2025 This and other issues of the Bulletin can be freely downloaded at: https://flmnhbulletin.com Send communications about this publication to: Editor of the Bulletin; Florida Museum of Natural History; University of Florida; P.O. Box 117800; Gainesville, FL 32611-7800 USA FAX: 352-846-0287; Email: bulletin@flmnh.ufl.edu URL: https://flmnhbulletin.com Cover image: Eupatagus antillarum holotype (left) and Eupatagus mooreanus (right); see caption for figure 99 within text. mailto:bulletin@flmnh.ufl.edu https://www.floridamuseum.ufl.edu/bulletin/home/ ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ Bulletin of the Florida Museum of Natural History vol. 61, no. 1, pp. 1 - 314 Bulletin of the Florida Museum of Natural History Paleogene Echinoids of Florida Adam S. Osborn1, Roger W. Portell2, and Rich Mooi3 11500 Lakeshore Drive, Camden SC, 29020, USA, macropneustes@netzero.com; 2Division of Invertebrate Paleontology, Florida Museum of Natural History, 1659 Museum Road, University of Florida, Gainesville, FL, 32611, USA, portell@flmnh.ufl.edu; 3Department of Invertebrate Zoology and Geology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA, 94118, USA, rmooi@calacademy.org Received: 4-3-2024 Accepted: 8-10-2024 Published: 2-17-2025 Keywords: • Echinoidea; • echinoids; • Florida; • Paleogene; • Eocene; • Oligocene. Sixty-five species and nine indeterminate taxa of Florida Paleogene echi- noids are discussed, and their geographic and stratigraphic distributions provided. These include 49 species documented from the Eocene and 16 from the Oligocene. Ten new species are described: Prionocidaris robertsi n. sp., Rhyncholampas mariannaensis n. sp., Rhyncholampas bao n. sp., Weisbordella inglisensis n. sp., Weisbordella libum n. sp., Durhamella tetrapora n. sp., and Brissus jonesi n. sp. from the Eocene; and Plagiobrissus cassadyi n. sp., Eupatagus dumonti n. sp., and Schiza- ster carlsoni n. sp. from the Oligocene. We reconsidered subjective junior synonyms of all species and resurrect Neolaganum archerensis, Echinocyamus macneili, and Eupatagus mooreanus. Furthermore, we updated the taxonomy for all included species and their known dis- tributions and provide emended diagnoses for the genera and species of Florida Neolaganidae. In addition, we herein report the occurrence of Porpitella micra in Cretaceous strata of the subsurface of Florida. This remarkable finding makes P. micra the earliest known of all the scutelloids. Echinoids within the Ocala Limestone are placed in five echinoid biozones, which are defined within, these include the Oligopy- gus phelani, Oligopygus haldemani, Oligopygus wetherbyi, Wythella eldridgei, and Haimea brooksi Zones. This document complements the Neogene (including the Quaternary) fossil echinoid fauna of Florida we published in 2020 and represents a compilation of the known Florida Paleogene echinoid record. The region is currently known to have the most speciose and diverse assemblage of Paleogene echinoids in the United States. Osborn, A. S., R. W. Portell, and R Mooi 2025. Paleogene echinoids of Florida. Bulletin of the Florida Museum of Natural History 61(1):1-314. https://doi.org/10.58782/flmnh.xqds7462 mailto:macropneustes@netzero.com mailto:portell@flmnh.ufl.edu mailto:rmooi@calacademy.org https://doi.org/10.58782/flmnh.xqds7462 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ TABLE OF CONTENTS Introduction............................................................................................................................2 Materials and Methods...........................................................................................................2 The Eocene.............................................................................................................................3 The Oligocene......................................................................................................................17 Systematic Paleontology ......................................................................................................23 Classification of Species Treated .......................................................................................296 Conclusions........................................................................................................................299 Acknowledgments ..............................................................................................................299 Literature Cited ..................................................................................................................300 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 2 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) INTRODUCTION The importance of echinoids in modern ecosystems underscores the need to understand how their diver- sity has changed through time. These paleontologi- cal benchmarks are crucial if we are to comprehend how echinoids, and indeed entire biomes in which they live, might respond to the global changes we are observing today. Therefore, this work attempts to provide the most accurate and comprehensive accounting to date of the diversity of the Florida Paleogene echinoid fauna. In the process, it demon- strates that the state has the most speciose and diverse assemblage of Paleogene echinoids in the eastern United States. An updated taxonomy for all known Paleo- gene echinoid species of Florida and document of their distributions is long overdue. The fauna in- cludes 49 species from the Eocene and 16 from the Oligocene, as well as nine additional taxa left in open nomenclature at or below genus level. We describe ten new species: Prionocidaris robertsi n. sp., Rhyn- cholampas mariannaensis n. sp., Rhyncholampas bao n. sp., Weisbordella inglisensis n. sp., Weisbor- della libum n. sp., Durhamella tetrapora n. sp., and Brissus jonesi n. sp. from the Eocene; and Plagio- brissus cassadyi n. sp., Eupatagus dumonti n. sp., and Schizaster carlsoni n. sp. from the Oligocene. With the abundance of material now avail- able, we reconsidered subjective junior synonyms for all species, and resurrect Neolaganum archerensis (Twitchell in Clark and Twitchell, 1915); Echinocya- mus macneili Cooke, 1959; and Eupatagus moore- anus Pilsbry, 1914. A review of the distinguishing characteristics of regional genera and species of neolaganids was required to inform taxonomic as- signment of three new species; emended diagnoses are included for these taxa. In addition, we rec- ognize Rhyncholampas lyelli (Conrad, 1850) as a subjective junior synonym of R. conradi (Conrad, 1850), and R. globosus (Fischer, 1951) to be syn- onymous with R. georgiensis (Twitchell in Clark and Twitchell, 1915). We examined hundreds of specimens of Oligocene species of Clypeaster and recognize three distinct forms that do not fall within the variation of any documented taxa. We were un- willing to name new species of this already overly split genus, and identify them as Clypeaster sp. A, B, and C. We also document what is likely the first occurrence of Prenaster in North American strata. In addition, we herein record the occurrence of Porpitella micra H. L. Clark, 1937, in Creta- ceous strata of the subsurface of Florida. This re- markable find makes P. micra the earliest known of all the scutelloids. Echinoids within the Ocala Limestone are placed in five echinoid biozones, which are defined within, these include the Oligopy- gus phelani, Oligopygus haldemani, Oligopygus wetherbyi, Wythella eldridgei, and Haimea brooksi Zones. Lastly, this paper serves as the companion to Osborn et al. (2020) which discussed the occur- rences of 43 species of echinoids from the Florida Neogene (including the Quaternary) and included a historical summary of the echinoid research in the state. MATERIALS AND METHODS The extensive collections in the Florida Museum of Natural History, Invertebrate Paleontology (FM-IP) Collection at the University of Florida (UF) were examined in detail, revealing new species and new stratigraphic records of Florida Echinoidea. Mate- rial cataloged into these collections is indicated by ”UF”, followed by a unique catalog number. We also indicate Florida Museum localities by ”FM-IP”. Other institutions to whose localities or material we refer include the United States Geological Survey (USGS), California Academy of Sciences’ Geology collections (CASG), and University of California Museum of Paleontology (UCMP). We also exam- ined specimens from the Smithsonian Institution’s National Museum of Natural History (USNM). Per- haps most importantly, material borrowed from numerous avocational collectors (see acknowledg- ments) was examined and specimens necessary for the completion of this work were donated to the FM-IP Collections. Unless otherwise indicated, all type and figured specimens are deposited at the FM-IP at the University of Florida in Gainesville, Florida. Standard caliper measurements in millimeters ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 3 include test length (TL), test width (TW), test height (TH), and test diameter (TD). Room Temperature Vulcanizing (abbreviated as ”RTV”) silicone rubber was used to make casts of some moldic specimens, as noted. THE EOCENE With 49 species and 54 distinct taxa (Table 1), Eocene strata of Florida (Fig. 1) contain the high- est diversity of Cenozoic echinoids in the eastern United States (Osborn et al., 2016). The greatest diversity occurs in the Upper Eocene Ocala Lime- stone (OLS), which contains 44 documented species and 49 taxa (Table 1). This is an increase of 12 taxa since the OLS echinoid fauna was last detailed by Osborn et al. (2016). This Upper Eocene peak of echinoid diversity in the region follows a global trend of an Eocene maximum bounded by much lower diversities in the Paleocene and Oligocene (McKinney et al., 1992), and aligns with the conclu- sion of Carter (1987a), who stated that the Upper Eocene (Jacksonian-Priabonian) strata of the south- east contain the most diverse echinoid fauna of the region. Influence Of The Gulf Trough On Echinoid Distribution In The Upper Eocene The boundary between the Gulf Coast and Florida provinces of Carter (1987a) coincides with an area of southern Georgia and the eastern panhan- dle of Florida known as the Gulf Trough. The Gulf Trough is a subsurface geological feature that was scoured by a strong marine current from the Late Cretaceous through the Oligocene (Fig. 2). The Gulf Trough is often referred to as the Suwannee Strait, or Suwannee Channel (e. g. Cheetham, 1963), but Huddlestun (1993) makes a distinction. The Suwan- nee Channel is a very broad feature, functional from the Late Cretaceous through Middle Eocene. The Gulf Trough was a much narrower channel (occupy- ing essentially the same area), and functional from the Middle Eocene to Middle Miocene, at which time the feature became sediment filled. The current that flowed through either Suwannee Channel or Gulf Trough is properly called the Suwannee Cur- rent. The Suwannee Strait is technically a descrip- tive, geographic term referring to the Cretaceous to Miocene marine passage that separated the continen- tal mainland from the shallow, carbonate producing Florida Bank. The Gulf Trough itself is a well- documented, subsurface, stratigraphic feature that has, at times, been implicated in influencing echi- noid distribution in the region. Most Upper Eocene echinoid species are typically restricted to one side or the other (Carter, 1987a). As discussed below, per- haps no taxon better represents this pattern than the oligopygids. Carter (1987a) suggested that the Suwannee Strait did not operate as a physical barrier to echinoid migration during Jacksonian times. He used the modern example of the Florida Strait, which is a deeper and wider barrier than the Suwannee Strait, and therefore presumably had a stronger current. He correctly noted that the Florida Strait do not serve as a barrier in echinoid distributions. Carter (1987a) and Carter and McKinney (1992) suggested these differences can, in part, be explained by stratigraphic mismatching, in which biozones in Florida are not stratigraphically preserved in the strata of Georgia, north of the strait. Therefore, no barrier is necessary to explain the faunal disparities. Carter (1987a) suggested that the most likely cause of the faunal differences between the Late Eocene of the Florida platform and remainder of the Gulf Coast is an environmental distinction, with depth disparities between the depositional environments of these two regions at least in part ac- counting for the contrasting echinoid faunas. Carter (1987a) noted that of those species which occur on both sides of the strait, most do not occur far from it, and in fact many species tend to cluster near the strait [Macropneustes mortoni (Conrad, 1850) and Weisbordella johnsoni (Twitchell in Clark and Twitchell, 1915) for example], suggesting that some environmental gradient straddled the strait. Carter (1987a) concluded that even if better collections of late Jacksonian age strata of Georgia were to reveal species currently restricted to the Florida platform, the differences in the relative abundance of the species between the two regions would remain. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 4 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Table 1: Distribution of Eocene echinoids of Florida. *The stratigraphic occurrence of taxa known only from deep wells is discussed in the remarks for each species herein. Avon Park Fm. OLS O. phelani Zone OLS Haimea brooksi Zone OLS O. haldemani Zone panhandle OLS O. haldemani Zone peninsula OLS O. wetherbyi Zone OLS Spirulaea/ Wythella Zone *From deep Wells Phyllacanthus mortoni (Conrad, 1850) X Prionocidaris robertsi n. sp. Euechinoidea gen. et sp. indet. Acanthechinus dixie (Cooke, 1941) X Brochopleurus pretiosus (Clark, 1915) X Lytechinus floralanus (Cooke, 1941) X unidentified aulodont Amblypygus americanus Michelin, 1856 X Haimea brooksi Osborn et al., 2016 Oligopygus haldemani (Conrad, 1850) Oligopygus phelani Kier, 1967 Oligopygus rotundus Cooke, 1942 Oligopygus wetherbyi de Loriol, 1887 X Eurhodia patelliformis (Bouvé, 1851) X Rhyncholampas conradi (Conrad, 1850) X Rhyncholampas ericsoni (Fischer, 1951) Rhyncholampas fontis (Cooke, 1942) X Rhyncholampas georgiensis (Twitchell, 1915) Rhyncholampas mariannaensis n. sp. Rhyncholampas trojanus (Cooke, 1942) Rhyncholampas bao n. sp. Echinolampas tanypetalis Harper and Shaak, 1974 X Echinocyamus macneili Cooke, 1959 Fibularia vaughani (Twitchell, 1915) X X Durhamella floridana (Twitchell, 1915) X Durhamella ocalana (Cooke, 1942) X Durhamella tetrapora n. sp. Neolaganum archerensis (Twitchell, 1915) Neolaganum dalli (Twitchell, 1915) X Neolaganum durhami Cooke, 1959 Weisbordella cubae (Weisbord, 1934) X Weisbordella inglisensis n. sp. Weisbordella johnsoni (Twitchell, 1915) Weisbordella libum n. sp. ? ? Wythella eldridgei (Twitchell, 1915) Periarchus floridanus Fischer, 1951 Periarchus quinquefarius (Say, 1825) ? ? Protoscutella pentagonium Cooke, 1942 X Gillechinus alabamensis (Cooke, 1942) Ova beckeri (Cooke, 1942) X Ova ocalanus (Cooke, 1942) X Schizaster armiger Clark, 1915 X Agassizia clevei Cotteau, 1875 X X aff. Prenaster sp. Brissopsis steinhatchee Cooke, 1942 X aff. Brissopsis sp. Brissus jonesi n. sp. Plagiobrissus curvus (Cooke, 1942) X X Plagiobrissus dixie (Cooke, 1942) X Macropneustes mortoni (Conrad, 1850) Eupatagus clevei (Cotteau, 1875) X Eupatagus mooreanus Pilsbry, 1914 Eupatagus ocalanus Cooke, 1942 X Eupatagus sp. A Total = 49 species and 54 taxa 2 18 15 14 12 20 13 3 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 5 Figure 1: Generalized Florida geologic map depicting occur- rences of Eocene and Oligocene sediments exposed in outcrop and shallow subcrop (sediments slightly more than 6 m depth). Modified from Scott et al. (2001). Note: in some areas where Eocene sediments only are shown both Eocene and Oligocene sediments may co-occur. Carter and McKinney (1992) noted that litho- logic differences between early Jacksonian strata of Florida and southern Georgia may partially explain the disparity of the faunas. Clastic rocks make up much of the earliest Jacksonian strata of Georgia, these contrasting with the carbonates of Florida. The middle Jacksonian strata of Georgia contain higher proportions of carbonate mud than the more calcareous limestones of the correlative OLS of central Florida. As noted by Carter (1989), most spatangoids prefer, or at least tolerate, significant carbonate mud in the substrate. As an example, the Muckalee Member of the OLS (member of the Williston Limestone per Hud- dlestun, 1981), well exposed near Albany, Georgia, has a higher percentage of carbonate mud than the equivalent Oligopygus haldemani Zone of the OLS of Florida (Carter and Hammack, 1989). Therefore, this mud contains a rich diversity of spatangoids, with only very rare occurrences of neolaganids or oligopygids that characterize the zone in central Florida. Comparison to habitats of laganids and cassiduloids, forms respectively similar to the taxa Figure 2: Approximate placement of the Late Cretaceous to Middle Eocene Suwannee Channel and the Middle Eocene through Middle Miocene Gulf Trough. These are the former areas through which the Suwannee Current flowed. Modified from Bryan et al. (2008). noted above, indicate that their tolerance of fine substrates is much less than that of spatangoids. The substrate preferences of Jacksonian echi- noids of the eastern Gulf Coast are discussed by Carter et al. (1989), and the echinoid faunas are further reviewed by McKinney and Zachos (1986), Carter (1987a, 1989, 1990), Carter and Hammack (1989), and Carter and McKinney (1992). Avon Park Formation The Middle Eocene Avon Park Formation is a largely subsurface unit underlying the OLS in cen- tral Florida (Table 2). Neolaganum dalli (Twitchell in Clark and Twitchell, 1915) is the only identifi- able echinoid documented from surface exposures of this unit and could be collected in abundance in the now inactive Gulf Hammock Quarry (FM- IP LV004, FM-IP LV039), Levy County. However, well cores often bring a profusion of specimens of N. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 6 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Table 2: Correlation of Florida Paleogene stratigraphic units referenced herein, modified from Scott et al. (2001). dalli to the surface (Cole and Ponton, 1932; Vernon, 1951), along with rarer specimens of Neolaganum archerensis (Twitchell in Clark and Twitchell, 1915). The holotypes of N. archerensis and N. dalli were collected by William Healey Dall in cuttings from a deep well near Archer early in the 20th century. Carter (1987a) noted that the depositional environ- ment of the shallow water carbonates of the Avon Park Formation would dictate low diversity and con- tain species able to tolerate ecological conditions not found in other regions, which likely explains the abundance of N. dalli, but the overall sparse fauna. The Avon Park Formation has limited sur- face exposures, largely restricted to two large, ir- regular shaped outcrops in central and southeast- ern Levy County, the latter extending into Cit- rus County along the Withlacoochee River (Ver- non, 1951). Small, irregular outcrops are present along the Withlacoochee River at the Florida Power Corporation dam where the contact of the Avon Park and overlying dolomites of the OLS (portion formerly referred to the Inglis Formation) can be seen. Vernon (1951) described the unconformable contact here evidenced by a rubble and pebble bed at the base of the OLS formed across eroded dolomite beds of the Avon Park Formation. Ivany et al. (1990) documented the occurrence of six poorly preserved (moldic) regular urchin spec- imens on Avon Park Formation seagrass blades from the Dolime Quarry (FM-IP CI009). Unfortunately, none can be assigned with confidence even to family level. Ocala Limestone (OLS) The term Ocala Limestone (hereafter referred to as the OLS) was proposed by Dall (1892) for lime- stone exposures near Ocala, Florida. Cooke (1916) defined it as Jacksonian in age (Upper Eocene) and that it underlies the Marianna Limestone (Table 2). Cooke and Mossom (1929) lumped all exposed Eocene sediments of Florida into the OLS and, for the first time, discussed its fauna at numerous lo- calities throughout the state, affirming the presence of 12 species of echinoid in the unit which were ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 7 first described by Twitchell in Clark and Twitchell (1915) (Cooke and Mossom, 1929:50). Applin and Applin (1944) subdivided the OLS into lower and upper members. Vernon (1951) proposed extending the Moodys Branch Formation from the Gulf Coastal Plain into Florida because of a continuity of biozones from the type locality near Jackson, Mississippi, and the gradual lateral change in facies from clastic to carbonate. For what he stated was Applin and Applin’s (1944) lower member of the OLS, Vernon (1951: 111) proposed the name Inglis Member and Williston Member for the overlying strata. Vernon (1951) therefore proposed restricting the designation of the OLS to only the upper-most portion of the OLS of Applin and Applin (1944). However, as discussed below, others have interpreted Applin and Applin’s lower member to include strata in both Vernon’s Inglis and Williston Members. Puri (1957) attempted to shed some light on this and provided the most complete foraminiferal zonation of the unit to date, asserted the OLS is distinct from the Moodys Branch Formation, raised the OLS to group status, and recognized three for- mations within it, from lowest to highest: the Inglis Formation, the Williston Formation, and a proposed Crystal River Formation (which was initially recog- nized in Puri [1953: 130]) for the upper beds of the unit. Puri (1957) also recognized eight foramaniferal faunal zones within the Ocala Group of Florida. Randazzo (1976) pointed out that the splitting and lumping of the OLS discussed above was done before the inception of the Code of Stratigraphic Nomenclature in 1961, which is published by the American Commission on Stratigraphic Nomencla- ture. Randazzo noted that geologists studying the OLS must identify biostratigraphic horizons rather than lithologic changes to distinguish between the Crystal River, Williston, and Inglis Formations and if the microfossils utilized in Puri’s (1957) zonation are missing or poorly preserved, users cannot appropri- ately identify these formations. He then asserted that splitting the OLS into three formations is not use- ful and misleading to all but a micropaleontolo- gist. Therefore, Randazzo (1976) proposed adopting the United States Geological Survey’s definition of the OLS, which divides the OLS into two mem- bers: a lower member (the Inglis and Williston Formations) and an upper member (the Crystal River Formation). He acknowledged that the litho- logic distinctions between the two members is subtle but proposed the Williston and Inglis Formations be treated as a micritic-skeletal limestone, lower mem- ber of the OLS, and the coarser-grained, skeletal- micritic limestone above these beds (Crystal River Formation) be considered the upper member of the OLS. Miller (1986) reiterated the points made by Randazzo and said Puri’s three formations cannot be recognized lithologically, even at their type sections, nor can they be differentiated in subsurface deposits. Subsequently, Scott (1991) stated the Florida Geo- logical Survey no longer recognized the formational designations of Puri (1957) and reduced the Ocala Group to formational status. The OLS, at this time, is therefore treated as a single lithostratigraphic unit (Table 2) but is informally divided into lower and up- per divisions: the lower OLS, which includes strata formerly included in the Inglis and Williston For- mations and the upper OLS, which includes strata formerly referred to the Crystal River Formation. Scott (1991) stated that the lower subdivision consists of a more granular limestone not present everywhere that may be partially to completely dolomitized in some regions. The upper subdivi- sion is composed of variably muddy (carbonate), granular limestone that is often very soft and friable with numerous large foraminifera. Scott (1991) also included the very late Eocene to early Oligocene Bumpnose Limestone within the upper OLS due to its lithological similarities to that unit. This was followed by Ellwood et al (2019) and we concur. For clarity, the Bumpnose Limestone, which is limited in outcrop to Jackson County, is discussed separately from the OLS in the Oligocene section below. However, as discussed by Randazzo (1976) and Miller (1986), differentiating the lithologies of the lower and upper divisions of the OLS remains difficult. Nevertheless, “finding one’s place” in the unit can be made easier by using index taxa. For instance, the lowermost portion of the OLS in Cit- rus and Levy Counties is a dolomitic horizon, the ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 8 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) lower portion of what was referred to as the Inglis Formation, and it contains characteristic internal and external molds of Eupatagus clevei Cotteau, 1875, and Periarchus floridanus Fischer, 1951. Im- mediately above this is a series of limestones with E. mooreanus and the much less common Oligopy- gus phelani Kier, 1967, of the Oligopygus zonation discussed below. This is the upper portion of what used to be called the Inglis Formation, and because of its echinoid fauna, this zone is also distinctive. Therefore, considering the difficulties differ- entiating the lithologies of the lower and upper divisions of the OLS, we will discuss the strati- graphic distribution of the echinoids within the OLS using Oligopygus zonations. Oligopygus Zonation In The Ocala Limestone Oligopygus de Loriol, 1887 is arguably the most stratigraphically important echinoid genus of the Florida Eocene (Carter, 1990; Croft and Shaak, 1985; McKinney and Jones, 1983; McKinney and Zachos, 1986). The type species is Oligopygus wetherbyi de Loriol, 1887. Oligopygus is known from the southeastern United States and throughout the Middle to Upper Eocene of the Caribbean re- gion as far south as Venezuela (Cooke, 1941b and 1961; Kier, 1967). In his review of the oligopygoids, Kier (1967) recognized 13 species of Oligopygus: O. nancei Cooke, 1941b; O. zyndeli Jeannet, 1928; O. jamaicensis Arnold and Clark, 1927; O. rotundus Cooke, 1942; O. kugleri Jeannet, 1928; O. wether- byi; O. pinguis Palmer in Sánchez-Roig, 1949; O. sanchezi Lambert, 1932; O. costuliformis Jeannet, 1928; O. haldemani (Conrad, 1850); O. putnami Israelsky, 1933; O. curasavica Mollengraaff, 1929, and the aforementioned O. phelani. He considered an additional six species (largely from Cuba) to be too inadequately described or figured, and for which he could not locate holotypes. Cooke (1959) and Kier (1967) recognized four species of Oligopygus in the region: O. rotundus (Middle to Upper Eocene), as well as O. halde- mani, O. phelani, and O. wetherbyi (Upper Eocene). Regional occurrences of the genus are largely con- fined to the OLS of Florida where it is a ubiquitous presence at most exposures and is only very rarely Table 3: Echinoid zonation in the Ocala Limestone of the Florida peninsula. represented elsewhere in the region: O. rotundus also occurs in southeastern Alabama and southwest- ern Georgia, and O. haldemani occasionally occurs in Georgia. Species of Oligopygus are well-documented index taxa for correlation within the OLS of Florida and Georgia (Table 3). Their suitability as index taxa is due to many factors that include: durability of the test, which lends itself to excellent preservation; ease in differentiating the species; abundance of the species within their respective horizons; and constrained zonation of most of the species. The bio- zonation is well documented in Carter (1990), Croft and Shaak (1985), McKinney and Jones (1983), McKinney and Zachos (1986), Zachos and Shaak (1978), and others. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 9 A summary of the zonation discussed in these works is as follows: O. wetherbyi is the youngest species and occurs in the upper portion of the upper OLS (the Crystal River Formation of Puri [1957]); O. haldemani largely occurs in the upper portion of the lower OLS (the Williston Formation of Puri [1957]), though it may extend into the lower portion of the upper OLS [McKinney and Jones, 1983], it does not overlap the range of O. wetherbyi; and O. phelani occurs in the underlying lower OLS (Inglis Formation of Puri [1957]), though not in the lower dolomitic portion of the unit, which contains Eupatagus clevei and Periarchus floridanus in Citrus and Levy Counties (Table 3). Oligopygus rotundus has the most complex stratigraphic distribution of the four regional species of the genus, occurring in the Middle Eocene of Geneva County, Alabama (type locality), Middle Eocene Lisbon Formation of Early County, Georgia (Toulmin, 1977), as well as with O. haldemani in the lower portion of the upper OLS in Jackson County, Florida (Osborn et al., 2016). Due to the lack of continuity in stratigraphic distribution of this species, O. rotundus is not suitable as an index taxon. McKinney and Jones (1983) stated that the overlap of the Oligopygus biozones is minimal and rarely, if ever, exceeds 1 m in thickness. We have not found O. haldemani and the younger O. wetherbyi occurring in situ in the same horizon, nor have we encountered O. haldemani and the older O. phelani occurring in situ together. Although it is feasible that there may be overlap with these two species, other than in mixed spoil in quarries on the Florida peninsula, we have not encountered them together in situ, despite intensive work throughout the region. The only definitive in situ sympatric occurrence of species within the genus occurs in Jackson County where O. rotundus and O. haldemani occur together, with Haimea brooksi Osborn et al., 2016, in the lowest bed currently quarried in the Brooks Quarry (FM-IP JA039), near Marianna (Fig. 3). Features differentiating O. phelani, O. wether- byi, and O. haldemani have very little overlap. The species are easily separated by periproct placement and peristome roundness when in bivariate plots (McKinney and Jones, 1983). McKinney and Jones (1983) did not include O. rotundus in their study. However, it is differentiated from the other three species (see remarks for O. rotundus herein). McK- inney and Jones (1983) also demonstrated that there was not a long-term evolutionary trend of these traits in these species. In other words, in Florida populations specifically, the species transitions are abrupt, with virtually no signs of overlap. The only area of difficulty for Oligopygus zonation appears to be at the upper and lower bound- aries of the unit itself. The issue with O. phelani, as an indicator of the lowermost stratum of the OLS, formerly included in the Inglis Formation, is that it Figure 3: Brooks Quarry (FM-IP JA020), Jackson County, Florida. Arrows indicate top and bottom of Bumpnose Limestone (Eocene/Oligocene boundary is within this unit). Overlying stratum is the Lower Oligocene Marianna Limestone and underlying stratum is the Upper Eocene Ocala Limestone. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 10 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) isn’t very common. However, Eupatagus mooreanus and Periarchus floridanus are ubiquitous (often as test fragments) at nearly every outcrop of the basal limestone of the OLS. Beneath this is a bed of dolomite that forms the base of the unit in Citrus and Levy Counties, with E. clevei and P. floridanus. So, E. mooreanus and P. floridanus serve well as an indicator of the lower portion of the OLS, that is, the Inglis Formation of Puri (1957). At the top of the OLS in northern Peninsu- lar Florida (excluding Jackson County) (Fig. 1) is a soft limestone that lies immediately above the O. wetherbyi Zone and directly below over- lying Oligocene strata. This horizon is, at least in part, the Spirolaea (now Rotularia) vernoni Zone of Puri (1957). Given the absence of Oligopy- gus in this bed, we herein use a different echi- noid and recognize this as the Wythella eldridgei Zone (Table 3). Hunter (1976) noted that Cheetham (1963), intentionally or not, raised the Spirolaea vernoni Zone of the OLS, at least in part, to an equiva- lent of the Lepidocyclina chaperi Zone of Jackson County, which is the Bumpnose Limestone (Fig. 3). Effectively, Cheetham (1963) considered the Bump- nose Limestone to be correlative of Puri’s (1957) Spirolaea vernoni Zone, without naming it. Cheetham’s (1963) correlation likely led to Hunter’s (1972, 1976, 1981) recognition of the Bumpnose Limestone (Puri’s Spirolaea vernoni Zone) in peninsular Florida where it is litholog- ically dissimilar to the type Bumpnose Limestone. However, the Rotularia vernoni Zone is clearly Up- per Eocene, as evidenced by its correlation to the Asterocyclina Zone when Puri (1957) erected his zonation, and reaffirmed by Nicol et al. (1976, 1984, 1989), Jones and Nicol (1989), and others. Bryan (1993) stated that the usage of the term Bump- nose Limestone in peninsular Florida, advocated by Hunter (1972, 1976, 1981), should be discon- tinued. Therefore, within Florida, we recognize the Bumpnose Limestone only in its type area near Mar- ianna (the unit does extend into southern Alabama), and recognize the Wythella eldridgei Zone as the uppermost bed of the OLS in northern peninsular Florida. The Wythella eldridgei Zone is characteristic throughout the northern peninsula, especially in Tay- lor, Lafayette (Fig. 4), Suwannee (Fig. 5), and Dixie Counties, and typically contains an abundance of Wythella eldridgei (Twitchell in Clark and Twitchell, 1915), a diversity of spatangoids including Schiza- ster armiger Clark in Clark and Twitchell, 1915; Brissopsis steinhatchee Cooke, 1942; Ova beckeri (Cooke, 1942); Eupatagus ocalanus Cooke, 1942; Plagiobrissus dixie (Cooke, 1942); Plagiobrissus curvus (Cooke, 1942), and other, rarer species. Lack- ing Oligopygus in this horizon, W. eldridgei serves as an ideal index taxon as it has not been docu- mented outside of this horizon. Above this zone are Oligocene strata. A classic exposure of this horizon is found along the west bank of the Suwannee River west of Dowling Park, as documented by Bryan (1993). Thus, other than the lack (or extreme rarity) of Oligopygus at the base and top of the OLS in peninsular Florida, the Oligopygus zonation dis- cussed above works well for correlating strata in the OLS of peninsular Florida. However, near Marianna, in the Florida panhandle, north of the Suwannee Strait (a feature discussed above), this zonation has drawbacks when compared to the foraminifera zona- tion of Puri (1957). Oligopygus haldemani occurs throughout the upper OLS in Jackson County, with the morphotype attributed by Miller et al. (2014) to O. colsoni Lambert, 1932 occurring up to the top of the OLS in Jackson County without the occurrence of O. wetherbyi between it and the overlying Lower Oligocene portion of the Bumpnose Limestone. However, this occurrence is within the Aste- rocyclina Zone, which Puri (1957) considered to reside at the top of the OLS. This should be the O. wetherbyi Zone according to Oligopygus zonation. It is important to note that the Asterocyclina Zone is not present in peninsular Florida (Puri, 1957) where Asterocyclina is very rare. Contrarily, O. wetherbyi is virtually absent north of the Suwannee Strait, being reported merely as rare specimens from resid- ual float in southwestern Georgia documented by Carter (1989: 190). As noted by Hunter (1976), the Asterocyclina Zone of the Florida panhandle has no recognized peninsular equivalent. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 11 Figure 4: Dell Limerock Mine (FM-IP LF001), Lafayette County, Florida. Exposed rocks are the Oligopygus haldemani and Oligopygus wetherbyi Zones of the Upper Eocene Ocala Limestone. This situation implies several possibilities. It could be that Puri’s assertion that the Asterocyclina Zone is the uppermost zone of the OLS is incorrect, with the Asterocyclina Zone marking the uppermost horizon of OLS that is present only in Jackson County, Florida (horizons above it implied to be absent). This is a possibility, as Puri (1957: 55) stated that the top of the Asterocyclina Zone is marked by an unconformity between it and the overlying latest Eocene to lowermost Oligocene strata (Lepidocyclina chaperi Zone, which resides within the Bumpnose Limestone near Marianna). Another possibility is that O. haldemani, while occurring stratigraphically below O. wether- byi in the Florida peninsula, appears either coeval to it, with O. wetherbyi absent in Jackson County, or above it north of the Suwannee Strait. This seems unlikely. Yet another prospect was discussed by Miller et al. (2014) when they asserted that populations of O. haldemani in Jackson County are O. colsoni (they proposed reinstating Lambert’s [1932] taxonomic assignment for these specimens). This assertion removes the stratigraphic ambiguity discussed above, as O. colsoni would then occur above the Oligopygus wetherbyi Zone, removing the conflict between the Oligopygus zonation and the foraminiferal zonation of Puri (1957). Miller et al. (2014) revised the Oligopygus zonation to proceed as follows, from earliest to latest: O. phelani, O. haldemani, O. wetherbyi and finally O. colsoni. This would be tenable if O. haldemani and O. colsoni could be consistently distinguished, which Cooke (1942, 1959) and Kier (1980) asserted was not possible. This issue is treated in detail below in the entry for O. haldemani. It could be that the zonation (sensu Carter, 1990; Croft and Shaak, 1985; McKinney and Jones, 1983; McKinney and Zachos, 1986) is best applied only south of the Suwannee Strait, where it has only been traditionally useful anyway, given the dearth of documented occurrences of O. wetherbyi and O. phelani north of the Suwannee Strait. Recognition of O. colsoni as a distinct species would only be useful ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 12 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 5: Denali Quarry (FM-IP SU003), Suwannee County, Florida. Exposed rocks are the Oligopygus wetherbyi and Wythella eldridgei Zones of the Upper Eocene Ocala Limestone. if it is readily distinguishable from O. haldemani, though even Miller et al. (2014) admitted that it is not. If O. colsoni is indeed a distinct but cryptic species, it could serve as an indicator for Miller et al.’s (2014) zonation scheme, although other explanations for the stratigraphic placement of O. haldemani are plausible, as discussed above (also, see Table 3). An alternative solution is to consider that the Oligopygus zonation used south of the Suwannee Strait cannot be applied in northern Florida. This is comparable to the foraminiferal zones of Puri (1957), which are not repeatable in their entirety south of the Suwannee Strait where Asterocyclina is no longer a dominant species of large foramin- fera (Puri, 1957; Hunter, 1976). This would imply that Puri was incorrect in replicating the northern foraminiferal zonations to peninsular Florida with- out differentiating between the two regions. Indeed, a zone of Asterocyclina does not form the uppermost bed of the OLS in peninsular Florida, where the soft pale limestone containing W. eldridgei, discussed above, occupies the uppermost portion of the unit. McKinney (1984) and Rice (1997) discussed the ontogeny and heterochrony of O. phelani, O. haldemani, and O. wetherbyi. McKinney (1984) studied this apparent lineage and considered it an excellent opportunity to examine the relationships between heterochrony and environmental change. Each species is similar to, but larger than, the pre- ceding one. He asserted that the morphological differences among these species may not have re- sulted from selection directed at those traits but were simply a secondary result of size increase or of changes that occurred to permit the size increase. McKinney’s assertion that each species is larger than the preceding one does not hold true when O. rotundus is added to the argument, since the latter is not smaller than O. haldemani. McKinney (1984) determined that charac- teristic test shape, peristome shape, and periproct position of the larger species are generally either simple extensions of preexisting ontogenetic trajec- tories related to size increase or result from other heterochronic processes that change the trajectory in such a way as to permit the larger size. Rice (1997) asserted that many of these changes occurred be- cause of changes in development, not just at the rate which the development proceeded. How a change in rate is not a change in development is not abundantly clear, and in this light, Rice (1997) points to issues ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 13 with the term heterochrony itself. Given that hete- rochrony is also defined in terms of ancestors and descendants, it is curious that no one has recognized the logical step of trying to discover phylogenetic re- lationships among species of Oligopygus. However, neither Rice nor the authors whose work he dis- cusses (notably McKinney, 1984) tried to perform phylogenetic analyses on the echinoids in question. Therefore, the value of heterochronic studies involv- ing Oligopygus has yet to be assessed on either side of the argument. What is clear, however, is that the species used to differentiate the Oligopygus zones have distinct identities and stratigraphy. Thus, herein, echinoid distribution of the OLS will be indicated based on the Oligopygus zonation discussed above: Oligopygus phelani Zone (lowermost OLS/Inglis Formation; perhaps best in- dicated by presence of E. mooreanus); Oligopy- gus haldemani Zone (upper portion of the lower OLS/Williston Formation, lowermost Crystal River Formation); Oligopygus wetherbyi Zone (upper OLS/Crystal River Formation); and Wythella el- dridgei Zone (top of OLS in peninsular Florida) (Table 3). This applies to peninsular Florida. The upper OLS in Jackson County, from the upper contact with the overlying Bumpnose Lime- stone, down to the Haimea brooksi Zone dredged up to 33 m below the top of the unit in the Brooks Quarry (FM-IP JA039) (Fig. 6), will be discussed dis- tinctly as either residing in the Haimea brooksi Zone (the most characteristic element of the lowermost strata quarried in Jackson County), or the overlying limestone better characterized by the presence of O. haldemani, all of which contains an abundance of Asterocyclina. The base of the Haimea Zone was not reached during quarrying in the Brooks Quarry, nor is it visible in Jackson Blue Spring, so the lower boundary of the unit is unknown. Care will be taken to be precise with the stratigraphic placement of species in Jackson County, given the number of new taxa, and its disparity from the echinoid fauna of peninsular Florida. Echinoid Distribution In The Ocala Limestone Although the OLS extends beyond Florida, into southwestern Georgia (Huddlestun and Hetrick, 1986) and southeastern Alabama (Toulmin, 1977), the greatest diversity of echinoids within the OLS is undoubtedly in Florida. Osborn et al. (2016: Table 2) documented 36 species of echinoid in the OLS of Florida, and herein we document 49 species and 54 distinct taxa in the unit (Table 1). The OLS is nearly always richly fossiliferous, and along with the characteristic foraminifera (Puri, 1957, discussed above) echinoids are also exceptionally important to the zonation of the unit within Florida (Table 3). Echinoid distribution in the Oligocene portion of the Bumpnose Limestone Member of the OLS is discussed below in the remarks for the Bumpnose Limestone. As shown, the uppermost portion of the OLS in peninsular Florida consists of a soft lime- stone, above the Oligopygus wetherbyi Zone, that is characterized by the presence of W. eldridgei, as well as Rhyncholampas trojana (Cooke, 1942) (which also appears to be restricted to this horizon), and a diversity of spatangoids including S. armiger; B. steinhatchee, O. beckeri, E. ocalanus, P. dixie, P. curvus, other, rarer species (Table 1). This zone is well exposed in Lafayette and Dixie Counties, as far west as Taylor County (Martin Marietta Perry Quarry), and as far south as Sumter County where the horizon is intermittently exposed in quarrying in the Cemex Quarry at Center Hill (FM-IP SM010). As noted above, this horizon is herein recognized as the Wythella eldridgei Zone (Table 3) and is overlain by Oligocene age strata wherever it occurs. Underlying the Wythella eldridgei Zone in peninsular Florida is the Oligopygus wetherbyi Zone, which is well exposed at many localities throughout the Ocala Arch region of central Florida (see occurrences in the discussion for O. wetherbyi). Oligopygus wetherbyi, Weisbordella cubae (Weis- bord, 1934), R. conradi, Amblypygus americanus Michelin, 1856, Ova ocalana (Cooke, 1942), and the rarer Echinolampas tanypetalis Harper and Shaak, 1974 (which appears to be restricted to this horizon), best characterize the unit, although additional species are present (Table 1). Within peninsular Florida, A. americanus appears to be restricted to the O. wetherbyi Zone, although in Jackson County it occurs in the uppermost OLS and Eocene portion ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 14 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 6: Brooks Quarry (FM-IP JA039), Jackson County, Florida. Exposed rocks are the Haimea brooksi Zone of the Upper Eocene Ocala Limestone, quarried by dragline below water level at a depth of up to 24 m, which is 29-30 m below the top of the Eocene exposures within the quarry. of the overlying Bumpnose Limestone, exposed in the Brooks Quarry (FM-IP JA039) (Fig. 3). The Oligopygus haldemani Zone, which underlies the Oligopygus wetherbyi Zone in peninsular Florida, often contains an abundance of Durhamella ocalana (Cooke, 1942), Durhamella floridana (Twitchell in Clark and Twitchell, 1915), W. cubae; Agassizia clevei Cotteau, 1875, Eurhodia patelliformis (Bouvé, 1851), R. conradi, and Phyl- lacanthus mortoni (Conrad, 1850), which is often represented by an abundance of spines and test frag- ments, in addition to many rarer species (Table 1). Other than O. haldemani, E. patelliformis appears to be the only echinoid restricted to this horizon in Florida. However, it occurs in Alabama and Georgia where it is not associated with O. haldemani, but Oligopygus is either absent or exceedingly rare in those exposures, so the Florida Oligopygus zonation does not apply. This zone is well exposed in many quarries in Florida, including the expansive Haile quarry complex in Alachua County (FM-IP AL004) (Fig. 7). The Oligopygus phelani Zone underlies the Oligopygus haldemani Zone and occupies the se- quence of limestone in the upper portion of what was formerly recognized as the Inglis Formation (Puri, 1957), largely exposed in Citrus and Levy Counties. Limestone exposures in this area are often richly fossiliferous. As discussed above, although O. phelani is typically rare, E. mooreanus (formerly ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 15 Figure 7: Limestone Products Quarries (FM-IP AL004), part of the Haile Quarries Complex, Alachua County, Florida. Exposed rocks are from the Upper Eocene Ocala Limestone. referred to Eupatagus antillarum Cotteau, 1875) is abundant at most exposures. The echinoid fauna of this unit was reviewed by Fischer (1951). although at the time this limestone was considered to belong to the Moodys Branch Formation. Fischer (1951) stated that some of the best-preserved echinoids in Florida were collected from this stratum, and while echinoids are often the most distinctive elements of the fauna and very well preserved, they are no better preserved than those of the upper OLS. Fischer (1951) documented 15 taxa in the Inglis Member (= Oligopygus phelani Zone), with most from the Withlacoochee River area. He named three new taxa: Periarchus lyelli floridanus (= P. floridanus), Cassidulus ericsoni [= R. ericsoni (Fischer, 1951), and Cassidulus globosus (= R. georgiensis). We herein document 18 taxa within this stratum (Table 1). This unit, formerly recognized as the Inglis Formation, consists of two distinct facies, an upper limestone (Oligopygus phelani Zone) and lower dolomitic facies, which, as discussed above, contains E. clevei and P. floridanus. The upper limestone is well-exposed along the Cross Florida Barge Canal south of Inglis (FM-IP CI003, CI012, CI015, CI020- CI022, CI038), especially west of HWY 19/98, the Withlacoochee River west of Inglis (FM-IP CI052, FM-IP LV021, FM-IP LV101, FM-IP LV104, FM-IP LV106), spoil islands offshore at the mouth of the Withlacoochee River (FM-IP CI004, FM-IP CI024, FM-IP LV035), and numerous small quarries in Levy County. Eupatagus mooreanus and P. floridanus are ever-present within the limestone horizon, with some lenses composed largely of fragmented tests of P. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 16 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) floridanus. This is the Oligopygus phelani Zone and a search of fine sediment and gravels often reveals an abundance of smaller species, especially O. phelani, Fibularia vaughani Twitchell in Clark and Twitchell, 1915, Weisbordella inglisensis n. sp., Durhamella tetrapora n. sp., D. floridana, and D. ocalana. The later two Durhamella have tests typically smaller than examples in the overlying Oligopygus haldemani Zone of the upper OLS. The lower dolomitic zone of the lower OLS is well-exposed along the Cross Florida Barge Canal south of Inglis, as well as the CEMEX and Red Level quarries (FM-IP CI014) south of Inglis. This zone typically contains internal and external molds of echinoid tests, but includes sporadic concentrations of the large spatangoid E. clevei and lesser num- bers of P. floridanus, and Rhyncholampas sp. It is interesting that when Fischer (1951) completed his detailed review of the echinoid fauna of the Inglis Member, he noted that a single specimen of E. clevei was collected. This specimen, which he figured, is beautifully preserved, retains its calcitic test, and was obtained from the limestone horizon along the Withlacoochee River. It appears that Fischer likely did not examine the dolomitic horizon, otherwise he would have noted the abundant fossils of this species which, although moldic, are nonetheless recognizable as E. clevei. The stratigraphy of the Inglis Formation was discussed by Puri (1957), Vernon (1951), Puri and Vernon (1964), and especially Puri (1970). The molluscan fauna was reviewed by Richards and Palmer (1953), the ostracods by Swain (1946), and the foraminifera by Puri (1957). The echinoid zonation reviewed above applies to peninsular Florida (south of the Suwannee Strait) (Table 3). However, north of the Suwannee Strait, in Jackson County, the OLS has a distinctly different fauna than that of peninsular Florida and contains species more commonly found in the muddier facies of the OLS of Georgia, as well as some elements of peninsular Florida. In the mining operation of Leon Brooks (FM-IP JA0009, (FM-IP JA018, FM- IP JA027, FM-IP JA031, FM-IP JA039), northwest of Marianna in Jackson County (Fig. 3), the OLS is overlain by the Bumpnose Limestone (which transi- tions across the Eocene and Oligocene boundary and is further discussed in the Oligocene section herein) and nearly 30.5 m of the OLS are quarried, with O. haldemani and the distinctive, large, star-shaped foraminiferan, Asterocyclina, occurring throughout the entire OLS sequence. The geology of Jackson County was reviewed by Moore (1955) and further discussed by Bryan et al. (2008) and Herbert (2012). Herbert (2012: 11) examined the uppermost bed of the OLS in the Brooks Quarry, and Bryan et al. (2008: 90) figured the OLS/Bumpnose contact (Fig. 3), as well as the overlying Bumpnose/Marianna contact. Diagnostic Upper Eocene echinoid species, such as Amblypygus americanus and Rhyncholampas conradi, occur in the lower (Upper Eocene) portion of the Bumpnose Limestone that were obtained in situ above the OLS/Bumpnose contact. Macropneustes mortoni, P. curvus, R. con- radi, Weisbordella johnsoni (Twitchell in Clark and Twitchell, 1915), O. haldemani, A. clevei, and other, rarer species are common in the upper 3 m of OLS in the Brooks Quarry. Below this horizon is a less fossiliferous zone that extends for at least 10 m and contains little except O. haldemani and W. johnsoni. In the northwestern most excavations within the Brooks Quarry complex (FM-IP JA039), the limestone brought up from the greatest depth with a dragline (˜29-30.5 m below the OLS/Bumpnose contact, Fig. 6) contains an interesting fauna that includes H. brooksi; O. rotundus, O. halde- mani, R. georgiensis, W. johnsoni, the herein described Rhyncholampas mariannaensis n. sp., Rhyncholampas bao n. sp., Brissus jonesi n. sp., and numerous rarer taxa (Osborn et al., 2016) (Table 1). This fauna not only includes numer- ous taxa seldom documented elsewhere, such as H. brooksi and the herein described B. jonesi n. sp., R. mariannaensis n. sp., and R. bao n. sp., but an exceptionally diverse echinoid fauna that is perhaps best characterized by the presence of H. brooksi. The presence of the latter species distinguishes it from the overlying strata that also contains O. haldemani, but none of the new taxa listed above, all of which appear to be restricted to the Haimea brooksi Zone. This horizon, with H. brooksi, is also exposed ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 17 Table 4: Distribution of Oligocene echinoids documented from Florida. Taxon Bumpnose Limestone Marianna Limestone Suwannee Limestone Bridgeboro Limestone Phyllacanthus mortoni (Conrad, 1850) X X X Phymotaxis mansfieldi Cooke, 1941 X Gagaria mossomi (Cooke, 1941) X Lytechinus floralanus (Cooke, 1941) X Clypeaster cotteaui Egozcue, 1897 X Clypeaster marinanus Jackson, 1937 X X Clypeaster oxybaphon Jackson, 1922 X Clypeaster rogersi (Morton, 1834) X Clypeaster sp. A X Clypeaster sp. B X Clypeaster sp. C X Rhyncholampas gouldii (Bouvé, 1846) X Echinolampas aldrichi Twitchell, 1915 X Schizaster americanus Clark, 1915 X X X Schizaster carlsoni n. sp. X Agassizia mossomi Cooke, 1942 X X X Brissus bridgeboroensis Carter, 1987 X Plagiobrissus cassadyi n. sp. X Eupatagus dumonti n. sp. X X Eupatagus sp. B X Total Species= 16; total taxa= 20 1 6 14 7 deep within the underground cavern system at Jack- son Blue Spring, east of Marianna, Jackson County (FM-IP JA033). Here, at over 33 m depth, certified cave divers collected H. brooksi, R. georgiensis, W. johnsoni, the herein described B. jonesi n. sp., Pri- onocidaris robertsi n. sp. (abundant spines of this species are present), and rarer taxa, including O. beckeri. Neither excavation in the Brooks Quarry or exposures in Jackson Blue Springs expose the base of the Haimea brooksi Zone, so the echinoid fauna of this portion of the OLS, as well as correlation of the Haimea brooksi Zone and the Oligopygus phelani Zone of peninsular Florida, is unknown. However, within Jackson County, the Haimea brooksi Zone is the lowest exposed portion of the OLS, residing beneath the zone of abundance of Oligopygus haldemani. THE OLIGOCENE The regional echinoid fauna experienced a dra- matic drop in diversity at the start of the Oligocene (Table 4), which was likely the result of a global- scale deterioration of the physical environment at the Eocene/Oligocene boundary. The ultimate cause of this deterioration is inferred to be climatic cooling in the Early Oligocene that was perhaps associated with the isolation of Antarctica (McKinney et al., 1992). This deterioration was most evident in the Early Oligocene, but likely continued through most of the epoch (see Prothero et al., 2003). Whereas the Middle to Upper Eocene strata of the region saw the extensive evolution of the protoscutellids (which includes many of the most important Claibornian and Jacksonian Stages index taxa within the region), the Early Oligocene saw the disappearance of such forms, and the proliferation of the clypeasteroids and other scutelloids in the region throughout the Oligocene. The greatest diversity of Florida Oligocene echinoids occurs in the Suwannee Limestone, which contains 14 taxa (Table 4). The most widespread unit in the southeastern United States is the Marianna Limestone, which comprises a thick sequence of soft limestone that extends from the Mississippi River southeastward to its type area in Jackson County (FM-IP JA084) in northern Florida. The Marianna Limestone contains the characteristic ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 18 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) species Clypeaster rogersi (Morton, 1834) in most exposures throughout the region, and within Florida, the Marianna Limestone contains six taxa (Table 4). McKinney et al. (1992) stated that Lower Oligocene strata of the region contained only Cy- claster drewryensis Cooke, 1942 from the Red Bluff Formation of southwestern Alabama. However, the Early Oligocene fauna is now considered to be more diverse, though only slightly, than previously under- stood (Table 4). Two basal Oligocene units contain echinoid faunas within the region: the Red Bluff Formation of southeastern Mississippi and south- western Alabama, and the Bumpnose Limestone of southeastern Alabama and northern Florida, espe- cially Jackson County. The Bridgeboro Limestone crops out in southwestern Georgia, northern Florida, and southeastern Alabama, and is currently known to contain at least seven echinoid species within Florida (Table 4). This fauna is further discussed below. Additional Oligocene units occur in the region but do not contain documented echinoids or have very sparse assemblages. The Oligocene portions of the Arcadia Formation of southern Florida do not contain any documented echinoids, although the Miocene portions of the unit does [see Osborn et al. (2020)]. Bumpnose Limestone The Bumpnose Limestone is best exposed near its type area in Jackson County, where it is exposed in its entirety in the Brooks Quarries, north- west of Marianna (FM-IP JA021, FM-IP JA025, FM-IP JA030) (Fig. 3). When Moore (1955) named the Bumpnose Limestone, he believed it to be upper- most Jacksonian in age, but it is now considered to straddle the Eocene and Oligocene Boundary (Ell- wood et al., 2019) (Table 2). At the Brooks Quarry, the unit is approximately 3.5 m thick (Herbert, 2012: 11). MacNeil (1944), Moore (1955), Huddlestun and Toulmin (1965), Miller (1986), and others noted that the Bumpnose Limestone correlates with the basal Oligocene Red Bluff Clay in Alabama (which also contains Clypeaster marinanus Jackson, 1937). Miller (1986) stated that gulfward, the Bumpnose Limestone merges with a thick sequence of unnamed limestone and finely crystalline dolomite. The unit pinches out southwestward in western Bay County. Without a doubt, the best exposures of the typically glauconitic Lepidocyclina chaperi-packstone of the Bumpnose Limestone are found near the type area in Jackson County. As noted above, Cheetham’s (1963) corre- lation likely led to Hunter’s (1972, 1976, 1981) recognition of the Bumpnose Limestone (for Puri’s Spirolaea vernoni Zone) in peninsular Florida. How- ever, the vernoni Zone is considered Upper Eocene, as evidenced with its correlation to the Asterocy- clina Zone when Puri (1957) erected his zona- tion, and reaffirmed by Nicol et al. (1976, 1984, 1989), Jones and Nicol (1989) and others. Bryan (1993) stated the usage of the term Bumpnose Lime- stone in peninsular Florida should be discontinued. However, the Bumpnose in Jackson County does straddle the Eocene/Oligocene boundary, so perhaps the equivalency is merely in the Eocene portion. The echinoids of the Rotularia vernoni Zone (Wythella eldridgei Zone) are Eocene in age, as discussed in the OLS section above. Therefore, within Florida, we recognize the Bumpnose Limestone only in its type area near Marianna (the unit does extend into southern Al- abama). The geology of Jackson County is reviewed by Moore (1955) and further discussed by Bryan et al. (2008), Ellwood et al. (2019), and Herbert (2012). Scott (1991) stated the Florida Geological Sur- vey no longer recognized the Bumpnose Limestone as a distinct formation, and due to its lithologi- cal similarities treated it as the uppermost bed of the OLS (effectively extending the OLS into the Oligocene). Ellwood et al. (2019) recognized the Bumpnose Limestone as the upper member of the Ocala Limestone. Huddlestun (1993) stated the Bumpnose is “Ocala-like” but glauconitic and clarified that there are no known deposits of Bumpnose Limestone correlative strata outside of the Gulf Trough in Georgia. Bryan et al. (2008) and Ellwood et al. (2019) figured the contact with the Bumpnose Limestone and underlying OLS in the Brooks Quarries (FM- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 19 IP JA018-JA021), as well as the contact with the overlying Marianna Limestone. Both contacts are sharp, and form indurated ledges that are visible as thick white bands in the quarry face. The Bumpnose Limestone therefore occurs between these two pale ledges. The Bumpnose Limestone does not appear to exceed 5 m in thickness in Jackson County (Moore, 1955). The Bumpnose Limestone is characterized by the large foraminiferan, Lepidocyclina chaperi Lemoine and Douville, 1904, and within Jackson County, the underlying OLS and lowermost Bump- nose Limestone are characterized by the large star- patterned foraminiferan, Asterocyclina. The upper indurated bed of the Bumpnose Limestone is glau- conitic, which is also characteristic of the unit in Jackson County. Just above the pale indurated ledge at the OLS/Bumpnose contact, the Bumpnose Lime- stone in the Brooks Quarry complex contains an often nodular, semi-indurated limestone with Amu- sium ocalanum Dall, 1898, and the characteris- tic Eocene echinoids Amblypygus americanus, and Rhyncholampas conradi, this being the Eocene por- tion of the Bumpnose Limestone. The remainder of the overlying lowermost Oligocene-portion of the Bumpnose Limestone con- sists of Lepidocyclina packstone with L. chaperi, Aturia alabamensis (Morton, 1834), Spondylus du- mosus, and other mollusks, as well Clypeaster mari- nanus, which is the only known echinoid in the Oligocene portion of the Bumpnose Limestone (Ta- ble 4). The overlying Marianna Limestone contains Clypeaster rogersi. Moore (1955: 38) stated that he asked C. Wythe Cooke to examine specimens of Clypeaster from the Bumpnose Limestone, and that Cooke deter- mined they represent a new species closely allied to C. rogersi. However, Cooke (1959) did not describe the taxon. Huddlestun (1993) also noted the Bump- nose Limestone contains an undescribed species of Clypeaster he assumed was likely ancestral to C. rogersi. We recognize this form with typically shorter petals and a thinner margin than C. rogersi as Jackson’s (1937) C. marinanus. Moore (1955) also stated the Bumpnose contains a new species of Anisopetalus (= Rhyncholampas). However, we have been unable to verify this occurrence (it is possible he was referring to large specimens of R. conradi, which occur in the Eocene portion of the lower Bumpnose Limestone). Herbert (2012) stated that the Bumpnose Limestone in the Brooks Quarries (FM-IP JA021, FM-IP JA025, FM-IP JA030) con- tains C. rogersi, which is likely a misidentification of C. marinanus, as we have only found C. rogersi in the overlying Marianna Limestone in this quarry. Marianna Limestone The Marianna Limestone was named by Mat- son and Clapp (1909) from exposures at Marianna in Jackson County (FM-IP JA084), which he defined as a soft, porous, light gray to white limestone character- ized by an abundance of Orbitoides mantelli Morton, 1833 (= Lepidocyclina mantelli). Cooke and Mos- som (1929) clarified that Matson and Clapp’s (1909) Marianna Limestone included strata then recognized within the OLS and refined understanding of the unit as the “white limestone or chimney rock” that over- lies the OLS at Marianna and containing L. mantelli and Pecten poulsoni Morton, 1834. The subsequent recognition of the Bumpnose Limestone by Moore (1955) modified Cooke and Mossom’s definition slightly and the Marianna Limestone of northern Florida is now recognized as residing above the Up- per Eocene-Lower Oligocene Bumpnose Limestone and below the Chattahoochee Formation in Jackson County, and Bridgeboro Limestone in Washington County at Duncan Church (FM-IP WG002). The Marianna Limestone is now considered to be Early Oligocene (Rupelian) and extends north- eastward from the type area to Pulaski County, Geor- gia (Huddlestun, 1993), and northwestward across southern Alabama and Mississippi to the Mississippi River, where the unit thins substantially (Dockery, 1982). The lithology of the Marianna in Georgia, northern Florida, Alabama, and eastern Mississippi is the typical, soft white limestone described by Matson and Clapp (1909), with C. rogersi being the characteristic echinoid of the unit (Dockery, 1980; 1982). In Jackson County, Schizaster americanus Clark in Clark and Twitchell, 1915 is also found with persistent collecting. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 20 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) South of Marianna, in southern Jackson County, quarries (e.g., FM-IP JA014) and natural exposures (e.g., FM-IP JA010), provide glimpses of a dolomitic limestone with a diverse echinoid fauna indicative of the Marianna Limestone, but the lithology is decidedly not like the soft pale lime- stone of the Marianna Limestone several kilometers northward at its type locality. A collectable expo- sure of this horizon is seen during low water levels along the banks of Dry Creek (FM-IP JA010) south- west of Marianna. Here, the tan to brown dolomitic limestone is variably indurated and contains an abun- dance of Lepidocyclina sp. and a diverse echinoid fauna preserved as internal and external molds, in- cluding abundant specimens of the characteristic Marianna Limestone species, Clypeaster rogersi and Schizaster americanus, as well as Plagiobrissus cas- sadyi n. sp., rarer Agassizia mossomi Cooke, 1942, and Phyllacanthus cf. P. mortoni. A quarry north of Altha (FM-IP JA014) provided extensive exposures of this dolomitic horizon, with an abundance of C. rogersi, Echinolampas aldrichi Twitchell in Clark and Twitchell, 1915 (the first documented occur- rence of this species in Florida), and S. americanus. Moore (1955) mapped the Suwannee Lime- stone in this area of Jackson County and described the unit here as tan to bluff colored limestone and dolomitic limestones residing above the Marianna Limestone and below the Tampa Limestone (now considered Chattahoochee Formation). This stratum is therefore certainly that which Moore (1955) re- ferred to the Suwannee Limestone. However, the lithology is decidedly not synonymous with the current understanding of the Suwannee Limestone (Huddlestun, 1993). Spencer and Loyd (1999: 10) also erroneously stated that the dolomitic limestone south of Marianna is Suwannee Limestone. Green et al. (2002) and Rupert and Means (2009) noted that the unit mapped as Suwannee Limestone in this area of Florida is likely either Bridgeboro Limestone or undifferentiated Mari- anna/Bridgeboro Limestone in cross sections. Bryan et al. (2008: 92) stated that south of Marianna the Marianna Limestone is a dolostone, as exposed in the quarries north of Altha. Clearly there is no consensus, but we include the Dry Creek echinoid assemblage in the Marianna Limestone in Table 4 with the understanding that the lithology is decid- edly distinct from either the typical Marianna or Suwannee Limestones. Suwannee Limestone The Suwannee Limestone is the dominant Oligocene unit of peninsular Florida. The name Suwannee Limestone was proposed by Cooke and Mossom (1929) for yellowish limestone ex- posed along the Suwannee River from Ellaville nearly to White Springs in northern Florida. This stratum had previously been referred to as the Hawthorn Formation (Matson and Clapp, 1909) and Tampa Limestone (Cooke and Mossom, 1929). Huddlestun (1993) made a persuasive argument for including the unit entirely within the Early Oligocene (Rupelian), which was validated by Brewster-Wingard et al. (1997) and Scott (2001). In peninsular Florida, the Suwannee Limestone crops out on the northwestern, northeastern, and southwestern flanks of the Ocala Platform and is absent on the eastern coast, either due to erosion, non-deposition, or both (Scott, 2001). Although Huddlestun (1993) recognized a dolostone directly below the typical Suwannee Limestone near Ellav- ille as a distinct unit, the Suwannee Limestone does have dolomitized and silicified beds throughout its distribution. Huddlestun (1993) refined the Suwannee Limestone by removing a few distinct mappable lithologies, several of which contain echinoids and are discussed below, especially near Ellaville, in northern Florida. This clarified the unit, and by in- cluding only the uppermost limestone beds along the Suwannee River at Ellaville within the Suwannee Limestone, Huddlestun (1993) revised the concept of the unit to be compatible with much of the lime- stone of west central and northern Florida that has traditionally been included within the Suwannee Limestone. In the type area of the Suwannee Lime- stone, and throughout its extent from southern Geor- gia to west central Florida, the unit can be described as a very pale orange, massive-bedded and structure- less to poorly but thickly stratified, coarse-grained, variably soft and hard limestone that is sporadically ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 21 richly macrofossiliferous (Huddlestun, 1993). Huddlestun (1993) erected the Ellaville Lime- stone for Lower Oligocene limestone that crops out along the Suwannee and (lower only) Withla- coochee River in Madison, Hamilton, and Suwan- nee Counties, and is present in the subsurface of southwestern Georgia. At the type locality of the unit, along the Suwannee River near Ellaville in Hamilton County, Florida, the Ellaville Lime- stone is a lithologically nondistinctive, sparsely but variably macrofossiliferous, relatively pure, mod- erately indurated limestone with minor compo- nents of dolomite and dolostone. Intraformational breccia (with mud rip-up clasts) occur in the Ellav- ille, indicating peritidal conditions. Cooke and Mossom (1929: 72, 73) referred these beds at the type locality of the Ellaville Lime- stone to the Glendon Limestone. Cooke (1945) in- cluded this stratum in the Byram Limestone, and Hunter (1972) referred this unit to the Bumpnose Limestone. The Ellaville Limestone overlies the Upper Eocene OLS and is overlain by the Lower Oligocene Suwannacoochee Dolostone (of Huddlestun, 1993) which gradationally and conformably overlies the Ellaville Limestone at Ellaville, Florida. Huddlestun (1993) erected the Suwannacoochee Dolostone for lithologically distinctive dolostone around the (lower only) Suwannee and Withlacoochee Rivers in Madi- son, Hamilton, and Suwannee Counties, Florida, as well as in the subsurface of southern Georgia. The type locality is along the Suwannee River at Ellaville, Florida where it immediately overlies the stratotype Ellaville Limestone. This dolostone has traditionally been included within the Suwannee Limestone. How- ever, Huddlestun (1993) proposed excluding it from that unit due to its distinctive, mappable, dolostone lithology. Scott (2001) stated that the Ellaville Lime- stone and Suwannacoochee Dolostone are not dif- ferentiated on the geological map of Florida (Scott et al. 2001) and are instead included within the Suwannee Limestone due to the limited data on the areal distribution of the two units. The Ellaville Limestone and Suwannacoochee Dolostone are not included in Table 4 merely to simplify the table, as both units are known to contain only Rhyncholam- pas gouldii (Bouvé, 1846). However, Clypeaster cf. C. marinanus occurs in dolomitic facies of the Suwannee Limestone exposed in the Cabbage Grove Quarry west of Perry (FM-IP TA001). This form has a much thinner test and shorter ambu- lacra than C. rogersi and has not hitherto been identified in the Suwannee Limestone of peninsular Florida. Rhyncholampas gouldii is the ubiquitous echi- noid of the Suwannee Limestone. However, other species are present, though less abundant (Table 4). In the Brooksville Rock Quarry (now Vulcan Quarry), northwest of Brooksville in Hernando County (FM-IP HE007, FM-IP HE012, FM-IP HE026, FM-IP HE028, FM-IP HE034), the regular echinoids Gagaria mossomi (Cooke, 1941a) and Phymotaxis mansfieldi Cooke, 1941a occur through- out the majority of the Suwannee Limestone, but their presence is greatly overshadowed by the abun- dant R. gouldii. A mollusk-mold and foraminifera- rich limestone is intermittently exposed in the lowest beds within this quarry that contains an echinoid fauna that is very distinct from the remainder of the unit (Fig. 8). This horizon contains an abun- dance of Clypeaster sp. B, which characterizes the fauna, as well as large specimens of R. gouldii, and rarer Agassizia mossomi, Phyllacanthus cf. P. mor- toni, and the herein described Schizaster carlsoni n. sp., and Eupatagus dumonti n. sp. This is the only documented occurrence of this echinoid faunal assemblage within the Suwannee Limestone, and as they have not excavated below this horizon in the Cargil Vulcan Quarry, it is not possible to determine how deep in the Suwannee Limestone this horizon occurs, though it is certainly quite low in the unit. Southwestward in the Sunwest Mine, near Aripeka, R. gouldii occurs with Phymotaxis mans- fieldi, Gagaria mossomi, and the much rarer Brissus bridgeboroensis Carter, 1987b. Bridgeboro Limestone The Bridgeboro Limestone, a rhodolithic lime- stone that occurs in southwestern Georgia, south- eastern Alabama, and northern Florida, was intro- duced by Huddlestun (1981), adopted by Manker and ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 22 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 8: Vulcan Brooksville Quarry (FM-IP HE038) Hernando County, Florida. Exposed rocks are the Lower Oligocene Suwannee Limestone. Limestone in foreground (excavated beneath typical floor of quarry) is the lowermost bed containing Clypeaster sp. B., Eupatagus dumonti n. sp., and Schizaster carlsoni n. sp. Carter (1987), and formally named by Huddlestun (1993). The type locality of the unit is the southern- most pit of the Bridgeboro Lime and Stone Company quarry, almost 10.5 km southwest of Bridgeboro in Mitchell County, Georgia, where Huddlestun (1993) documented R. gouldii, Clypeaster cotteaui Egozcue in Cotteau, 1897 (which is the characteris- tic echinoid of the unit throughout its extent), and, questionably, C. rogersi. Manker and Carter (1987: table 2) also documented an undescribed Brissus and an undescribed Prionocidaris(?) (= Phyllacan- thus cf. P. mortoni?) from the rhodolith facies of the Bridgeboro Limestone. Huddlestun (1993) designated the Duncan Church Quarry (FM-IP WG002) (Fig. 9), in Wash- ington County, Florida, as a reference section for the lower boundary of the unit where it gradationally overlies the Marianna Limestone. Huddlestun (1993) erected the Florala Lime- stone Member of the Bridgeboro Limestone for a fossiliferous, Lepidocyclina-rich, non-rhodolithic facies of the Bridgeboro Limestone. The type local- ity for this member is the Stovall Quarry, located about 11 km east of Florala, southern Covington County, Alabama (FM-IP ZA023), very near the Alabama/Florida border (Huddlestun, 1993). The Florala Limestone Member appears to be an off- shore, likely deeper, quieter water facies of the typical Bridgeboro Limestone. It contains a more diverse echinoid fauna than the typical unit. Huddlestun (1993) documented C. cotteaui; C. rogersi; B. bridgeboroensis, Lytechinus floralanus (Cooke, 1941a), and Macropneustes mortoni from the Florala Limestone Member at its type locality. See the remarks for M. mortoni for discussion on this occurrence in the Florala Member. At the Duncan Church Quarry (FM-IP WG002) (Fig. 9) in the northern Florida panhan- dle, the Bridgeboro Limestone conformably over- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 23 Figure 9: Duncan Church Quarry (FM-IP WG002), Washington County, Florida. Exposed rocks are the Lower Oligocene Marianna and Bridgeboro Limestones. lies the Marianna Limestone (which contains C. rogersi) and contains a much greater diversity of echinoids than the typical Bridgeboro Lime- stone, with an abundance of C. cotteaui, as well as additional rarer species, including A. mos- somi, L. floralanus, C. rogersi, Phyllacanthus cf. P. mortoni, S. americanus, and E. dumonti n. sp (Table 4). The Bridgeboro Limestone at Duncan Church contains rare rhodoliths and a greater con- centration of Lepidocyclina than the typical Bridge- boro Limestone. The fauna and lithology of the Bridgeboro Limestone at Duncan Church is there- fore more like that of the Florala Member at its type locality than to that of the typical Bridgeboro Lime- stone. However, Huddlestun (1993: 55) included the Duncan Church beds in the undivided Bridgeboro Limestone. Bryan (1993) referred to the Bridge- boro Limestone at Duncan Church as the Duncan Church beds. SYSTEMATIC PALEONTOLOGY Classification follows Kroh (2020), the World Echi- noidea Database (Kroh and Mooi, 2022), and Stokes and Kroh (2022). Species within the genera are listed alphabetically after type species of the genus, if included, and new species are described in al- phabetical order at the end of the treatment for the relevant genus. Synonymies of commonly cited species are abbreviated, consisting of unique formu- lations of names referring to the taxon in question, as well as names occuring in major monographs or most recent works. Descriptions are provided for new species only. Readers interested in descriptions of previously named taxa can reference original descriptions listed in the synonymy for each species and consult our images. Diagnoses contain charac- ters pertinent to recognition of the new taxon to the species level. However, due to the nature of the taxa and the limited knowledge concerning them, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 24 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) these characters are not always autapomorphies, and we must rely on unique combinations of multiple characters. Usage of aff. and cf. in taxonomic desig- nations follows Lucas (1986). Terminology follows that of Mortensen’s ”Monograph of the Echinoidea” (volumes spanning 1925 to 1952), and Smith and Kroh (2011). Class ECHINOIDEA Schumacher, 1817 Subclass CIDAROIDEA Smith, 1984 Order CIDAROIDA Claus, 1880 Family CIDARIDAE Gray, 1825 Genus Phyllacanthus Brandt, 1835 Phyllacanthus mortoni (Conrad, 1850) (Figs. 10-15) Cidarites mortoni Conrad, 1850. p. 40. pl. 1, fig. 13. ?Cidaris blandus Gregorio, 1890. p. 253. pl. 44, fig. 5. (Typographical error for blandis) ?Cidaris moerens Gregorio, 1890. p. 252. pl. 43, figs. 22, 23. ?Cidaris modestus Gregorio, 1890. p. 253. pl. 43, fig. 26. ?Cidaris ordinatus Gregorio, 1890. p. 252. pl. 44, fig. 1. ?Cidaris perdubius Gregorio, 1890. p. 253. pl. 44, fig. 2. ?Cidaris moereus (Gregorio). Clark and Twitchell, 1915. p. 115. (Typographical error for mo- erens) ?Cidaris modestus (Gregorio). Clark and Twitchell, 1915. p. 115. ?Cidaris ordinatus (Gregorio), Clark and Twitchell, 1915. p. 115. ?Cidaris perdubius (Gregorio). Clark and Twitchell, 1915. p. 116. ?Cidaris blandis (Gregorio). Clark and Twitchell, 1915. p. 116. Cidaris mortoni (Conrad). Clark and Twitchell, 1915. p. 157. pl. 73, fig. 1. Cidaris georgiana Clark, in Clark and Twitchell, 1915. p. 158. pl. 73, fig. 4. Cidaris (Dorocidaris) georgiana (Clark). Lambert and Thiéry, 1925. p. 560. Phyllacanthus mortoni (Conrad). Cooke, 1941a. p. 3. pl. 1, figs. 16-18. Cidaris (Phyllacanthus) mortoni (Conrad). Fischer, 1951. p. 55. Phyllacanthus mortoni (Conrad). Cooke, 1959. p. 11. pl. 1, figs. 15-17. Phyllacanthus mortoni (Conrad). Osborn et al., 2016. tbl. 2. Occurrence.—Phyllacanthus mortoni occurs in Florida throughout the OLS, although it is most abundant in the upper portion of the unit where it is often associated with O. wetherbyi in quarries north- west of Mayo, Lafayette County (FM-IP LF001); the Branford 01A Quarry (FM-IP SU003) north of Branford, Suwannee County and west of Dowling Park, Lafayette County (FM-IP LF010), among other many other localities. Persistent searching in the Oligopygus wetherbyi Zone will usually reveal at least test fragments or spines of this distinctive species, although complete tests are rare. The species also occurs in the Oligopygus haldemani Zone at the Brooks Quarries (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031), northwest of Marianna, Jackson County, and a quarry northwest of Mayo (FM-IP LF001), Lafayette County. This species is occasionally found in the Oligopygus phelani Zone, along the Withlacoochee River west of Yankeetown (FM-IP LV024) and the Cross Florida Barge Canal south of Inglis in Citrus and Levy Counties (e.g., FM-IP CI001). Phyllacanthus mortoni is widely distributed in Upper Eocene strata in Alabama, where it occurs in the OLS and Yazoo Clay, as well as Georgia where it appears in the OLS. We also document the occurrence of partial tests of a cidarid in Oligocene strata that cannot be differentiated from P. mortoni in the Duncan Church beds of the Bridgeboro Limestone at Duncan Church (FM-IP WG002), Washington County and in the dolomitic facies of the Marianna Limestone at Dry Creek (FM-IP JA010) south of Marianna, Jackson County. Fragments also occur in the Marianna Lime- stone at a limestone quarry southeast of Brooklyn, Alabama (FM-IP ZA147). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 25 Figure 10: Phyllacanthus mortoni (UF 3345), 58 mm TD, 37 mm TH, Upper Eocene, Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A. aboral. B: oral. C, D: lateral. Cooke (1961) documented P. mortoni in his work on the echinoids of Trinidad and Venezuela. However, Cutress (1980) referred that specimen to Prionocidaris loveni (Cotteau, 1875). Discussion.—Conrad (1850) described Cidarites mortoni from Eocene strata near New Palmyra, Lee County, Georgia. However, four years earlier, Morton (1846) described Cidaris alabamensis, but did not figure it, from the Eocene of Washington County, Alabama. Clark and Twitchell (1915), in asserting that Morton very inadequately described C. alabamensis, also stated that no material could be obtained that can be regarded as belonging to that species. Subsequently, Cooke (1941a) placed Cidarites mortoni in Phyllacanthus but only questionably placed C. alabamensis into synonymy with P. mortoni, again stating that it had not been figured or distinguished using Morton’s original description. Having done so, though, Cooke implicitly recognized priority of the name C. alabamensis Morton, 1846, even after correctly stating that should it prove to be identical with P. mortoni, its name would have priority. We do not include Morton’s C. alabamensis in our synonymy. If C. alabamaensis can be shown to be a valid taxon ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 26 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 11: Phyllacanthus cf. P. mortoni (UF 3347), 69 mm TD, 55 mm TH, Upper Eocene, Ocala Limestone, Alachua County, Florida (FM-IP 2613). A. aboral. B: oral. C: lateral. identical with P. mortoni, the latter would become a subjective junior synonym of C. alabamensis. Nevertheless, without types of C. alabamensis, and given the meager description and locality information of Morton (1846), we, like Clark and Twitchell (1915) and Cooke (1941a, 1959), cannot presently conceive of a situation where this would be warranted. Clark in Clark and Twitchell (1915), named Cidaris georgiana from Muckafoonee Creek, Georgia, in strata he referred to the Oligocene, which is now considered Late Eocene. Asserting that it comes from beds known to contain P. mortoni, Cooke (1941a) placed C. georgiana into synonymy with P. mortoni, an action with which we agree. Cooke (1959) considered Phyllacanthus caro- linensis (Emmons, 1858) and Phyllacanthus mitchel- lii (Emmons, 1858) to be subjective junior synonyms of P. mortoni. However, Cutress (1980) and Kier (1980) disputed this assertion and treated these species as distinct. We have examined specimens of P. carolinensis from the OLS of Georgia and the Castle Hayne Formation of North Carolina and find differentiation of the species not at all troublesome. The broad area of coarse granulation in the inter- ambulacrals and less prominent scrobicular ring of P. carolinensis readily differentiate the two species. In addition, P. carolinensis has nonconjugate pores, whereas P. mortoni has clearly conjugate pores. We follow Kier (1980) and retain P. carolinensis as a distinct species. We also concur with Cutress (1980) and Kier (1980) and recognize P. mitchellii as a species distinct from P. mortoni. As noted by Kier (1980), P. mitchellii differs from P. mortoni in having its peristome smaller than its apical system, the in- terambulacral granular zone is wider in P. mitchellii and there are 11 to 11.5 ambulacral plates adjacent to each interambulacral plate at the midzone, whereas there are only seven to eight in P. mortoni. In the OLS of Florida, P. mortoni is most associated with O. wetherbyi, though it is present throughout the unit. This large cidarid could only be confused with Prionocidaris robertsi n. sp., which is readily differentiated by the granules of the interam- bulacral area being arranged in clear, longitudinal rows in P. mortoni, but not so in Pr. robertsi n. sp. In addition, the spines associated with Pr. robertsi n. sp. are heavily thorned and often flare at their apex, whereas spines associated with P. mortoni lack these prominent thorns or flared distal end and are relatively smooth or corrugated with rows of fine to coarse granulations. Phyllacanthus mortoni can- not be confused with any other echinoid currently documented from the Eocene of Florida. We figure a very large internal mold of a cidarid (UF 3347) that is likely P. mortoni, but ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 27 Figure 12: Phyllacanthus mortoni spines (UF 338012), none is complete but show variation in ornamentation; Oligopygus haldemani Zone of Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA086). A, B: 21.6 x 3.7 mm. C, D: 23.3 x 3.4 mm. E-G: 26.5 x 4.1 mm. H, I: 27.8 x 3.6 mm. J-L: 30.2 x 3.7 mm. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 28 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 13: Phyllacanthus mortoni spines (UF 338013) none is complete but show variation in ornamentation; Oligopygus wetherbyi Zone of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A, B: 21.6 x 5.3 mm. C, D: 26.4 x 3.9 mm. E, F: 29.6 x 5.6 mm. G, H: 30.8 x 4.9 mm. I, J: 31.4 x 6.1 mm. K, L: 32.8 x 5.9 mm. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 29 Figure 14: Phyllacanthus cf. P. mortoni (UF 337994), 36 mm x 35 mm, Oligocene Duncan Church Beds, Bridgeboro Limestone, Washington County, Florida (FM-IP WG002). A: interambulacrum. B, E: medial area showing granular-like tubercles arranged in rows, as in P. mortoni. C, D: close-up of non-crenulated, perforate primary tubercle. F: pore-pairs of ambulacrum and oblique view of interambulacral tuberculation. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 30 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 15: Phyllacanthus cf. P. mortoni (UF 337991) 21 mm x 28 mm, external mold in matrix from dolomitic portion of lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). which lacks surface details that could definitively place it in this species. It measures 69 mm in diameter, and 55.5 mm in height (Fig. 11), and was collected from FM-IP 2613. As noted in the discussion above, P. mor- toni has historically been referred to as Oligocene, based on a few subjective junior synonyms. However, these instances are either based on assumptions, or occurrences in strata that have subsequently been referred to the Eocene. Therefore, no definitive occurrences of Phyllacanthus species have previ- ously been documented from the Oligocene of the region. We document incomplete Phyllacanthus tests from the Oligocene Duncan Church beds of the Bridgeboro Limestone at the Duncan Church (Fig. 14), south of Chipley, Washington County (UF 337994), and from the dolomitic facies of the Mari- anna Limestone at Dry Creek (FM-IP JA010) south of Marianna, Jackson County (UF 337991) (Fig. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 31 15). Given the incompleteness of the tests, we ex- ercise caution and only refer to these as Phylla- canthus cf. P. mortoni. These fossils are not sep- arable from P. mortoni, but more complete ma- terial would need to be collected from in situ Oligocene deposits before we can assert that P. mortoni crosses from the Late Eocene into the early Oligocene. Genus Prionocidaris A. Agassiz, 1863 Prionocidaris robertsi n. sp. (Figs. 16-19) Diagnosis.—Prionocidaris with small tuber- cles in medial areas of interambulacra not aligned in distinct horizontal rows, ambulacra at ambitus only 36% width of interambulacra, and four columns of tubercles in interporiferous area of ambulacra. Asso- ciated spines adorned with pronounced thorns that are not verticillate (formed in whorls), with coronate cup-like cluster of thorns at tip; up to nearly twice as long (175%) as greatest diameter of only known complete test. Description.—Test small, only one known complete test (holotype: UF 147100) 28.3 mm TD, 16.2 mm TH; low: TH 57% TD. Aboral surface flattened; apical system missing, opening 13.7 mm x 13.6 mm. Peristomial plating unknown, opening larger than apical opening which is 11.9 mm x 12.9 mm. Ambulacra almost straight, very slightly sin- uous, average of all five ambulacra 4.1 mm wide at ambitus, 1.5 mm at apical plate, 2.6 mm wide at peristomial edge, total number of pore pairs or ambulacral plates indeterminate due to damage; am- bulacra on average 36% interambulacral width at ambitus. Poriferous areas slightly depressed, on av- erage 1.6 mm wide at ambitus; pores oval, conjugate, on average 0.4 mm wide at ambitus, on average 57 pore pairs per poriferous area, gap between pore pairs on average 3.7 mm. Interporiferous areas on average 1.4 mm wide at ambitus, 87% width of poriferous areas; with two pairs of distinct columns of small tubercles, each pair of columns separated by narrow naked medial area 40% width of inter- poriferous area. Interambulacra on average 11.3 mm wide at ambitus, 275% width of ambulacra at ambitus, two columns of primary tubercles per in- terambulacrum, five to seven tubercles per column; areolas ovate, large, on average 4.1 mm wide and 3.7 mm high at ambitus, 36% width of interam- bulacra at ambitus; rings of scrobicular tubercles distinct and well-separated at ambitus, intersect- ing, with rings becoming less distinct orally and aborally, areoles often not separated by distinct scrobicular rings furthest from ambitus. Primary tubercles large, moderately high, platforms smooth, not crenulated, mamelon perforate. Medial areas covered by small tubercles, not aligned in distinct horizontal rows. Numerous associated spines available (paratype: lot UF 117489: 17 spines); longest complete spine 49.8 mm in length (175% TD of only known complete test), 4.5 mm wide at coronate distal end, 5.0 mm wide at widest thorn; with distinct thorns, not verticillate, thorns perpendicular to spine, becoming inclined distally toward distal end of spine; thorn length up to 55% greatest width of spine, often lacking in medial area of spine, replaced by small thorns or nodules often in vertical ridge-like columns; slightly tapered; coronate at tips, rimmed by large spinules in a cup-like arrangement. Zoobank Nomenclatural Act.—C7C32F9A- C284-4EF6-B39C-B6B6AE8D060A Discussion.—While studying the only known complete test of Prionocidaris robertsi n. sp. (UF 147100) (Fig. 16) and associated spines (UF 117489) (Fig. 17), it became clear that although this taxon was readily distinguishable as a new species, assigning it to a genus (either Phyllacanthus or Prionocidaris) is not simple. Modern classification of the Cidaroida is largely based on work by Mortensen (1903; 1928, especially) which was, as was much of Mortensen’s work, largely based on the pedicellariae. For this reason, Lambert and Thiéry (1909, 1910) and Gig- noux (1933) rejected Mortensen’s classification, but as noted by Philip (1963), although Mortensen’s sys- tem is not perfect, it is preferred over any previously proposed system. However, the fact remains that it is difficult to apply this classification to fossil forms lacking the appendages and less often preserved ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 32 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 16: Prionocidaris robertsi n. sp., holotype (UF 147100), 28.3 mm TD, 16.2 mm TH, Upper Eocene, Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033). A: aboral. B: lateral, interambulacrum center. C: ambulacrum. D: interambulacrum. E: oral. F: tilted aboral viewpoint, interambulacrum center. G: close-up of primary tubercles (very worn) and ambulacrum near ambitus. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 33 Figure 17: Prionocidaris robertsi n. sp., spines (UF 117489), Upper Eocene, Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033). regions used in Mortensen’s classification. Cooke (1946, 1953, 1957, 1959) was an opponent of apply- ing Mortensen’s classification to fossil specimens due to the inadequacy of characters seen in many fossils (typically tests and disassociated spines). Another approach, notably seen in the works of Chapman and Cudmore (1934), Fell (1954, 1966), and Philip (1963) was to attempt to recognize affini- ties in fossil forms that give an indication of the modern genus to which the echinoid could be as- signed. However, as cautioned by Philip (1963), names extracted from the list of living genera should be employed only as “form genera” with their identi- fication based on but a single attribute (spines with cup-shaped terminations = Goniocidaris; spines with thorns = Prionocidaris; spines with flanges = Chondrocidaris, etc.). As Philip (1963) stated, this can lead to unsatisfactory taxonomy, but it is still often possible to ascertain genus affinities of Ceno- zoic species from a detailed study of living cidarids. Philip (1963) accomplished his review of the Aus- tralian fossil cidarids by making comparisons with the modern forms of the Indo-Pacific region. He emphasized that because of the inherent uncertainty of cidarid taxonomy, this method usually leads to a very broad interpretation of some genera. This was noted by Cutress (1980) when she stated that her use of Cidaris (for species with nonconjugate pores, noncrenulate tubercles, and simply tapered, ridged spines) is not to be recognized in the strict sense of the Recent genus. Fell (1966) stated that although the usual keys to the identification of cidaroids are difficult to apply to fossils due to the lack of preserved pedicellariae, if attention is paid to the sum of available characters of the spines and test, most fossils can be classified into taxa that are unlikely to differ substantially from characters used for the living species. Fell (1966) provided diagnoses using spine and test characteris- tics for all fossil and modern cidarid genera. It is also important to note that in many cases, the problem of higher taxon determination can be greatly reduced ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 34 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 18: Prionocidaris robertsi n. sp., (UF 341763), 18 mm x 14 mm, test fragment with half an interambulacrum and half an ambulacrum, Upper Eocene, Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033). by using features that clearly eliminate possible genus assignments for the fossil in question. The test of Pr. robertsi n. sp. is very similar to that of Phyllacanthus, possessing noncrenulate, perforate, primary tubercles, well-separated areoles, and conjugate pore pairs. However, the test of Pr. robertsi n. sp. is readily distinguished from P. mor- toni, which occurs in overlying strata of the OLS in Jackson County, because the granulations of the interambulacra are not latitudinally arranged. In P. mortoni these granules are latitudinally (horizon- tally) arranged in rows, whereas in Pr. robertsi n. sp. they are not aligned in rows. Fortunately, many spines are associated with Pr. robertsi n. sp. and are exceptionally abundant on the floor of Jackson Blue Spring (Fig. 17). These spines are usually adorned with sharp thorns whose length often exceeds the thickness of the shaft, and complete spines are often flared at the dis- tal end (coronate sensu Cutress, 1980) with flared thorns protruding in a cup-like shape (Figs. 17, 19). These spines help confirm the genus assignment to Prionocidaris for this species and are very simi- lar to those of the modern Prionocidaris bispinosa (Lamarck, 1816) of the Indo-Pacific region (see Mortensen, 1928: pl. XLIV, fig. 2). However, in Pr. bispinosa, the lowermost thorns on the spine are often verticillate, whereas the thorns in Pr. robertsi are not. Fell’s (1966: U330) diagnosis for Prionocidaris described the spines as: “primary spines long, tapering with coarse thorns in longitu- dinal series, less commonly cylindrical, smooth or widened distally, or with thorns arranged in whorls.” No evidence of any other cidarid exists in this horizon (the Haimea brooksi Zone of the OLS is also exposed in a quarry of Leon Brooks, 8 mi. to the west), so attributing these spines to the single, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 35 Figure 19: Prionocidaris robertsi n. sp., spines, none of which is complete (UF344344-46), Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, Briar Cave (FM-IP MR018), south of Ocala, Marion County, Florida (SL= spine length, SD = widest diameter of spine). A-C: 15.2 mm SL, 5.4 mm SD. D, E: 21.6 mm SL, 5.4 mm SD. F, G: 37.9 mm SL, 3.3 mm SD. H-J: 44.7 mm SL, 2.9 mm SD. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 36 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) complete, test of this species that was collected amongst them seems justified. Although P. mortoni occurs in strata above this horizon in the Brooks Quarry (FM-IP JA039), the spines of P. mortoni are typically thick and cylindrical with a shaft covered in fine granules that are typically arranged in lon- gitudinal series. They may develop larger granules that resemble “bumps” but never with prominent thorns and coronate tips as seen in Pr. robertsi n. sp. Fell’s (1966: U330) diagnosis for Phyllacanthus describes the spines as: “primary spines cylindrical, thick, robust, with fine granules arranged in regular longitudinal series on shaft.” However, as cautioned by Cutress (1980), cidarids are highly variable, and though she asserted the spines may in some instances provide better taxonomic characters than the test, they can be highly variable, especially in Prionocidaris. Cutress (1980) stated that it was discouraging that the mod- ern species of Prionocidaris were among the most variable of the species she studied, particularly Pr. bispinosa and Pr. hawaiiensis (Agassiz and Clark, 1907). She stated these two species had variation in measurements that ranged as high as 36% and in the less variable species of the genus, the range was still 12 to 18%. Given this variability, Cutress (1980) cautioned against isolating taxa based on disconti- nuities in her measured data, which she provided in tables. Variability of the test of Pr. robertsi n. sp. cannot be determined until additional specimens are collected. However, spine variability is demonstrated in Figs. 17 and 19. The typical spines of this species are heavily thorned, and when complete, usually show the flared coronate tip. Non-thorned spines are much rarer, as seen in specimens of the modern Pr. bispinosa (Mortensen, 1928: pl. XLIV, fig. 2). Although it is evident this species is not comparable to any other species from Cenozoic strata in North America, many cidarids occur in the Caribbean region that require comparisons. Thank- fully, Cutress (1980) compiled all known occur- rences of fossil cidarids of the Caribbean area, pro- viding a useful series of figures and an updated tax- onomy. She identified six species in Prionocidaris: Pr. cojimarensis (Lambert and Sánchez-Roig in Sánchez-Roig, 1926), from the Miocene to Pliocene throughout the Caribbean area; Pr. spinidentatus (Palmer in Sánchez-Roig, 1949), from the Oligocene to Miocene of Cuba, Antigua, and Puerto Rico; Pr. katherinae Cutress, 1980, from the Miocene of Cuba and Puerto Rico; Pr. clevei (Cotteau, 1875), from the Miocene of Anguilla; Pr. cookei Cutress, 1976, from the Miocene of Florida and Cuba, and Pr. loveni (Cotteau, 1875), from the Eocene of St. Bartholomew, Cuba, and Jamaica. Cutress (1980) noted that the tubercles of Pr. cojimarensis are crenulate, whereas they are not in Pr. robertsi. Cutress also stated that spines of Pr. co- jimarensis are exceptionally variable, ranging from evenly spinose to distinctly verticillate, nodulose, or ridged, yet none exhibits thorns as pronounced as those of Pr. robertsi, nor are the distal ends flared and coronate. In fact, the thorny, coronate spines of Pr. robertsi n. sp. distinguishes it from all other rep- resentatives of the genus in the Caribbean region except for Pr. spinidentatus. The spines of these two species are very similar (see Cutress, 1980: pl. 8), though the thorns are less densely arranged on the spines of Pr. robertsi. However, the tests of these two species are very dissimilar. The small tubercles in the medial area of the interambulacral of Pr. spindentatus are aligned in horizontal rows as opposed to the haphazard arrangement of these small granule-like tubercles in Pr. robertsi. At its type locality, 30 m below the surface of Jackson Blue Spring east of Marianna, Jackson County, Florida (FM-IP JA033), Pr. robertsi n. sp. occurs with abundant specimens of H. brooksi, O. haldemani, and R. georgiensis, as well as W. john- soni, B. jonesi n. sp., rare specimens of O. beckeri, and an undetermined species of regular urchin dis- cussed herein as Euechinoidea gen. et sp. indet. Spines of Pr. robertsi n. sp. occurs in the O. phelani Zone of the lower OLS at Briar Cave (FM- IP MR018), in Marion County. These (UF 344344- UF 343446; Fig. 19) are found with the typical fauna of the Oligopygus phelani Zone, including E. mooreanus, P. floridanus, Durhamella ocalana, and Weisbordella inglisensis n. sp. Etymology.—This species is named in honor ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 37 Figure 20: Acanthechinus dixie (UF 66559), 45 mm TD, partial specimen with preserved Aristotle’s Lantern, Upper Eocene, upper portion of Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A, B: lateral viewpoints. C: oral. of Sean W. Roberts, who (along with Rebecca Roberts and Melissa Marzahn) collected the Jackson Blue material used in this study through recreational SCUBA diving. Materials and Occurrence.—Holotype UF 147100 (test) and paratypes (UF 117489), 17 associ- ated spines, from OLS, associated with H. brooksi in Jackson Blue Spring (FM-IP JA033), Jackson County, Florida. Subclass EUECHINOIDEA Bronn, 1860 Infraclass AULODONTA Jackson, 1912 Aulodonta gen. et sp. indet. Discussion.—This taxon is represented by a single test (UF 332884), which is complete but broken into two pieces, with preserved Aristotle’s lantern and some associated spines. It was collected from the upper portion of the OLS (either Oligopy- gus wetherbyi Zone or overlying Wythella eldridgei Zone), in a now-inactive and waterfilled quarry (FM- IP TA004) near Tennille, Taylor County, Florida. The specimen is remarkable for several reasons. It is large (˜52 mm TD) with a thin, fragile, test, and remarkable retention of delicate features as indicated above. An extensive search of regional collections, both institutional and of avocational col- lectors, failed to locate even a fragment of a similar taxon. The specimen is so unlike any known fossil echinoid from the eastern Americas and Caribbean region (see Table 5), that it requires study that will go beyond the time frame for the present work. Aulodonts are generally so rare in the fossil record of the region, a stand-alone publication will be necessary to place the fossil in the correct taxon and perform a full revision of the group to which it might belong, depending on how this find impacts the systematics of that group. To be as inclusive as possible, yet not delay publication of the present work, we refer to it as an unidentified aulodont. Infraclass CARINACEA Kroh and Smith, 2010 Order PHYMOSOMATOIDA Mortensen, 1904 Family PHYMOSOMATIDAE Pomel, 1883 Genus Acanthechinus Duncan and Sladen, 1882 Acanthechinus dixie (Cooke, 1941a) (Figs. 20, 21) Phymosoma dixie Cooke, 1941a. p. 17. pl. 2, fig. 15; pl. 4, figs. 1, 2, 9. Dixieus dixie (Cooke). 1948a. p. 607. Phymosoma dixie (Cooke). Fell and Pawson in Moore, 1966. p. 396. Dixieus dixie (Cooke). Mortensen, 1951. p. 558, fig. 281 (after Cooke). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 38 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 21: Acanthechinus dixie (UF 342232), 61.8 mm TD, 26 mm TH (maximum measurable TH due to matrix), Upper Eocene upper Ocala Limestone, Jackson County, Florida (FM-IP JA018). A. aboral. B: oral. C, D: lateral. Dixieus dixie (Cooke). Cooke, 1959. p. 24. pl. 5, figs. 8-10. Dixieus dixie (Cooke). Kier, 1980. p. 21. pl 5, figs. 1-3. Dixieus cf. D. dixie (Cooke). Kier, 1997. p. 3. pl. 1, figs. 1-4. Dixieus dixie (Cooke). Osborn et al., 2016. tbls. 1, 2. Acanthechinus dixiei (Cooke). Kroh and Mooi, 2023. p. 1. Occurrence.—Acanthechinus dixie is present, though typically rare, at numerous localities in the Upper Eocene OLS, including quarries northwest of Mayo (FM-IP LF001), Lafayette County; the Bran- ford 01A Quarry (FM-IP SU003) north of Branford, Suwannee County, and south of Tennille (FM-IP DI001), Dixie County, among other localities. Cooke (1959) designated the type locality of A. dixie as a quarry east of Steinhatchee River north of U.S. Highway 19 near Clara, in Dixie County, Florida ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 39 Figure 22: Phymotaxis mansfieldi (UF 113858), 43 mm TD, 17 mm TH, Lower Oligocene Suwannee Limestone, Hernando County, Florida (FM-IP HE017). A: aboral. B: oral. C, D: lateral. (type locality-USGS 12747). It is most frequently encountered in the uppermost OLS where it occurs in the Wythella eldridgei Zone. The species is also documented in the Mid- dle Eocene Clinchfield Sand in southwest Georgia (Carter and Hammack, 1989) and Castle Hayne Limestone of North Carolina (Kier, 1980). Discussion.—Cooke (1941a) initially de- scribed this species as Phymosoma dixie but sub- sequently (Cooke, 1948a) designated it as the type of his new genus Dixieus which he stated differs from Phymosoma koenigi (Mantell, 1822), the type species of Phymosoma, in the greater number of zygopores (twice as many), the biserial part of its poriferous zones, the smaller size and more equi- lateral shape of its apical system, its more tumid, less discoid shape, its larger, more deeply notched peristome, and its smaller tubercles, which exhibit more even gradation in size. Fell and Pawson (1966) considered Dixieus a subjective junior synonym of Phymosoma. How- ever, this assertion has not been followed by most echinoid workers of the region (Kier, 1980; 1997). Smith and Kroh (2011) considered Dixieus to be a subjective junior synonym of Acanthechinus (Dun- can and Sladen, 1882). We agree that no genus-level differences are apparent between D. dixie and Acan- thechinus pelorius (Arnold and Clark, 1927) from ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 40 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 23: Phymotaxis mansfieldi (UF 118239), 44 mm x 41 mm TD, 21 mm TH, Lower Oligocene, Suwannee Limestone, Hernando County, Florida (FM-IP HE007). A: aboral. B: oral. C, D: lateral. the Eocene of Jamaica, or Acanthechinus abnor- male (Duncan and Sladen, 1882: 32) from the Late Paleocene-Early Eocene of Pakistan. In the World Echinoidea Database, Kroh and Mooi (2023) refer to this species as A. dixiei. We are unable to support this change, since the taxon is named not for a person but for a county, much as for another of Cooke’s (1942) species, Plagiobrissus dixie. Acanthechinus dixie is most common in the uppermost horizon of the upper OLS of Florida where it occurs with W. eldridgei, S. armiger, B. steinhatchee, E. ocalanus, P. dixie, and other species, above the horizon of O. wetherbyi in the Wythella eldridgei Zone. We figure an exceptional specimen (UF 66559) that retains its Aristotle’s lantern (Fig. 20). Order STOMOPNEUSTOIDA Kroh and Smith, 2010 Family STOMOPNEUSTIDAE Mortensen, 1903 Genus Phymotaxis Lambert and Thiéry, 1914 Phymotaxis mansfieldi Cooke, 1941a (Figs. 22, 23) Phymotaxis mansfieldi Cooke, 1941a. p. 18. pl. 4, figs. 6-8. Phymotaxis mansfieldi (Cooke). Cooke, 1959. p. 24. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 41 pl. 6, figs. 4-8. Occurrence.—Phymotaxis mansfieldi is not documented outside of the Lower Oligocene, Suwannee Limestone. Cooke (1941a) designated Brooksville, Florida as the type locality of the species, where it can still be collected in numer- ous quarries west of Brooksville, Hernando County (FM-IP HE019). Discussion.—Phymotaxis mansfieldi occurs with the very abundant R. gouldii, as well as G. mossomi and other, rarer echinoid species, in the Lower Oligocene Suwannee Limestone. Smaller specimens of P. mansfieldi can be confused with G. mossomi in the field. However, they can be readily differentiated by the poriferous zones, which are straight in G. mossomi and in arcs in P. mansfieldi, which gives the poriferous zones in the latter a sinuous appearance. Order CAMARODONTA Jackson, 1912 ”Triplacidiids” temporary family (Kroh and Mooi, 2022) Genus Gagaria Duncan, 1889 Gagaria mossomi (Cooke, 1941a) (Figs. 24-26) Thylechinus (Gagaria) mossomi Cooke, 1941a. p. 13. pl. 2, fig.16; pl. 3, figs. 6-9; pl. 4, figs. 3-5. Gagaria mossomi (Cooke). Cooke, 1959. p. 17. pl. 3, figs. 10-14. Gagaria mossomi (Cooke). Fell and Pawson, 1966. pp. U430, U431. figs. 319.1a-c. Gagaria mossomi (Cooke). Pickering, 1970. pp. 20, 29. Gagaria mossomi (Cooke). Kier, 1997. p. 5. pl. 2, figs. 1-5. Gagaria mossomi (Cooke). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 6. Occurrence.—Within Florida, G. mossomi is not documented outside of the Suwannee Limestone, where it is commonly associated with R. gouldii near Brooksville (FM-IP HE019), the now defunct Morrell Pit in Pasco County (FM-IP PA002), and Terramar Quarry in Polk County (FM-IP PO017). The species is also documented in the Oligocene of Georgia (Pickering, 1970) and North Carolina (Kier, 1997). Discussion.—When Cooke (1941a) initially described G. mossomi, he designated two syntypes (USNM 372885) but no holotype. The syntypes consist of a fragment and complete test (Cooke, 1941a: pl. 4, figs. 3-5). However, Cooke (1959), in the captions for plate 3, figures 10-14, identi- fied the figured specimens (USNM 498884a-c) as paratypes. Yet, these specimens were not identi- fied as paratypes when Cooke (1941a) erected the species. Paratypes cannot be named as such after the original description, even if they were part of the same suite of specimens used in the description. To further complicate matters, these three specimens are now cataloged in the USNM database as syn- types with distinct catalog numbers: USNM 498884, 648532, 648533. Cooke appears to have included fragmentary material of what was likely P. mansfieldi in his orig- inal description (Cooke, 1941a). In that description, Cooke (1941a) described the poriferous zones of G. mossomi as nearly straight but confused the is- sue by noting that each group of three pore pairs slightly curves around a large tubercle, which is often the case in poriferous zones of many camarodonts, among other non-irregular echinoid groups. He cor- rected his description of G. mossomi in his 1959 monograph and removed reference to arc-forming pore pairs. Specimens of this small species, usually less than 2.5 cm in diameter, can be confused with juvenile specimens of the usually larger P. mansfieldi, as both typically occur in the same stratigraphic horizons. As stated by Cooke (1959) and well-illustrated by Kier (1997: pl. 2, fig. 5), closer examination will easily differentiate them as, the poriferous zones of G. mossomi are straight, whereas in P. mansfieldi, the pore pairs are arranged in arcs that give the poriferous zones a sinuous appearance. We figure (Fig. 24) a very large specimen (UF 28244) from the Suwannee Limestone of Pasco County (FM-IP PA002) that measures 40.7 mm ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 42 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 24: Gagaria mossomi (UF 28244), 40 mm TD, 22 mm TH; very large specimen for the species, Lower Oligocene Suwannee Limestone, Pasco County, Florida (FM-IP PA002). A: aboral. B: oral. C, D: lateral. A: aboral. B: oral. C, D: lateral. maximum TD and 21.2 mm TH. Family TRIGONOCIDARIDAE Mortensen, 1903 Genus Brochopleurus Fourtau, 1920 Brochopleurus pretiosus (Clark in Clark and Twitchell, 1915) (Fig. 27) Orthechinus pretiosus Clark in Clark and Twitchell, 1915. p. 118. pl. 56, figs. 2a-b. Gagaria americana Clark in Clark and Twitchell, 1915. p. 159. pl. 73, figs. 8a-b. Thylechinus americanus (Clark). Lambert and Thiéry, 1925. p. 567. Progonechinus pretiosus (Clark). Cooke, 1941a. p. 15. pl. 2, figs. 4-6. Pseudodiadema(?) americanum (Clark). Cooke, 1941a. p. 8. Brochopleurus pretiosus (Clark). Cooke, 1959. p. 18. pl. 4, figs. 4-6. Occurrence.—Within Florida, B. pretiosus is only known from the Upper Eocene OLS in the Brooks Quarry, near Marianna, Jackson County (FM-IP JA031). Brochopleurus pretiosus is more ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 43 Figure 25: Gagaria mossomi (UF 322466), 26 mm TD, 12 mm TH, Lower Oligocene, Suwannee Limestone, Hernando County, Florida (FM-IP 6340). A: aboral. B: oral. C, D: lateral. widespread in Upper Eocene strata of Alabama and Georgia (Cooke, 1959). Discussion.—This is the first documenta- tion of B. pretiosus from Florida. Its minute tu- bercles readily differentiate it from other regu- lar urchins in the region. The lone documented Florida specimen of B. pretiosus was collected with M. mortoni, R. conradi, O. haldemani, and other, rarer species in the upper portion of the up- per OLS exposed in the Brooks Quarries (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031, FM-IP JA039), northwest of Marianna, Jackson County. Mortensen (1943) stated that Brochopleurus is likely a junior synonym of Scolechinus Lam- bert (in Lambert and Thiéry, 1925), and Nisiyama (1966) stated the genus was very closely related to Javanechinus (Jeannet in Lambert and Jeannet, 1935) from the Miocene of Java and Fiji, and may be identical to it. Nisiyama (1966) also remarked on the similarity between Brochopleurus and the modern genus Desmechinus (H. L. Clark, 1923). However, Brochopleurus was considered by Smith and Kroh (2011) and Kroh and Mooi (2022) as a junior syn- onym of Ortholophus. However, Borghi et al. (2022) disagreed and stated Temnechinus stellatus Duncan and Sladen, 1886 (the type species of Brochopleurus) has smooth, non crenulate, tubercles, differentiating ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 44 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 26: Gagaria mossomi (UF 110863), 23 mm TD, 12 mm TH, external mold in silicified limestone with RTV cast, Lower Oligocene Suwanee Limestone, Alachua County, Florida (FM-IP AL030). A: aboral. B: oral. C: lateral. D: external mold. E: RTV cast of external mold. it from Scolechinus and Ortholophus. Borghi et al. (2022) therefore maintained Brochopleurus as dis- tinct from both Scolechinus and Ortholophus. In the absence of further data, we maintain Brochopleurus for this species. We figure a specimen (UF 68923) from the Upper Eocene Yazoo Formation near Perdue Hill, Alabama (FM-IP ZA026) that has excellent preser- vation of plate details to aid identification of this taxon (Fig. 27). Family TOXOPNEUSTIDAE Troschel, 1872 Genus Lytechinus A. Agassiz, 1863 Lytechinus floralanus (Cooke, 1941a) (Figs. 28, 29) Psammechinus floralanus Cooke, 1941a. p.15. pl. 3, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 45 Figure 27: Brochopleurus pretiosus (UF 68923), 11 mm TD, 6 mm TH, Upper Eocene Yazoo Formation, Perdue Hill, Monroe County, Alabama (FM-IP ZA026). A: aboral. B: oral. C, D: lateral. figs. 10, 11. Psammechinus(?) ocalanus Cooke. 1941a. p.16. pl. 2, figs. 9-11. Lytechinus floralanus (Cooke). Cooke. 1959. p. 15. pl. 2, figs. 17-21. Lytechinus floralanus (Cooke). Kier. 1974. txt. fig. 10A. Lytechinus floralanus (Cooke). Osborn et al. 2016. tbl. 2. Occurrence.—Lytechinus floralanus is one of the rare species that occurs in both the Upper Eocene and Lower Oligocene strata of the region. Within Florida, it is known from the Upper Eocene, uppermost portion of the OLS in the Brooks Quar- ries near Marianna, Jackson County (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031) and the Lower Oligocene Bridgeboro Limestone at Duncan Church, Washington County (FM-IP WG002). This species is commonly found in the Oligocene Marianna and Bridgeboro Limestones of Alabama, especially at its type locality in the Florala Limestone Member of the Bridgeboro Limestone, Florida Lime Products Quarry, east of Florala, Cov- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 46 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 28: Lytechinus floralanus (UF 61439), 28 mm TD, 12 mm TH, Oligocene Bridgeboro Limestone, Washington County, Florida (FM-IP WG002). A: aboral. B: oral. C, D: lateral. ington County, Alabama. The type of Psammech- inus(?) ocalanus was collected near Bainbridge Georgia, in what Cooke (1941a) called “question- ably Eocene strata”. Discussion.—Lytechinus floralanus occurs with C. cotteaui, and other, rarer species, in the Oligocene Bridgeboro Limestone at Duncan Church. We herein figure (Fig. 28) a large specimen from Duncan Church (UF 61439) (Fig. 9) that measures 28.7 x 28.4 mm TD, and 12.6 mm TH. Osborn et al. (2016: table 2) documented L. floralanus in the Upper Eocene OLS of Florida, where it occurs with M. mortoni, W. johnsoni, R. conradi, and other species, just below the con- tact with the overlying Bumpnose Limestone in the Brooks Quarries (FM-IP JA021, FM-IP JA025, FM-IP JA030). The proportionately large tubercles readily distinguish L. floralanus from B. pretiosus, which it occurs within the OLS within the Brooks Quarry. We herein figure (Fig. 29) a specimen from ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 47 Figure 29: Lytechinus floralanus (UF 226884), 20 mm TD, 11 mm TH, Upper Eocene, upper Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C, D: lateral. this horizon that measures 20.1 x 20.2 mm TD, 11.7 mm TH (UF 226884). Euechinoidea gen. et. sp. indet. (Fig. 30) Occurrence.—Upper Eocene OLS, (occurs with H. brooksi), Jackson Blue Spring (FM-IP JA033), east of Marianna, Jackson County. Discussion.—This taxon is represented by two test fragments (UF 341758, UF 341759) from deep within the Jackson Blue Spring cave system. The specimens were associated with H. brooksi, R. georgiensis, and other species, and though both specimens are corroded and incomplete, they are sufficient to distinguish the taxon from any known regular urchin in North American deposits. However, they are too incomplete for assignment, even to any higher taxon. The specimens clearly display large, imper- forate, non-crenulate tubercles with pore pairs in arcs of five. Large tubercles form a single column within each ambulacrum. The known fragments do not exhibit a relatively complete interambulacrum ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 48 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 30: Euechinoidea gen. et. sp. indet. Two test fragments of an undetermined genus of regular urchin from the Ocala Limestone (associated with Haimea brooksi) in Jackson Blue Spring (FM-IP JA033), east of Marianna, Jackson County, Florida. A, B: UF 341759; 25.7 mm x 22.7 mm; fragment includes portions of an ambulacrum and interambulacrum. B: close-up of ambulacrum showing pore series. C-H: UF 341758; 34.3 mm x 33.0 mm; two test fragments adhered together in matrix with portions of ambulacra and interambulacra. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 49 or ambulacrum. The largest specimen (UF 341758) represents two fragments in the same block of ma- trix, which measures 34 mm x 33 mm, indicating this was a rather large species. It is likely the largest non-cidarid regular urchin in Eocene deposits of North America. The specimens are readily distinguishable from either A. dixie from Florida or Acanthechinus peloria (Clark in Arnold and Clark, 1927), which is from the Eocene of Jamaica, because these taxa have only two columns of ambulacral or interambulacral primary tubercles. The specimens of the unknown echinoid are unlike the smaller L. floralanus. Ceno- zoic regular urchins are not abundant in the remain- der of the eastern Americas and Caribbean Region, and we were able to compare these specimens to most of these taxa. The single column of tubercles in the ambulacrum is like Triadechinus multiporus Clark in Arnold and Clark, 1927, from the Eocene of Jamaica, but the triserial pore pair arrangement of T. multiporus is very different. Likewise, the fragments cannot be confused with Stenechinus regularis Clark in Arnold and Clark, 1927; Stenechinus perplexus Clark in Arnold and Clark, 1927; or Hebertia sim- plex Hawkins, 1924, from the Eocene of Jamaica. These three forms, all now placed in Echinopsis, have much wider ambulacra, different tuberculation and pore pairs arranged in straight columns, not arcs. Clark in Arnold and Clark (1927) also described Scoliechinus axiologus from the Eocene of Jamaica, which Smith and Kroh (2011) asserted belongs in Lytechinus. Leiosoma chondra Clark in Arnold and Clark, 1927, from the Eocene of Jamaica, and Leiosoma guadalupense Sánchez-Roig, 1949, from the Eocene of Cuba, are now placed in Trochalosoma (Fell and Pawson, 1966), and have triserial pore pairs orally and biserial aborally, readily distinguishing these Trochalosoma from the Florida material. Diadema principeana Weisbord, 1934, the holotype of which is housed at the Paleontologi- cal Research Institute (PRI 3807) is readily distin- guished from the Florida specimens by its straight ambulacra. Stomopneustes pristinus Jackson, 1937, from the Oligocene of Veracruz, Mexico, has very large tubercles and triserial pore pairs that readily dis- tinguish this species from the Florida fragments at hand. The remaining Paleogene Euechinoidea of the Caribbean are either too poorly figured and under- stood (mostly Cuban species of Sánchez-Roig), or too dissimilar from these specimens to warrant com- parison. These taxa include Amblypneustes corrali Lambert and Sánchez-Roig in Sánchez-Roig, 1949 (Eocene? of Cuba); Echinopedina cubensis Cotteau, 1881 (Cuban strata now considered Cretaceous); Gauthieria sanchezi Lambert, 1921 (Eocene? of Cuba); Leiopedina cienagensis Sánchez-Roig, 1949 (Eocene of Cuba); Pedina eocenica Sánchez-Roig, 1949 (Eocene? of Cuba); Phymosoma conceptionis Sánchez-Roig, 1952c (Eocene? of Cuba); Phymo- soma gigantea Sánchez-Roig, 1953b (Eocene? of Cuba); and Pseudorthopsis rojasi Sánchez-Roig, 1953b (Eocene of Cuba). In addition, Cooke (1961) described Phymosoma trinitensis from the Pale- ocene of Trinidad, which Smith and Jeffery (2000) assert belongs in Trochalosoma. We include the above discussed taxon in a table of fossil Euechinoidea (non-cidarid regular urchins) from Cenozoic deposits of the remain- der of the eastern Americas and Caribbean Region (Table 5). After the above considerations, we can only refer this taxon to the Euechinoidea Bronn, 1860, which includes all non-cidarid echinoids. Bather (1909: 7) stated: “As regard the application of species names to such fragments, it seems to me that if our descriptions are to be of practical service to stratigraphers, then they must be accompanied by names. A description without a name is soon lost sight of, whereas a name compels attention until at last it finds its proper place, if only as a synonym.” We disagree when insufficient diagnostic features exist in fragments to allow attribution even to order or infraclass. However, to bring attention to the possible existence of a large, unidentified euechinoid in the Florida Paleogene, we include these specimens as a previously undocumented taxon. Given the scarcity of regular urchins in the OLS, it is hoped that additional material will become available to permit formal description in future. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 50 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Table 5: A listing of fossil Euechinoidea (non-cidarid regular echinoids) described from the Cenozoic of the eastern Americas and Caribbean region, outside of the eastern United States. Locality provided is the type area for the species. Amblypneustes corrali Lambert and Sánchez-Roig, 1949 Oligocene, Cuba Echinopedina cubensis Cotteau, 1881 strata is now considered Cretaceous (Broderman, 1949) Ellipsechinus palmeri Lambert, Sánchez-Roig, 1949 strata is now considered Cretaceous (Broderman, 1949) Diadema principeana Weisbord, 1934 Eocene, Cuba, likely not Diadema per Weisbord (1934) Gauthieria sanchezi Lambert, 1921 Eocene, Cuba Hebertia jacksoni Sánchez-Roig, 1949 Oligocene Cuba = Echinopsis (Fell and Pawson, 1966) Hebertia simplex Hawkins, 1924 Eocene Jamaica = Echinopsis (Fell and Pawson, 1966) Irenechinus rosei Poddubiuk, 1987 Oligocene, Antigua = Ortholophus (Smith and Kroh, 2011) Leiopedina cienagensis Sánchez-Roig, 1949 Eocene, Cuba Leiosoma chondra Clark, 1927 Eocene, Jamaica = Trochalosoma (Fell and Pawson, 1966) Leiosoma guadalupense Sánchez-Roig, 1949 Eocene, Cuba = Trochalosoma (Fell and Pawson, 1966) Pedina eocenica Sánchez-Roig, 1949 Eocene, Cuba Phymosoma peloria Clark, in Arnold and Clark, 1927 Eocene, Jamaica, = Acanthechinus (Smith and Kroh, 2011) Phymosoma conceptionis Sánchez-Roig, 1952 Eocene, Cuba Phymosoma gigantea Sánchez-Roig, 1952 Eocene, Cuba Phymosoma trinitensis Cooke, 1961 Paleocene, Trinidad = Trochalosoma (Smith and Jeffery 2000) Psammechinus anguillensis Poddubiuk, 1987 Miocene, Anguilla Pseudorthopsis rojasi Sánchez-Roig, 1953 Eocene, Cuba Scoliechnius axiologus Arnold and Clark, 1927 Eocene, Jamaica = Lytechinus (Smith and Kroh, 2011) Stenechinus perplexus Arnold and Clark, 1927 Eocene, Jamaica = Echinopsis (Fell and Pawson, 1966) Stenechinus regularis Arnold and Clark, 1927 Eocene, Jamaica = Echinopsis (Fell and Pawson, 1966) Stomopneustes pristinus Jackson, 1927 Oligocene, Mexico Triadechinus multiporus Arnold and Clark, 1927 Eocene, Jamaica; type species of genus Tripneustes (Eotripneustes) tintamarrensis Poddubiuk, 1987 Lowest Miocene, Tintamarre Tripneustes tobleri Jeannet, 1928 Miocene, Venezuela Infraclass IRREGULARIA Latreille, 1825 Subterclass NEOGNATHOSTOMATA Smith, 1981 Order ECHINONEOIDA H.L. Clark, 1925 Family ECHINONEIDAE Agassiz and Desor, 1847 Genus Amblypygus L. Agassiz, 1840 Amblypygus americanus Michelin, 1856 (Figs. 31-33) Amblypygus americanus Michelin, 1856. p. 222. Amblypygus americanus (Michelin). Desor, 1858. p. 256. Amblypygus americanus (Desor). Duncan and Sladen, 1883. p. 13. Amblypygus merrilli Twitchell in Clark and Twitchell 1915. p. 165. pl. 76, figs. 1a-d. pl. 57, figs. 1a-b. pl. 78, figs. 1a-b. Amblypygus americanus (Desor). Lambert and Thiéry, 1921. p. 341. Amblypygus merrilli (Twitchell). Lambert and Thiéry, 1921. p. 341. Amblypygus americanus (Desor). Jackson, 1922. p. 55. Amblypygus americanus (Desor). Arnold and Clark, 1927. p. 25. pl. 3, figs. 1-3. Amblypygus americanus (Desor). Cooke, 1942. p. 10. Amblypygus americanus (Desor). Cooke, 1959. p. 27. pl. 7, figs. 8, 9. Amblypygus americanus (Desor). Toulmin, 1977. pp. 338-339. pl. 64, figs. 4, 5. Amblypygus americanus (Michelin). Donovan and Lewis, 1993. p. 189. Amblypygus americanus (Michelin). Donovan, 1993. pp. 382-383. fig. 9.1. Amblypygus americanus (Desor). Osborn et al., 2016. tbl. 2. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 51 Figure 31: Amblypygus americanus (UF 3297), 90 mm TD, 55 mm TH, Upper Eocene Oligopygus wetherbyi Zone of Ocala Limestone, Alachua County, Florida (FM-IP AL002). A: aboral. B: oral. C: left side. D: right side. Occurrence.—Within Florida, A. americanus occurs most commonly with O. wetherbyi in the upper portion of the Upper Eocene OLS in northern and central peninsular Florida; especially in quarries northwest of Mayo, Lafayette County (FM-IP LF001), and north of Ocala, Marion County. The type locality of Amblypygus merrilli (USNM 164934 = A. americanus) is Kendrick, Marion County, Florida (USGS 4246). Amblypygus americanus also occurs in Jackson County, in the Florida panhandle, where it rarely occurs in the Oligopygus haldemani Zone of the upper OLS in the Brooks Quarries, northwest of Marianna (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031, FM-IP JA039), and Eocene portion of the overlying Bumpnose Limestone in the same quarry. This species is also documented from the Up- per Eocene of Georgia (Cooke, 1959) and Jamaica (type locality of A. americanus: Michelin, 1856; Desor, 1858; Donovan, 1993; Donovan and Lewis, 1993). Discussion.—Desor (1858) described this ro- bust species from the Paleogene of Jamaica based on a specimen collected by Michelin and credited it ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 52 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 32: Amblypygus americanus (UF 2213), 105 mm TD; oral surface, Upper Eocene Ocala Limestone, Alachua County, Florida (FM-IP 2432). to “Michelin, mnscr”. Duncan and Sladen (1883), Jackson (1922), Arnold and Clark (1927), Cooke (1942, 1959), and most subsequent authors credit the species to Desor (1858), likely following Duncan and Sladen (1883), ignoring Desor’s attribution of the species name to Michelin. Clark and Twitchell (1915), Donovan (1993), and Donovan and Lewis (1993) are a few exceptions to this, attributing the name to Michelin as quoted in Desor (1858). However, Michelin (1856) did indeed name the species two years prior to Desor’s (1858) work. Michelin presented a plaster cast of the holotype of A. americanus to the Société Géologique de France, stating (1856: 222, translated from the orig- inal French): “this genus is remarkable by a very ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 53 Figure 33: Amblypygus americanus (UF 2218), 91 mm TD, 52 mm TH, internal mold, Upper Eocene Ocala Limestone, Alachua County, Florida (FM-IP 2434). A: aboral. B: oral. C: left side. D: right side. elongated and very large mouth almost like Echi- noneus. The American species differs from that in that it is rounded and very thick. . . ten centimeters in diameter.” Michelin then stated that the cast bears the name Amblypygus americanus. Thus, as stated by Desor (1858) himself, A. americanus should be attributed to Michelin (1856) and not Desor (1858). In a departure from Cooke (1959) and several other major monographs, we cite Michelin’s (1856) usage as the senior synonym. Twitchell in Clark and Twitchell (1915), de- scribed Amblypygus merrilli from the OLS, which he believed to be Oligocene in age, in the Kendrick Lime quarries near Ocala, Florida. However, Cooke (1959) asserted there was little doubt that the Ja- maican species (A. americanus) and American species (A. merrilli) were synonymous, an asser- tion that has been upheld by subsequent authors. Amblypygus americanus most commonly oc- curs in the upper portion of the OLS of peninsular Florida, where it is often associated with O. wether- byi and other species. A. americanus occurs more rarely in the panhandle of Florida and southwestern Georgia, where it is associated with O. haldemani and the large, star-patterned foraminiferan, Astero- cyclina. The largest specimen examined, from the Mayo Quarry (FM-IP LF001), Lafayette County, Florida, was 129 mm TL. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 54 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Superorder LUMINACEA Mongiardino Koch et al., 2022 Order CLYPEASTEROIDA A. Agassiz, 1872 Family OLIGOPYGIDAE Duncan, 1889 Genus Haimea Michelin, 1851 Haimea brooksi Osborn et al., 2016 (Figs. 34, 35) Haimea brooksi Osborn et al., 2016. pp. 18-23. figs. 11, 12. Occurrence.—This species has not been docu- mented outside of the Upper Eocene OLS of Jackson County, Florida. Osborn et al. (2016) described this taxon from the Brooks Quarry (FM-IP JA039), Sills Pit (FM-IP JA086), and Jackson Blue Spring (FM-IP JA033). Discussion.—Haimea brooksi is common at its type locality (FM-IP JA039) where it occurs with R. georgiensis, O. haldemani, O. rotundus, W. johnsoni, and many other, rarer echinoid species, in- cluding new taxa described herein: Rhyncholampas mariannaensis n. sp., Rhyncholampas bao n. sp., and Brissus jonesi n. sp. As discussed by Osborn et al. (2016), the precise stratigraphic placement of H. brooksi within the OLS cannot be determined, as neither excavation in the Brooks (FM-IP JA039) or Sills quarries (FM-IP JA086), nor exposures in Jack- son Blue Spring (FM-IP JA033) expose the base of the Haimea brooksi Zone. Within the Brooks Quarry, the zone was quarried by dragline below the water level at a depth of up to 24 m, which is 29-30 m below the top of the Eocene exposures within the quarry (Fig. 6). This horizon falls within the Asterocyclina Zone named for this abundant, large, star-shaped, foraminiferan, and below the zone of abundance of Oligopygus haldemani. Haimea brooksi can only be confused with O. haldemani, with which it occurs at all known localities. However, Haimea and Oligopygus are only superficially similar. Haimea is readily differ- entiated from Oligopygus in having a pentagonal to subpentagonal peristome that is not situated in a transverse circumoral depression, the infundibu- lum (sensu Durham [1955]). In addition, the sutures in Oligopygus are sinuous (corrugated sensu Kier, 1967), whereas they are smooth and straight in Haimea. Haimea brooksi is further differentiated from O. haldemani by its much more globose test. Genus Oligopygus de Loriol, 1887 Oligopygus haldemani (Conrad, 1850) (Figs. 36-40) Discoidea haldemani Conrad, 1850. p. 40. pl. 1, fig. 12. Discoidea haldemani (Conrad). Conrad, 1865. p. 75. Discoidea haldemani (Conrad). Conrad, 1866. p. 22. Discoidea haldemani (Conrad). Agassiz, 1883. p. 88. Oligopygus haldermani (Conrad). Clark and Twitchell, 1915. p. 167. pl. 78, figs. 4a-d, 5a-d. Oligopygus haldermani (Conrad). Cooke and Mos- som, 1929. pl. 3, figs. 3a-b. Oligopygus colsoni Lambert, 1932. p. 290. pl. 17, figs. 1-4. Oligopygus haldemani (Conrad). Cooke, 1942. p. 8. Oligopygus haldemani (Conrad). Cooke, 1945. fig. 5, no. 3. Oligopygus haldemani (Conrad). Cooke, 1959.p. 29. pl. 8, figs. 6-8. Oligopygus haldemani (Conrad). Kier, 1967. pp. 83-88. pl. 3, figs 1, 2; pl. 11, figs. 1-6; pl. 23. txt figs. 4, 8, 13-16, 19, 24, 37-39. Oligopygus haldemani (Conrad). Toulmin, 1977. p. 343. pl. 67, figs. 4-6. Oligopygus haldemani (Conrad). McKinney and Jones, 1983. pp. 21-29. 8 figs. 1 tbl. Oligopygus haldemani (Conrad). McKinney, 1984. pp. 407-419. 12 figs. 3 tbls. Oligopygus haldemani (Conrad). Rice, 1997. pp. 907-912. fig. 3. Oligopygus haldemani (Conrad). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 3. Oligopygus haldemani (Conrad). Miller et al., 2008. p. 66. Oligopygus colsoni (Lambert). Miller et al., 2014. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 55 Figure 34: Haimea brooksi (UF 329688), 25 mm TL, 21 mm TW, 20 mm TH, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. pp. 145-160. Oligopygus haldemani (Conrad). Osborn et al., 2016. tbl. 2. Occurrence.—The greatest abundance of this species is undoubtedly in the OLS of Florida (strata formerly included within the Williston Formation of Puri [1957]), although it does occur much more rarely in the Upper Eocene of Georgia. In peninsu- lar Florida, O. haldemani occurs stratigraphically below the occurrence of O. wetherbyi. Localities for this commonly encountered echinoid are too numerous to list, but notably include quarries such as northwest of Mayo in Lafayette County (FM- IP LF001), the Cemex Quarry west of Center Hill in Sumter County (FM-IP SM010), and the Haile ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 56 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 35: Haimea brooksi (UF 329689), 23 mm TL, 19 mm TW, 19 mm TH, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. quarry complex (FM-IP AL004) in Alachua County. Oligopygus haldemani is also very abundant in the OLS of the Florida panhandle, where it occurs at numerous localities in Jackson County, including the Brooks Quarries (FM-IP JA009) and Sills Pit (FM-IP JA086). Discussion.—Kier (1967) stated that O. halde- mani and O. wetherbyi are commonly found to- gether. However, this implied association is likely due to occurrences in mixed spoil from multiple horizons of the OLS often encountered in the limestone quarries of Florida. The two species are not documented together in situ in the same horizon. Oligopygus haldemani and the usually larger O. wetherbyi are only superficially similar. The ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 57 Figure 36: Oligopygus haldemani (UF 329695), 21 mm TL, 19 mm TW, 11 mm TH; uppermost bed of Upper Eocene Ocala Limestone in the Marianna Lime Quarry (FM-IP JA031), Jackson County, Florida. A: aboral. B: oral. C: left side. D: right side. periproct of O. wetherbyi is typically located much closer to the peristome than in O. haldemani, and this trait is consistent even in smaller specimens of O. wetherbyi, which could otherwise be confused with the much smaller O. haldemani. In addition, Kier (1967) clarified that the periproct of O. wetherbyi is more circular, there are more pore pairs in petals III and V, the test is slightly narrower, and the greatest width is posterior, especially in larger specimens. Oligopygus haldemani is more superficially like O. phelani from the lower portion of the OLS, with specimens of which it had been confused (Fis- cher, 1951; Cooke, 1959) until Kier (1967) erected the species O. phelani for these specimens. The two species are readily differentiated by the deeper in- fundibulum and more posteriorly situated periproct in O. haldemani. Oligopygus haldemani occurs with O. ro- tundus (the only documented occurrence of strati- graphic range overlap of Oligopygus species in the ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 58 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 37: Oligopygus haldemani (UF 329696), 16 mm TL, 15 mm TW, 8 mm TH; uppermost bed of Upper Eocene Ocala Limestone in the Marianna Lime Quarry (FM-IP JA031), Jackson County, Florida. A: aboral. B: oral. C: left side. D: right side. region), as well as the oligopygoid H. brooksi, in the lower beds of the OLS in the Brooks Quarry (FM-IP JA039) northwest of Marianna, Jackson County (Os- born et al., 2016). Oligopygus rotundus has longer petals and the posterior slope of its infundibulum is much steeper (Cooke, 1959; Kier, 1967). Two specimens referred in our treatment of O. haldemani from near Marianna, Jackson County, were described by Lambert (1932) as O. colsoni. He stated that O. colsoni is like O. haldemani but dif- fered in being less elongate, with a more pronounced ”oral sulcus” (to which we refer as an infundibulum, likely homologous with that of clypeasteroids), less inflated interporiferous zones in the petals, a lower density of pores in the petals, and shorter petals. Cooke (1942) placed O. colsoni in synonymy with O. haldemani. Cooke (1959) subsequently sur- mised that O. colsoni was a juvenile specimen of O. haldemani, attributing the decreased pore density to the small size of the former. Kier (1967) agreed with Cooke (1959) noting that although he was unable to locate the type of O. colsoni (Kier stated that it is not in the Lambert Collection at the Sorbonne, France), he asserted that the specimen ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 59 Figure 38: Oligopygus haldemani (UF 329700), 19 mm TL, 17 mm TW, 10 mm TH, Haimea brooksi Zone, Upper Eocene Ocala Limestone, Marianna Lime Quarry (FM-IP JA039), Jackson County, Florida. A: aboral. B: oral. C: left side. D: right side. Lambert figured is indistinguishable from small O. haldemani. Miller et al. (2008) compared four morpholog- ical measurements from 42 specimens of O. halde- mani from the Florida panhandle and 148 specimens from peninsular Florida, stating that the only dis- tinction that could be found was when the number of pore pairs in ambulacrum I is plotted against other test measurements. However, they indicated ambiguity in assigning single specimens to either O. haldemani or O. colsoni, considering the degree of overlap between the populations. Miller et al. (2014) reversed their earlier (2008) opinion and concluded that O. colsoni is a valid taxon and that all populations near Marianna, Jackson County, should be given that name. In doing so, they continued to acknowledge their specimens showed considerable overlap of fea- tures and that comparing an individual O. colsoni to any given O. haldemani (when referencing speci- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 60 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 39: Oligopygus haldemani (UF 337978), 21 mm TL, 18 mm TW, 10 mm TH, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. mens outside of the type material) is problematic. They indicated that of the five traits Lambert (1932) used to distinguish O. colsoni from O. haldemani, neither test shape, peristomial infundibulum, in- flation of poriferous zones, nor petal length can be used to differentiate the species. However, con- cerning the density of pore pairs, they calculated a mean pore density of 3.62 for the specimens they recognized as O. colsoni and 4.127 for specimens they identified as O. haldemani. Again, they stated that overlap of this feature implies that some indi- viduals from either species could be misidentified. It is consistent with the biology of echinoids to suggest that populations of O. haldemani in Jackson County are characterized by a slightly higher density of pore pairs. Whether or not this feature, which Miller et al. (2014) admitted has considerable over- lap with populations unambiguously assigned to O. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 61 Figure 40: Oligopygus haldemani (UF 337980), 15 mm TL, 13 mm TW, 8 mm TH, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. haldemani, is sufficient to distinguish a new species is debatable. Moreover, the proposed Oligopygus zonation of Miller et al. (2014) would only be useful if O. colsoni is separable from O. halde- mani, and even Miller et al. (2014) acknowledged it is not. Kier (1967) noted that of the 140 specimens of O. haldemani he examined, the gonopores were small on some individuals and large on others, which led him to propose that the species is sexually di- morphic. This was the first suggestion of a sexually dimorphic oligopygid. Oligopygus phelani Kier, 1967 (Figs. 41, 42) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 62 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 41: Oligopygus phelani (UF 343885), 13 mm TL, 11 mm TW, 7 mm TH, lower portion of Upper Eocene Ocala Limestone, Levy County, Florida (FM-IP LV016). A: aboral. B: oral. C: left side. D: right side. not Oligopygus haldemani (Conrad). Fischer, 1951. p. 56 (not O. haldemani). Oligopygus phelani Kier, 1967. pp. 81-83. pl. 22, txt fig. 9. Oligopygus phelani (Kier). McKinney and Jones, 1983. pp. 21-29. 8 figs. 1 tbl. Oligopygus phelani (Kier). McKinney, 1984. pp. 407-419. 12 figs. 3 tbls. Oligopygus phelani (Kier). Rice, 1997. pp. 907-912. fig. 3. Oligopygus phelani (Kier). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 2. Oligopygus phelani (Kier). Osborn et al., 2016. tbl. 2. Occurrence.—This is the common oligopy- gid of the lowermost portion of the OLS, in strata formerly known as the Inglis Formation. The type locality is spoil along the Cross Florida Barge Canal south of Inglis. This species is present, though typ- ically never as common as the oligopygids of the upper OLS, at nearly all exposures of the lower OLS, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 63 Figure 42: Oligopygus phelani (UF 343886), 11 mm TL, 10 mm TW, 5 mm TH, lower portion of Upper Eocene Ocala Limestone, Levy County, Florida (FM-IP LV016). A: aboral. B: oral. C: left side. D: right side. especially along the Withlacoochee River west of Yankeetown (FM-IP LV024) and small quarries in Levy County (e.g., FM-IP LV114). Oligopygus phelani occurs very rarely in the Upper Eocene of Georgia (Carter and Hammack, 1989; Carter and McKinney, 1992). Discussion.—Oligopygus phelani is superfi- cially similar to O. haldemani from higher in the OLS, with which it had been historically confused (Fischer, 1951; Cooke, 1959). However, upon close examination the two species are readily differenti- ated by the deep circum-peristomial infundibulum and more posteriorly situated periproct of O. halde- mani (Kier, 1967). Oligopygus phelani appears to be restricted to the lowermost OLS (formerly referred to as the Inglis Formation) in its natural outcropping area near Inglis, in northern Citrus and Levy Counties, Florida. It is present at most exposures of the lower OLS, but never as abundant as the later species O. wetherbyi and O. haldemani in the horizons within which they occur. As such, although O. phelani is recognized as an indicator species, it is perhaps not as good an indicator as the much more abundant E. mooreanus, which is also restricted to the lowermost OLS. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 64 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 43: Oligopygus rotundus (UF 329691), 50 mm TL, 45 mm TW, 22 mm TH, Haimea brooksi Zone, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. Oligopygus rotundus Cooke, 1942 (Figs. 43, 44) Oligopygus rotundus Cooke, 1942. p. 9. pl. 2, figs. 1-3. Oligopygus rotundus (Cooke). Cooke, 1959. p. 29. pl. 8, figs. 1-5. Oligopygus rotundus (Cooke). Cooke, 1961. p. 12. pl. 3, figs. 4-6. (not O. rotundus per Kier, 1967: 69). Oligopygus rotundus (Cooke). Kier, 1967. pp. 67-69. pl. 17, figs. 3-5. txt figs. 24, 31, 38, 39. Oligopygus rotundus (Cooke). Toulmin, 1977. p. 306. pl. 51, figs. 8, 9. Oligopygus rotundus (Cooke). Buitrón and Sánchez, 1979. p. 123. figs. 2a-c, Oligopygus rotundus (Cooke). Osborn et al., 2016. tbls. 2, 3. Occurrence.—Within Florida, this species has only been documented in the OLS in the Brooks Quarry (FM-IP JA039), northwest of Marianna, Jackson County, where it occurs with O. haldemani and H. brooksi. The type locality of the species is Geneva ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 65 Figure 44: Oligopygus rotundus (UF 329692), 44 mm TL, 40 mm TW, 23 mm TH, Haimea brooksi Zone, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. County, Alabama (Cooke, 1942), where it is rare in the Lisbon Formation. Toulmin (1977) documented the occurrence of O. rotundus in the Lisbon Forma- tion in Early County, Georgia. The species was also documented in the Eocene of Mexico (Buitrón and Sánchez, 1979). Discussion.—Cooke (1942) initially de- scribed this species from Middle Eocene strata at Koons Mill, in Geneva County, Alabama. Subse- quently, Cooke (1959) added two additional locali- ties in Alabama. Cooke (1961) also documented specimens of O. rotundus from the Eocene of Trinidad and Venezuela. However, Kier (1967) referred these specimens to O. zyndeli Jeannet, 1928. Kier (1967: 69) stated that O. rotundus is very similar to O. zyndeli. However, the peristomial infundibulum in O. rotundus is slightly larger and deeper longitudinally. Additional specimens of O. rotundus from Jackson County (FM-IP JA039) prove Kier’s assessment to be accurate, although the two forms are obviously very closely related. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 66 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Oligopygus rotundus was only recently docu- mented in the Upper Eocene OLS in Florida (Osborn et al., 2016), where it occurs in the Brooks Quarry (FM-IP JA039) with a diverse echinoid assemblage including: H. brooksi, R. georgiensis, O. haldemani, W. johnsoni, and others from a horizon within the Asterocyclina Zone (sensu Puri, 1957), roughly 24 to 29 m below the contact of the upper OLS and the overlying Bumpnose Limestone within the quarry (Fig. 6). The occurrence in Jackson County, Florida is the first documentation of more than one species of Oligopygus occurring in the same horizon in the southeastern United States, where the succession within peninsular Florida is well known: O. phelani (lowermost OLS) to O. haldemani, and finally to O. wetherbyi (uppermost OLS) (Table 3). This strati- graphic succession is now modified to include O. rotundus as overlapping with the lower portion of the range of O. haldemani in the northern Florida panhandle. The significant number of specimens from the OLS of Jackson County, allows for a much bet- ter understanding of this species. The specimens from Florida are not identical with the holotype of O. rotundus, being much larger. The largest speci- men is 54.1 mm TL, 49.8 mm TW, 26.3 mm TH; more than twice as large as the holotype. Given this larger size and considering the ontogeny of species within the genus described by Kier (1967), it should come as no surprise that the general form of the test would change throughout growth. The smaller forms are more similar to the holotype. However, as the specimens become larger, they become more pen- tagonal and often develop shallow indentations in the interambulacra at the ambitus. There is also variation in periproct placement, as some specimens have a periproct much closer to the margin than is typical. This variation is not evident in the three spec- imens previously available from the type area in Alabama. If additional material of O. rotundus from the type area becomes available, it could provide for a more comprehensive comparison of the popula- tions from Alabama and Florida, but until then, we consider the Florida population to be conspecific. Oligopygus rotundus has the greatest strati- graphic range of the four species of Oligopygus currently documented from North America, span- ning from the Middle Eocene (Lisbon Formation of Alabama) to the Upper Eocene (OLS in Jackson County, Florida). In contrast, O. haldemani, O. phe- lani, and O. wetherbyi are not documented outside of Upper Eocene strata within the region (though O. wetherbyi does appear to extend into the Middle Eocene in Jamaica [Donovan, 1993]). Oligopygus rotundus is superficially like O. haldemani in adapical view. However, the species are readily differentiated because the periproct of O. rotundus is further from the margin, and its infundibulum is smaller and deeper than in O. halde- mani. Oligopygus rotundus can be differentiated from O. wetherbyi because the periproct of O. ro- tundus is closer to the margin, and it has a smaller and deeper infundibulum. Oligopygus rotundus dif- fers from O. phelani in having a broader, deeper infundibulum. Oligopygus wetherbyi de Loriol, 1887 (Figs. 45, 46) Oligopygus wetherbyi de Loriol, 1887. p. 396. pl. 17, figs. 7-7d. Oligopygus wetherbyi (de Loriol). Clark and Twitchell, 1915. p. 166. pl. 78, figs. 2a-d, 3a-b. Oligopygus floridanus Twitchell in Clark and Twitchell, 1915. p. 169. pl. 79, figs. 1a-f. Oligopygus wetherbyi (de Loriol). Cooke and Mos- som, 1929. pl. 3, figs. 2a-b. Oligopygus wetherbyi (de Loriol). Lambert, 1932. p. 290. Oligopygus wetherbyi (de Loriol). Cooke, 1942. p. 8. Oligopygus floridanus (Twitchell). Cooke, 1942. p. 8. Oligopygus wetherbyi (de Loriol). Cooke, 1945. fig. 5, no. 2. Oligopygus wetherbyi (de Loriol). Mortensen, 1948. p. 247, fig. 247. Oligopygus wetherbyi (de Loriol). Cooke, 1959. p. 28. pl. 8, figs. 9-12. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 67 Figure 45: Oligopygus wetherbyi (UF 329702), 47 mm TL, 40 mm TW, 21 mm TH, Upper Eocene upper Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. Oligopygus wetherbyi (de Loriol). Wagner and Durham, 1966. p. U448. fig. 334.5. Oligopygus wetherbyi (de Loriol). Kier, 1967. pp. 54-59. pl.1, figs. 3-5; pl. 4, figs. 1, 2, 4; pl. 5. pl. 8, figs. 1-3; pl. 10, figs. 1, 3-5; pls, 13, 14. pl.16, fig 8. txt. figs. 2, 7a, 11a, 17, 24-26, 30, 38, 39. Oligopygus wetherbyi (de Loriol). Kier, 1969. txt fig. 2. Oligopygus wetherbyi (de Loriol). Toulmin, 1977. p. 343. pl. 68, figs.1-3. Oligopygus wetherbyi (de Loriol). McKinney and Jones, 1983. pp. 21-29. 8 figs. 1 tbl. Oligopygus wetherbyi (de Loriol). McKinney, 1984. pp. 407-419. 12 figs, 3 tbls. Oligopygus wetherbyi (de Loriol). Donovan, 1993. p. 384. figs. 9.9, 9.10. Oligopygus wetherbyi (de Loriol). Rice, 1997. pp. 907-912. fig. 3. Oligopygus wetherbyi (de Loriol). Oyen and Portell, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 68 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 46: Oligopygus wetherbyi (UF 329703), 43 mm TL, 36 mm TW, 20 mm TH, Upper Eocene upper Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. 2001. pp. 193-218. pl. I, fig. 4. Oligopygus wetherbyi (de Loriol). Osborn et al., 2016. tbl. 2. Occurrence.—This species is a ubiquitous presence in the upper portion of the OLS of peninsu- lar Florida (strata formerly attributed to the Crystal River Formation), especially in quarries north of Ocala, Marion County; north of Branford, Suwan- nee County (FM-IP SU002); northwest of Mayo, Lafayette County (FM-IP LF001); and west of Dowl- ing Park, Lafayette County (FM-IP LF002). Oligopygus wetherbyi rarely occurs in the Upper Eocene of Georgia (Carter and McKinney, 1992). It is also documented from the Eocene of Jamaica (Donovan, 1993) and Cuba (Sánchez-Roig, 1949). Discussion.—Oligopygus wetherbyi is one of the key biostratigraphic indicator species of the upper portion of the OLS of Florida where the Oligopygus wetherbyi Zone overlies the Oligopygus haldemani Zone. Oligopygus wetherbyi is often the dominant species in the horizon in which it occurs, commonly occurring with the same echinoid ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 69 assemblage throughout its range. This assemblage includes P. mortoni, W. cubae, E. tanypetalis, A. americanus, R. conradi, and other, rarer species. Oligopygus wetherbyi and O. haldemani are superficially similar. The periproct of O. wetherbyi is always located much closer to the peristome than in O. haldemani, and this trait is consistent even in smaller specimens of the former, which could otherwise be confused with the much smaller O. haldemani. Kier (1967) showed that the periproct of O. wetherbyi is more circular, there are more pore pairs in petals III and V, the test is slightly narrower, and the greatest width is more posterior (especially in larger specimens) than in O. halde- mani. In addition, the interambulacral plates are commonly more tumid. Oligopygus wetherbyi also attains a greater size than O. haldemani. The largest specimen of the former documented by Kier (1967) out of 140 specimens available for his study is 54 mm TL, whereas the largest O. haldemani, out of 120 specimens he measured, was 37 mm TL. Palmer (in Sánchez-Roig, 1949) erected a variety of Oligopygus floridanus (= O. wetherbyi), called O. floridanus var. laevis, for some specimens from the Late Eocene of Cuba. However, he did not figure the specimens, so Kier (1967) did not feel comfortable adding them to his synonymy of O. wetherbyi, given that the occurrence of O. wetherbyi in Cuba is also questionable. Cooke (1961) con- sidered O. kugleri from the Eocene of Trinidad to be a subspecies of O. wetherbyi. However, Kier (1984) asserted that O. kugleri is a distinct species because O. kugleri and O. wetherbyi are very dis- tinct, with O. kugleri having a much higher test, and a much narrower and less depressed peristomial infundibulum. Suborder CLYPEASTERINA L. Agassiz, 1835 Family CLYPEASTERIDAE L. Agassiz, 1835 Genus Clypeaster Lamarck, 1801 The State of Clypeaster Systematics in the Region Clypeaster is abundantly represented in Ceno- zoic faunas of the eastern Americas and Caribbean Region, as well as globally. In endeavoring to de- scribe all Florida forms of Clypeaster from the Paleogene, we were compelled to perform a detailed review of the regional taxa (Table 6). More than 400 nominal species of fossil and extant Clypeaster have been described and grouped into a variety of subgenera distinguished mainly on gross test mor- phology (Mortensen, 1948). These subgenera have been recognized as being of little taxonomic value (Durham, 1966; Hopkins, 1988; Mihaljevic et al., 2011; Mooi and van Noordenburg, 2021; Roman, 1952). It is readily apparent that Clypeaster is grossly over-split based on the number of available charac- ters, let alone on fossils. This is especially evident when reviewing the Mediterranean faunas, but it is likewise apparent among the fossil species of the West Indies. In describing Clypeaster topilanus from the Oligocene of Mexico, Jackson (1937: 232) expressed frustration over the number of species of Clypeaster described at that time: “With all the species assigned to Clypeaster, it is annoying to make still another.” Yet, he did. Ali (1983) re- ported that at the time of his writing, 83 species of Clypeaster had been documented in Cenozoic strata of the Caribbean Province, which included the southeastern United States. Although this num- ber is undoubtedly inflated (see below), there re- mains a significant number of species from this region. A great many of these species exist only as single specimens or from single occurrences, which makes assessing their validity even more challenging. Ali (1983) documented 35 species of Clypeaster in the Oligocene of the Caribbean Province, with 26 of these from Cuba alone. An- other 17 species occurred in the Miocene of Cuba, five in additional West Indian localities, four from Venezuela (Jeannet, 1928; Casanova, 1955; Kier, 1963), four from the Miocene of Brazil (March- esini Santos, 1958; Brito and Ramires, 1974), and eight from the Miocene of Florida. Ali (1983) cited Jackson (1917), Gordon (1963) and Kier (1963) to develop his tallies, but this latter number is obviously incorrect, as during the time of his work, only Clypeaster concavus Cotteau, 1875 (sensu ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 70 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Table 6: A listing of fossil Clypeaster species described from the eastern Americas and Caribbean region. Locality provided is the type area for the species; NA=North America. Distribution of species further discussed herein is provided within the remarks for each species. Durham (1966) placed the genera Anomalanthus, Bunactus, Coronanthus, Herrerasia, Oranthus, Paratinanthus, Rhaphidoclyphus, Rojaster, Tholeopelta, and Zanolettia in synonymy with Clypeaster. Some species designations would have to be resolved if Durham’s (1966) synonymies are accepted (such as Bunactus sanchezi and Rojasaster hernandezi), as these species names are already occupied in the genus Clypeaster. Clypeaster abruptus Sánchez Roig, 1926; Miocene, Cuba C. petersonorum Osborn et al., 2020; Miocene, Florida C. antillarum Cotteau, 1875; Miocene, Antigua C. pileus Israelsky, 1924; Oligocene, Mexico C. batheri Lambert, 1915; Miocene, Antigua C. pinarensis Lambert and Sánchez Roig, 1934; Miocene, Cuba C. brodermanni Sánchez Roig, 1949; Oligocene, Cuba C. placentoides Jackson, 1922; Oligocene, Cuba C. canimarensis Palmer, 1949; Pleistocene, Cuba C. planipetalus Cotteau, 1897; Oligocene, Antigua C. caudatus Jackson, 1922; Miocene, Dominican Republic C. planus Sánchez Roig, 1949; Oligocene, Cuba C. chiapasensis Mullerried, 1951; Oligocene, Mexico C. platygaster Jackson, 1922; Oligocene, Cuba C. concavus Cotteau, 1875; Miocene, Anguilla C. polygonalis Sánchez Roig, 1949; Oligocene, Cuba C. concavus puertoricanus Gordon, 1963; Miocene, Puerto Rico C. profundus Sánchez Roig, 1949; Oligocene, Cuba C. cotteaui Egozcue, in Cotteau, 1897; Oligocene, Cuba C. rogersi (Morton, 1834); Oligocene, southeastern USA C. cryptopetalus Jackson, 1922; Oligocene, Antigua C. romani Kier, 1964; Pliocene-Pleistocene, SE USA, C. cubensis Cotteau, 1875; Miocene, Cuba C. rosaceus (Linnaeus, 1758); Pliocene-recent, FL and Caribbean, C. dalli (Twitchell, 1915); Pleistocene, Florida C. sanchezi Lambert, in Sánchez Roig, 1926; Oligocene, Cuba C. densus Sánchez Roig, 1949; Oligocene, Cuba C. sandovali Sánchez Roig, 1949; Oligocene, Cuba, C. dondolii Fischer, 1985; Miocene, Costa Rica C. sanrafaelensis Palmer, 1949; Oligocene, Cuba, C. duchassaingi Michelin, 1861; Pliocene, Guadaloupe C. staubi Lambert, 1928; Miocene, Mexico, C. douvillei Stefanini, 1911; Oligocene, Mississippi C. subdepressus Gray, 1825; Pliocene-recent; SE USA Caribbean C. elevatus Sánchez Roig, 1949; Miocene, Cuba C. sunnilandensis Kier, 1963; Pliocene, Florida C. ellipticus Michelin, 1861, Miocene, Antilles C. tenuicoronae Palmer, 1949; Oligocene, Cuba C. eurychorus Arnold and Clark, 1934; Cenozoic, Jamaica C. topilanus Jackson, 1937; Oligocene, Mexico C. gatuni Jackson, 1917; Miocene, Panama Anomalanthus elevatus Sánchez Roig, 1952; Oligocene, Cuba C. guadalupense Sánchez Roig, 1952; Oligocene, Cuba A. gigas Sánchez Roig, 1953; Oligocene, Cuba C. guillermi Sánchez Roig, 1952; Oligocene, Cuba A. guadalupense Sánchez Roig, 1952; Oligocene, Cuba C. hernandezi Sánchez Roig, 1949; Oligocene, Cuba A. oligocenicus Sánchez Roig, 1949; Oligocene, Cuba C. herrerae Sánchez Roig, 1926; Miocene, Cuba A. rojasi Sánchez Roig, 1952; Oligocene, Cuba C. julli Roman 1952; Miocene, Antigua A. zanoletti Sánchez Roig, 1952; Oligocene, Cuba C. kugleri Jeannet, 1928; Miocene, Venezuela Bunactus aguayoi Sánchez Roig, 1952; Oligocene, Cuba C. lamegoi Marchesini Santos, 1958; Miocene, Brazil B. sanchezi Lambert, 1926; Miocene, Cuba, C. lamprus H. L. Clark, 1914; Recent, (Pleistocene, Jamaica?) B. sanchezi altus Sánchez Roig, 1952; Miocene, Cuba C. lanceolatus Cotteau, 1897; Oligocene, Cuba B. sanchezi gigantea Sánchez Roig, 1952; Miocene, Cuba C. lopezriosi Sánchez Roig, 1953; Oligocene, Cuba B. santanae Sánchez Roig, 1952; Oligocene, Cuba C. maoadentroensis Kier, 1992; Pliocene, Dominican Republic Coronanthus artilesi Sánchez Roig, 1952; Oligocene, Cuba C. maribonensis Sánchez Roig, 1949; Oligocene, Cuba Co. conceptionis Sánchez Roig, 1952; Oligocene, Cuba C. marinanus Jackson, 1937; Oligocene, Mexico Herrerasia profundus Sánchez Roig, 1952; Miocene, Cuba C. meridanensis Michelin, 1850; Miocene, Guadeloupe Paratinanthus lamberti Sánchez Roig, 1952; Oligocene, Cuba C. moronensis Sánchez Roig, 1951; Miocene-Olig., Cuba Rhaphidoclyphus rojasi Sánchez Roig, 1952; Oligocene, Cuba C. ovatus Palmer, 1949; Oligocene, Mexico R. armadilloensis Sánchez Roig, 1953; Oligocene, Cuba C. oxybaphon Jackson, 1922; Oligocene, Antigua R. costulatus Sánchez Roig, 1952; Oligocene, Cuba C. palmeri Sánchez Roig, 1949; Miocene, Cuba Rojasaster hernandezi Sánchez Roig, 1952; Oligocene Cuba, C. paraensis Brito, 1979; Miocene, Brazil R. camagueyanus Sánchez Roig, 1952; Oligocene, Cuba C. parrae Desmoulins, 1835; Neogene, Cuba and Guadeloupe Tholeopelta herrerae Sánchez Roig, 1951; Oligocene, Cuba C. parvus Michelin, 1861; Miocene, Guadeloupe Zanolettia zanolettii Sánchez Roig, 1951; Oligocene, Cuba C. paulinoi Marchesini Santos, 1958; Miocene, Brazil Z. gigantea Sánchez Roig, 1952; Oligocene, Cuba Cooke, 1959) (= Clypeaster petersonorum Osborn et al., 2020), was documented from the Miocene of Florida. Ali (1983) noted that the Pliocene strata of the Caribbean Province contained only six species from the southeastern United States and Cuba (Cotteau, 1897; Jackson, 1922; Engel, 1961; Kier, 1963), and four species from Cuba and Florida (Sánchez-Roig, 1949, 1952b; Kier, 1963). Correcting for the error in count for the Miocene of Florida leaves 75 species of Clypeaster in the province by Ali’s (1983) count. We have re- searched 86 species of fossil Clypeaster described from eastern North America southward throughout the eastern Americas and eastward through the West Indies and Caribbean region (Table 6). Sánchez-Roig (1949) documented the occur- rence of 35 species of Clypeaster in the Ceno- zoic of Cuba, as well as two species he assigned to Anomalanthus, which Durham (1966) reas- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 71 signed to Clypeaster. Sánchez-Roig (1951-1953) subsequently documented another 23 species of Clypeaster, as well as several new genera he erected, such as Herrerasia, Rojaster, and Zanolettia, all which Durham (1966) also synonymized with Clypeaster. This still left a questionable total of nearly 60 species of Clypeaster within Oligocene to Pleistocene strata of Cuba alone. Most of these species were poorly figured, insufficiently described, with type material poorly designated, lost, or un- available. Many were likely described without con- sidering variation within species, comparison with other candidate taxa in the region, let alone globally, and most of the type specimens figured in Sánchez- Roig’s various works are incomplete and lack detail, especially considering the types of characters now suggested to be necessary for full diagnoses (Mooi and van Noordenburg, 2021), as indicated below. Poddubiuk’s (1985) reevaluation of type ma- terial of the Upper Oligocene to Middle Miocene species of Clypeaster from the Caribbean Islands east of Cuba suggested that the occurrences should be grouped into no more than seven species: C. batheri; C. caudatus Jackson, 1922; C. concavus; C. cubensis Cotteau, 1875; C. julii Roman, 1952, and C. oxybaphon (he listed six, but stated there should be no more than seven valid species). Poddubiuk’s PhD thesis (Poddubiuk, 1987) contains discussions on potential junior synonyms of C. batheri, C. con- cavus, and C. oxybaphon, and is discussed in the remarks for C. oxybaphon below. Poddubiuk (1985, 1987) attributed over-splitting to previous workers describing species based on small sample sizes with few characters and without considering the intraspe- cific variation (a problem clearly not confined to the Caribbean faunas). Poddubiuk (1985, 1987) stated that the prob- lem was compounded among Caribbean mid- Cenozoic faunas by two additional factors: 1) most early workers were in Europe (although, most of these species were described by Sánchez Roig in Cuba), basing their species designations on small col- lections sent to them by collectors and accompanied by minimal locality and/or stratigraphic data; and 2) type specimen repositories are scattered across the Caribbean, USA, and Europe, forcing many workers to rely on poor illustrations and incomplete and often inaccurate descriptions. While we agree with the overall message of these assertions, Poddubiuk’s claims concerning this taxonomic history remain unsupported by evidence. Furthermore, it is virtu- ally impossible to apply his findings to the Florida Paleogene occurrences. This is in spite of the fact that several additional papers, including Poddubiuk and Rose (1984), Poddubiuk (1985), and Rose and Poddubiuk (1987) continued to comment, without data concerning synonymies, on the systematics and evolution of Clypeaster in the Caribbean region. Therefore, faced with a multitude of poorly de- scribed species, anyone describing a new species of Clypeaster from the eastern United States is forced to rely on previous work that is difficult if not impos- sible to penetrate. For example, such an endeavor proved problematic in describing C. petersonorum (Osborn et al., 2020), and even for description of the extant Clypeaster brigitteae (Mooi and van Noor- denburg, 2021). Mooi and van Noordenburg (2021: 1 and 2) also discussed some of these issues while comparing fossils to extant forms: ”... there [are] in excess of 350 nominal fossil Clypeaster found in strata from the Eocene to the Recent ... an un- wieldy level likely to cause considerable challenges for a complete revision of the group. Some attempts to deal with what appears to be rampant splitting among fossil Clypeaster taxa have been made on a local level ... but a full assault on the mountain of paleontological nomenclature on a global scale has yet to be made. With that in mind ... most neontolo- gists have generally ignored the seemingly endless job of dealing with the fossil forms when describing new extant species discovered since Mortensen’s (1948) monograph ...” This underscores our sense that providing taxa in open nomenclature is the best that can be done for some of the species we are attempting to document for the Florida Paleogene. Jackson (1922) provided a key to 19 Clypeaster species documented from the Caribbean region at the time. These included: C. rosaceus Linnaeus, 1758; C. concavus, from the Oligocene of Anguilla and Puerto Rico; C. caudatus, from the Miocene of the Dominican Republic; C. dalli ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 72 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Twitchell in Clark and Twitchell, 1915, herein recog- nized as C. rosaceus, from Florida; C. cubensis from the Miocene of Cuba and Puerto Rico; C. parrae Desmoulins, 1837, from the Cenozoic of Cuba and Guadeloupe; C. lanceolatus Cotteau, 1897 from the Oligocene of Cuba, which Cooke (1959) considered a synonym of C. concavus; C. antillarum Cotteau, 1875, from Miocene of Anguilla and Antigua, which Cooke (1959) considered a synonym of C. concavus; C. planipetalous Cotteau, 1875, from the Miocene of Antigua; C. cryptopetalus Jackson, 1922, from the Miocene of Antigua; C. cotteaui from the Oligocene of Cuba and Florida; C. batheri from the Miocene of Antigua and Oligocene of Florida; C. parvus Michelin, 1861, from the Miocene of Cuba and Antigua; C. duchassaingi Michelin, 1861, from an unstated age in Guadeloupe; C. ambigenus Lamarck, 1816, from Guadeloupe, which is undoubtedly a misidentifi- cation since C. ambigenus is a subjective junior synonym of C. humilis (Leske, 1778) (Jackson stated Michelin documented it from the Eocene of Georgia, which is also clearly erroneous); C. placentoides Jackson, 1922, from the Oligocene of Cuba; C. meridianensis Michelin, 1861, from the Miocene of Guadeloupe; C. oxybaphon from the Oligocene of Antigua, Puerto Rico and Florida; and C. platygaster Jackson, 1922, which Cooke (1959) considered to be a synonym of C. oxybaphon, from Cuba. Poddubiuk (1987) provided a key to differen- tiate the fossil Clypeaster species of Anguilla and Antigua: C. batheri, C. julii, C. oxybaphon, and C. concavus. Donovan (1993) provided a very useful key to the Cenozoic species of Clypeaster of Jamaica, which included C. cotteaui; C. rosaceus; C. con- cavus; C. lanceolatus, and C. eurychorus Arnold and Clark, 1934. Arnold and Clark (1927) also considered two specimens from the Cenozoic of Jamaica to represent Clypeaster antillarum Cotteau, 1875. As noted in Osborn et al. (2020), C. antillarum has been considered a subjective junior synonym of C. concavus (Donovan, 1983: 386; Poddubiuk, 1985: 76). The remaining species of Clypeaster doc- umented from the Cenozoic deposits of the eastern Americas are summarized in Table 6. Within the Cenozoic deposits of the eastern United States, the genus is best represented within the Oligocene, with C. cotteaui, C. marinanus, C. oxybaphon, C. rogersi, and three taxa discussed below in open nomenclature: Clypeaster sp. A (re- ferred to C. batheri by Oyen, 2001); Clypeaster sp. B and Clypeaster sp. C, both from the basal Suwannee Limestone of Hernando County, Florida. For the record, the Miocene contains only C. petersonorum from the Chipola Formation of north Florida; the Pliocene contains C. sunnilandensis Kier, 1963, the Plio-Pleistocene C. romani Kier, 1964; and the Pleistocene C. rosaceus and C. subdepressus (Gray, 1825). It is therefore difficult to assign with confi- dence any of the newly discovered forms to more than open nomenclature, as they could fall within one of the 86 taxa from the remainder of the east- ern Americas (Table 6), and more likely into some of the 38 species from the Oligocene of Cuba de- scribed in one of several works by Sánchez-Roig outlined above, the majority of which will likely remain nomena nuda. However, we can say that none of Sánchez Roig’s figured specimens appear similar to Clypeaster sp. B from the Suwannee Limestone. However, we emphasize again that his figures generally do not show views necessary to assess features used classically for the genus, let alone those advocated by more recent workers. For example, Nisiyama (1968: 34) stated that test shape, position of the periproct, form and relative length of petals, tuberculation, and character of the buccal membrane are of importance in classifying species of Clypeaster. Mooi and van Noordenburg (2021) developed these ideas further in reviewing the species of Clypeaster found in the Philippines. Their study showed that value can be placed on fea- tures seldom observable on fossils, such as spines or spicules. However, other characters such as tubercle and pore pair densities in the poriferous zone, den- sity of tubercles on the aboral surface outside the petals, degree of plate occlusion at the ends of the petals, depth of the infundibulum, certain aspects of the oral plate pattern, detailed morphometerics of periproct position, petal dimensions, TH, TW, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 73 Figure 47: Clypeaster cotteaui (UF 337958), 64 mm TL, 50 mm TW, 18 mm TH, Oligocene Bridgeboro Limestone, Washington County, Florida (FM-IP WG002). A: aboral. B: oral. C: left side. D: right side. internal structure, and other features seldom used to differentiate members of can and should be assessed in fossil taxa. It is our hope to do so with additional fossil material from the Paleogene of Florida so that better characterizations can be made of those taxa we present in open nomenclature, but that we were intent on including so that the diversity of Clypeaster in the region would be more accurately represented than in the past. We provide a tabular key to the Florida species of Paleogene Clypeaster in Table 7. Clypeaster cotteaui Egozcue in Cotteau, 1897 (Figs. 47, 48, Table 7) Clypeaster cotteaui Egozcue, in Cotteau, 1897. p. 40, pl. 10, figs. 1-1. Clypeaster cotteaui (Egozcue). Lambert, 1915. p. 24. Clypeaster cotteaui (Egozcue). Jackson, 1922. p. 41, pl. 6, figs. 6-8. Clypeaster cotteaui (Egozcue). Cooke, 1942. p. 13. Clypeaster cotteaui (Egozcue). Sánchez-Roig, 1949. p. 68. Clypeaster cotteaui (Egozcue). Cooke, 1959. pp. 36-37. pl. 12, figs. 1-3. Clypeaster cotteaui (Egozcue). Pickering, 1970. pp. 20, 58. Clypeaster cotteaui (Egozcue). Rose and Poddubiuk, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 74 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) 1987. p. 466. fig. 1. tbl. 1. Clypeaster cotteaui (Egozcue). Donovan, 1993. p. 388. Occurrence.—Clypeaster cotteaui is the com- mon clypeasteroid of the Bridgeboro Limestone, which outcrops in southwestern Georgia, southeast- ern Alabama, and Washington County (e.g., FM-IP WG002) in northern Florida. This species is also documented from the type locality in Cuba (Cotteau, 1897). Donovan (1993) tentatively identified it from the Oligocene of Jamaica. Discussion.—Jackson (1922) recognized C. cotteaui from Jamaica and Arnold and Clark (1927) reported a test of uncertain geologic age that they referred to this species from a river near Port Maria, Parish of St. Mary, Jamaica. How- ever, Donovan (1993), in his review of Jamaica’s Cenozoic echinoid fauna, suggested that C. cotteaui occurred there. Poddubiuk and Rose (1984: table 2) indicated that specimens of C. cotteaui listed from Antigua (Lambert, 1915) are conspecific with Clypeaster batheri Lambert, 1915. Clypeaster cot- teaui is therefore undoubtedly represented only in the Oligocene strata of the southeastern United States and Cuba. Clypeaster cotteaui is the dominant species in the Bridgeboro Limestone (especially the Florala Member) in southwestern Georgia and southeast- ern Alabama, and in the Duncan Church beds at the Duncan Church Quarry in Washington County. Its very swollen test, with thick rounded margins, deeply concave oral surface, and petals that are wide open distally readily distinguish C. cotteaui from any other fossil species in the fauna of the eastern United States (Table 7). Clypeaster cotteaui is readily differentiated from C. rogersi, which is often found with C. cotteaui in mixed spoil of the Marianna and Bridgeboro Limestones in the Dun- can Church Quarry (FM-IP WG002), Washington County, Florida, by the much thicker margin, more concave oral surface, and petals that are wide open distally. It is also distinguished from Clypeaster oxybaphon Jackson, 1922, with which it also occurs but much more rarely, by the thicker margin, more concave oral surface, and lack of an inframarginal depression on the aboral surface in C. cotteaui (Table 7). Jackson (1922) stated the widely open petals, nearly straight line of inner pores of the pore pairs, and the four pore pairs situated distal to the ends of each column or pore pairs of the petals are features useful in distinguishing C. cotteaui from similar Clypeaster species of the Caribbean region. Clypeaster marinanus Jackson, 1937 (Figs. 48-51, Table 7) Clypeaster marinanus Jackson, 1937. p. 231. pl. 12, fig. 2; pl. 13, fig. 1. Clypeaster marinanus (Jackson). Caso, 1957. p. 508. Clypeaster rogersi (Morton). Cooke, 1959. pp. 36- 37 (in part, Cooke believed C. marinanus to be a synonym of C. rogersi). Clypeaster marinanus (Jackson). Zachos and Mo- lineux, 2007. pp. 79-91. figs. 6 a-b, e. Occurrence.—Within Florida, this is the com- mon echinoid of the lowermost Oligocene portion of the Bumpnose Limestone in Jackson County, es- pecially in Brooks Quarry near Marianna (FM-IP JA021). A single specimen likely attributable to this species was collected in the Suwannee Limestone at the Cabbage Grove Quarry west of Perry, Taylor County (FM-IP TA001). Clypeaster marinanus has not been documented from the Suwannee Lime- stone of peninsular Florida. Zachos and Molineux (2007) reported it from the Oligocene of Texas. The type locality for the species is the Oligocene Meson Formation in Tamaulipas, Mexico (Jackson, 1937). Discussion.—This is the first report of the species in Florida. However, the occurrence of a species of Clypeaster that is distinct from C. rogersi in the Bumpnose Limestone is well docu- mented. Moore (1955: 39) asked C. Wythe Cooke to examine Clypeaster specimens from the Bumpnose Limestone. Cooke agreed that the specimens repre- sented a new species closely related to C. rogersi. Cooke (1959) failed to mention this occurrence of a Clypeaster from the Bumpnose Limestone, even as he placed Jackson’s C. marinanus in synonymy with C. rogersi. Cooke (1959) also stated that spec- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 75 Ta bl e 7: Ta bu la rk ey of th e Pa le og en e C ly pe as te r of Fl or id a. K ey fo llo w sM oo ia nd va n N oo rd en bu rg (2 02 1) . Sp ec ie so f C ly pe as te r D ist r ib ut io n1 Te s t ou tli ne Av er ag e TW as % of TL Av er ag e m ar gi n he ig ht as % TH 2 Av er ag e TH as % TL D eg re e to w hi ch pe ta ls ar e op en di sta lly W id th pe ta l III as % pe ta l le ng th 3 Le ng th pe ta l II Ia s% of TL M ar gi n in fla te d, w ith pr ox im al de pr es si on of ab or al su rfa ce 4 O ra ls ur fa ce co nc av ity # Pr im ar y tu be rc le sb et w ee n in ne ra nd ou te rp or e of re sp ira to ry tu be fo ot 5 co tte au i FL ,G A ,C B O va te - su bp en ta go na l 83 .6 (7 8- 89 ) 66 .3 (6 3- 69 ) 29 .4 (2 6- 32 ) A ll w id e op en 57 .2 (5 3- 63 % ) 34 .2 (3 3- 36 ) N o D ee pl y co nc av e 5- 7 m ar in an us A L, FL ,M X , TX Su pe nt ag on al 90 .1 (8 6- 92 ) 35 .4 (3 1- 48 ) 16 .6 (1 4- 20 ) A ll so m ew ha t op en 58 (5 0- 65 ) 26 .5 (2 3- 28 ) N o N ea rly fla t 5- 7 ox yb ap ho n6 FL ,C B O va te - su bp en ta go na l 70 -9 0 50 -9 0 14 -2 2 A ll ne ar ly cl os ed 45 -6 0 25 -3 5 Ye s Va ria bl y co nc av e 9- 12 ro ge rs i7 A L, G A ,F L, M S, N C O va te - su bp en ta go na l 89 .1 (8 2- 95 ) 53 .4 (4 5- 67 ) 22 .3 (1 7- 28 ) A ll us ua lly w id e op en 8 53 .4 (4 7- 68 ) 31 .8 (2 8- 34 ) N o N ea rly fla t 5- 8 sp .A FL El on ga te su bp en ta go na l 75 .4 44 21 A ll w id e op en 9 42 .9 34 .7 N o C on ca ve 6- 8 sp .B FL Su bp en ta go na l 91 .1 (8 5- 95 ) 36 .4 (3 3- 41 ) 19 .6 (1 8- 26 ) A ll w id e op en 53 .4 (4 1- 59 ) 32 .9 (3 1- 34 ) N o Sl ig ht ly co nc av e 7- 9 sp .C FL Su bp en ta go na l 91 .6 41 .2 13 .6 A ll w id e op en 60 .3 31 .3 N o N ea rly fla t N ot di sc er na bl e 1 S ta nd ar d U .S .p os ta ls er vi ce sta te ab br ev ia tio n, pl us ”C B ” = C ar ib be an ,“ M X ” = ea ste rn M ex ic o. 2 M ea su re m en tt ak en at an te rio rm ar gi n as in fig ur e, lo w er rig ht . 3 M ea su re m en tt ak en at w id es tp oi nt of pe ta lI II in am bu la cr um II I. 4 I n C .o xy ba ph on ,t he m ar gi n is in fla te d re la tiv e to th e re st of th e di sta lp ar to ft he ab or al su rfa ce so th at th e di sta le nd so ft he pe ta ls ar e be lo w th e le ve lo ft he m ar gi n. 5 N um be ro ft ub er cl es in po rif er ou sz on e be tw ee n ad ja ce nt po re pa irs ne ar m id po in to fp et al II Ia ,w he re po rif er ou sz on e is ge ne ra lly w id es t. 6 D at a su pp le m en te d w ith m ea su re m en ts fr om Po dd ub iu k (1 98 7) ,o nl y ra ng e pr ov id ed ,b ut no ta ve ra ge s. 7 P op ul at io ns sa m pl ed fr om th e M ar ia nn a Li m es to ne in FL ,M S, an d A L. 8 P et al in am bu la cr um II Is om et im es sl ig ht ly cl os ed di sta lly . 9 P et al II Is lig ht ly m or e cl os ed th an ot he rp et al s. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 76 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 48: Comparison of Oligocene species of Clypeaster from Florida. Aboral, oral and leftside views shown for all. A: Clypeaster rogersi, (UF 337962) Marianna Limestone, Jackson County, (FM-IP JA029). B: Clypeaster ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 77 (Fig. 48 caption cont.) marinanus (UF 337995) Bumpnose Limestone, Jackson County, (FM-IP JA021). C: Clypeaster cotteaui (UF 337958) Bridgeboro Limestone, Washington County, (FM-IP WG002). D: Clypeaster sp. B. (UF 338008) Suwannee Limestone, Hernando County, (FM-IP HE038). E: Clypeaster cf. C. oxybaphon (UF 342868) Bridgeboro Limestone, Washington County, (FM-IP WG002). F: Clypeaster sp. C. (UF 338011) Suwannee Limestone, Hernando County, (FM-IP HE019). G: Clypeaster sp. A. (UF 2546) Suwannee Limestone, Suwannee County, (FM-IP SU002). H: Clypeaster rogersi (UF 27190) Suwannee Limestone, Polk County (FM-IP PO017). Figure 49: Clypeaster marinanus (UF 337995), 49 mm TL, 45 mm TW, 8 mm TH, Lower Oligocene Bumpnose Limestone, Jackson County, Florida (FM-IP JA021). A: aboral. B: oral. C: left side. D: right side. imens from the Suwannee Limestone and Meson Formation of Mexico (= C. marinanus for the Mex- ican occurrence) tend to have slightly wider inter- poriferous zones and a thinner margin than the typi- cal Marianna Limestone specimens of the species (he did not mention the distinctly shorter petals of C. marinanus). Huddlestun (1993: 66, 67) reiterated the point made by Moore (1955: 39) and stated the Bumpnose Limestone does not contain C. rogersi, but rather contains a smaller, undescribed, probably ancestral species. Osborn et al. (2016) tentatively considered this an undescribed species. It was during the pro- cess of assessing that assumption and examining a significant number of specimens from the Bumpnose Limestone near Marianna, Florida, and Brooklyn, Alabama, that we realized these specimens cannot be differentiated from C. marinanus. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 78 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 50: Clypeaster marinanus (UF 337996), 41 mm TL, 38 mm TW, 9 mm TH, Lower Oligocene Bumpnose Limestone, Jackson County, Florida (FM-IP JA021). A: aboral. B: oral. C: left side. D: right side. Jackson (1937) described C. marinanus as closest in form to C. rogersi, but different in having a flatter test, thinner margin, less sunken infundibulum around the peristome, and shorter, narrower, and less curved ambulacra. These attributes are here recognized as features distinguishing C. marinanus from C. rogersi (Table 7). While C. marinanus is the characteristic echi- noid of the Bumpnose Limestone, it also occurs in the correlative Red Bluff Formation of southwestern Alabama and the Suwannee Limestone in the Cab- bage Grove Quarry west of Perry, Taylor County (FM-IP TA001). It has not been documented in the typical Suwannee Limestone of peninsular Florida. Clypeaster oxybaphon Jackson, 1922 (Figs. 48, 52, Table 7) Clypeaster oxybaphon Jackson, 1922. p. 44. pl. 7, figs. 3, 4; pl. 8, figs. 1-3. Clypeaster oxybaphon (Jackson). Cooke, 1942. p. 13. pl. 8, fig. 7. Clypeaster oxybaphon (Jackson). Sánchez-Roig, 1949. p. 81. ?Clypeaster polygonalis Sánchez-Roig, 1949. p. 84, pl. 7, figs. 1, 2. ?Clypeaster pinarensis Lambert and Sánchez-Roig in Sánchez-Roig, 1949. p. 86. pl. 12, figs. 1, 2. ?Clypeaster profundus Sánchez-Roig, 1949. p. 91. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 79 Figure 51: Clypeaster marinanus (UF 232289), 77 mm TL, 71 mm TW, 12 mm TH, Lower Oligocene Suwannee Limestone, Taylor County, Florida (FM-IP TA001). A: aboral. B: oral. C: left side. D: right side. pl. 6, figs. 1-3. Clypeaster oxybaphon (Jackson). Caso, 1957. p. 505. figs. 6-8. Clypeaster oxybaphon (Jackson). Cooke, 1959. p. 35. pl. 11, fig. 1. Clypeaster oxybaphon (Jackson). Gordon, 1963. pp. 636-637. txt fig. 2d. pl. 79, figs. 3, 4. Clypeaster oxybaphon (Jackson). Rose and Poddu- biuk, 1987. p. 466. fig. 1. tbl. 1. Clypeaster oxybaphon (Jackson). Poddubiuk, 1987. pp. 58-60, 278-279. pls. 91-94. Clypeaster oxybaphon (Jackson). Donovan, 2004. pp. 144-145. figs. 2, 3, 4. Clypeaster cf. oxybaphon (Jackson). Zachos and Molineux, 2007. pp. 79-91. Occurrence.—Within Florida, this species only rarely occurs in the Bridgeboro Limestone at the Duncan Church Quarry, Washington County (FM-IP WG002). Zachos and Molineux (2007) tentatively referred fragmentary material from the Oligocene at Damon Mound, northwestern Brazoria County, Texas to this species. Clypeaster oxybaphon is much more commonly distributed in Oligocene strata throughout the Caribbean region, including Antigua (type locality: Jackson, 1922), Cuba, Ja- maica, Panama, and Puerto Rico (Cooke, 1959; Gordon, 1963). Discussion.—Cooke (1959) first documented the presence of C. oxybaphon in the Oligocene of Washington County (FM-IP WG002), noting that the shape of the petals, thickened margin, broad ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 80 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 52: Clypeaster cf. C. oxybaphon (UF 342868), 74 mm TL, 69 mm TW, 16 mm TH, Oligocene Bridgeboro Limestone, Washington County, Florida (FM-IP WG002). A: aboral. B: distal end of petal III. C: oral. D: posterior. E: anterior. F: left side. G: tilted aboral viewpoint from posterior. H: tilted aboral viewpoint from anterior. I: tilted oral viewpoint from posterior. J: oblique oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 81 posterior truncation, and nearly flat lower surface were consistent features of the species. Although there is a submarginal, aboral depression in all the specimens Cooke (1959) had available, he noted that the degree of inflation of the apical region is variable. The depressed submargin and lower test of C. oxybaphon readily differentiate it from the more inflated C. cotteaui, which also has much thicker mar- gins but without the periperal depression between the margin and the petaloid area. Additionally, the petals of C. cotteaui are wide open distally, whereas they are nearly closed in C. oxybaphon (Table 7). The petaloid area is variably slightly inflated, as evidenced by the Bridgeboro Limestone specimens, and noted by Gordon (1963) concerning Puerto Rican specimens. The Bridgeboro Limestone tests have petals that are more open distally than typical C. oxybaphon, but not nearly as widely open as those of C. rogersi. Cooke (1959) considered C. platygaster Jack- son, 1922; C. polygonalis Sánchez-Roig, 1949; C. pinarensis Lambert and Sánchez-Roig, 1934; and C. profundus Sánchez-Roig, 1949, to be subjective junior synonyms of C. oxybaphon. However, Cooke (1959) stated that the three species of Sánchez-Roig, all from Cuba, are too poorly figured for complete assessment, and Gordon (1963) agreed, and Pod- dubiuk (1987) agreed with all these synonymies except for C. platygaster. Poddubiuk (1987:279) noted: “A numerical study of Cuban and Puerto Rican material available to the author suggests that although C. oxybaphon and C. platygaster are very closely related, the latter can be distinguished by greater petal closure, if not absolutely on the basis of oral surface concavity, Jackson’s original diagnostic character.” We follow Poddubiuk and do not include C. platygaster in the synonymy of C. oxybaphon above. Caso (1957) documented the presence of C. oxybaphon in Mexico, but Gordon (1963) disagreed with her identification. Caso’s designation should remain questionable until the specimen is reexam- ined. Within the Bridgeboro Limestone of Wash- ington County, C. oxybaphon occurs with the much more common C. cotteaui, as well as L. floralanus, A. mossomi, and other, rarer species. Clypeaster rogersi (Morton, 1834) (Figs. 48, 53-60, Table 7) Scutella rogersi Morton, 1834. p. 77. pl. 13, fig. 3. Lagana rogersi (Morton). Agassiz, 1840. p. 6. not Scutella rogersi (Morton). Agassiz, 1841. p. 85. pl. 19a, figs. 1-4 (= Periarchus quinquefarius). Scutella jonesi Forbes in Lyell, 1845. p. 574. txt fig. not Mortonia rogersi (Morton). Desor, 1858. p. 231. (=Periarchus quinquefarius). ?Mortonia turgida Conrad, 1865. p. 184. Clypeaster rogersi (Morton). Conrad, 1866. p. 22. Clypeaster jonesi (Forbes). Conrad, 1866. p. 22. Mortonia tumidus (Conrad). Conrad, 1866. pp. 22, 37. Clypeaster tumidus (Conrad). Conrad, 1866. pp. 22, 37. Scutella (Mortonia) rogersi (Morton). Gregorio, 1890. p. 250. pl. 43, fig. 16 (not description or figs. 17-20 = Periarchus quinquefarius). Clypeaster douvillei Stefanini, 1911. p. 682. pl. 22, figs. 1a-c. Clypeaster rogersi (Morton). Clark and Twitchell, 1915. p. 136. pl. 64, figs. 2a-d, 3a-d (includes additional references). Clypeaster rogersi (Morton). Cooke, 1926. pl. 97, fig. 7. Clypeaster rogersi (Morton). Cooke and Mossom, 1929. pl. 7, fig. 7. Clypeaster rogersi (Morton). Cooke, 1942. p. 12. Clypeaster brodermanni Sánchez-Roig, 1949. p. 74. pl. 9, figs. 1-3. Clypeaster aff. rogersi (Morton). Sánchez-Roig, 1949. pp. 82, 83. pl. 8, fig. 4. Clypeaster rogersi (Morton). Cooke, 1959. pp. 36- 37. pl. 12, figs. 4-6. Clypeaster rogersi (Morton). Pickering, 1970. pp. 20, 57. Clypeaster rogersi (Morton). Dockery, 1980. p. 191. pl. 80, figs. 6, 7. Clypeaster rogersi (Morton). Kier, 1997. pp. 8, 10. pl. 6, figs. 3, 4. Clypeaster rogersi (Morton). Oyen and Portell, 2001. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 82 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 53: Clypeaster rogersi (UF 27190), 59 mm TL, 52 mm TW, 17 mm TH, Lower Oligocene Suwannee Limestone, Polk County, Florida (FM-IP PO017). A: aboral. B: oral. C: left side. D: right side. pp. 193-218. pl. I, fig. 7. Occurrence.—This is the ubiquitous sand dollar of the Lower Oligocene Marianna Lime- stone throughout the Gulf Coastal Plain, commonly collected in the Marianna Limestone in the Florida panhandle, especially in Jackson and Washing- ton Counties. Notable localities include the Dun- can Church Quarry (FM-IP WG002), Washington County, and the Brooks Quarries (FM-IP JA013, FM-IP JA019, FM-IP JA026, FM-IP JA029) north- west of Marianna, Jackson County. Moldic spec- imens are frequent in the dolomitic facies of the Marianna Limestone in the bed and banks of Dry Creek, south of Marianna, Jackson County (FM-IP JA010). The only specimens that appear to belong to this species from the Suwannee Limestone are from the Terramar Quarry in Polk County (FM- IP PO017). Specimens from the lower Suwannee Limestone in the Brooksville Rock Quarry (now Vulcan Quarry) northwest of Brooksville (FM- IP HE038), Hernando County, are discussed as Clypeaster sp. B. Clypeaster rogersi is very abundant in the Marianna Limestone of Alabama, the Marianna and Glendon Limestones of Mississippi, and Lower ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 83 Figure 54: Clypeaster rogersi (UF 31970), 67 mm TL, 61 mm TW, 11 mm TH, Lower Oligocene Suwannee Limestone, Polk County, Florida (FM-IP PO017). A: aboral. B: oral. C: left side. D: right side. Oligocene strata of Georgia. Kier (1997) recorded three fragments of this species from the River Bend Formation (sensu Ward, 2007) of North Carolina. Clypeaster rogersi has also been reported from the Oligocene of Cuba (Sánchez-Roig, 1949; Cooke, 1959). Discussion.—Clypeaster rogersi is the most widespread species in Oligocene deposits of the eastern United States, distributed eastward from the banks of the Mississippi River in Vicksburg, Mississippi, southward to Florida, and northward to North Carolina. It is also the characteristic echinoid of the Marianna Limestone, the dominant Lower Oligocene unit of the Gulf Coast. Described and figured by Morton (1834), it was among the first echinoids documented from the Cenozoic deposits of North America. It should come as no surprise that a species with a lineage that dates back nearly 200 years should have some amount of taxonomic compli- cation. Louis Agassiz (1841: pl. 19a, figs. 1-4) created the first instances of this when he figured a specimen of Periarchus quinquefarius Say, 1825 as C. rogersi. Desor (1858) repeated this error, com- pounding it by founding the new genus Mortonia for what was P. quinquefarius but citing Morton’s figure of C. rogersi. Gregory (1892) first explicated the errors of Agassiz (1841) and Desor (1858), and Clark and Twitchell (1915: 137) stated the confusion between C. rogersi and Say’s P. quinquefarius “was found to be in a tangle requiring significant labor to unravel”. Thankfully, Clark and Twitchell (1915) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 84 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 55: Clypeaster rogersi (UF 337998), 66 mm TL, 57 mm TW, 14 mm TH, Lower Oligocene Marianna Limestone, Washington County, Florida (FM-IP WG033). A: aboral. B: oral. C: left side. D: right side. persisted and corrected the synonymies of both C. rogersi and P. quinquefarius. Forbes, in Lyell (1845), described Scutella jonesi from specimens Lyell collected near Jones- boro, Georgia, and his figures of S. jonesi clearly depict C. rogersi. Subsequently, Conrad (1865) de- scribed Mortonia turgida from Jasper County, Mis- sissippi, which he stated was allied to C. rogersi, but was larger, more elevated medially, and with a greater depression around the peristome. However, he did not figure the form. Conrad (1866) likely had the intent to change the name of his M. turgida, as within a year of describing the species he listed M. tumida (Conrad) from Mississippi, and not M. turgida. Conrad (1865: 184) described M. turgida from Jasper County, Mississippi. Considering the species has not been further described or discussed since, it is worth reproducing Conrad’s descrip- tion and comments on the species here: “Mortonia turgida: suboval or subpentagonal, swelling medi- ally, with a convex outline; thin on the submarginal portion of the disc; ambulacra elliptical. Allied to C. rogersi, but larger, thinner round the central promi- nence, more elevated medially, the depression about the mouth greater, and the anus smaller. It bears about the same relation to Cl. rogersi as Sismondia alta bears to S. lyelli.” In his notes (Conrad, 1866: 37) he listed Mortonia tumida in reference to the work in which he described M. turgida. He then subsequently changed ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 85 Figure 56: Clypeaster rogersi (UF 337962), 63 mm TL, 53 mm TW, 13 mm TH, Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA029). A: aboral. B: oral. C: left side. D: right side. the name to C. tumidus (Conrad, 1866: 22). This is not to be confused with the modern Pacific form Clypeaster tumidus Tenison-Woods, 1878, which is apparently a homonym of C. tumidus Conrad, 1866, which seems to be an available name despite the confusion Conrad himself introduced. Conrad (1866) did not provide a rationale for his species redesignation from turgida to tumida, then finally to tumidus. Presumably, he was attempting to introduce agreement in gender with the genus name, which is masculine. The fact that he gave his species two different names does not alter the fact of homonymy suggested above. In any case, variation within collections of C. rogersi was starting to be reflected in taxonomic usage. When Conrad (1866) developed his checklist, he documented three species from three states based on minimal differences among the three: C. rogersi from Alabama, C. jonesi from Georgia, and C. tu- midus from Mississippi, all from strata that were, at that time, considered to be Eocene in age. All are now recognized as C. rogersi. Stefanini (1911) described Clypeaster dou- villei from strata now recognized as the Marianna Limestone in Jasper County, Mississippi (the same locality information given by Conrad for his M. turgida discussed above). Clark and Twitchell (1915) and Cooke (1942, 1959) considered C. jonesi, C. douvillei, and C. turgida junior synonyms of C. rogersi and emphasized the great variability of C. rogersi. How- ever, C. turgida (which, given the account above, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 86 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 57: Clypeaster cf. C. rogersi (UF 337990), 55 mm TL, 52 mm TW, 13 mm TH, internal mold showing internal structure, dolomitic portion of Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: aboral. B: oral. C: left side. D: right side. should probably be emended to C. turgidus) and C. douvillei were described for specimens with a higher test, deeper infundibulum, and thinner margin than typical C. rogersi. Considering the proximity of the type localities, and relatively similar descriptions (both distinguished from C. rogersi by similar traits), they undoubtedly described the same form from the population of Clypeaster in the Glendon Limestone that overlies the Marianna Limestone in Mississippi and contains typical C. rogersi. If subsequent workers resurrect Conrad’s Mortonia turgida for these forms with a thinner margin, more elevated test and often more concave oral surface, Stefanini’s C. douvillei would likely be a subjective junior synonym of C. turgida, and not of C. rogersi. These subjective junior synonyms of C. rogersi are further discussed in the remarks for Clypeaster sp. B, below. Clark and Twitchell (1915) stated that C. rogersi is the most abundant American Clypeaster and documented the variability in marginal out- line, tumidity of the aboral surface, and concav- ity of the oral surface. Cooke (1959) noted that though the shape and size of C. rogersi is variable, the shape of the petals is relatively constant. How- ever, the variability expressed by Cooke included the thinner-margined and shorter-petaled C. mari- nanus from northern Florida and Mexico, as well as Clypeaster sp. B from the Suwannee Limestone of ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 87 Figure 58: Series of Clypeaster rogersi showing variation in test thickness and oral surface concavity, from the Lower Oligocene Marianna Limestone, quarry southeast of Brooklyn (FM-IP ZA181), Conecuh County, Alabama. A-E: UF 338015; 49.2 mm TL, 43.2 mm TW, 11.9 mm TH; A: aboral. B: oral. C: tilted aboral viewpoint from anterior. D: tilted oral viewpoint from posterior; concavity around peristome deeper than typical. E: left side. F-J: UF 338016; 55.6 mm TL, 50.9 mm TW, 9.4 mm TH; F: aboral. G: oral. H: tilted aboral viewpoint from anterior. I: tilted oral viewpoint from anterior. J: lateral. K-O: UF 338014; 58.8 mm TL, 52.6 mm TW, 14.5 mm TH; K: aboral. L: oral. M: tilted aboral viewpoint from posterior. N: tilted oral viewpoint from posterior. O: left side. P-T: UF 338017; 61.2 mm TL, 56.1 mm TW, 13.6 mm TH; P: aboral. Q: oral. R: tilted aboral viewpoint from posterior. S: oblique oral viewpoint from posterior. T: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 88 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 59: Series of Clypeaster rogersi showing variation in test thickness and oral surface concavity, from the Lower Oligocene Marianna Limestone, Smith County Lime Quarry, MS (FM-IP ZZ104). A-F: UF 338020; 39.1 mm TL, 32.4 mm TW, 7.4 mm TH; A: aboral. B: oral. C: tilted aboral viewpoint from anterior. D: anterior. E: oblique oral viewpoint from anterior. F: left side. G-L: UF 338019; 44.8 mm TL, 40.0 mm TW, 8.2 mm TH; G: aboral. H: oral. I: tilted aboral viewpoint from posterior. J: posterior. K: tilted oral viewpoint from posterior. L: left side. M-R: UF 338018; 50.7 mm TL, 46.5 mm TW, 9.1 mm TH; M: aboral. N: oral. O: tilted aboral viewpoint from anterior. P: posterior. Q: oblique oral viewpoint from posterior. R: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 89 Figure 60: Two specimens of Clypeaster rogersi from the Lower Oligocene Marianna Limestone, St. Stephens, Washington County, Alabama (FM-IP ZA192). A-G: UF 338022, 50.9 mm TL, 10.9 mm TW, 10.9 mm TH; A: aboral. B: oral. C: anterior. D: tilted aboral viewpoint from anterior. E: oblique lateral viewpoint from posterior. F: left side. G: periproct. H-N: UF 338021, 64.8 mm TL, 56.7 mm TW, 11.7 mm TH; H: aboral. I: distal end of petaloid portion of ambulacrum III. J: oral. K: anterior. L: tilted aboral viewpoint from posterior. M: oblique lateral viewpoint from posterior. N: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 90 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Florida. Cooke (1959) stated that the specimen of C. rogersi from the Suwannee Limestone (Clypeaster sp. B) and Meson Formation of Mexico (herein referred to C. marinanus) have shorter petals, a thinner margin, and often have a more inflated apical area than typical C. rogersi. The first two of these characteristics are true of C. marinanus and the second two are accurate for the population from the lower Suwannee Limestone in the Vulcan Quarry west of Brooksville (FM-IP HE038) here referred to Clypeaster sp. B. When these populations are removed from the synonymy, C. rogersi remains variable, but more restrictedly so (Table 7). The typical form is a thick-margined, relatively flat, cookie-like form with a nearly flat to very slightly concave oral surface, and as such, the species is a common element in the Marianna Limestone of northern Florida, Alabama, and Mississippi. As mentioned above, the Glendon Limestone of Mississippi contains a Clypeaster intermediate between typical C. rogersi and what we discuss as Clypeaster sp. B. Within the Suwannee Limestone, the only specimens that appear to belong to C. rogersi are from the Terramar Quarry in Polk County (FM-IP PO017) (Figs. 53, 54). Clypeaster rogersi has a thicker margin with longer petals than C. marinanus. It lacks the aboral submarginal depression of C. oxybaphon and has a lower test with a much less concave oral sur- face than C. cotteaui (Fig. 48) (Table 7). Twitchell in Clark and Twitchell (1915), indicated that C. cotteaui has straighter and more widely divergent poriferous zones than C. rogersi. See the remarks for Clypeaster sp. B below for features differentiating it from C. rogersi. Clypeaster sp. A (Figs. 48, 61, Table 7) not Clypeaster batheri (Lambert). Oyen, 2001. p. 96. figs. 3-12, F-G. Occurrence.—Suwannee Limestone north of Branford, in Suwannee County (FM-IP SU002). Discussion.—A single specimen (UF 2546) from the Lower Oligocene Suwannee Limestone of Suwannee County, Florida is distinct from all other species of Clypeaster described from North America. The specimen was initially discussed by Oyen (2001) as Clypeaster batheri Lambert, 1915. However, this fossil is clearly not conspecific with C. batheri, as further discussed below. Oyen (2001) also identified another specimen (UF 5341) as C. batheri, but this specimen consists of a test fragment that cannot be attributed to this species with confidence. Oyen (2001) briefly described the complete specimen (UF 2546) but his description is nearly verbatim from Jackson’s (1922) description for C. batheri. The specimen has a heavily eroded oral surface, but it is sufficiently preserved to be distin- guished from C. batheri. The holotype of C. batheri (NHM E12860) measures 52 mm TL, 44 mm TW, and 15 mm TH (Jackson, 1922). The specimen from the Suwannee Limestone (UF 2546) is very large for an Oligocene Clypeaster (113 mm TL, 85 mm TW, and 23 mm TH), and is much narrower (width is 75% length vs 84.6% in the type of C. batheri), and lower (height is 21% length vs 28.8% in the type of C. batheri) than C. batheri. Clypeaster sp. A is also much more peaked apically, as the maximum height rises from a much thinner margin than in C. batheri (Table 7). The thicker margin of C. batheri also distinguishes these two forms, as do the petals of C. batheri, which are not as widely open distally. Clypeaster batheri is more like C. cotteaui than Clypeaster sp. A. Clypeaster sp. A is also much larger, narrower, with a much more “peaked” apical system than C. rogersi, and it does not have a swollen margin with depressed submarginal periphery on the aboral sur- face like C. oxybaphon. Clypeaster sp. A also lacks the thin margin and short petals of C. marinanus. Furthermore, it is much narrower, with a more con- cave oral surface than Clypeaster sp. B, from the lowermost Suwannee Limestone, Hernando County, Florida (Table 7). The single specimen was collected in 1969 from an area that is still actively quarried (O’Brien Mine: FM-IP SU002). However, current operations have exposed only a thin horizon of Suwannee Limestone with an abundance of R. gouldii, and ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 91 Figure 61: Clypeaster sp. A., (UF 2546), 114 mm TL, 86 mm TW, 24 mm TH, Lower Oligocene Suwannee Limestone, Suwannee County, Florida (FM-IP SU002). A: aboral. B: oral. C: left side. D: right side. although the site has been extensively collected, no additional specimens of Clypeaster sp. A have been discovered. Lacking additional material to determine the variability of the taxon, we can only document its existence in the hope that future col- lecting will provide further specimens adequate to determine if it is indeed a new species, or conspe- cific with one of the myriad Clypeaster taxa already known from New World fossil and Recent localities. In addition, a much better understanding is critically needed of the 40 or so taxa of Cuban Clypeaster described in the works of Sánchez-Roig alone (Table 6), as it is possible that Clypeaster sp. A might be assigned to one of those. Clypeaster sp. B (Figs. 48, 62-66, Table 7) not Clypeaster rogersi (Morton). Cooke, 1959. pp. 36-37 (in part, discussion of C. rogersi in Suwannee Limestone is likely Clypeaster sp. B). Occurrence.—This form occurs in the lower beds of the Suwannee Limestone in the Vulcan Quarry (FM-IP HE038), Hernando County, Florida (Fig. 8). Discussion.—Specimens of Clypeaster are very common in the lowermost beds of the Suwan- nee Limestone in the quarries west of Brooksville, Hernando County, especially in the Vulcan Quarry (FM-IP HE038). A collector familiar with specimens ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 92 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) typical of C. rogersi from the Marianna Limestone (holotype ANSP 1070), especially from the type area of the species west of Claiborne, Alabama, will quickly notice some of the other specimens from there are very different. These examples, herein re- ferred to in open nomenclature as Clypeaster sp. B, typically have a thinner margin, are more tumid centrally on the aboral surface, and a more concave oral surface than typical C. rogersi (Table 7). The concavity of the oral surface is most pronounced in smaller specimens (as in UF 338001, 54 mm TL, 50 mm TW) but much less so in larger specimens (UF 337999, 82 mm TL, 73 mm TW). The widest point of Clypeaster sp. B is ante- rior of the apical area, the distal end of ambulacrum III being on average 66.4% of the distance from the center of the apical area to the anterior margin (the same average for C. marinanus is 55%). How- ever, the distance of the periproct from the posterior margin is highly variable, ranging from 5.5% to 9% TL. Cooke (1942) recognized the differences be- tween the Florida specimens and typical C. rogersi, though he still considered them synonymous. How- ever, he noted the Florida specimens have a thinner margin and are more tumid centrally than is custom- ary for C. rogersi. Nevertheless, Cooke (1942: 12 and 13) stated: “these features can be matched in selected individuals from Alabama and Mississippi”. This is true, but specimens of C. rogersi from the Marianna Limestone of the Gulf Coast only rarely display these traits, whereas Clypeaster sp. B is dominant and typical for the specimens in the basal Suwannee Limestone west of Brooksville. Cooke (1959) stated specimens of C. rogersi from the Suwannee Limestone (that is, Clypeaster sp. B) and Meson Formation of Mexico (referred here to C. marinanus) have shorter petals, a thinner margin, and often have a more inflated apical area than typical C. rogersi. The first two of these char- acteristics are true of C. marinanus and the second two are accurate for the population from the lower Suwannee Limestone in the Vulcan Quarry (FM-IP HE038) referred to Clypeaster sp. B. As noted in the remarks for C. marinanus and C. rogersi above, removal of these forms from the range of variation in C. rogersi, the species remains more variable than other regional species of Clypeaster, but readily recognizable as the typical flat, thick, cookie-like form of the Marianna Limestone of the Gulf Coast (reference holotype: ANSP 1070). While determining the identity of the thin specimens with short ambulacra from the Bumpnose Limestone was rather straight forward given their similarity to C. marinanus, assigning a species name to Clypeaster sp. B is less straightforward, as the forms in synonymy with C. rogersi would attest. Scutella jonesi, from Lower Oligocene strata in eastern Georgia (Forbes in Lyell, 1845), is clearly a typical C. rogersi, as noted by Clark and Twitchell (1915) and Cooke (1942, 1959), being recognized as a junior synonym of C. rogersi since Clark and Twitchell (1915). As noted above, Conrad (1866) intended to emend his M. turgida to C. tumidus. Although Con- rad’s statement that M. turgida is more elevated medially than C. rogersi implies an affinity with the Suwannee Limestone specimens, as was typical for Conrad, he didn’t provide catalog information for a type, potential or otherwise, and it is not listed in the catalog of type specimens in the Philadelphia Academy of Sciences, which houses most of Con- rad’s collection. The type is therefore unavailable for examination (Richards, 1968). Conrad’s species has been recognized as a subjective junior synonym of C. rogersi since Clark and Twitchell (1915), and has not been further described, discussed, or figured. While Conrad’s description is tantalizingly close to Clypeaster sp. B, M. turgida is otherwise unrecognizable and is only questionably included in our synonymy of C. rogersi (following Clark and Twitchell, 1915; Cooke, 1942, 1959). Stefanini (1911) stated that M. turgida is a nomen nudum and then described another Clypeaster from the same general area (Jasper County, Mississippi) as Conrad’s M. turgida, which does not clarify the issue. Stefanini (1911) subsequently described what would become recognized as yet another junior synonym of C. rogersi (Clark and Twitchell, 1915; Cooke, 1942, 1959), when he named C. douvillei from strata now recognized as the Marianna Lime- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 93 Figure 62: Clypeaster sp. B., (UF 337999), 82 mm TL, 74 mm TW, 16 mm TH, Lower Oligocene, basal beds of Suwannee Limestone, Vulcan Quarry (FM-IP HE038), west of Brooksville, Hernando County, Florida. A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: distal end of petaloid portion of ambulacrum III. E: posterior. F: left side. G: oblique lateral viewpoint from posterior. H: tilted oral viewpoint from posterior. I: pore-pairs at widest point of ambulacrum III. J: anterior. K: periproct. L: oblique oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 94 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 63: Clypeaster sp. B., (UF 338000), 59 mm TL, 53 mm TW, 13 mm TH, Lower Oligocene, basal beds of Suwannee Limestone, Vulcan Quarry (FM-IP HE038), west of Brooksville, Hernando County, Florida. A: aboral. B: oral. C: anterior. D: posterior. E: left side. F: oblique lateral viewpoint from posterior. G: oblique oral viewpoint from posterior. H: tilted oral viewpoint from posterior. I: tilted aboral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 95 Figure 64: Clypeaster sp. B., (UF 338001), 54 mm TL, 50 mm TW, 12 mm TH, Lower Oligocene, basal beds of Suwannee Limestone, Vulcan Quarry (FM-IP HE038), west of Brooksville, Hernando County, Florida. A: aboral. B: tilted aboral viewpoint from posterior. C: posterior. D: tilted aboral viewpoint from anterior. E: anterior. F: oral. G: tilted oral viewpoint from posterior. H: oblique oral viewpoint from posterior. I: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 96 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 65: Clypeaster sp. B., (UF 338008), 49 mm TL, 45 mm TW, 9 mm TH, Lower Oligocene, basal beds of Suwannee Limestone, Vulcan Quarry (FM-IP HE038), west of Brooksville, Hernando County, Florida. A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: distal end of petaloid portion of ambulacrum III. E: posterior. F: tilted aboral viewpoint from anterior. G: anterior. H: left side. I: periproct. J: tilted oral viewpoint from posterior. K: oblique lateral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 97 Figure 66: Clypeaster sp. B., (UF 338009), 64 mm TL, 60 mm TW, 13 mm TH, Lower Oligocene, basal beds of Suwannee Limestone, Vulcan Quarry (FM-IP HE038), west of Brooksville, Hernando County, Florida. A: aboral. B: posterior. C: tilted aboral viewpoint from posterior. D: anterior. E: tilted aboral viewpoint from anterior. F: left side. G: oral. H: tilted oral viewpoint from posterior. I: oblique oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 98 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) stone in Jasper County, Mississippi. Unfortunately, neither Conrad nor Stefanini provided more specific locality information than Jasper County, Mississippi for their respective specimens. The specimen Stefanini figured is incom- plete but shows a C. rogersi with a slightly thin- ner than typical margin, and more elevated apical area (the same traits Conrad used to distinguish his C. turgida/tumidus from C. rogersi). Subsequent recognition of Conrad’s M. turgida would likely require Stefanini’s C. douvillei to be a subjective junior synonym of C. turgida, and not of C. rogersi. According to Stefanini (1911), the holotype of C. douvillei is in the École Nationale Supérieure des Mines de Paris. However, we have not yet been able to examine this material, and this needs to be done before assigning a species name to Clypeaster sp. B. We only questionably include C. douvillei in the synonymy of C. rogersi above (following Clark and Twitchell, 1915; Cooke, 1942, 1959). Clypeaster sp. C (Figs. 48, 67, Table 7) Occurrence.—Lower Oligocene Suwannee Limestone, Vulcan Quarry (FM-IP HE019), north- west of Brooksville, Hernando County, Florida. Discussion.—A single (UF 338011) speci- men of Clypeaster with a distinctly thin, flat, test with broad petals that are wide open distally, was col- lected in the zone of R. gouldii roughly 10 m above the basal Suwannee Limestone (zone of Clypeaster sp. B) in the Vulcan Quarry (FM-IP HE019) northwest of Brooksville, Hernando County. Clypeaster sp. C is characterized by a low, nearly flat test that cannot be confidently attributed any species listed in Table 6. It occurs in the same quarry as Clypeaster sp. B, but from a soft limestone, rich in R. gouldii, roughly 10 m above the basal Suwannee Limestone bed containing Clypeaster sp. B. Clypeaster sp. C is readily distinguished from C. rogersi and Clypeaster sp. B by its exceptionally thin, flat, test (Table 7). The specimen measures 83.5 mm TL, 76.5 mm TW, 11.4 mm TH. The flat test is reminiscent of C. marinanus, but the latter has much shorter petals, with ambulacrum III on average equal to 26.5% TL vs 31.3% in Clypeaster sp. C., which also has much broader, distally open petals than C. marinanus (Table 7). Additional specimens are necessary to fully describe this species. Considering that the Suwannee Limestone is heavily collected in the Brooksville area, it is possible that specimens have already been collected and await recognition. We hope that by highlighting this specimen as distinct from any we have been able to study, collectors will examine their material and bring additional examples to light. Order ECHINOLAMPADACEA Mongiardino Koch et al., 2018 Suborder CASSIDULOIDA Agassiz and Desor, 1847 Family EURHODIIDAE Souto et al., 2019 Genus Eurhodia Haime in d’Archiac and Haime, 1853 Eurhodia patelliformis (Bouvé, 1851) (Fig. 68) Catopygus patelliformis Bouvé, 1851. p. 2. 2 txt figs. Cassidulus patelliformis (Bouvé). Desor, 1858. p. 290. ”Cassidulus” patelliformis (Bouvé). Cotteau, 1888. p. 521. Cassidulus patelliformis (Bouvé). Stefanini, 1911. p. 700. Cassidulus (Rhynchopygus?) patelliformis (Bouvé). Clark and Twitchell, 1915. p. 143. pl. 66, figs. 3a-d. Cassidulus (Rhynchopygus) depressus Twitchell in Clark and Twitchell, 1915. p. 144. pl. 66, figs. 4a-e, 5a-b. Procassidulus patelliformis (Bouvé). Lambert and Thiéry, 1921. p. 362. Procassidulus depressus (Twitchell). Lambert and Thiéry, 1921. p. 362. Eurhodia patelliformis (Bouvé). Cooke, 1942. p. 35. pl. 5, figs. 1-4. Cassidulus (Rhynchopygus) depressus Twitchell. Sánchez-Roig, 1949. p. 141. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 99 Figure 67: Clypeaster sp. C., (UF 338011), 83 mm TL, 76 mm TW, 11 mm TH, Lower Oligocene Suwannee Limestone, ˜10 m above quarry floor, Vulcan Quarry (FM-IP HE019) west of Brooksville, Hernando County, Florida. A: aboral. B: tilted aboral viewpoint from posterior. C: posterior. D: anterior. E: tilted posterior viewpoint from anterior. F: oral. G: oblique oral viewpoint from posterior. H: periproct. I: peristome. J: ambulacrum V. K: oblique lateral viewpoint from posterior. L: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 100 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 68: Eurhodia patelliformis (UF 308037), 27 mm TL, 18 mm TW, 11 mm TH, upper portion Upper Eocene Ocala Limestone, Dixie County, Florida (FM-IP DI013). A: aboral. B: oral. C: left side. D: right side. E: posterior. Eurhodia patelliformis (Bouvé). Cooke, 1959. p. 64. pl. 22, figs. 7-10. Eurhodia patelliformis (Bouvé). Toulmin, 1977. p. 341. pl. 65, fig. 4-6. Eurhodia patelliformis (Bouvé). Osborn et al., 2016. tbl 2. Occurrence.—This is an Upper Eocene species that occurs throughout Oligopygus halde- mani and Oligopygus wetherbyi Zones of the OLS, but has not been documented from the Oligopygus phelani Zone in the lower portion of the unit. Eu- rhodia patelliformis occurs at numerous localities ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 101 in Jackson County, notably the Brooks Quarries northwest of Marianna (FM-IP JA009, JA018, FM- IP JA027, FM-IP JA031, FM-IP JA039). It is also present west of Dowling Park, Lafayette County (FM- IP LF002); south of Tennille (FM-IP DI001), Dixie County, and the O’Brien Quarry north of Branford, Suwannee County (FM-IP SU002), among other localities. Eurhodia patelliformis also occurs in the Ya- zoo Formation of Alabama (Toulmin, 1977) and the upper OLS of Georgia. The type locality of E. patel- liformis and Cassidulus depressus Twitchell in Clark and Twitchell, 1915 (subjective junior synonym of E. patelliformis) is in Baker County, Georgia. Sánchez- Roig (1949) documented specimens he attributed to Cassidulus (Rhynchopygus) depressus Twitchell in Clark and Twitchell, 1915, in Cuban strata, which Cooke (1959) included in the synonymy of E. patel- liformis. Discussion.—Bouvé (1851) described Catopygus patelliformis from the Paleogene strata of Georgia and though his description is somewhat limited, he figured the species well. Twitchell in Clark and Twitchell (1915), better described the species and stated the holotype was likely collected in Baker County, Georgia. This small species of Eurhodia is commonly found in northern Florida where it is often associated with O. haldemani and W. johnsoni in Jackson County. We figure a specimen (UF 308037) from Brooks Quarry (FM-IP JA009) that measures 27 mm TL, 18.5 mm TW, and 11 mm TH (Fig. 68). Family CASSIDULIDAE L. Agassiz and Desor, 1847 Genus Rhyncholampas A. Agassiz, 1869 The State of Rhyncholampas Systematics in the Region Many species of Rhyncholampas are recog- nized in the Cenozoic strata of the eastern United States. Here, a discussion concerning Rhyncholam- pas is provided to clarify what we consider to be a member of this genus. Smith and Kroh (2011) noted that Rhyn- cholampas is usually distinguished from Cassidulus by its larger test, broader, lanceolate petals, generally more posterior periproct, and more developed phyl- lodes with more occluded plates. However, Smith and Kroh (2011) also stated that these differences are largely size related, and it was not clear that there is sufficient evidence to merit keeping the two genera separate. Souto et al. (2019: 651) brought some clarity by providing emended diagnoses for Cassidulus and Rhyncholampas. Their diagnosis for Rhyncholam- pas was: “Small to large cassidulids of varying test shape. Anterior paired petals tulip or leaf-shaped; posterior petals bowed. Paired petals with unequal number of pores in a and b columns (difference up to four pore pairs in anterior paired petals, up to ten pore pairs in posterior paired petals). Poriferous zones with three or more reduced primary tubercles. Periproct transverse, with prominent aboral hood. Posterior region of test sometimes truncated. Six to ten interambulacral plates between basicoronal plate 5 and base of periproct. Interambulacrum 5 naked zone usually wide, finely pitted, although pits vary from absent to large. Bourrelets bulged or pointed; bourrelet 5 often convex, projecting towards peris- tome. Outer column of anterior phyllodes with five to 12 phyllopores per half; posterior phyllodes with eight to 12 phyllopores per half; occluded plates often scattered. Phyllopores in large specimens often disorganized throughout phyllodes. Sphaeridia con- cealed by thin layer of stereom (fossils could have lost pit covering or pits in some species possibly open).” Mooi (1990a) pointed out that as with Cas- sidulus, any treatment of fossil species must take into consideration comparison with the Recent type species of the genus. Although Kier (1962) accom- plished a substantial revision of the cassiduloid echi- noids, including examination of types, considerable work remains to assign members of the Cassiduli- dae, which has been a dumping ground for species and genera of uncertain affinities. The phylogenic analysis of Souto et al. (2019) concluded by mention- ing species they considered of undetermined genus affinities, such as R. conradi and R. carolinensis (Twitchell in Clark and Twitchell, 1915), which we ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 102 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) retain in Rhyncholampas. Kier (1962: 174 and 180) moved Cassidu- lus sabistonensis (Kellum, 1931), C. georgiensis, C. alabamensis (Twitchell in Clark and Twitchell, 1915), C. ericsoni, and C. gouldii to Rhyncholam- pas. Cooke (1959) considered C. conradi, with its subspecies C. conradi carolinensis, to belong to the subgenus Plagiopygus. Kier (1962: 186) considered Plagiopygus a synonym of Rhyncholampas, which consequently moved R. conradi into Rhyncholampas. Kier (1962) left Cassidulus trojana in Cassidulus from the fossil fauna of the region. However, Carter and Beisel (1987) placed C. trojana in Eurhodia, in part because of the lack of naked areas on the oral surface, larger oral tubercles, and the presence of pits on the oral surface near the peristome. Never- theless, Souto et al. (2019) subsequently placed this species in Rhyncholampas, with which we agree. The result is that the Cenozoic deposits of eastern North America contain no species in Cas- sidulus. Under older usage of the name [i.e., pre-Kier (1962) and Carter and Beisel (1987)], Cassidulus was very well represented with seven species and three subspecies (Cooke, 1959). Therefore, there is a dramatic difference between the cassiduloid tax- onomy of the present monograph and that of Cooke (1959). The genus Rhyncholampas, as currently rec- ognized, contains 15 species in the eastern United States, ranging from the Middle Eocene to Upper Pleistocene: R. alabamensis (Twitchell in Clark and Twitchell, 1915), Oligocene; R. ayresi Kier, 1963, Pleistocene; R. carolinensis (Twitchell in Clark and Twitchell, 1915), Eocene; R. chipolanus Oyen and Portell, 1996, Miocene; R. conradi (Conrad, 1850), Eocene; R. ericsoni (Fischer, 1951), Eocene; R. ev- ergladensis (Mansfield, 1932), Pliocene; R. fontis (Cooke, 1942), Eocene; R. georgiensis (Twitchell in Clark and Twitchell, 1915), Eocene; R. gouldii, Oligocene; R. meansi Osborn et al., 2020, Pleis- tocene; R. sabistonensis (Kellum, 1931), Pleis- tocene; R. trojana (Cooke, 1942), Eocene; and Rhyncholampas bao n. sp., which occurs in the same strata as R. mariannaensis n. sp. Therefore, the Cenozoic fauna of the eastern United States currently includes 15 species of Rhyn- cholampas, and the Cenozoic strata of the remainder of the eastern Americas and Caribbean region poten- tially includes at least 11 additional species. Many of these taxa, especially the Cuban forms described by Sánchez-Roig, are poorly described and figured, most with inaccessible or lost holotypes that will likely remain poorly understood. Remarks on many of these species are included in the discussion below for Rhyncholampas mariannaensis n. sp. Rhyncholampas conradi (Conrad, 1850) (Figs. 69-71, 88) Catopygus conradi Couper MS, Conrad, 1850. p. 39, pl. 1, fig. 9. Nucleolites lyelli Conrad, 1850. p. 40, pl. 1, fig. 14. Cassidulus lyelli (Conrad). Conrad, 1865. p. 75. Pygorhynchus lyelli (Conrad). Cotteau, 1888. p. 550. Cassidulus conradi Couper in Conrad. Stefanini, 1911. p. 700. Cassidulus (Pygorhynchus) conradi (Conrad). Clark and Twitchell, 1915. p. 145. pl. 67, figs. 1a-f. Cassidulus (Rhynchopygus) lyelli (Conrad). Clark and Twitchell, 1915. p. 141. pl. 65, figs. 4a-d. Rhyncholampas lyelli (Conrad). Lambert and Thiéry, 1921. p. 370. Rhyncholampas conradi (Conrad). Lambert and Thiéry, 1921. p. 370. Cassidulus (Paralampas) conradi (Conrad). Cooke, 1942. p. 33. Cassidulus (Paralampas) lyelli (Conrad). Cooke, 1942. p. 33. Cassidulus conradi lyelli (Conrad). Cooke, 1959. p. 60. pl. 25, figs. 1, 2. Cassidulus (Plagiopygus) conradi (Conrad). Cooke, 1959. p. 59-60. pl. 25, figs. 3-6. Rhyncholampas conradi (Conrad). Kier, 1962. p. 186 (considered Plagiopygus a synonym of Rhyncholampas). Cassidulus conradi (Conrad). Toulmin, 1977. pp. 399-340. pl. 65, figs.1-3. Rhyncholampas conradi (Conrad). Osborn et al., 2016. tbl. 2. Occurrence.—Within Florida, R. conradi oc- curs throughout the Oligopygus haldemani and ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 103 Figure 69: Rhyncholampas conradi (UF 329690), 46 mm TL, 40 mm TW, 24 mm TH, Upper Eocene upper Ocala Limestone in Marianna Lime Quarry, Jackson County, Florida (FM-IP JA009). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 104 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 70: Rhyncholampas conradi (UF 338003), 41 mm TL, 35 mm TW, 19 mm TH, Oligopygus wetherbyi Zone Upper Eocene, upper Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 105 Figure 71: Rhyncholampas conradi (UF 338004), 32 mm TL, 27 mm TW, 20 mm TH, Oligopygus wetherbyi Zone Upper Eocene, upper Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 106 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Oligopygus wetherbyi Zones of the OLS and is especially abundant in Jackson County (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031, FM-IP JA039) where it occurs with O. haldemani. The species also occurs south of Tennille (FM- IP DI001), Dixie County, west of Dowling Park, Lafayette County (FM-IP LF002); northwest of Mayo, Lafayette County (FM-IP LF001), and north of Branford near O’Brien (FM-IP SU002), to name only a few localities. Rhyncholampas conradi also occurs in the Upper Eocene of Alabama (Toulmin, 1977) and Georgia (Carter, 1987a, 1989; Cooke, 1959). Discussion.—This species was named in honor of Timothy Abbott Conrad, the individual to whom the name is also attributed. However, Con- rad (1850) named the species from the white lime- stone of Palmyra, Lee County, Georgia, stating that the collector, J. Hamilton Couper, Esq., specifically requested the species name. Conrad (1850) was merely honoring the request of the collector, and as it remains only practice, and not an ICZN rule, not to name a new species after oneself, this was not unusual for the time. Conrad (1850) attributed the species name to an unpublished manuscript of Mr. Couper. Conrad (1850) also described Nucleolites lyelli from an internal mold collected in burrstone in Baker County, Georgia. Cooke (1942) indicated that specimens of Rhyncholampas lyelli Conrad, 1850 from Florida are higher in front than behind, thus differentiating them from those in Georgia, which are uniformly inflated. He stated that this difference is likely not of specific importance as other species in the genus show considerable variation in tumidity. This is accurate, as discussed in the remarks for R. gouldii and R. georgiensis below. Cooke (1959) considered R. lyelli to be a subspecies of R. conradi and stated that although R. lyelli is more ovate and less truncate posteriorly than R. conradi, it is otherwise indistinguishable from R. conradi. We do not recognize R. lyelli as a distinct species, nor do we consider it worth retaining it as a subspecies as asserted by Cooke (1959). Both R. conradi and R. lyelli share the same stratigraphic horizon in the OLS and specimens of both morphotypes (specimens with distinctly constricted posteriors and more ovate specimens) are often found at the same locality from the same horizon, with smaller specimens tending to be more nearly ovate. It is possible, therefore, to make a substan- tial collection of Rhyncholampas from this stratum, select more ovate, smaller forms and call them R. lyelli, as well as separate out specimens that are more truncated posteriorly and somewhat larger that would be called R. conradi, but such selectivity would ignore clear intermediates between the forms throughout the rest of the population. As implied by Cooke (1959) when he made R. lyelli a subspecies, R. conradi lyelli, there is no empirical support to maintain R. lyelli as distinct species or subspecies, so we consider it a subjective junior synonym of R. conradi. Fischer (1951) documented the occurrence of R. conradi lyelli in the Oligopygus phelani Zone of the lower portion of the OLS in Levy County. How- ever, Cooke (1959) referred those specimens to R. georgiensis. We are unaware of verified occurrences of R. conradi from the Oligopygus phelani Zone. Rhyncholampas ericsoni (Fischer, 1951) (Figs. 72, 73, 88) Cassidulus ericsoni Fischer, 1951. p. 65. txt figs. 6-10; pl. 2, figs. 1, 2; pl. 3, figs. 1-3. Cassidulus ericsoni (Fischer). Cooke, 1959. pp. 57- 58. pl. 24, figs. 13-16. Rhyncholampas ericsoni (Fischer). Kier, 1962. pp. 174, 180. Rhyncholampas ericsoni (Fischer). Toulmin, 1977. pp. 345-346. pl. 69, figs.7-9. Rhyncholampas ericsoni (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—This species has not been doc- umented outside the Oligopygus phelani Zone of the OLS of Florida (formerly the Inglis Formation). The type locality is along the Withlacoochee River in Levy County, but the species also occurs westward at the mouth of the river (FM-IP LV024) and is found ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 107 Figure 72: Rhyncholampas ericsoni (UF 247899) holotype, 48 mm TL, 43.5 mm TW, 32 mm TH, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, Citrus County, Florida (FM-IP CI052). A: aboral. B: oral. C: right side. D: posterior. along the banks of the Cross Florida Barge Canal south of Inglis (FM-IP CI001) and in numerous small quarries in Levy County (FM-IP LV114). Discussion.—Rhyncholampas ericsoni is sim- ilar in general appearance to R. gouldii but its periproct is much farther towards the anterior (Cooke, 1959). Rhyncholampas ericsoni occurs with R. georgiensis (form described by Fischer [1951] as R. globosus), but its lower, less rotund, and slightly more elongate test readily differentiates it from R. georgiensis. See the discussion for R. mariannaensis n. sp. for distinction of these two taxa. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 108 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 73: Rhyncholampas ericsoni (UF 322471), 44.7 mm TL, 42.1 mm TW, 27.5 mm TH, Oligopygus phelani Zone, Upper Eocene lower Ocala Limestone, Citrus County, Florida (FM-IP 6341). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 109 Rhyncholampas fontis (Cooke, 1942) (Figs. 74) ?Cassidulus (Galerolampas) fontis Cooke, 1942. p. 35. pl. 2, figs. 26-29. Echanthus georgiensis (Twitchell). Cooke, 1959. pp. 62-63. pl. 26, figs. 13-16 (in part, Cooke con- sidered R. fontis a subj. jr. syn. of E. georgien- sis). Rhyncholampas fontis (Cooke). Souto et al., 2019. pp. 643, 652. tbl. 4. Occurrence.—This species is not documented outside of its type locality, 487 m below the surface in a well drilled near Falling Waters, 4 mi. south of Chipley (Station 8872g, E. C. Butler collector), in Washington County (Cooke, 1942). Cooke (1942) surmised the specimen came from Lower Eocene or Paleocene strata. Discussion.—The type (USNM 498997) is the only known specimen of the species, and al- though it is exceptionally unlikely it would be found while surface collecting in Florida, we include it herein for completeness. Cooke (1959) subsequently placed Cassidulus fontis in synonymy with Gitolampas georgiensis, stating that the periproct of C. fontis was likely vertical prior to being crushed and distorted. However, as indicated by Zachos (2017), the type of C. fontis has better developed bourrelets and a transverse peristome, with no evidence of it being crushed along a vertical axis as stated by Cooke (1959). Cassidulus fontis is therefore not conspecific with G. georgiensis, and we recognize this form as a distinct species within Rhyncholampas. Souto et al. (2019) confirmed this genus assignment. The holotype of R. fontis (USNM 498997) was recovered from well washings originating from an unknown formation and was included in a larger lot of fossils that Zachos (2017) stated is now lost, so associated faunas are not available to assist in stratigraphic assignment. Additional material of this species may be collected in the future, enabling better understanding of this form. Although the holotype is slightly crushed, it is not severe enough to distort its overall test morphology. The specimen is much lower than any other Rhyncholampas currently recognized within the Paleogene strata of the region. With TW equal 92% of TL, the species is also more nearly circular in aboral view than any other Rhyncholampas known from the Paleogene of the region. Rhyncholampas georgiensis (Twitchell in Clark and Twitchell, 1915) (Figs. 75-79, 88) Cassidulus (Pygorhynchus) georgiensis Twitchell in Clark and Twitchell, 1915. p. 170, pl. 79, figs. 2a-d. Rhyncholampas georgiensis (Twitchell). Lambert and Thiéry, 1921. p. 370. Cassidulus (Paralampas) lyelli (Conrad). Cooke, 1942. p. 33 (in part, Cooke believed R. georgiensis a subjective junior synonym of C. lyelli). Cassidulus (Paralampas) globosus Fischer, 1951. p. 71. pl. 4, figs. 1-5. txt figs. 8-10. Cassidulus georgiensis globosus (Fischer). Cooke, 1959. p. 61. pl. 25, figs.11-13. Cassidulus (Plagiopygus) georgiensis (Twitchell). Cooke, 1959. pp. 60-61. pl. 25, figs. 14-17. Rhyncholampas georgiensis (Twitchell). Kier, 1962. p.180. Rhyncholampas georgiensis (Twitchell). Osborn et al., 2016. fig. 2. Occurrence.—Apart from the type locality near Bainbridge, Georgia, R. georgiensis occurs abundantly in two distinct localities in Florida. It occurs with O. haldemani in the OLS of north- ern Florida, especially near Marianna, Jackson County (FM-IP JA039). It also occurs in the Oligopygus phelani Zone near the base of the OLS (C. globosus of Fischer [1951]) in Levy County, notably 3.2 km southwest of Gulf Ham- mock (type locality of C. globosus), and along the Withlacoochee River west of Yankeetown (FM-IP LV024). Discussion.—When Twitchell in Clark and Twitchell (1915), described R. georgiensis, he noted ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 110 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 74: Holotype of Cassidulus (Galerolampas) fontis (= Rhyncholampas fontis) (USNM 498997), 38.7 mm TL, 35.8 mm TW, 16 mm TH, likely Early Eocene or Paleocene (per Cooke, 1942), well near Falling Waters, 4 mi. south of Chipley, Washington County, Florida. A: aboral. B: oral. C: left side. D: right side. E: posterior. its rarity (represented by three specimens at the time), and provided two occurences for the species: the Flint River near Bainbridge, Georgia and 4 km north of Cuthbert, Georgia. However, the exact position of the second locality is unknown. It is unfortunate that he formed its name from a state in which it is far less abundant and widespread than in the OLS of Florida. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 111 Figure 75: Rhyncholampas georgiensis (UF 308036), 32 mm TL, 30.5 mm TW, 28 mm TH, Oligopygus phelani Zone of Upper Eocene lower portion of Ocala Limestone, Levy County, Florida (FM-IP LV114). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 112 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 76: Rhyncholampas georgiensis (UF 329686), 28 mm TL, 26 mm TW, 23 mm TH, Haimea brooksi Zone of Upper Eocene, Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 113 Figure 77: Rhyncholampas georgiensis (UF 329687), 24.5 mm TL, 22 mm TW, 19.5 mm TH, Haimea brooksi Zone of Upper Eocene, Ocala Limestone, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 114 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 78: Holotype of Cassidulus globosus (= Rhyncholampas georgiensis), UF 248491 (with original catalog number of I-5346 still on specimen), Oligopygus phelani Zone, Upper Eocene lower Ocala Limestone, Levy County, Florida (FM-IP LV014). A: aboral. B: oral. C: left side. D: posterior. Rhyncholampas georgiensis largely occurs in two areas within the region: in the Oligopygus haldemani Zone of the OLS of northern Florida (especially Jackson County) and southwestern Geor- gia (the typical R. georgiensis), and the more southerly area in northern Citrus and southern Levy Counties, especially near Inglis, on the western Florida Peninsula (occurrences previously attributed ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 115 Figure 79: Comparison of various dimensions of 71 specimens of Rhyncholampas georgiensis from the Haimea brooksi Zone of the Ocala Limestone in the Brooks Quarry (FM-IP JA039), Marianna, Jackson County, Florida, and 41 specimens of Rhyncholampas globosus from the Oligopygus phelani Zone of the lower Ocala Limestone near Inglis (FM-IP LV114), Levy County, Florida. These similarities prompted us to herein designate R. globosus as a subjective junior synonym of R. georgiensis. to R. georgiensis globosus in the Oligopygus phelani Zone of the lowermost OLS). When Fischer (1951) described R. globosus, it appears he was unaware of R. georgiensis as he did not mention it in his consideration of other Rhyn- cholampas from the region (he compared his R. er- icsoni to R. trojana, R. alabamensis, R. gouldii, and R. lyelli). He stated R. globosus is most like R. lyelli, from which he differentiated it by its greater size, relative height, and inflation. Had Fischer (1951) compared his R. globosus with R. georgiensis, he would likely have realized their similarities. Subse- quently, Cooke (1959) designated R. globosus as a subspecies of R. georgiensis and noted that R. globo- sus is much more highly inflated than the type of R. georgiensis, but little more so than many specimens from Bainbridge, Georgia, the type locality of the species. We present comparisons of TH, TW, TL, and distance of periproct above the margin of these two forms (Fig. 79). This variation in TH among different, widely separated populations is not unique in Rhyncholam- pas. McKinney et al. (2014) documented variation in TH within populations of R. gouldii. We likewise recognize the variation in R. georgiensis and do not recognize the subspecific designation of globosus for the Levy County specimens. Rhyncholampas georgiensis is readily differ- entiated from any of its congeners in the Upper Eocene of the region by its proportionately higher test and very inflated appearance, sharply truncated anterior margin, and an oral surface that is not at all concave or depressed. See the remarks for R. mar- iannaensis n. sp., for discussion on distinguishing these two species. Rhyncholampas gouldii (Bouvé, 1846) (Figs. 80-82) Pygorhynchus gouldii Bouvé, 1846. p. 192. Pygorhynchus gouldii (Bouvé). Bouvé, 1847a. p. 437. Pygorhynchus gouldii (Bouvé). Bouvé, 1847b. p. 142. Nucleolites mortoni Conrad, 1850. p. 40. pl. 1, fig. 11. Pygorhynchus gouldii (Bouvé). Bouvé, 1851. p. 2. 2 txt figs. Pygorhynchus gouldii (Bouvé) Desor, 1858. p. 299. Ravenelia gouldii (Bouvé). McCrady, 1859. p. 282. Pygorhynchus gouldii (Bouvé). Cotteau, 1888. p. 550. Nucleolites mortoni (Conrad). Boyle, 1893. p. 200. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 116 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Pygorhynchus gouldii (Bouvé). Stefanini, 1911. p. 696. Cassidulus (Pygorhynchus) gouldii (Bouvé). Clark and Twitchell, 1915. p. 171, pl. 79, figs. 3a-d; pl. 80, figs. 1a-f, 2a-d. Procassidulus (Bardouinia) gouldii (Bouvé). Lam- bert and Thiéry, 1921. p. 363. Cassidulus (Cassidulus) gouldii (Bouvé). Cooke, 1942. p. 31. Cassidulus (Cassidulus) gouldii (Bouvé). Cooke, 1959. pp. 57-58. pl. 24, figs. 5-12. Rhyncholampas gouldii (Bouvé). Kier, 1962. pp. 174, 180. Cassidulus gouldii (Bouvé). Pickering, 1970. pp. 29, 36, 66. tbl. 1. Rhyncholampas gouldii (Bouvé). Carter and Beisel, 1987. pp. 1080-1083. fig. 1. 3. Rhyncholampas gouldii newbernensis Kier, 1997. pp. 6-8. fig. 3. pl. 5, figs. 1-7. Rhyncholampas gouldii (Bouvé). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 8. Rhyncholampas gouldii (Bouvé). McKinney et al., 2014. pp. 215-219. Occurrence.—This species is exceptionally abundant within the Suwannee Limestone at nu- merous Florida localities in Alachua, Columbia, Gadsden, Hamilton, Hernando, Hillsborough, Jeffer- son, Pasco, and Suwannee Counties. In the northern portion of the state, Rhyncholampas gouldii is com- monly preserved as molds in the dolomitic facies of the Suwannee Limestone and equivalent units (Huddlestun, 1993). This is the most ubiquitous echinoid of the Suwannee Limestone. Rhyncholampas gouldii also occurs in the Oligocene of Georgia (Huddlestun, 1993), North Carolina (Kier, 1997), and Mississippi (McKinney et al., 2014). Discussion.—Bouvé (1846) described Py- gorhynchus gouldii from Paleogene strata of Geor- gia but did not figure the species. This led to confusion, as Conrad (1850) subsequently de- scribed Nucleolites mortoni from an intenal mold from Palmyra, in Lee County, Georgia, which is clearly the same species. Bouvé (1851) quickly remedied this by redescribing and figuring his species while explaining that Conrad’s species was synonymous. Clark and Twitchell (1915) subsequently also figured Bouvé’s type specimen and noted that though this species was considered rare at the time it was documented, it was by then known to be one of the most seen of all American Cenozoic echinoids, especially so in Florida where they noted its abun- dance and highly variable form. In fact, Clark and Twitchell (1915) stated this species is so variable that extreme specimens differ in so many details that they would likely be described as distinct species if not for the intermediate forms found within large quantities of material, just as noted above for taxa such as R. conradi. Kier (1997) described a subspecies, R. gouldii newbernensis, from the Oligocene Trent Formation near New Bern, North Carolina. This stratum has since been referred to the Upper Oligocene River Bend Formation (Ward, 2007). Kier distinguished R. gouldii newbernensis from R. gouldii gouldii by the following: 1) longer petals; 2) peristome slightly higher and wider; 3) lower test; and 4) higher periproct. Nevertheless, Kier (1997) used the overlap in most of these characters between the two taxa to prevent their consideration as separate species. As discussed by McKinney et al. (2014) in their study of populations of R. gouldii from five locations in Florida, Mississippi, and North Carolina, this species demonstrates extraordinary variation in test shape among and within separated populations. Given this fact, we see no value in retaining the subspecies newbernensis, as there is no evidence whatsoever that it a separately evolving lineage worth recognizing. In the Suwannee Limestone of peninsular Florida, R. gouldii is often present in profusion and associated with G. mossomi and P. mansfieldi throughout much of the unit. Rhyncholampas trojana (Cooke, 1942) (Figs. 83, 84) Cassidulus (Cassidulus) trojanus Cooke, 1942. p. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 117 Figure 80: Rhyncholampas gouldii (UF 337955), 38 mm TL, 36.5 mm TW, 21 mm TH, Lower Oligocene Suwannee Limestone, Hernando County, Florida (FM-IP HE038). 32. pl. 2, figs. 22-25. Cassidulus trojanus (Cooke). Cooke, 1959. pp. 58- 59. pl. 24. figs. 1-4. Cassidulus trojanus (Cooke). Kier, 1962. p. 175. Eurhodia trojana (Cooke). Carter and Beisel, 1987. pp. 1080-1083. Eurhodia trojanus (Cooke). Osborn et al., 2016. tbl. 2. Rhyncholampas trojanus (Cooke). Souto, 2018. p. 59. Cassidulus trojanus (Cooke). Souto et al., 2019. pp. 625, 651. figs. 11-13, 16. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 118 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 81: Rhyncholampas gouldii (UF 337956), 30.5 mm TL, 29 mm TW, 14 mm TH (a very low specimen), Lower Oligocene Suwannee Limestone, Hernando County, Florida (FM-IP HE038). A: aboral. B: oral. C: left side. D: right side. E: posterior. Rhyncholampas trojana (Cooke). Souto et al., 2019. p. 651. Occurrence.—Rhyncholampas trojana is most commonly found in the uppermost beds of the OLS where it is associated with W. eldridgei in the Rotularia vernoni Zone, especially west of Dowl- ing Park, Lafayette County (FM-IP LF002); north- west of Mayo, Lafayette County (FM-IP LF001), Cemex Quarry near Center Hill, Sumter County ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 119 Figure 82: Rhyncholampas gouldii (UF 337957), 35 mm TL, 33 mm TW, 20 mm TH, Lower Oligocene Suwannee Limestone, Hernando County, Florida (FM-IP HE038). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 120 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 83: Rhyncholampas trojana (UF 329704), 28.5 mm TL, 24 mm TW, 13.5 mm TH, uppermost portion of Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: right side. E: posterior. (FM-IP SM010), and north of Branford, Suwannee County (FM-IP SU002). The type locality is along the Suwannee River below Troy Springs, Lafayette County. Rhyncholampas trojana also occurs in the OLS of Georgia (Cooke, 1959). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 121 Figure 84: Rhyncholampas trojana (UF 329705), 27.5 mm TL, 23 mm TW, 13 mm TH, uppermost portion of Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: right side. E: posterior. Discussion.—Rhyncholampas trojana cannot be confused with the much smaller and narrower E. patelliformis, with which it has been considered a congener, and often occurs within the OLS of northern Florida. Eurhodia patelliformis has a more sharply truncated posterior margin. Rhyncholampas trojana is more easily confused with R. gouldii, from the overlying Lower Oligocene Suwannee Lime- stone. The two species often co-occur in mixed spoil in northern Florida quarries, or in outcrops ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 122 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 85: Rhyncholampas bao n. sp., holotype (UF 342094), 60.3 mm TL, 52.1 mm TW, 24.9 mm TH, Upper Eocene, Ocala Limestone, Haimea brooksi Zone, Brooks Quarry, northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: apical area. C: oral. D: posterior. E: oblique lateral viewpoint from posterior. F: anterior. G: tilted aboral viewpoint from posterior. H: oblique oral viewpoint from posterior. I: left side. J: distal end of petaloid portion of ambulacrum I. K: periproct. L: pore-pairs at widest point of ambulacrum I. M: peristome and phyllodes. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 123 in very close proximity to the Eocene/Oligocene boundary, where the Suwannee Limestone often immediately overlies the Wythella eldridgei Zone in the uppermost OLS. Carter and Beisel (1987) discussed the impor- tance of the large, deep pits in the interambulacrum of the nearly flat oral surface of R. trojana. These pits are absent in the more gently concave oral sur- face of R. gouldii. Species of Rhyncholampas have a prominent naked area on the oral surface that extends from near the peristome to very near the posterior margin that is diagnostic in comparing R. gouldii to R. trojana. In contrast, R. trojana has an oral surface that is rather evenly adorned with comparatively large tubercles and no such naked area. As discussed by Carter and Beisel (1987), misidentification of these two species can lead to confusion on the part of paleontologists and geolo- gists attempting to make age determinations of the strata with which they are concerned. The differ- ences can be subtle, so this confusion can lead to errors because R. gouldii is considered an indicator of the Oligocene strata of the region. The traits discussed above also led Carter and Beisel (1987) to assign this species to Eurhodia. Mooi (1990b: 696) stated that re-examination of Carter and Beisel’s characters, with comparison to the type species of Cassidulus and Rhyncholampas, is needed before R. trojana can be correctly placed, an assertion applicable to many cassiduloids, which is overall, a poorly studied group with uncertain phylogenetic relationships. Phylogenetic analyses of Recent species, combined with fossil taxa, especially those that are the type species for their genera (Souto et al., 2019), is essential to establish a basis for ongoing treatment of the fossil species. Previous attempts, such as that of Suter (1994a, b) failed largely due to a lack of understanding of homologies within the group, as well as evident non-monophyly that makes identification of appropriate outgroups extremely problematic. Souto et al. (2019) stated that Carter and Beisel’s (1987) assertion that R. trojana belongs in Eurhodia was not correct. The traits they used (i.e., deep pits in the naked zone and concave oral surface) are not present in the type species of Eu- rhodia. Souto et al. (2019) then placed trojana into Rhyncholampas, and we follow this assignment. Rhyncholampas trojana has traits that suggest placement in Eurhodia, Rhyncholampas, and Cassidulus and debate will continue among some echinologists who weigh one characteristic more heavily than another. Nevertheless, the species itself is relatively easy to identify, and appears to be restricted to the Wythella eldridgei Zone in the uppermost OLS. Rhyncholampas bao n. sp. (Figs. 85-88) Diagnosis.—Large Rhyncholampas (TL av- erage 56.1 mm TL, largest 60.8 mm TL) with low test, TH on average 45.3% TL and 54% TW, posterior margin sharply truncated with oral edge of periproct on average 18.7% above the posterior margin; posterior end of prominent aboral hood over periproct forms furthest point of posterior margin of test, with long, straight, narrow, only slightly widened halfway along, but petals extending nearly to ambitus. Description.—Based on holotype (UF 342094), four paratypes (UF 342095, UF 342096, UF 342097, UF 342098), and non-type specimens. Test large, holotype largest specimen: 60.3 mm TL, 52.1 mm TW, 24.9 mm TH; smallest specimen (UF 342098): 29.1 mm TL, 25.3 mm TW, 13.3 mm TH; ten additional specimens range from 49 to 60 mm TL (avg. 56.1 mm TL). Test wide, width 86.4% TL in holotype, average 84%; low: TH 41.2% TL and 47.7% TW in holotype, average TH is 45.3% TL and 54% TW; highest point posterior of apical system, greatest width posterior of apical system, point of greatest length on posterior is above ambitus at prominent aboral hood over periproct which overhangs posterior margin of test. Posterior ambitus sharply truncated, shallow sinus extends from periproct to posterior ambitus. Apical system monobasal with four gonopores, anterior (center of apical system on average 69.3% TL from posterior margin). Petals narrow, long, nearly straight (only ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 124 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 86: Rhyncholampas bao n. sp., paratype (UF 342095), 58.9 mm TL, 50.1 mm TW, 23.6 mm TH, Upper Eocene, Ocala Limestone, Haimea brooksi Zone, Brooks Quarry, northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: posterior. D: oblique lateral viewpoint from posterior. E: anterior. F: tilted aboral viewpoint from posterior. G: oblique lateral viewpoint from posterior. H: peristome. I: left side. J: oblique oral viewpoint from posterior. K: apical area. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 125 Figure 87: Rhyncholampas bao n. sp., paratype (UF 342096), 58.6 mm TL, 46.5 mm TW, 26.1 mm TH, Upper Eocene, Ocala Limestone, Haimea brooksi Zone, Brooks Quarry, northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: oral. C: posterior. D: tilted aboral viewpoint from posterior. E: apical area. F: oblique oral viewpoint from posterior. G: anterior. H: ambulacrum I. I: left side. J: oblique lateral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 126 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 88: Comparison of test height and test width (top) and test length (bottom) of Eocene Rhyncholampas species from the eastern United States. slightly widened at their midpoint; most pronounced in smaller specimens), pore pairs almost parallel, open distally; average petal length as a percentage of TL is: I: 44.0%, II: 33.5%, III: 34.2%, IV: 35.2%, V: 43.5% (petal length taken from distal end of longest pore-pair column, if columns are of unequal length); pore pair columns of petal I, II, IV and V usually of unequal length, petal I and V more so; anterior pore pair columns of petal I and IV longer; posterior pore pair columns of petal II and IV longer; pore pair columns of petal III usually equal: anterior pore pair columns of petal I and V have, on average, six more pore pairs than the posterior series; pore pair columns of petal II and IV have, on average, four more pore pairs than the anterior series; both pore pair columns of petal III usually equal; ambulacra with unipores between end of petals and peristome. Periproct transversely elongate, width on av- erage 49.7% its height, with prominent aboral hood that usually extends over posterior margin of test. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 127 Adoral edge of periproct on average 18.7% above the posterior margin, shallow sinus below periproct leads to truncated posterior margin. Oral surface nearly flat, only in smallest spec- imen (29 mm TL) is there slight, but noticeable, concavity in peristomial area; larger tubercles than aboral surface; narrow naked area in interambu- lacrum V from peristome to posterior margin with minimal pitting, greatest width of naked area on average 8.9% TW, 8.4% TW in holotype, naked area tapers posteriorly. Peristome pentagonal, wider than high, peri- stome height on average 63.5% width; anterior, posterior edge of peristome on average 55.8% from posterior margin. Five pronounced, vertical bour- relets, each convex towards peristome. Phyllodes with unipores, on average 39% width of peristome at widest point, occluded plates usually present; two buccal pores present. Zoobank Nomenclatural Act.—F527F287- DB04-4585-859A-A91D25229B67 Discussion.—Rhyncholampas bao n. sp. has not been documented outside of its type locality in the northwesternmost pit in the Brooks Quarry (FM-IP JA039) northwest of Marianna, Jackson County, Florida, where it occurs in the OLS in material dredged from 24 m below the top of the Eocene stratum in the quarry (Fig. 6). It is found in association with H. brooksi, O. haldemani O. rotundus, R. georgiensis, W. johnsoni, and other, rarer species including Brissus jonesi n. sp. and Rhyncholampas mariannaensis n. sp. While attempting to place these specimens into R. conradi, we quickly realized this was unten- able, and they could not be placed with confidence in any other currently described species from the Amer- icas. It is possible that if these specimens occurred with otherwise typical specimens of R. conradi, we could consider them outliers of R. conradi’s typical form. This taxon occurs stratigraphically below the occurrence of R. conradi within the Brooks quarry and is typically much larger than the R. conradi which occur above it (other than one small specimen that is 29 mm TL, ten additional specimens range from 49 to 60 mm TL vs 81 R. conradi that ranged from 15.4 to 55.1 mm TL). It is further distinct from R. conradi by its much lower test (TH on average 41.2% TL and 47.7% TW in Rhyncholampas bao n. sp., compared to 54.7% TL and 63% TW in 81 specimens of R. conradi we measured) (Fig. 88). TW as a percentage of TL is similar in these two forms, being on average 86.2% in R. conradi and 84% in Rhyncholampas bao n. sp. (Fig. 88). In addi- tion, the periproct of R. conradi is on average 28% above the posterior margin of the test, whereas in Rhyncholampas bao n. sp. it is significantly lower at 18.7%. Other than the remarkably low test of this species and long narrow petals that are typically only very slightly bowed (widened medially), the most striking characteristic is the very posterior high point of the test that in many larger specimens gives the test a striking “humped” posterior portion of the test, visible from a lateral viewpoint. The oral surface is nearly flat, and only in the smallest specimen (29 mm TL) is there a slight, but noticeable, concavity in the peristomial area. The low test of this species is like that of R. fontis, which has an even lower test (though the holotype and only known specimen is slightly crushed) with a TH 41% TL and 44% TW. However, R. fontis is a nearly circular species when viewed aborally, with a TW 92% TL. We complete a review of species of Rhyn- cholampas, and potential Rhyncholampas, from the remainder of the eastern Americas in the discussion for R. mariannaensis n. sp., below and of these forms, Rhyncholampas bao n. sp. is most like R. ellipticus (Clark in Arnold and Clark, 1927) which Clark designated as the type species of his new genus Anisopetalus (subsequently recognized as a subjective junior synonym of Rhyncholampas). Though the holotype of this species is 50 mm in length, Clark noted the remaining six specimens he had available were much smaller, from 26 to 41 mm in length. Rhyncholamaps ellipticus has a low test (holotype TH 40% TL and 51% TW), but it is narrower than Rhyncholampas bao n. sp. (TW 78% TL vs 84% in R. bao n. sp.), and the highest point of the test is much more anterior. The figures of the species provided by Arnold and Clark (1927) also appear to show a much more concave oral surface ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 128 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 89: Rhyncholampas mariannaensis n. sp., holotype (UF 338027), 49.8 mm TL, 38.8 mm TW, 32.6 mm TH, Upper Eocene Ocala Limestone, Haimea brooksi Zone, Leon Brooks Quarry (FM-IP JA039), Marianna, Jackson County, Florida. A: aboral. B: apical area. C: oral. D: tilted aboral viewpoint from posterior. E: oblique posterior viewpoint. F: posterior. G: left side. H: tilted oral viewpoint from posterior. I: oblique oral viewpoint from posterior. J: anterior. K: peristome. L: petaloid portion of ambulacrum III. M: petaloid portion of ambulacrum V. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 129 Figure 90: Rhyncholampas mariannaensis n. sp., paratype (UF 342090), 56.7 mm TL, 46.5 mm TW, 33.4 mm TH, Upper Eocene Ocala Limestone, Haimea brooksi Zone, Leon Brooks Quarry (FM-IP JA039), Marianna, Jackson County, Florida. A: aboral. B: oral. C: posterior. D: anterior. E: left side. F: tilted oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 130 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) in R. ellipticus than R. bao n. sp. Rhyncholampas bao n. sp. is not comparable to any other species of Rhyncholampas currently documented from the eastern Americas or Caribbean region, as reviewed in the remarks for R. mariannaensis n. sp. Etymology.—The new species is named for their resemblance to bao, a type of Chinese steamed meat or vegetable filled bun. Material and Occurrence.—Holotype (UF 342094), four paratypes (UF 342095, UF 342096, UF 342097, UF 342098), from the type locality in the OLS, where it occurs with H. brooksi and O. haldemani in material dredged from 24 m below the top of the Eocene stratum in the most northwestern pit in the quarry complex of Leon Brooks, north- west of Marianna, Jackson County, Florida (FM-IP JA039). Rhyncholampas mariannaensis n. sp. (Figs. 88-91) Diagnosis.—Rhyncholampas with TW on av- erage 80.1% TL; highest point posterior of apical system; TH on average 65.4% TL and 84% TW; adoral edge of periproct on average 33.9% TH above posterior margin; naked area along oral interambu- lacrum 5 on average 15.5% TW. Description.—Description based on holotype (UF 338027), three paratypes (UF 342090, UF 342091, UF 342092), and several non-type speci- mens. Test large (holotype 49.8 mm TL, 38.8 mm TW, 32.6 mm TH, largest specimen [UF 342091] 64.6 mm TL, 52.0 mm TW, 39.6 mm TH); elon- gate, narrow (holotype TW 77.9 % TL; average 80.1%), high (holotype TH 65.4% TL, average 61.5% TL); highest point posterior of apical sys- tem; greatest width along transverse line poste- rior of apical system; posterior ambitus sharply truncated, anterior vertical to slightly overhung. Api- cal system monobasal, four gonopores, anterior (cen- ter of apical system 63.5% TL from posterior margin of test in holotype; 63.8% average). Petals lanceolate, moderately open distally, posterior petals widened about halfway their length; petals I and V longest, III shortest (petal length taken from distal end of longest pore-pair column, if columns are of unequal length), average petal length as a percentage of TL is: ambulacrum I: 43.5%; II: 36.8%; III: 36.4%; IV: 37.2%; V: 43.2%. Columns of pore pairs in petals of ambulacra I, II, IV and V usually of unequal lengths, most pronounced in petals I and V; anterior pore pair column of ambulacra I and IV longer; posterior columns of pore pairs in ambulacra II and IV longer; pore pair columns of petal in ambulacrum III usually equal; pore counts of petals of holotype follows: I: 44 and 47, II: not counted, III: 44 and 44, IV: 34 and 42, V: 49 and 41; ambulacra single-pored beyond petals. Periproct transversely elongate, width on av- erage 50.8% of its height (upper portion of periproct of holotype damaged), with prominent aboral hood. Adoral edge of periproct on average 33.9% TH above the posterior margin, 34.6% TH in holotype; shal- low sinus below periproct proceeding to truncate posterior margin. Oral surface flat, not concave, with larger tu- bercles than on aboral surface; broad naked area in interambulacrum 5 from peristome to posterior margin with minimal pitting, greatest width of naked area on average 15.5% TW, 15.7% TW in holotype. Peristome pentagonal, wider than long, 4.6 mm wide, 3.3 mm long on holotype, peristome height on average 71.7% width; anterior, posterior edge of peristome 60.4% TL in holotype, average 58.5%. Five pronounced, vertically oriented bourrelets, con- vex towards peristome. Phyllodes with unipores, on average 47% width of peristome at widest point, occluded plates usually present; two buccal pores present. Zoobank Nomenclatural Act.—F3FBFDF8- 5D2F-48AB-98A3-9EF6FDF36638 Discussion.—This species has not been doc- umented outside of its type locality in the Brooks Quarry in Jackson County, Florida (FM-IP JA039) (Fig. 6). Here, it occurs in material dredged from 24 m below the top of the Eocene stratum in the northwesternmost pit in the quarry complex, in an horizon of Asterocyclina-rich limestone with a diverse echinoid assemblage, initially discussed in Osborn et al. (2016). This assemblage includes H. brooksi, O. haldemani, O. rotundus, W. john- soni, R. georgiensis, Brissus jonesi n. sp., Rhyn- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 131 Figure 91: Rhyncholampas mariannaensis n. sp., paratype (UF 342091), 64.6 mm TL, 52.0 mm TW, 39.6 mm TH, Upper Eocene Ocala Limestone, Haimea brooksi Zone, Leon Brooks Quarry (FM-IP JA039), Marianna, Jackson County, Florida. A: aboral. B: apical. C: oral. D: posterior. E: anterior. F: distal end of petal V. G: left side. H: oblique lateral viewpoint from posterior. I: oblique oral viewpoint from posterior. J: naked medial area of oral surface just posterior of peristome. K: tilted aboral viewpoint from posterior. L: peristome. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 132 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) cholampas bao n. sp., and many other, rarer species (Table 1). Within this assemblage, R. mariannaensis n. sp. could only be confused with R. georgiensis, from which it is readily distinguished by its much greater size (known specimens of R. mariannaensis n. sp. range from 48-64 mm TL, and the largest specimen of R. georgiensis known to us is 33 mm TL) and narrower test (average TW of R. georgien- sis is 88.5% TL; average TW R. mariannaensis n. sp. is 80.1% TL; holotype is 77.9% TL). However, as shown in Fig. 88, TW vs TL is not a very use- ful feature for distinguishing regional species of Eocene Rhyncholampas. Perhaps most significantly, the periproct of R. mariannaensis n. sp. is on aver- age 33% TH above the posterior margin, whereas in R. georgiensis it is on average 40% TH above the posterior margin (Fig. 79). Differences in TH vs TW are also apparent in these two species: TH is on average 83.5% TW in R. georgiensis and 73.7% in R. mariannaensis n. sp. (Fig. 88). In addition, TH is on average 74% TL in R. georgiensis, and 61.5% in R. mariannaensis n. sp. (Fig. 88). The large, high test of R. mariannaensis n. sp. is distinguished from the relatively lower test of R. conradi, which occurs in strata immediately above the horizon of R. mariannaensis n. sp. (TH on average 54.7% TL and 63% TW in R. conradi and 61.5% TL and 73.7% TW in R. mariannaensis n. sp.). Furthermore, the periproct is on average only 28% TH above the posterior margin in R. conradi, whereas it is 33.9% TH in R. mariannaensis n. sp. Other than R. conradi and R. georgiensis, of the regional species R. mariannaensis n. sp. could only be confused with R. ericsoni, which occurs in the lowermost portion of the OLS near Inglis in Citrus and Levy Counties. Rhyncholampas ericsoni has a smaller and proportionately lower and wider test: TH is 66.6% TL in the holotype of R. ericsoni and on average 61.5% TL in R. mariannaensis n. sp. (Fig. 88). In addition, TH is 73.5% TW in the holotype of R. ericsoni and on average 83.9% TW in R. mariannaensis n. sp. (Fig. 88). However, the test of R. ericsoni is quite variable: the range of 12 measured specimens revealed a test width from 52.5- 66.8% TL, with an average of 59.8%; TH ranged from 52.5-66.5% TL (average of 59.8%) and 60.7- 73.9% TW (average of 68.6%). The average TH of R. mariannaensis n. sp. is therefore significantly higher relative to width than in R. ericsoni. In addition, the periproct of R. mariannaensis n. sp. is lower (on average 33.9 % TH above the margin) than R. ericsoni (on average 40.1% TH above margin). Rhyncholampas contains comparatively few species in the remainder of the eastern Ameri- cas and Caribbean region, and we will endeavor to compare R. mariannaensis n. sp. to all of them. These species include R. candidoi Garrafielo and Carreira, 1994, from the Miocene of Brazil; R. cervantesi Sánchez-Roig, 1949, from the Oligocene of Cuba, and R. rodriguezi Lambert and Sánchez-Roig in Sánchez-Roig, 1926, from the Oligocene to Miocene of Cuba (Souto et al., 2019). The large, high, narrow test of R. marian- naensis n. sp. readily distinguishes it from these small Cuban taxa. Five species of Anisopetalus have been de- scribed from the eastern Americas and this genus has since been considered a junior synonym of Rhyn- cholampas (Kier, 1966b: U515), requiring compari- son with R. mariannaensis n. sp.: Anisopetalus bro- dermanni Sánchez-Roig, 1952c; A. cookei Sánchez- Roig, 1952c (Souto et al. [2019] stated this species is likely not a Rhyncholampas but did not reassign it), and A. caobaense Sánchez-Roig, 1952c, all three of which are from the Eocene of Cuba. These taxa were figured adequately enough by Sánchez-Roig (1952c) to readily distinguish them from R. mariannaensis n. sp. Anisopetalus ellipticus Clark in Arnold and Clark, 1927 (not to be confused with Cassidulus ellipticus Kew, 1920, from the Eocene of California and Mexico which Souto et al., 2019, stated should be reassigned to Eurhodia), was described from the Eocene of Jamaica, but the low test of this species readily differentiates it from R. mariannaensis n. sp. Anisopetalus oliveirae Marchesini Santos, 1958, was described from the Miocene of Brazil but it can not be confused with R. mariannaensis n. sp. Echinanthus parallelus Azpeitia in Cotteau, 1897, from the Eocene of Cuba, and E. antillarum Cotteau, 1875, St. Bartholomew, and Cuba, were ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 133 placed in Parapygus by Jackson (1922) and subse- quently reported from the Eocene of Jamaica by Arnold and Clark (1927). Donovan (1993) question- ably attributed both species to Rhyncholampas, but stated they require further study. Judging by the fig- ures of these two species, Donovan was correct, but given their dissimilarity to Rhyncholampas, we do not place them in that genus, and they are likewise not comparable to R. mariannaensis n. sp. Six species from the Oligocene-Miocene of Cuba were named to Procassidulus in the works of Sánchez-Roig. Kier (1962, 1966b) considered Procassidulus a junior synonym of Rhynchopygus, which Kier (1962) stated is largely distinguished from Cassidulus by its tetrabasal api- cal system, whereas Cassidulus and Rhyncholam- pas have monobasal apical systems. As shown by Smith and Kroh (2011), Rhynchopygus has strongly developed bourrelets, whereas Cassidulus has bour- relets largely confined to the vertical sides of the peristome, and Rhyncholampas has very weakly developed bourrelets. Smith and Kroh (2011) state Procassidulus to be very similar to Rhynchopygus in petal form, phyllode structure, and in having an invaginated periproct. It differs in lacking the strongly pronounced tongue-like projection above the periproct opening. Sánchez-Roig did not charac- terize the apical system, nor did he figure Procas- sidulus avilensis Palmer in Sánchez-Roig, 1949, or P. habanensis Sánchez-Roig, 1949, both from the Oligocene of Cuba. However, P. brodermanni Sánchez-Roig, 1949, from the Oligocene, P. circularis Palmer in Sánchez-Roig, 1949, also from the Oligocene, and P. jeanneti Sánchez-Roig, 1949, from the Miocene, are figured well enough to determine they likely do not belong in Procassidulus or Rhynchopygus and are potentially Rhyncholampas. Though we did not examine any of these specimens, and we are not prepared to assert the genus placement for any of them, we compared them with R. mariannaensis n. sp., and there is no way to confuse R. mariannaensis n. sp. with any of these small forms. Judging by Sánchez-Roig’s understanding of Procassidulus, it is likely P. avilensis, P. habanensis, and P. echevarriai Sánchez-Roig, 1953b, from the Oligocene of Cuba could also be Rhyncholampas, along with Paralampas conceptionis Sánchez-Roig, 1953b, from the Eocene of Cuba. None of these taxa could be confused with R. mariannaensis n. sp. Although Cassidulus is not currently rec- ognized in the Cenozoic faunas of the eastern United States, many species have been placed in the genus in the remainder of the eastern Americas and Caribbean region. These will require additional study to determine if they should be reassigned to Rhyncholampas, and include: C. mestieri Kier, 1966a and C. senni Kier, 1966a, two small species that appear to be good Cassidulus, from the Eocene of Barbados. These bear no resemblance to R. mari- annaensis n. sp. Rhyncholampas falconensis (Jeannet, 1928), from the Miocene of Venezuela, was originally placed in Eurhodia, but considered a Cassidulus by Cooke (1961) and subsequently Rhyncholampas by Souto et al. (2019). We agree that this species appears to be a Rhyncholampas, though it bears no resemblance to R. mariannaensis n. sp. Cassidulus (Rhynchopygus) zanolettii Sánchez-Roig, 1952d, was described from the Eocene of Cuba and may belong to Eurhodia, as does Nucleopygus tamarindensis Sánchez-Roig, 1952d, also from the Cuban Eocene. Cassidulus rojasi Sánchez-Roig, 1953b, from the Oligocene of Cuba, as well as C. sphaeroides Arnold and Clark, 1934 and C. platypetalus Arnold and Clark, 1934, both from the Eocene of Jamaica, could belong in Rhyncholampas. Sánchez-Roig (1949) described Cassidulus (Pygorhynchus) riveroi from the Oligocene (potentially Miocene according to Souto et al. [2019]) of Cuba, and which Souto et al. (2019) asserted is a Rhyncholampas; and Catopygus rodriguezi Lambert and Sánchez Roig in Sánchez Roig (1926) from the Miocene (and potentially Oligocene according to Souto et al. [2019]) of Cuba, which Souto et al. (2019) also asserted is a Rhyncholampas. Perhaps a few of these taxa can be assigned to Rhyncholampas, but none of these comparatively small taxa bears resemblance to R. mariannaensis n. sp. Etymology.—Named in honor of the city of Marianna, Jackson County, Florida. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 134 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 92: Echinolampas cf. E. aldrichi (UF 112504), 67 mm TL, 58 mm TW, 33.5 mm TH, internal mold from the dolomitic portion of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA014). A: aboral. B: oral. C: left side. D: oblique posterior viewpoint tilted upward from oral surface showing periproct placement. Material and Occurrence.—Holotype (UF 338027) and paratypes (UF 342090 and UF 342091), from the type locality in the OLS, where it oc- curs with H. brooksi and O. haldemani in material dredged from 24 m beneath the top of the Eocene stratum in the most northwestern pit in the quarry complex of Leon Brooks, northwest of Marianna, Jackson County, Florida (FM-IP JA039). Suborder ECHINOLAMPADOIDA Kroh and Smith, 2010 Family ECHINOLAMPADIDAE Gray, 1851 Genus Echinolampas Gray, 1825 Echinolampas aldrichi Twitchell in Clark and Twitchell, 1915 (Fig. 92) Echinolampas aldrichi Twitchell in Clark and Twitchell, 1915. p. 173. pl. 81, figs. 1. a-d, 2. Echinolampas (Miolampas) aldrichi (Twitchell). Lambert and Thiéry, 1921. p. 383. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 135 Figure 93: Echinolampas tanypetalis (UF 5464), 70 mm TL, 65 mm TW, 34 mm TH, upper portion of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: right side. E: tilted oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 136 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 94: Echinolampas tanypetalis (UF 271053), 50 mm TL, 44.5 mm TW, 23 mm TH, Upper Eocene Ocala Limestone, Alachua County, Florida (FM-IP AL004). A: aboral. B: oral. C: left side. D: tilted oral viewpoint from posterior. Echinolampas aldrichi (Twitchell). Cooke, 1942. p. 38. Echinolampas aldrichi (Twitchell). Cooke, 1959. pp. 55-56. pl. 22, figs. 11-13. Echinolampas aldrichi (Twitchell). Gordon, 1963. p. 637. pl. 81, figs. 1, 2. Echinolampas sp. cf. E. aldrichi (Twitchell). Dono- van et al., 2005. p. 323. fig. 1. Echinolampas aldrichi (Twitchell). Osborn and Ciampaglio, 2014. p. 142. Echinolampas aldrichi (Twitchell). Martinez-Melo, 2019. pp. 5-8. figs. 6, 7. Echinolampas aldrichi (Twitchell). Buitrón-Sánchez et al., 2019. p. 53, figs. 2a, b, c. Occurrence.—Specimens of E. aldrichi (e.g., UF 112389) were collected from a dolomitic facies of the Marianna Limestone, in the Oakdale Quad- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 137 rangle between Marianna and Altha in southern Jackson County (FM-IP JA014). This is the first documented record of this species in Florida. The species is found in greatest abundance in the Upper Oligocene Chickasawhay Limestone in southwestern Alabama and southeastern Mississippi, and rarely occurs in the Upper Oligocene River Bend Formation of North Carolina. The type locality is in the Chickasawhay Limestone, near Gainestown, Clarke County, Alabama. Echinolampas aldrichi also occurs in the Oligocene of Chiapas, Mexico (Durham, 1969); the Miocene of Palenque, Chiapas, Mexico (Martinez- Melo, 2019); Oligocene of Veracruz, Mexico (Buitrón-Sánchez, 2019); the Oligocene of Puerto Rico (Cooke, 1959; Gordon, 1963), and the Middle Miocene of Belize (Donovan et al., 2005). Discussion.—Twitchell in Clark and Twitchell (1915), described this robust species from the holotype collected by T. H. Aldrich in the St. Stephens Limestone (now considered to be the Chickasawhay Limestone) near Gainestown, Alabama. However, as the occurrences noted above attest, this species is now known to have a much broader distribution. In Florida, E. aldrichi is known only from internal molds collected in dolomitic facies of the Marianna Limestone, south of Marianna, where it occurs with C. rogersi and S. americanus. This is the only species of Echinolampas in the Oligocene strata of the region and cannot be confused with any other element of the fauna. We figure one specimen (UF 112504) from this deposit that measures 66 mm TL, 58 mm TW, 33 mm TH (Fig. 92), but the collections include numerous specimens. Echinolampas tanypetalis Harper and Shaak, 1974 (Figs. 93-96) Echinolampas tanypetalis Harper and Shaak, 1974. pp. 166-169. txt fig. 2. tbl. 1, pl. 1. Echinolampas tanypetalis (Harper and Shaak). Os- born et al., 2016. tbl. 2. Occurrence.—This species has not been doc- umented outside of the Oligopygus wetherbyi Zone of the OLS of Florida. The type locality is a quarry west of Dowling Park (FM-IP LF002), in Lafayette County. The species also occurs in quarries north- west of Mayo (FM-IP LF001), Lafayette County and south of Tennille (FM-IP DI001), Dixie County. Discussion.—Harper and Shaak (1974) differ- entiated E. tanypetalis and the Oligocene E. aldrichi by the lower test and more anterior apical system of E. tanypetalis, and its longer petals that have pore pairs almost perpendicular to the petal axis. The pore pairs of E. aldrichi are distinctly oblique (ap- proximately 35° towards the ambitus from the petal axis). Furthermore, the peristome of E. tanypetalis is further forward. This species is rare, which would account for it remaining undocumented for so long in the OLS (an otherwise historically heavily collected unit), but persistent collecting in the Oligopygus wetherbyi Zone of the OLS will usually provide specimens, especially in the more northerly exposures of the Oligopygus wetherbyi Zone in Lafayette and Dixie Counties. We figure four specimens, the largest of which (UF 5464) is from the type locality, and measures 70 mm TL, 65 mm TW, 34 mm TH (Fig. 93). Suborder SCUTELLOIDA Mongiardino Koch et al., 2018 Infraorder LAGANIFORMES Desor in L. Agassiz and Desor, 1847 Family FIBULARIIDAE Gray, 1855 Genus Echinocyamus van Phelsum, 1774 Echinocyamus macneili Cooke, 1959 (Figs. 97-100) Echinocyamus macneili Cooke, 1959. p. 32. pl. 9, figs. 6-8. Echinocyamus macneili (Cooke). Kier, 1966a. p. 7, figs. 10A, B; 11A, B; 13A. not Echinocyamus parvus (Emmons). Zachos, 2005. p. 220. figs. 2.4-6. not Echinocyamus parvus (Emmons). Osborn et al., 2016. tbl. 3. Occurrence.—Within Florida, E. macneili is ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 138 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 95: Echinolampas tanypetalis (UF 271052), 32 mm TL, 29 mm TW, 17 mm TH (small specimen for the species), Upper Eocene Ocala Limestone, Alachua County, Florida (FM-IP AL004). A: aboral. B: oral. C: right side. D: tilted oral viewpoint from posterior. known only from the subsurface, where it was ob- tained from the OLS in Florida Geological Survey core W-19663 between 244 and 256 m depth (FM- IP OK005) in Okaloosa County. The type local- ity for the species is in the basal Upper Eocene Moodys Branch Formation where it is associ- ated with Periarchus lyelli (Conrad, 1834) along a creek (presumably Fall Creek) flowing into the Conecuh River in the NE¼ sec. 32, T. 4 N, R. 15 E., southwest of Andalusia, Covington County, Alabama. Discussion.—The holotype of E. macneili (USNM 862297; Fig. 99) was collected in the basal Upper Eocene Moodys Branch Formation, as noted above. The species is abundant at the type locality near the upper portion of the “Scutella bed” (dense ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 139 Figure 96: Echinolampas tanypetalis (UF 4901), 44.5 mm TL, 40 mm TW, 18 mm TH, specimen with some oral surface plate details, Upper Eocene Ocala Limestone, Levy County, Florida (FM-IP LV005). A: aboral. B: oral. C: left side. D: right side. concentration of P. lyelli) atop the bank just north of Fall Creek. The recognition of E. macneili from the OLS in the Okaloosa County well material discussed herein is the first documentation of Echinocyamus in Florida and the first known occurrence of E. mac- neili outside of its type locality. Although internal structure of this species seems not to have ever been mentioned in the literature, preparation of a pair of specimens show that it has strongly developed inter- nal supports, and the species is therefore attributable to Echinocyamus, and not Fibularia. Cooke (1959) stated that the lateral profile of E. macneili is like that of Echinocyamus parvus Emmons, 1858, but the upper surface is higher and more rounded. Unfortunately, Cooke compared E. macneili to specimens of Echinocyamus from the Thomas Farm site north of Jacksonville, Onslow County, NC that are not definitively E. parvus (Em- mons, 1858). With issues such as this, and to ensure clarity concerning our comparative material, a short summary of the taxonomy of the genus in the Eocene of North America is necessary. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 140 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 97: Echinocyamus macneili (UF 297039 #12), Upper Eocene Ocala Limestone, from 800-840 ft. depth in core from Fred Gannon Rocky Bayou State Park (FM-IP OK005), Okaloosa County, Florida. A: aboral. B: oral. C: right side. Figure 98: Echinocyamus macneili (UF 297039 #13), Upper Eocene Ocala Limestone, from 800-840 ft. depth in core from Fred Gannon Rocky Bayou State Park (FM-IP OK005), Okaloosa County, Florida. A: aboral. B: oral. C: right side. Echinocyamus parvus Emmons, 1858, the first Echinocyamus reported from North America, was described from the Eocene of Craven County, North Carolina, in strata now known as the Middle to Upper Eocene Castle Hayne Limestone. Meyer (1886) described Echinocyamus huxleyanus from a single damaged specimen collected at Claiborne, Alabama, in strata subsequently recognized as the Middle Eocene Gosport Sand (Cooke, 1942). The holotype of E. huxleyanus (USNM 559484) has been broken, but examination of the fragments remaining shows that they have no relationship with E. parvus. The type of E. huxleyanus has apparently not been consulted in revisionary works such as Clark and Twitchell (1915), Cooke (1942), and more recently, Zachos (2005), leading to incorrect synonymy with E. parvus. Determination of what E. huxleyanus, a species from Alabama, represents falls outside the scope of the present work except to note that it is not part of the synonymies of any Echinocyamus discussed here, and that it is almost undoubtedly not attributable to the genus. Cooke (1959) described E. macneili from the basal Upper Eocene Moodys Branch Formation ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 141 Figure 99: Comparison of aboral surfaces of female specimens of regional Eocene Echinocyamus emphasizing gonopore arrangement. A: E. macneili, holotype (USNM 562297), 5.6 mm TL, 4.0 mm TW, 2.4 mm TH, Upper Eocene Moodys Branch Formation, creek flowing into Conecuh River, Covington County, Alabama. B-E: E. macneili (from lot UF 338023) Upper Eocene Moodys Branch Formation, screened from Periarchus lyelli bed, Conecuh River at mouth of Fall Creek, above Brooklyn Rd. (very near the type locality of the species) west of Andalusia, Covington County, Alabama (FM-IP ZA145); all females. B: 3.2 mm TL, 2.1 mm TW, 1.3 mm TH. C: 4.2 mm TL, 2.9 mm TW, 1.5 mm TH. D: 4.9 mm TL, 3.6 mm TW, 1.9 mm TH. E: 5.3 mm TL, 3.9 mm TW, 1.6 mm TH. F-I: Echinocyamus cf. E. macneili (from lot UF 297039) Upper Eocene Ocala Limestone, from 800-840 ft. depth in core from Fred Gannon Rocky Bayou State Park (FM-IP OK005), Okaloosa ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 142 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) (Fig. 99 caption cont.) County, Florida. F: 2.6 mm TL, 1.8 mm TW, 1.1 mm TH. G: 2.3 mm TL, 1.5 mm TW, .9 mm TH. H: 4.5 mm TL, 3.2 mm TW, 1.4 mm TH. I: 5.3 mm TL, 3.7 mm TW, 1.9 mm TH. J-L: Echinocyamus cf. E. parvus, (from lot UF 338024) upper sequence three (of Zullo and Harris, 1987), late zone of Kier (1980) of the Middle to Upper Eocene Castle Hayne Limestone, Lanier Pit south of Maple Hill, Pender County, North Carolina (FM-IP ZN039). J: 3.4 mm TL, 2.5 mm TW, 1.3 mm TH. K: 3.5 mm TL, 2.5 mm TW, 1.4 mm TH. L: 3.6 mm TL, 2.6 mm TW, 1.3 mm TH. M, N: Echinocyamus cf. E. parvus; specimens identified as E. parvus in Kier (1980), Lanier Quarry (see F-H above); M: USNM 264054. N: USNM 264055. O: E. bisexus, holotype (USNM 650722), a male, Middle Eocene, Lake City Formation; USGS test well number 5, near Brunswick, Glynn County, Georgia. P: E. bisexus (USNM 650720) from Kier (1968); female, from the type locality, test well level 1130-1135 ft., Middle Eocene Lake City Formation, Glynn County, Georgia. Q-T: Echinocyamus cf. E. bisexus, (from lot UF 338025) upper sequence three (of Zullo and Harris, 1987), late zone of Kier (1980) of the Middle to Upper Eocene Castle Hayne Limestone, Martin Marietta Richland Quarry, near Catherine Lake, Onslow County, North Carolina (FM-IP ZN126). Q: 6.0 mm TL, 4.2 mm TW, 2.2 mm TH. R: 6.4 mm TL, 4.2 mm TW, 2.5 mm TH. S: 6.6 mm TL, 4.5 mm TW, 2.1 mm TH. T: 7.6 mm TL, 5.5 mm TW, 2.7 mm TH. U: Echinocyamus cf. E. bisexus, (from lot UF 338026), upper sequence three (of Zullo and Harris, 1987), late zone of Kier (1980) of the Middle to Upper Eocene Castle Hayne Limestone, Trent River, boat ramp at Trenton, Jones County, North Carolina (FM-IP ZN125): 5.4 mm TL, 3.4 mm TW, 1.6 mm TH. in Alabama. Kier (1968) described Echinocyamus bisexus from a Georgia test well in the Middle Eocene Lake City Formation (subsumed into the Avon Park Formation by Miller 1986). We discuss the implications of these descriptions in more detail, below. Zachos (2005) suggested that Echinocyamus meridonalis Meyer, 1887 was a juvenile P. lyelli, and went on to assert that E. bisexus, E. huxleyanus, and E. macneili are junior synonyms of E. parvus. This action effectively assigned all Eocene Echinocyamus in North America to one highly variable species (but see errors noted above with respect to E. huxleyanus, which we will not consider further). It became im- portant to compare our test well material to each form. While doing so, differences among the species became evident (except for E. parvus as discussed below), leading us to maintain E. macneili as a separate species. Echinocyamus parvus is likely the least un- derstood Echinocyamus in North America, making synonymies challenging. Emmons (1858) described E. parvus from a single specimen (the holotype was uncataloged, poorly figured, and has since been lost) collected in the Eocene of Craven County, North Carolina, in strata now referred to the Middle to Upper Eocene Castle Hayne Limestone. His fig- ure, a hand drawing of the oral surface done at natural size, lacks detail but could be interpreted as the oral surface of an Echinocyamus. At the time of his description, the sense of Echinocyamus and Fibularia were reversed from the presently ac- cepted nomenclature, so this adds yet another layer of difficulty over what Emmons meant with his drawing. Distinction between the two genera is diffi- cult (though not impossible), without reference to the internal structure. Emmons leaves us to guess what that might have been for his E. parvus, and we will likely never be sure what Emmons meant by the name ”Echinocyamus”. Emmons’ (1858: 307) description of E. parvus is: “Test small, oval, with rounded sides; avenues dorsal; mouth sub-central, rounded, large, with a crenulated margin; vent between the mouth and hinder margin; genital pores apparently four. The mouth is large in proportion to the size of the body and the vent is situated half way between the mouth and margin.” This description, though inade- quate for determing his concept of Echinocyamus, is almost sufficient to distinguish it from other small taxa found in the Castle Hayne Limestone. However, it is clearly not sufficient to distinguish it from sub- sequently described species of Echinocyamus from the region. Clark and Twitchell (1915) stated that the holotype of E. parvus was lost. They did not figure the species and merely reiterated Emmons’ descrip- tion. Kellum (1926) documented E. parvus from the Castle Hayne Limestone at Thomas Farm, 10 mi. northwest of Jacksonville, Onslow County, North Carolina. Kellum did not figure the specimens, nor indicate what led him to the identification, but con- sidering that Emmons’ description is not diagnostic, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 143 nor is his figure, and the holotype of E. parvus had been lost, it is likely Kellum made his specific identification solely based on what he perceived to be the genus designation and locality. That is, an Echinocyamus from the Castle Hayne Limestone must be E. parvus. Cooke (1942) reported on Kellum’s speci- mens (he documented five) from Onslow County, North Carolina, and figured one (USNM 499002). Cooke cautioned the figured specimen could not be compared to the type specimen, because it appeared to be lost. He noted that his image showed a speci- men wider posteriorly and narrower anteriorly than Emmons’ figure. Cooke (1959) figured the same specimen, and again listed Kellum’s locality as the only lo- cality for E. parvus in North Carolina. Cooke (1959) provided the most complete description of what workers continued to call ”E. parvus”, but it must be remembered that at this point, the only specimens available for study since Emmons’ de- scribed the type specimen, was the material in the USNM first documented by Kellum (1926) from Onslow County. Whether or not this is E. parvus remains an open question. Kier (1968) described E. bisexus from the Middle Eocene Lake City Formation (now Avon Park Formation) in a Georgia test well and noted that E. bisexus most resembled the concept of E. parvus in use at the time. He stated that both species have a similar shape, periproct position, and peristome size, but that E. bisexus differs in having longer petals with more pore pairs and a smaller madreporite (he did not mention differences in gonopore position). Again, Kier was comparing his new species with material in the USNM identified as E. parvus by Kellum (1926). Kier (1980) subsequently recorded 67 speci- mens of E. parvus and 13 specimens of E. bisexus from the Castle Hayne Limestone in the Lanier Quarry, near Maple Hill, Pender County, North Carolina. He redescribed “Echinocyamus parvus” from this material, whether or not it actually rep- resents E. parvus of Emmons (1858). Kier (1980) also documented disparity in gonopore size, with some specimens having very small gonopores and others very large ones, interpreting this as sexual dimorphism. Kier (1980: pl. 10, figs. 5-10) figured three specimens (USNM 264053-55) he identified as E. parvus from the Lanier Quarry. Of these, two (USNM 264054 and 264055) appear to be females with large gonopores and are distinguished from E. macneili by their gonopores being situated further apart, though not partway adoral along the inter- ambulacrum like E. bisexus (Fig. 99). We refer to the specimens in Kier’s (1980: pl. 10, figs. 5-10) as Echinocyamus cf. E. parvus in view of the analysis above. Kier (1980: 34) lists USNM 499002 (the same specimen from Onslow County collected by Kellum [1926] and figured by Cooke [1942, 1959]) as one of his figured specimens, however, it does not seem to appear among his figures. The stratigraphy and locality-based identifi- cation of Kellum (1926), followed by Cooke (1942, 1959) and Kier (1968, 1980) was reasonable prior to Kier (1980) because only E. parvus had been documented from the Castle Hayne Limestone. That is only the specimens from Thomas Farm, north of Jacksonville, North Carolina were known, and they all appeared to belong to one species. However, when Kier (1980) documented both E. bisexus and another that he attributed to E. parvus by compar- ing it to the Thomas Farm material, in the Castle Hayne Limestone at the Lanier Quarry, this circum- stantial approach to identification of E. parvus was rendered invalid. We can no longer ascertain that the holotype of E. parvus is distinct from what Kier (1968) subsequently named E. bisexus. We figure the latter species from two additional localities in the Castle Hayne Limestone of North Carolina (Fig. 99) and note that E. bisexus is far more common in the Castle Hayne Limestone than the form Kellum (1926) identified as E. parvus from Thomas Farm. The only solution, other than finding the miss- ing holotype, is to discover and describe new ma- terial from the type locality of E. parvus, possibly by additional collecting. Emmons (1858) stated the type locality of E. parvus is Craven County, North Carolina, which is vague, but he provided clues to perhaps two specific localities. Within his report, Emmons discussed two exposures of Eocene “marl” ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 144 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) in Craven County. One is on the south bank of the Neuse River on the Biddle Plantation (Biddle Land- ing on the Neuse River) and the other is on the plantation of William B. Wadsworth at Core Creek. He also indicated the fossil fauna associated with E. parvus, although he did not state what additional echinoids might be a part of that fauna. Interestingly, he noted that the small comatulid crinoid Microcri- nus conoideus Emmons, 1858 was collected with E. parvus. Unfortunately, our research has not yet yielded records of comutalid crinoids (or Echinocyamus) at Biddle Landing. This leaves the Wadsworth Marl at the Wadsworth Plantation at Core Creek. Emmons (1858: 105) stated that a crinoid is abundant in the Wadsworth Marl, which also lends evidence that the type locality of E. parvus is at the Wadsworth Plantation at Core Creek, Craven County, North Carolina. Wadsworth wrote a letter to Emmons (1858), included in the latter’s report, detailing how he utilized the marl dug on his plantation as fertilizer, suggesting that the marls was not a natural exposure on Core Creek, but excavations on the Wadsworth Plantation near Core Creek. Wadsworth’s Plantation is shown south of Core Creek, roughly midway between Rock Land- ing and Biddle Landing on the Neuse River, on a beautiful map hand-drawn by Jeremy Francis Gilmer (1818-1883) during the Civil War (between 1861 and 1865: ”Reconnaissance of the country between Newbern and Goldsboro between the Trent and the Neuse Rivers”). Unfortunately, there are no current marl pits or other exposures of the Castle Hayne Limestone in this area. Therefore, although we have a suggestion as to where to search for additional specimens in the type area, until exposures become available, or the holotype is discovered, we see no viable way to identify positively any specimen as E. parvus. As a result, we maintain that specimens referred to as E. parvus from the Castle Hayne Limestone by Kellum (1926), Cooke (1942, 1959), and Kier (1980) (Fig. 99) can only be referred to as Echinocyamus cf. E. parvus. Consequently, no other species can be placed in junior synonymy. Test width (TW) as a percentage of test length (TL) is very similar in these taxa: 70.2% for Rich- lands Quarry E. bisexus, 70.8% for Lanier Quarry Echinocyamus cf. E. parvus, 70.1% for type local- ity E. macneili, and 72.1% for Florida E. macneili. However, as noted by Cooke (1959), the test of E. macneili is higher than that of Echinocyamus cf. E. parvus (Fig. 100). The average TH as a percent- age of TL for type locality E. macneili is 40%, and Florida E. macneili is 40.9%; whereas the average of E. bisexus from the Richlands Quarry was 36%, and Echinocyamus cf. E. parvus from the Lanier quarry was 38.3%. TH vs TW revealed similar results, with TH of E. macneili from the type locality being on av- erage 57.2% TW. The Florida E. macneili is 59.3%, but in Richland Quarry E. bisexus TH is only 51.4% TW; specimens of E. parvus from the Lanier quarry were closer to E. macneili with TH 53.5 % TW. However, TH is not the characteristic that best distinguishes E. macneili, Echinocyamus cf. E. parvus, and E. bisexus. The gonopores of females of E. macneili are closer together than in Echinocya- mus cf. E. parvus, and much closer together than that of E. bisexus (Fig. 99). We compared female specimens (specimens with larger gonopores) of regional Echinocyamus, including the available holo- types. This includes E. macneili from its type lo- cality, Echinocyamus cf. E. parvus from the Castle Hayne Limestone, E. bisexus from the Castle Hayne Limestone, and specimens we recognize as E. mac- neili from Okaloosa County, to explore differences in gonopore arrangement of these species (Fig. 99). This comparison confirms we have more than one species of Echinocyamus in the regional Eocene strata. To reiterate, Echinocyamus macneili is readily distinguished from E. bisexus in not only having a higher test, but by differences in gonopore arrange- ment in females of these two species. In E. macneili, the four enlarged gonopores are closer together in the apical area than any other regional Eocene species of Echinocyamus. In E. bisexus, the gonopores are placed well outside the apical area part of the way down their respective interambulacra (Fig. 99). Cooke (1959) alluded to possible sexual di- morphism in E. macneili in his description. Due to the large gonopores, he considered the holotype of E. macneili (Fig. 99) to be a female. The species ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 145 seems very likely to be sexually dimorphic (females with large gonopores, males with gonopores that are barely discernable without magnification) when comparing multiple specimens from the type lo- cality and from the specimens we studied from the Okaloosa County Well Core (UF 297039, UF 307764). Osborn et al. (2016) followed Zachos (2005) in recognizing E. parvus (and not E. macneili) in the Moodys Branch Formation of Alabama (Osborn et al., 2016: table 3). In retrospect, this was an error that we have corrected in the synonymy above. As mentioned above, although E. macneili is not known from surface exposures in Florida, it was abundant in Florida Geological Survey Core W-19663, being represented by two lots: UF 297039 (14 tests) and UF 307764 (15 tests). Genus Fibularia Lamarck, 1816 Fibularia vaughani (Twitchell in Clark and Twitchell, 1915) (Figs. 101-104) Echinocyamus vaughani Twitchell in Clark and Twitchell, 1915. p. 160, pl. 74, figs. 1a-f. Echinocyamus vaughani (Twitchell). Lambert and Thiéry, 1925. p. 576. Echinocyamus vaughani (Twitchell). Cooke and Mossom, 1929. pl. 3, figs. 4a-b. Fibularia vaughani (Twitchell). Cooke, 1942. p. 7. Fibularia vaughani (Twitchell). Cooke, 1945. fig. 5, no. 4. Fibularia vaughani (Twitchell). Fischer, 1951. p. 55. txt fig. 1. Fibularia vaughani (Twitchell). Cooke, 1959. p. 30. pl. 9, figs. 23-27. Fibularia vaughani (Twitchell). Osborn et al., 2016. tbl. 2. Occurrence.—This is the common micro- echinoid of the OLS of Florida, and more rarely in Georgia. It occurs in sporadic abundance in the Oligopygus haldemani Zone of the OLS, es- pecially northwest of Mayo, Lafayette County (FM- IP LF001) (Figs. 101, 102). It is common in the Oligopygus phelani Zone in the lower portion of the OLS, especially along the banks of the With- lacoochee River (FM-IP LV024), Levy County (Figs. 103, 104); Cross Florida Barge Canal (FM-IP CI001), south of Inglis, Citrus County, and in small pits near Inglis, Levy County. Discussion.—Twitchell in Clark and Twitchell (1915), described this species as Echinocyamus vaughani from strata believed to be lower Oligocene at the time, but now recognized as the Upper Eocene OLS, along the Flint River in Georgia. Fibularia vaughani appears to be restricted to the OLS of Florida and Georgia, and it cannot be confused with any other species within the OLS. Detailed examination of the type, includ- ing plate patterns of both the aboral and oral sur- faces, strongly suggest that this species should not be placed in Fibularia. Aspects of these patterns strongly ally F. vaughani with members of Fibu- lariella Mortensen, 1948, presently considered to be rotulids. A revision of these minute taxa is in progress, but it is almost certain that placement in Fibularia is not the final word on the systematics of F. vaughani. Family NEOLAGANIDAE Durham, 1954 The State of Neolaganid Systematics in the Region Durham (1954) erected the family Neola- ganidae to contain New World laganids whose basi- coronal plates are arranged in a pentagon and not in a star-like pattern (i.e., no points on the distal part of the interambulacral basicoronals, and with large ambulacral basicoronals almost equal in length to the interambulacral basicoronals). He also erected numerous new genera to contain many species of laganids including Cubanaster Sánchez Roig, 1952e, Neolaganum Durham, 1954, Weisbordella Durham, 1954, and Wythella Durham, 1954. Kier (1967) subsequently erected Pentedium but left its family attribution in question, stating it likely belonged to a new family. Pentedium is distinguished from the neo- laganids in having five, instead of four genital pores. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 146 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 100: Comparison of regional Eocene Echinocyamus: comparisons of test length, width, and height of Echinocyamus macneili from the Moodys Branch Formation at the type area, Conecuh River west of Andalusia, Covington County, Alabama. (FM-IP ZA145); Echinocyamus cf. E. bisexus from the Castle Hayne Limestone, Martin Marietta Quarry near Catherine Lake, Onslow County, North Carolina (FM-IP ZN126); Echinocyamus cf. E. parvus from the Castle Hayne Limestone, Lanier Pit, Pender County, North Carolina (FM-IP ZN039), and E. macneili, Upper Eocene Ocala Limestone, from 800-840 ft. depth in core from Fred Gannon Rocky Bayou State Park (OK005), Okaloosa County, Florida. Although test width (TW) as a percentage of test length (TL) is very similar in these specimens: 70.2% for Richlands Quarry E. bisexus, 70.8% for Lanier Quarry Echinocyamus cf. E. parvus, 70.1% for type locality E. macneili, and 72.1% for Florida E. macneili; the test of E. macneili is higher than its congeners. The average test height (TH) as a percentage of TL for type locality E. macneili is 40%, and Florida E. macneili is 40.9%; whereas the average of E. bisexus from the Richlands Quarry is 36%, and Echinocyamus cf. E. parvus from the Lanier quarry is 38.3%. TH vs TW revealed similar results, with the TH of E. macneili from the type locality on average 57.2% TW, and the Florida E. macneili is 59.3%, and in Richland Quarry E. bisexus the height is only 51.4% TW; specimens of Echinocyamus cf. E. parvus from the Lanier quarry were closer to E. macneili with a TH on average equal to 53.5 % TW. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 147 Figure 101: Fibularia vaughani (UF 337969), 4.5 mm TL, 3.5 mm TW, 2.5 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. However, a year later Kier erected Durhamella Kier, 1968, and placed it in the Neolaganidae without hes- itation, although two species of that genus have five genital pores. We recognize both Durhamella and Pentedium in the Neolaganidae, and further discuss Durhamella below. Without providing a formal statement or rea- soning, Kroh and Smith (2010) demoted the Ne- olaganidae to a subfamily within the Laganidae, subdividing the Laganidae into two subfamilies: the Neolaganinae (with a strong presence in the fos- sil record of the region), and the Laganinae (with no known representatives in the fossil record of the eastern United States). This designation was apparently adopted by Kroh (2020), as the family Neolaganidae does not appear in that work. Smith and Kroh (2011) followed Durham (1954) in stating that the Neolaganinae have basicoronal plates in the shape of a pentagon, adding that the petals usually have compound plating. The coordinate subfam- ily Laganinae have basicoronal ambulacral plates in a stellate arrangement with ambulacrals form- ing points of the star in the aboral direction, and petaloid plating that is usually simple, made en- tirely of primary plates. We follow Durham (1954) in maintaining the Neolaganidae as a family along with the Laganidae, pending deeper taxon sampling than provided in Kroh and Smith (2010). This is ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 148 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 102: Fibularia vaughani (UF 337970), 5 mm TL, 3.5 mm TW, 2.5 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. because their phylogenetic analysis, based on a sin- gle laganid and a single neolaganid, provided no evidence for or against placing the Neolaganidae into the Laganidae. Moreover, the latter family is likely not monophyletic (Mooi et al., 2001). In con- trast, there is good evidence to suggest that the Neolaganidae is monophyletic. While determining taxonomic assignment of the new neolaganid species described below, it be- came apparent that the diagnoses of many neo- laganid genera were not, in fact, diagnostic. The greater number of available specimens revealed that many characteristics used by Durham (1954) and Kier (1968) to distinguish the genera are either not consistent or not diagnostic, being present in nu- merous other genera. Study of oral plate maps for all regional genera (Fig. 105), as well as the plat- ing in the petals (Fig. 106), revealed characteristics that facilitate identification of some of the genera. Genus diagnoses still require comparison of numer- ous features, such as oral surface plate arrangement, concavity of the oral surface, presence and structure of hydropore grooves, and petaloid structure. Therefore, revised diagnostic criteria are pro- vided in revisions of Durhamella, Neolaganum, and Weisbordella below. Even with these revised diag- noses, it seems clear that the Neolaganidae likely contains too many genera, as there remain few di- agnostic characteristics to distinguish them. Future revisions will have to take this into account, as well ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 149 Figure 103: Fibularia vaughani (UF 337988), 6.5 mm TL, 5 mm TW, 3.5 mm TH, Oligopygus phelani Zone, Upper Eocene lower Ocala Limestone, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: left side. D: right side. as the many species from the Caribbean that fall well outside the study region. Genus Neolaganum Durham, 1954 When Durham (1954) erected Neolaganum, he designated it as the type genus of his new family Neolaganidae. He characterized the genus by the moderate number of plates on the oral surface and presence of a hydropore groove, neither of which are diagnostic, either in combination or as unique fea- tures. Durham (1954) stated the hydropore groove, the more pentagonal shape, especially posteriorly, and the flat or nearly flat oral surface distinguish this genus from Weisbordella. Durham (1954: 681) stressed the importance of the presence of hydropore grooves in this genus but noted: “if these differences should be found to be of lesser value, Neolaganum should have priority over Weisbordella”. Our anal- ysis indicates the presence of hydropore grooves ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 150 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 104: Fibularia vaughani (UF 337989), 5 mm TL, 3.5 mm TW, 3 mm TH, Oligopygus phelani Zone, Upper Eocene lower Ocala Limestone, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: left side. D: right side. is widespread in the neolaganids, including Weis- bordella, and therefore not of great taxonomic sig- nificance except in some cases. However, further analysis indicated that we should not consider Weis- bordella a junior synonym of Neolaganum at this time. Durham (1954) designated N. archerensis as the type species for Neolaganum. Confusion between N. archerensis and N. durhami, discussed further in the remarks for N. archerensis, indicated that further analysis of the oral plate architecture of N. archerensis, not previously considered, needed to be performed. The importance of oral plate architecture in neolaganids was recognized by Durham (1954) when establishing the family, and in describing some of the taxa he placed therein. Therefore, we provide oral plate maps of all three species of Neolaganum: N. archerensis, N. dalli, and N. durhami (Fig. 105). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 151 Figure 105: Comparison of oral surface details of Florida neolaganids. Ambulacrum III is towards the top, the interambulacral plates are shaded, and peristome and periproct are solid black (see table 7). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 152 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 106: Plate patterns in petals of Florida neolaganid species. For each, perradius (Lovénian numbering) in which petal is situated is indicated, along with specimen repository. Plating shown for section of b column near approximate midpoint of petal’s length (except D. tetrapora, for which entire length of b column is depicted, including ocular). Pore pairs indicated by solid black, conjugation of pores by light gray line. Ambitus downward for each image. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 153 Figure 107: Laganum archerensis holotype (= Neolaganum archerensis) (USNM 164667A), 18 mm TL, 16 mm TW, 3.5 mm TH, Middle Eocene Avon Park Formation, likely from a well, near Archer, Florida. (images from Smithsonian NMNH online database). A: aboral. B: oral. Neolaganum archerensis, N. dalli, and N. durhami all tend to have a greater number of plates on the oral surface, especially in the ambulacra, than Weisbordella (Fig. 105; Tables 8, 9). Furthermore, all three Neolaganum species have at least one plate more in each of the anterior paired regions (i.e., interambulacral 2 and 3) than in the posterior paired (interambulacral 1 and 4). Therefore, not all the interambulacral regions have the same number of plates in each individual, as in both columns a and b of the anterior pair, there are more oral plates in Neo- laganum than in Weisbordella. This kind of variation suggests that a total number of oral plates among all interambulacra would be helpful to distinguish these taxa. Among all species of Weisbordella, the total number or oral interambulacral plates ranges from 41 to 46, whereas in Neolaganum, the range is non-overlapping, at 50 to 52. For ambulacral plates, the respective numbers are 54 to 59 in Weisbordella, but 66 to 78 in Neolaganum (Table 9). The flat oral surface of Neolaganum further distinguishes it from the typically concave oral sur- face of Weisbordella, though in the latter, the con- cavity can be very shallow. Emended Diagnosis.—Neolaganids that com- bine: presence of five gonopores with a branched hydropore groove; almost planar oral surface with between 66 and 78 ambulacral oral plates and 50 to 52 interambulacral plates; poriferous zones equal to or slightly greater than half the width of the porifer- ous zone as measured from the perradial to abradial suture about two thirds of the length of the petal; at most, one demiplate every third plate in the petal. Neolaganum archerensis (Twitchell in Clark and Twitchell, 1915) (Figs. 105-113; Tables 8, 9) Laganum archerensis Twitchell in Clark and Twitchell, 1915. p. 161. pl. 75, figs. 1a-d. Echinodiscus archerensis (Twitchell). Lambert and Thiéry, 1925. p. 581. Laganum archerensis (Twitchell). Cooke and Mos- som, 1929. pl. 3, figs. 6a-b. Rumphia archerensis (Twitchell). Cooke, 1942. p. 26. In part, not pl. 2, figs. 11-13. Rumphia archerensis (Twitchell). Cooke, 1945. fig. 5, no. 6. Neolaganum archerensis (Twitchell). Durham, 1954. pp. 680, 681. not txt fig. 28 which is N. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 154 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 108: Neolaganum archerensis (UF 344751), 8.8 mm TL, 8.0 mm TW, 2.7 mm TH, Middle Eocene Avon Park Formation, in a well core at 50 ft. depth (in a 200 ft. deep well) near Lake Wales, Polk County, Florida. A: aboral. B: oral. C: left side. D: right side. durhami. Neolaganum archerensis (Twitchell). Durham, 1955. p. 146. not txt figs. 15f, 30b which are N. durhami. not Neolaganum dalli (Twitchell). Cooke, 1959. pp. 51-52 (in part). not pl. 21, figs. 1-4, which is N. dalli. Neolaganum archerensis (Twitchell). Durham, 1966. p. 475. (in part) not fig. 365, 1a-1d which are N. durhami. Neolaganum archerensis (Twitchell). Mooi, 1989. p. 37. (in part) not figs. 9b or16c which are N. durhami. Occurrence.—This species has only been re- covered from well cuttings in subsurface deposits of the Middle Eocene Avon Park Formation, Florida. It is not known from surface exposures. The type locality is near Archer, Alachua County, where the holotype (USNM 164667A) was presumably ob- tained from well cuttings at an unknown depth. A few notable lots of this species within the FM-IP collections are UF 12892, from 15.2 m depth in a well near Lake Wales, Polk County, and UF 56401, from 512-518 m depth in a well near Belle Meade, Collier County. Discussion.—The complicated synonymy, as well as that for N. durhami, indicates this species has been misunderstood since its description by Twitchell in Clark and Twitchell (1915). William Healy Dall collected the holotype of Laganum archerensis Twitchell in Clark and Twitchell, 1915, as well as the associated holotype of Laganum dalli Twitchell in Clark and Twitchell, 1915, from near ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 155 Figure 109: Neolaganum archerensis (UF 344752), 14.8 mm TL, 12.5 mm TW, 4.1 mm TH, Middle Eocene Avon Park Formation, in a well core at 50 ft. depth (in a 200 ft. deep well) near Lake Wales, Polk County, Florida. A: aboral. B: oral. C: left side. D: right side. Archer, Alachua County, Florida. It was presumed by Cole and Ponton (1932) and Cooke (1959) that these taxa were both obtained from well cuttings, as no exposures of the Avon Park Formation exist at the surface near Archer. Twitchell in Clark and Twitchell (1915), de- scribed L. archerensis from a single specimen col- lected in strata that were recognized as Oligocene at the time. Twitchell noted its similarity to L. dalli, which was collected from the same well cuttings, but distinguished it by a concave ring encircling the petaloid region on the upper sur- face, narrower petals, less depressed and narrower poriferous zones, and a periproct that was closer to the margin. The holotypes of both species are clearly different from each other (Figs. 107, 114). This is further evidenced by analysis of oral plate architecture. The oral plate arrangement of N. archerensis and N. dalli (Fig. 105) demonstrates the first post- basicoronals in interambulacrum 5 (plates 5.b.2 and 5.a.2) of N. archerensis are significantly longer ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 156 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 110: Neolaganum archerensis (UF 344753), 18.8 mm TL, 16.7 mm TW, 4.2 mm TH, Middle Eocene Avon Park Formation, in a well core at 50 ft. depth (in a 200 ft. deep well) near Lake Wales, Polk County, Florida. A: aboral. B: oral. C: left side. D: right side. than those in N. dalli. However, even though the periproct of N. archerensis is much nearer the posterior margin than in N. dalli, the periproct in both species is first in contact with plates 5.b.2 and 5.a.2. Twitchell assumed that L. archerensis and L. dalli were associated, as both were collected from the same well. However, there are clear differences in preservation and color of the two holotypes. It is likely they were not obtained from the same depth in the well, as discussed below. Cole and Ponton (1932) conducted a study of 266 specimens they identified as Laganum dalli from numerous deep wells throughout Florida. They noted that no specimens of L. dalli had been doc- umented from surface exposures. Their intent was to demonstrate the great variability, especially in periproct placement, found among specimens they ascribed to L. dalli. Apparently, their first inclina- tion was to describe a new species for the forms with a more marginal periproct, and an examina- tion of specimens in the FM-IP Collections reveals a few specimens (UF 249008 and UF 249006) with the name “Laganum polkensis Ponton” on ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 157 Figure 111: Neolaganum archerensis (UF 249008), 12.8 mm TL, 11.3 mm TW, 3.2 mm TH, Middle Eocene Avon Park Formation, 240-250 ft. depth in the Davenport city well, Polk County, Florida. Specimen was deposited at the Florida Geological Survey (now in the Florida Museum collections) with original labels identifying it as Laganum polkensis, attributing the name to Ponton; likely the potential new species indicated by Cole and Ponton (1932). A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: tilted oral viewpoint from posterior. E: peristome. F: left side. G: apical area and petaloid portion of ambulacrum III. H, I: original Florida Geological Survey labels. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 158 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) the labels (Fig. 111). This was the name Cole and Ponton intended to give to these forms. Interest- ingly, Cole and Ponton (1932) did not discuss L. archerensis and its more posterior periproct, instead concluding that periproct placement of L. dalli is highly variable but situated closer to the posterior margin in smaller specimens, even though their data show considerable overlap with periproct position in their “L. polkensis” (Cole and Ponton, 1932: ta- ble 2). Cole and Ponton (1932) did not distinguish their specimens by population and/or well depth. If specimens of both L. dalli and L. archerensis exist in these samples (which they likely do), perhaps even from different depths in the well, the finding of variation they ascribed to a single species, L. dalli, would be compromised, since there could be two distinct taxa in their samples – whether or not one of these was already named. Regrettably, Cole and Ponton (1932) did not assign catalog numbers to the specimens they mea- sured or indicate them in some way. However, their specimens were deposited in the collections of the Florida Geological Survey, whose material now re- sides in the FM-IP Collections at the University of Florida. Specimens from this material are further discussed below. Cooke (1942) assigned L. archerensis to Rumphia because of its four genital pores and flat oral surface with ambulacral grooves. He also as- signed L. dalli to Peronella (curiously, a very dif- ferent genus than the one to which he assigned L. archerensis) because of its four genital pores, subpentagonal peristome, buccal pores, and punc- tate extrapetalous ambulacral regions, which Cooke felt resembled those of Peronella peronii (Agassiz, 1841), the type species of Peronella. Cooke (1941a) mentioned the great variation in periproct place- ment among L. dalli also noted by Cole and Ponton (1932). However, Cooke (1942) made an additional decision that would have consequences – confus- ing species concepts of L. archerensis and L. dalli for many decades. Cooke (1942) modified the con- cept of L. archerensis by including specimens col- lected from surface exposures of what would become known as the Inglis Formation, and subsequently lower OLS, at numerous localities in Levy County, Florida. As discussed below, this taxon from the lower OLS would later be described as a mem- ber of a new genus, Neolaganum Durham, 1954, as N. durhami (Cooke, 1959), but misidentifica- tion of specimens of N. durhami as N. archerensis would continue, largely by those following Durham (1954), for example, Durham (1955, 1966), and Mooi (1989). Durham (1954) erected Neolaganum and des- ignated L. archerensis as the type species. Making matters worse was that to represent the genus, he fig- ured a specimen, UCMP 33296, that he designated as a hypotype of N. archerensis. This specimen was collected from the lower OLS at the mouth of the Withlacoochee River (FM-IP LV024) west of Inglis, in Levy County, Florida. Neolaganum archerensis has never been documented at this site, nor are there exposures of the Avon Park Forma- tion in the vicinity, but what is now known as N. durhami is very abundant. Cooke (1959) rec- ognized this and postulated a plausible scenario in which Durham (1954) based Neolaganum on a misidentified specimen that was N. durhami, and not the L. archerensis described originally from a well near Archer. Luckily, Durham (1954) indicated that there were no genus-level differences between the taxa, so Durham’s description of the genus still ap- plies. However, Durham’s (1954) figures of the oral plate arrangement of N. durhami (UCMP 33296), erroneously identified as N. archerensis, were repli- cated by Durham, (1955, 1966) and Mooi (1989). As far as we can determine, figures of the oral plating of the holotype of N. archerensis (USNM 164667A) have not been published until the present work. Cooke (1959) recognized N. archerensis as a subjective junior synonym of N. dalli. He did not provide a rationale for this assertion, but he did refer to the study of Cole and Ponton (1932) while stating that the periproct placement of L. dalli is highly variable. As the more posterior periproct of N. archerensis is the most reliable characteristic distinguishing it from N. dalli, Cooke (1959) likely based his decision on the variability of this feature in N. dalli suggested by Cole and Ponton (1932), as ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 159 Ta bl e 8: Ta bu la rk ey fo rg en er a in th e fa m ily N eo la ga ni da e D ur ha m ,1 95 4, pl us sp ec ie so cc ur rin g in stu dy ar ea . Ta xo n D ist rib ut io n1 Te st sh ap e Te st le ng th (m m )2 O ra l su rfa ce co nc a v ity Pe rip ro ct po si tio n A bo ra ld e- pr es si on s Pe ta ls su nk en G on op or e nu m be r H yd ro po re s O ut er pe ta l po re pa ir po rif er ou s zo ne s4 D em ip la te s in pe ta l5 O ra l am bu la cr al pl at e #6 O ra li nt er - am bu la cr al pl at e #7 C ub an as te r Sá nc he z Ro ig , 19 52 E co as t Pa na m a, M X El on ga te o v al , th ic k m ar gi n 70 A bs en t C lo se rt o pe ris to m e th an am bi tu s N on e Sl ig ht ly 4 In br an ch ed gr oo ve Lo ng , sl it- lik e < 1/ 2 R ar e? 7 – 9 4 – 6 Sa nc he ze lla D ur ha m ,1 95 4 C ub a Ve ry el on ga te , th ic k m ar gi n 50 A bs en t C lo se rt o pe ris to m e th an am bi tu s N on e D ee pl y 4 In br an ch ed gr oo ve Lo ng , sl it- lik e > > > 1/ 2 R ar e 9 – 11 5 – 6 Ne ol ag an um D ur ha m , 19 54 FL ,C B Pe nt ag on al , lo w m ar gi n 40 A bs en t N ea ro rf ar fr om pe ris to m e N on e N o 5 In br an ch ed gr oo ve Sl ig ht ly el on ga te to sl it- lik e = or > 1/ 2 A bs en to r ev er y fe w pl at es 6 – 8 4 – 6 Ne ol ag an um ar ch er en si s (T w itc he ll, 19 15 ) FL Pe nt ag on al , lo w m ar gi n 20 A bs en t C lo se rt o am bi tu st ha n pe ris to m e N on e N o 5 In br an ch ed gr oo ve Lo ng , sl it- lik e > 1/ 2 A bs en t 6 – 7 5 – 6 Ne ol ag an um da lli (T w itc he ll, 19 15 ) FL ,C B Pe nt ag on al , lo w m ar gi n 20 A bs en t C lo se rt o pe ris to m e th an am bi tu s N on e N o 5 In br an ch ed gr oo ve Sh or t, on ly sl ig ht ly el on ga te > 1/ 2 A bs en t 6 – 7 4 – 5 Ne ol ag an um du rh am i C oo ke ,1 95 9 FL Pe nt ag on al , lo w m ar gi n 40 A bs en t C lo se rt o am bi tu st ha n pe ris to m e N on e N o 5 In br an ch ed gr oo ve Lo ng , sl it- lik e = 1/ 2 Ev er y 2n d or 3r d pl at e 7 – 8 4 – 5 W ei sb or de lla D ur ha m , 19 54 A L, FL ,G A , C B El on ga te ov al , th ic k or lo w m ar gi n ∼8 0 Va ria bl y pr es en t C lo se rt o am bi tu st ha n pe r is to m e N on e N o 4 In lo ng , w in di ng gr oo ve Ro un d to lo ng , s lit -li ke < 𝑜 𝑟 < < < 1/ 2 A lw ay s pr es en t 5 – 6 3 – 4 W ei sb or de lla cu ba e (W ei sb or d, 19 34 ) FL ,G A El on ga te ov al , lo w m ar gi n 40 D ee p C lo se rt o am bi tu st ha n pe r is to m e N on e N o 4 In lo ng , w in di ng gr oo ve Ro un d < < < 1/ 2 M an y, pr im ar ie s ab se nt 5 – 6 3 – 4 W ei sb or de lla jo hn so ni (T w itc he ll, 19 15 ) A L, FL ,G A El on ga te ov al , lo w m ar gi n 60 Va ria bl e C lo se rt o am bi tu st ha n pe ris to m e N on e N o 4 In lo ng , w in di ng gr oo ve A lm os t ro un d < < < 1/ 2 M an y, pr im ar ie s ra re ∼6 ∼4 W ei sb or de lla in gl is en si sn . sp . FL El on ga te ov al , lo w m ar gi n 25 A bs en t C lo se rt o am bi tu st ha n pe ris to m e N on e N o 4 In lo ng , w in di ng gr oo ve Sh or t, on ly sl ig ht ly el on ga te < < < 1/ 2 Ev er y 2n d pl at e 5 3 – 4 W ei sb or de lla lib um n. sp . FL El on ga te ov al , th ic k m ar gi n ∼8 5 Sh al lo w C lo se rt o am bi tu st ha n pe ris to m e N on e N o 4 In co m pl ex br an ch ed gr oo ve s Lo ng , sl it- lik e Sl ig ht ly < 1/ 2 M an y, pr im ar ie s ra re ∼6 ∼4 W yt he lla D ur ha m ,1 95 4 FL El on ga te pe nt ag on al , ve r y th in m ar gi n 10 0 A bs en t C lo se rt o am bi tu st ha n pe r is to m e N on e N o 4 In sh or t, un br an c h ed gr oo ve Lo ng , sl it- lik e = 1/ 2 M an y, pr im ar ie s ra re 7 – 8 4 – 6 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 160 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Ta bl e 8 C on t. Ta bu la r k ey fo r g en er a in th e fa m ily N eo la ga ni da e D ur ha m , 1 95 4, p lu s s pe ci es o cc ur rin g in st ud y ar ea . Ta xo n D ist rib ut io n1 Te st sh ap e Te st le ng th (m m )2 O ra l su rf ac e co nc av ity Pe rip ro ct po si tio n A bo ra l d e- pr es si on s Pe ta ls su nk en G on op or e nu m be r H yd ro po re s O ut er p et al po re p ai r po rif er ou s zo ne s4 D em ip la te s in p et al 5 O ra l am bu la cr al pl at e #6 O ra l i nt er - am bu la cr al pl at e #7 W yt he lla e ld ri dg ei FL El on ga te pe nt ag on al , ve ry th in m ar gi n 10 0 A bs en t C lo se r t o am bi tu s t ha n pe ris to m e N on e N o 4 In sh or t, un br an ch ed gr oo ve Lo ng , sl it- lik e = 1/ 2 M an y, pr im ar ie s ra re 7 – 8 4 – 6 FL , G A O va l o r pe nt ag on al , lo w m ar gi n 40 A bs en t C lo se r t o am bi tu s t ha n pe ris to m e Pr es en t o r ab se nt N o 5 In sh or t, un br an ch ed gr oo ve R ou nd = 𝑜 𝑟 < < < 1/ 2 R ar e or ne ar ly ab se nt 4 – 6 2 – 4 FL , G A O va l o r pe nt ag on al , lo w m ar gi n 25 A bs en t C lo se r t o am bi tu s t ha n pe ris to m e N on e N o 5 In sh or t, un br an ch ed gr oo ve R ou nd < < < 1 /2 V er y ra re , ev er y 5 to 10 p la te s 5 – 6 3 – 4 FL O va l o r pe nt ag on al , lo w m ar gi n 40 St ro ng ly de ve lo pe d al on g su tu re s C lo se r t o am bi tu s t ha n pe ris to m e N on e N o 5 In sh or t, un br an ch ed gr oo ve R ou nd < < < 1 /2 V irt ua lly ab se nt 5 – 6 3 – 4 FL El on ga te o va l, lo w m ar gi n 10 Pr es en t on ly in so m e in di vi du al s C lo se r t o am bi tu s t ha n pe ris to m e N on e N o 4 In sh or t, un br an ch ed gr oo ve R ou nd = 1/ 2 V irt ua lly ab se nt 4 – 6 2 – 3 G A O va l, th ic k m ar gi n 5 A bs en t C lo se r t o am bi tu s t ha n pe ris to m e In te ra m bu l- ac ra l m ar su pi a N o 5 Si ng le , n o gr oo ve R ou nd ? ∼4 ∼3 (T w itc he ll, 1 91 5) D ur ha m el la K ie r, 19 68 D ur ha m el la fl or id an a (T w itc he ll, 1 91 5) D ur ha m el la o ca la na (C oo ke , 1 94 2) D ur ha m el la te tr ap or a n. sp . Pe nt ed iu m K ie r, 19 67 Te tr ad ie lla 3 Li ao & L in , 19 81 C hi na O va l, al m os t ro un d 5 A bs en t, sl ig ht ly co nv ex C lo se r t o am bi tu s t ha n pe ris to m e A da pi ca l m ar su pi a N o 4 Sc at te re d, no g ro ov e R ou nd ? > 1/ 2, re du ce d > 1/ 2? , re du ce d ? ? ? 1 S ta nd ar d po st al se rv ic e st at e ab br ev ia tio n, p lu s ” C B ” = C ar ib be an a nd M X = M ex ic o. 2 T es t l en gt h, a pp ro xi m at e up pe r s iz e lim it. 3 T oo li ttl e is k no w n of th is g en us to b e ce rta in it is a n eo la ga ni d. 4 M ea su re m en t i s b as ed o n th e di st an ce a cr os s o nl y ha lf th e pe ta l ( i.e ., di st an ce fr om p er ra di al to a br ad ia l s ut ur e) a t a p oi nt a bo ut 2 /3 th e le ng th o f t he p et al fr om th e ap ic al sy st em , a s d ep ic te d in F ig . 1 06 . 5 S ee F ig . 1 06 . 6 C ou nt ed in si ng le c ol um n of a nt er io r p ai re d am bu la cr a II o r I V, se e Fi g. 1 05 . 7 P os t-b as ic or on al s o nl y, c ou nt ed in si ng le c ol um n of a nt er io r p ai re d in te ra m bu la cr a 2 or 4 , s ee F ig . 1 05 . ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 161 Table 9: Total oral surface plate counts for typical adult specimens, including basicoronals, of all species of Neolaganum and Weisbordella except W. libum, n. sp. Plate system Neolaganum Weisbordella archerensis dalli durhami cubae johnsoni inglisensis Ambulacral 66 69 78 58 59 54 Interambulacral 51 52 50 42 46 41 Ambulacral range for genus 66 – 78 54 – 59 Interambulacral range for genus 50 – 52 41 – 46 discussed below. Cooke (1959: 52) asserted that N. dalli should have priority “because dalli has been correctly iden- tified and is a well-known species, whereas archeren- sis has been repeatedly confused with the species herein described as Neolaganum durhami”. Not only is this an inapplicable criterion for nomenclatural decisions, but Cooke did not mention that his own work (Cooke, 1942) was largely the only source of that confusion. We disagree that L. dalli (N. durhami as in- dicated by Cooke [1959]), should be recognized as the type species of Neolaganum. Durham (1954) clearly designated N. archerensis as the type of his new genus, and as we demonstrate, differentiating it from N. dalli is not difficult. Misidentifications after Cooke (1942) notwithstanding, systematists should be able to distinguish among N. archerensis, N. dalli, and N. durhami. To verify Cole and Ponton’s (1932) assertion of variability in periproct position in N. dalli, we sought out the lots they examined in the FM-IP Collections. In addition, we examined numerous specimens of N. dalli from in situ surface exposures of the Avon Park Formation at the Gulf Hammock Quarry (FM-IP LV004, FM-IP LV039) in Levy County. Specimens from this locality (Figs. 115, 117, 118) clearly do not display the great variation in periproct placement noted by Cole and Ponton (1932). The specimens are indistinguishable from the holotype of N. dalli (USNM 164667B) and consistently display a periproct situated further from the margin than that of N. archerensis. We measured the periproct position (distance of posterior edge of periproct from posterior margin of the test) in 109 specimens of N. dalli from the upper portion of the Avon Park Formation in the Gulf Hammock Quarry. The periproct averages 23.04% TL from the posterior margin (range 17.8-28.3%) (Figs. 114, 115). Additional specimens of N. dalli were ob- tained from the upper portion of the Avon Park For- mation by SCUBA divers in Blue Spring (VO001), Volusia County, Florida, and these specimens are likewise indistinguishable from either the holo- type of N. dalli or the specimens from Gulf Ham- mock, again not exhibiting the variation in periproct position suggested by Cole and Ponton (1932) (Fig. 117). At least in these two populations, mor- phology of N. dalli is consistent with characteristics of the holotype. Specimens with a more posterior periproct are found in certain well cuttings (all within the Avon Park Formation), such as lot UF 296335, from 117 m depth in a well at the Palmetto Phosphate Company near Tiger Bay, Polk County, Florida [site I-1601 of Cole and Ponton, (1932) and FM-IP PO050] and UF 56401 from 1680-1700 ft. depth in a well near Belle Meade, Collier County (FM-IP CR013). The specimens from Tiger Bay were identified as Laganum dalli by Cooke on August 30, 1941, and the lot of 193 specimens contains many that are inseparable from N. archerensis, as well as some more reminiscent of N. dalli. Specimens as large as the holotype of L. dalli (16 mm TL, 15 mm TW, 4.5 mm TH) possess periprocts in a position identical to that of N. archerensis. Therefore, a more posterior periproct cannot be attributed to ontogenetic effects, contrary to the assertions by Cole and Ponton (1932). This observation alone undermines the conclusions ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 162 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) of Cole and Ponton (1932), which were, in turn, the basis for Cooke’s (1959) subsequent decision to synonymize N. archerensis and N. dalli. In addition, a lot of 66 tests (UF 12892) from 15.2 m depth in a well near Lake Wales, Polk County (FM-IP 3573), also show a more posterior position of the periproct (Figs. 110-112, 115), very similar to that seen in the holotype of N. archerensis, but very unlike the situation in the holotype of N. dalli. In fact, this lot does not contain any specimens with more anterior periproct positions as seen in N. dalli, and all appear to be N. archerensis. We measured periproct placement of 34 tests from the Lake Wales well that had preserved oral surfaces (Fig. 113) and the periproct averages 13.52% TL from the posterior test margin. An additional lot of 28 specimens (UF 56401) from 512-518 m depth in a well near Belle Meade, Collier County (FM-IP CR013) show simi- larly placed periprocts, and again, all appear to be N. archerensis. Thirteen specimens from the Belle Meade material had preserved oral surfaces and the periproct of these specimens averaged 15.8% TL from the posterior margin. When plotted with the holotypes of both N. dalli and N. archerensis, a clear distinction of the two species is readily apparent (Fig. 113). Taking all the data into account, the periproct of N. archerensis averages 14.1% TL from the posterior margin, with a range of 9.5-20.9% TL, but the periproct of N. dalli averages 23.04% TL from the posterior margin, with a range of 17.8-28.3%. There is some overlap in these percentage values, but the bivariate plots show that when test size is considered, it is easy to distinguish these two forms based on periproct position alone. Our data indicate that variation in periproct position of N. dalli documented by Cole and Ponton (1932) does not exist in populations of N. dalli in the upper portion of the Avon Park Formation ex- posed in the Gulf Hammock Quarry, Levy County, and Blue Spring, Volusia County. Furthermore, the assertion of Cole and Ponton that the periproct is nearer the posterior margin only in small speci- mens of L. dalli is not evidence that L. dalli and L. archerensis are the same taxon. The holotype of L. archerensis (18 mm TL, 16 mm TW, 3.5 mm TH) is larger than the holotype of L. dalli (16 mm TL, 15 mm TW, 4.5 mm TH), yet its periproct is much closer to the margin than that of L. dalli (Fig. 107). That some of the well cuttings contain exam- ples of forms attributable to both N. archerensis and N. dalli (e.g., UF 56401, UF 296335) is explained by the fact that both species can occur together (as sur- mised by Clark and Twitchell, 1915), and possibly by mixing of specimens while fossils were collected from the well cuttings. The latter might be verified by more strict sampling of future or existing cores of Avon Park Formation strata. We therefore conclude that N. archerensis is distinct from N. dalli as distinguished its more pos- terior periproct and should remain the type species of Neolaganum as designated by Durham (1954). Emended Diagnosis.—Neolaganum with: thick test with almost parallel oral and aboral sur- faces, the latter slightly depressed just proximal to the ambitus; long, slit-like outer pores in the petals. Neolaganum dalli (Twitchell in Clark and Twitchell, 1915) (Figs. 105, 106, 112-117; Tables 8, 9) Laganum dalli Twitchell in Clark and Twitchell, 1915. p. 164. pl. 75, figs. 4a-d. Echinodiscus dalli (Twitchell). Lambert and Thiéry, 1925. p. 581. Laganum dalli (Twitchell). Cole and Ponton, 1932. p. 23. figs. 1a-12b. Peronella dalli (Twitchell). Cooke, 1942. p. 26. Peronella dalli (Twitchell). Cooke, 1948b. p. 91. pl. 22, figs. 5, 6. Peronella dalli (Twitchell). Fischer, 1951. p. 57. Neolaganum dalli (Twitchell). Cooke, 1959. pp. 51- 52. pl. 21, figs. 1-4. Neolaganum dalli(Twitchell). Mooi, 1989. fig. 20a. cf. Neolaganum dalli (Twitchell). Donovan, 1993. p. 389. fig. 12. Neolaganum dalli (Twitchell). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 5. Occurrence.—Neolaganum dalli is very abun- dant in a soft limestone bed of the Middle Eocene ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 163 Figure 112: Comparison of periproct placement in Neolaganum dalli and N. archerensis, including holotypes of each species. Vertical axis is distance, in millimeters, of posterior edge of periproct from posterior margin of test (ambitus). Horizontal axis is test length, in millimeters. N. dalli are from the Avon Park Formation in the Gulf Hammock Quarry in Levy County, Florida, and consists of 109 specimens from lots UF 114745, 136349, 156333, 336832, 336853, 336031, and 337030. N. archerensis are UF 12892 (50 ft. depth in a well near Lake Wales, Polk County, Florida) and UF 56401 from 1680-1700 ft. depth in a well near Belle Mead, Collier County, Florida. Avon Park Formation in a now inactive quarry east of Hwy 19/98 in Gulf Hammock (FM-IP LV004, FM-IP LV039), Levy County. Specimens of N. dalli have also been obtained from the Avon Park For- mation by SCUBA in Blue Spring (FM-IP VO001), Volusia County. Other than these two occurrences, N. dalli is documented only from the Avon Park For- mation in well cores (e.g., FM-IP PO050). The type locality is a well near Archer in Alachua County. Donovan (1993) documented N. dalli in the Eocene of Jamaica, and Cooke (1948b) noted its occurrence in the Eocene of Panama. Discussion.—Twitchell in Clark and Twitchell (1915), initially described this species as Laganum dalli from a well near Archer, Florida. Twitchell noted that it is most closely related to Laganum archerensis from the same well, which Twitchell described a few pages previously in the same work. Twitchell noted that L. archerensis is like L. dalli but is distinguished from it by the concave ring on its upper surface, narrower petals, less depressed and narrower poriferous zones, and a periproct that is closer to the posterior margin. We figure the holotypes of both species (Figs. 107, 114). As noted above, Cole and Ponton (1932) reviewed L. dalli through study of specimens obtained from wells within Florida. Cole and Ponton discussed variation, especially in periproct position. However, as discussed in the remarks for N. archerensis, we believe the specimens Cole and Ponton examined included both N. dalli and N. archerensis. Specimens of N. dalli from the upper portion of the Avon Park Formation in the Gulf Hammock Quarry (FM-IP LV004) and from Blue Springs in Volusia County (FM-IP VO001) show minimal variation in periproct po- sition and are consistent with the holotype of N. dalli. Specimens with more posteriorly placed periprocts (N. archerensis) are present only in deep well cores, as discussed above in the remarks for N. archerensis. Cooke (1942) reassigned Twitchell’s (in Clark and Twitchell, 1915) L. dalli to Peronella, and L. archerensis to Rumphia, neither of which are presently regarded as like any neolaganid. Durham (1954) subsequently erected Neolaganum and named N. archerensis the type species of his new genus. As noted above, Cooke (1959) considered N. dalli and N. archerensis to be synonymous (likely informed by the work of Cole and Ponton [1932], ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 164 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 113: Comparison of oral surfaces showing periproct position in Neolaganum archerensis (columns A-C; UF 12892 Avon Park Formation, 50 ft. depth in a well near Lake Wales, Polk County, Florida) and Neolaganum dalli (columns D-F; UF 136349 Avon Park Formation, Gulf Hammock Quarry, Levy County, Florida). Test length provided in millimeters from top to bottom of each column: Column A: 8.9, 9.8, 10.1, 10.5, 10.8, 11.4, 11.6, 12.1. Column B: 12.6, 12.8, 12.9, 13.1, 13.5, 14.3, 14.8, 14.8. Column C: 14.9, 15.2, 15.7, 16.9, 17.1, 17.8, 18.4, 18.7. Column D: 9.2, 10.5, 10.7, 10.8, 11.0, 11.3, 11.7, 11.7. Column E: 12.1, 12.2, 12.4, 12.7, 13.5, 13.8, 13.9, 14.0. Column F: 14.1, 14.7, 14.8, 15.2, 15.6, 17.0, 18.6, 20.1. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 165 Figure 114: Laganum dalli (= Neolaganum dalli) holotype (USNM 164667B), 15.9 mm TL, 15 mm TL, 4.6 mm TH, Middle Eocene Avon Park Formation, likely from a well, near Archer, Alachua County, Florida (images from Smithsonian NMNH online database). A: aboral. B: oral. all of which they erroneously identified as N. dalli). Cooke (1959) asserted, that the species name dalli is preferred to archerensis because dalli had been correctly identified and was a well-known species, whereas archerensis has been confused with Ne- olaganum durhami (Cooke, 1942; Durham 1954, 1955). We disagree with Cooke (1942), and recog- nize both species as distinct. Past confusion is not a criterion for enshrining Cooke’s taxonomic error, and in any case, we do not find the “confusion” to be sufficient to suppress N. archerensis. Further- more, we follow the recent stipulation by the IZCN, which states, “... if an author discovers that the type species fixation of a genus-group taxon was based on a misidentification of the type species, the author may, in the interests of stability and without making application to the Commission, fix as type species either the taxonomic species actually involved or the misidentified nominal species fixed previously.” In accordance, we follow Durham’s (1954) desig- nation of Laganum archerensis Twitchell in Clark and Twitchell, 1915 as the type species of the genus- group taxon, Neolaganum Durham, 1954. Neolaganum dalli is characteristic of the Mid- dle Eocene Avon Park Formation and undocumented outside of that unit within the United States. Neola- ganum dalli is also the only echinoid known from surface exposures of the Avon Park Formation (Fis- cher, 1951), which is restricted in outcrop to Levy County. However, in the Gulf Hammock Quarry (FM-IP LV004, FM-IP LV039), spoil often contains a mixture of both the Avon Park and lower OLS, so a collection from the locality can create confusion by containing specimens from both units. The swollen, subpentagonal test of N. dalli is very consistent in shape throughout growth, and this readily differentiates it from the thinner, often much larger test of N. durhami, which occurs in the lower OLS, immediately overlying the occurrence of N. dalli in the Avon Park Formation. The presence of four gonopores in Neolaganum readily distinguishes it from members of Durhamella, which have five, except for D. tetrapora n. sp., which differs in having a much thinner, very small test and, in presumed females, depressions in the aboral surface just prox- imal to the ambitus. As stated above, N. dalli can readily be dis- tinguished from N. archerensis, which also occurs ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 166 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 115: Neolaganum dalli (UF 337953), 16.5 mm TL, 15.5 mm TW, 5.5 mm TH, Middle Eocene Avon Park Formation, Levy County, Florida (FM-IP LV004). A: aboral. B: oral. C: left side. D: right side. in the Avon Park Formation (but only from well cuttings), because of the more posterior position of the periproct in N. archerensis (see treatment of this species, above). Neolaganum archerensis also tends to have a few more pore pairs in the petals for any given size. The oral plate arrangements of N. archerensis, N. dalli, and N. durhami (Tables 7, 8) demonstrate that the first post-basicoronals in inter- ambulacrum 5 (plates 5.b.2 and 5.a.2) of N. dalli are significantly shorter than in either N. archerensis or N. durhami. Nevertheless, even though the periproct of N. dalli is much further from the posterior margin than in its congeners, the periproct in all three species is first in contact with plates 5.b.2 and 5.a.2. Emended Diagnosis.—Neolaganum with: thick test with almost parallel oral and aboral sur- faces, the latter slightly depressed just proximal to the ambitus; only slightly elongated outer pores in the petals; no demiplates in the petals; periproct closer to peristome than the ambitus. Neolaganum durhami Cooke, 1959 (Figs. 105, 106, 118; Tables 8, 9) Rumphia archerensis (Twitchell). Cooke, 1942. p. 27 (in part), pl. 2, figs. 11-13. Peronella archerensis (Twitchell). Fischer, 1951. p. 58. pl. 2, fig. 3; txt figs. 2, 3. Neolaganum archerensis (Twitchell). Durham, 1954. txt fig. 28 (in part, figure is N. durhami). Neolaganum archerensis (Twitchell). Durham, 1955. txt figs. 15f, 30b (in part, figures are N. durhami). Neolaganum durhami Cooke, 1959. p. 52. pl. 21. figs. 5-7. Neolaganum archerensis (Twitchell). Durham, 1966. p. 475. fig. 365, 1a-1d. Neolaganum durhami (Cooke). Kier, 1970. txt. fig. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 167 Figure 116: Neolaganum dalli (UF 337954), 14.5 mm TL, 13.5 mm TW, 5 mm TH, Middle Eocene Avon Park Formation, Levy County, Florida (FM-IP LV004). A: aboral. B: oral. C: left side. D: right side. 1F. Neolaganum durhami (Cooke). Toulmin, 1977. p. 343. pl. 67, figs. 1, 2. Neolaganum durhami (Cooke). Mooi, 1989. fig. 18b. (also 9b and 16c as N. archerensis). Neolaganum durhami (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—This species is abundant in the Oligopygus phelani Zone of the lower OLS at numer- ous localities in Levy County, including the Cross Florida Barge Canal west of Hwy 19/98 south of Inglis (FM-IP CI001), and along the Withlacoochee River near its mouth (FM-IP LV024), west of Yan- keetown. The type locality is along the Suwannee River south of Sulphur Spring, 24 km south of Ellaville, Florida. Discussion.—The taxonomy of this species is entangled with the complicated nomenclatural issues associated with the other two species, N. archerensis and N. dalli. Cooke (1942) initially discussed specimens of what would eventually be assigned to this species under the name Rumphia archerensis, believing the specimens he was exam- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 168 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 117: Neolaganum dalli (UF 46461), 13.1 mm TL, 12.6 mm TW, 3.2 mm TH, Middle Eocene Avon Park Formation, collected by divers in Blue Springs, Volusia County, Florida (FM-IP VO001). A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: posterior. E: apical area, note very pronounced hydropore groove. F: peristome and periproct. G: petaloid area, note small pores in ambulacrum outside petaloid portion of ambulacrum III. This is what Cooke (1942) described as: punctate extrapetalous ambulacral regions, also seen in the holotype of L. dalli. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 169 Figure 118: Neolaganum durhami (UF 337984), 33.5 mm TL, 31 mm TW, 5.5 mm TH, Oligopygus phelani zone of Upper Eocene lower Ocala Limestone, Citrus County, Florida (FM-IP CI001). ining were synonymous with Twitchell’s (in Clark and Twitchell, 1915) L. archerensis. However, the localities Cooke (1942) provided for his specimens of R. archerensis, other than the type locality in a well near Archer, Florida, are largely localities in Levy County where N. durhami is found in the Oligopygus phelani Zone of the lower OLS. The specimen Cooke (1942) figured is from an unknown depth in a well in Polk County, Florida, and this specimen (USNM 498994) has been established to be what would subsequently become recognized as N. durhami, not N. dalli. Fischer (1951), following Cooke (1942), dis- cussed the occurrence of N. durhami (as Peronella archerensis) in the Oligopygus phelani Zone in Citrus and Levy Counties, Florida and noted that the type of the species illustrated by Twitchell in Clark and Twitchell (1915) is an unusually flattened specimen. Fischer (1951) was unaware that he was comparing specimens of a different species with Twitchell’s form (= N. archerensis), but Fischer’s figured and referred specimens of P. archerensis are also recognizable as N. durhami. Durham (1954) erected Neolaganum and de- scribed N. archerensis as the type species of the genus. However, Cooke (1959) asserted that Durham (1954) mistakenly based the genus on misidentified specimens that he insisted were his N. durhami. Al- though Cooke (1959) didn’t provide a rationale for his assertion that Durham misidentified N. durhami as N. archerensis, it appears he was correct. Durham (1954) considered UCMP 33296 a hypotype of N. archerensis. Durham’s figures of N. archerensis in Durham (1954, 1955, 1966) are of this specimen, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 170 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 119: Holotype of Laganum ocalanum (= Durhamella ocalana) (USNM 372873), 22.3 mm TL, 19.5 mm TW, 3.0 mm TH, Upper Eocene upper Ocala Limestone, two miles NE of Sumterville, Sumter County, Florida. A: aboral. B: oral. and the specimen is clearly N. durhami. In addition, the locality for this specimen is the mouth of the Withlacoochee River west of Yankeetown (FM-IP LV024), which has exposures of the lower OLS containing N. durhami, but no trace of exposures of the Avon Park Formation that would contain N. archerensis, which has also never been identified in surface exposures. We conclude that Durham (1954) based Neolaganum on a specimen (UCMP 33296) he called N. archerensis but was actually N. durhami. Cooke (1959) later described and named this form as N. durhami and included the previous designations in the synonymy of his new species. Nevertheless, as stated above in reference to the ICZN’s recommendation concerning misidentified types of a species group, N. archerensis is here maintained as the type species of Neolaganum. Neolaganum durhami typically attains a size about twice that of N. dalli, is proportionately thin- ner, and has denser tuberculation (Cooke, 1959). These two species are often found together in mixed spoil of the lowest OLS and underlying Avon Park Formation in the Gulf Hammock Quarry (FM- IP LV004). There remains potential to confuse N. durhami with members of Weisbordella in mixed spoil of the OLS, but its flattened test, five gono- pores, and flat oral surface distinguish N. durhami from both species of Weisbordella known to occur in the region. Neolaganum durhami is more likely to be confused with Durhamella floridana (Twitchell in Clark and Twitchell, 1915), from which it can be readily differentiated because N. durhami has four gonopores instead of five. Neolaganum durhami has much narrower petals than the lanceolate petals of W. inglisensis n. sp., with which it occurs within the type area near Inglis, and again, W. inglisensis n. sp. has only four gonopores. Neolaganum durhami is much smaller than the very elongate W. eldridgei, which also has a very thin margin and only four gonopores that further distinguish the two species. In addition, these two species are not typically found together. Wythella eldridgei occurs in the Wythella eldridgei Zone in the uppermost OLS and N. durhami appears to be restricted to the Oligopygus phelani Zone near the base of the OLS. Fischer (1951) noted that N. durhami (as Per- onella archerensis) has considerable variation in ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 171 Figure 120: Durhamella ocalana (UF 337975), 27.5 mm TL, 25 mm TW, 3 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. outline and thickness of the margin. The margin of this species is usually thick, but a collection of specimens will undoubtedly contain some individ- uals with relatively thin margins. In addition, the ambitus typically has truncations in the posterior ambulacra that are less expressed anteriorly, and this gives the test what Cooke (1959) described as a subdecagonal outline. More rarely, specimens can have a nearly oval or even nearly circular outline. The five gonopores and relatively large number of oral plates (Tables 8, 9) distinguish it from other neolaganids with a similar overall test shape. Though Fischer (1951: table 1) showed N. durhami (as Peronella archerensis) to occur through- out the OLS, we have only found N. durhami in the Oligopygus phelani Zone, where it is commonly found with D. ocalana, E. mooreanus, P. floridanus, and other, rarer species. Neolaganum durhami is not otherwise documented from the upper OLS, and a search of the FM-IP collections revealed no speci- mens of N. durhami from the upper OLS. Therefore, we do not recognize N. durhami above the Oligopy- gus phelani Zone of the OLS in Florida. Emended Diagnosis.—Neolaganum with: low test; very elongate, slit-like outer pores in the petals; demiplate every two or three plates in the petals; periproct closer to peristome than the ambitus; large number of plates (usually about 78 in total) in the oral ambulacra. Genus Durhamella Kier, 1968 Kier (1968) erected Durhamella in honor of J. Wyatt Durham, and designated D. ocalanum as the type species, noting the occurrence of what he called “pseudocompound plates” in the petals. Kier ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 172 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 121: Durhamella ocalana (UF 337976), 23.5 mm TL, 21 mm TW, 4 mm TH, Oligopygus haldemani Zone Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. stated that the presence of these plates, coupled with basicoronal plates arranged in a pentagon, indicate Durhamella should be placed in the Neolaganidae. He did not provide a diagnosis for the genus but did state that it is distinguished from other neolaganid genera by the presence of 5 gonopores, fewer pseu- docompound plates in the petals, and the outer pore of the petals not being in a pronounced slit. Like Durhamella, Weisbordella has a reduced number of oral plates, but differs in that Durhamella often has depressions along the sutures in the margins of the aboral regions. In addition, Weisbordella has very strongly expressed demiplates in the petals that are very poorly expressed or completely absent in Durhamella (Fig. 106). The latter also lacks the long, sinuous, or complexly branched hydropore groove of Weisbordella. Emended diagnosis.—With the inclusion of Durhamella tetrapora n. sp., Durhamella is now un- derstood to include taxa with both four and five gono- pores. Durhamella is best characterized by a com- bination of: depressions in the aboral submarginal region; short, unbranched, hydropore grooves; flat oral surface made up of a reduced number of plates, notably in the interambulacra; very elongate inter- ambulacral first post-basicoronals; and round outer pores in the petal pore pairs. Durhamella ocalana (Cooke, 1942) (Figs. 105, 106, 119-123; Table 8) Laganum ocalanum Cooke, 1942. p. 23. pl. 2, figs. 7-10. Laganum ocalanum (Cooke). Fischer, 1951. p. 57. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 173 Figure 122: Durhamella ocalana (UF 337985), 11 mm TL, 10 mm TW, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: left side. D: right side. Laganum ocalanum (Cooke). Cooke, 1959. p. 51. pl. 20, figs. 11-15. Durhamella ocalana (Cooke). Kier, 1968. pl. 6, figs. 1-5; pl. 7, figs. 1-3; pl. 8, figs. 2, 3. figs. 28-30, 32-34. Laganum ocalanum (Cooke). Toulmin, 1977. p. 342. pl. 64, fig. 6, 7. Durhamella ocalana (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—Present in both the Oligopygus phelani and Oligopygus haldemani Zones of the OLS of Florida, and absent only from the uppermost portion of the unit. Durhamella ocalana is sporadi- cally common in the Oligopygus haldemani Zone of the OLS, especially in quarries near St. Catherine and Center Hill in Sumter County (FM-IP SM010), and northwest of Mayo (FM-IP LF001) in Lafayette County, as well as the type locality of the species in a pit 3.2 km NE of Sumterville. It is more abundant in the Oligopygus phelani Zone in the lower OLS, especially along the banks of the Withlacoochee River, Levy County (FM- IP LV024) and Cross Florida Barge Canal, Citrus County (FM-IP CI001). Discussion.—Cooke (1942) initially de- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 174 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 123: Durhamella ocalana (UF 337986), 7.5 mm TL, 6.5 mm TW, 2.0 mm TH, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, Levy County, Florida (FM-IP LV024). scribed this species as Laganum ocalanum from the OLS at numerous localities in Florida. Kier (1968) named D. ocalanum the type species of his new genus Durhamella and emended the species name from ocalanum to ocalana. In addition, with over 100 tests available, Kier (1968) redescribed the species in much greater detail than previously. Durhamella ocalana appears to be restricted to the OLS of Florida and in its adult stages, is a very distinct species that cannot be confused with any other members of the echinoid fauna of the region except for its congener, D. floridana. Durhamella ocalana is perhaps best characterized by the pres- ence of depressed sutures in the submarginal area on the aboral surface, giving the appearance that the plates surrounded by these sutures are tumid. The depressions, which are absent or only slightly dis- cernable in D. floridana, are present in D. ocalana in both the adapical interambulacral plates and the am- bulacral plates beyond the petals. These two species of Durhamella are otherwise very similar, having similar petals, length to width, length to height ra- tios, similar peristomes, and similar adoral plate arrangement (Kier, 1968). Although, gonopores are found in the smallest identifiable specimens (3.5 mm TL) of D. ocalana, depressed sutures in the periphery of the aboral surface are only consistently discernable in specimens greater than 5 mm TL. Therefore, distinguishing small (¡5 mm TL) speci- mens of D. floridana and D. ocalana is done with ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 175 Figure 124: Holotype of Laganum floridanum (= Durhamella floridana), (USNM 137884), 19.6 mm TL, 17.7 mm TW, 4.2 mm TH, Upper Eocene lower Ocala Limestone, Johnson’s Sink, Levy County, Florida. A: aboral. B: oral. difficulty. Durhamella ocalana is readily distinguished from D. tetrapora n. sp. by the presence of five gonopores and depressed plate sutures near the margins of the aboral surface. As discussed in the remarks for D. floridana, D. ocalana and D. floridana are often found in associ- ation in both the Oligopygus phelani and Oligopygus haldemani Zones of the OLS of Florida. Specimens of D. ocalana and D. floridana rarely exceed 10 mm in diameter in the Oligopygus phelani Zone where it occurs in abundance with D. floridana, D. tetrapora n. sp., E. mooreanus, N. durhami, W. inglisensis n. sp., and P. floridanus. In the O. haldemani Zone, D. ocalana and D. floridana achieve greater sizes, exceeding 35 mm in TL. See the remarks for D. flori- dana for additional comments on the association of these two taxa and possible sexual dimorphism. Emended Diagnosis.—The only Durhamella with a combination of: five gonopores; strongly expressed depressions along the sutures in the pe- ripheral region of the aboral surface; poriferous zones that are much less than half the width of the poriferous zone as measured from the perradial to abradial suture about two thirds of the length of the petal. Durhamella floridana (Twitchell in Clark and Twitchell, 1915) (Figs. 105, 106, 124-128; Table 8) Laganum floridanum Twitchell in Clark and Twitchell, 1915. p. 163. pl. 75, figs. 3a-d. Echinodiscus floridanus (Twitchell). Lambert and Thiéry, 1925. p. 581. Laganum floridanum (Twitchell). Cooke, and Mos- som, 1929. pl. 3, figs. 5a-b. Laganum floridanum (Twitchell). Cooke, 1942. p. 23. Laganum floridanum (Twitchell). Cooke, 1945. fig. 5, no. 5. Laganum floridanum (Twitchell). Cooke, 1959. pp. 50-51. pl. 20, figs. 8-10. Durhamella cf. D. floridana (Twitchell). Kier, 1968. pp. 29-38. pls. 8.1, 9.1-5, 10.1-3. figs. 27, 31, 34-43. Laganum floridanum (Twitchell). Toulmin, 1977. pp. 341-342. pl. 67, fig. 3. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 176 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 125: Durhamella floridana (UF 338007), 20 mm TL, 17.5 mm TW, 3 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. Durhamella floridanum (Twitchell). Mooi, 1989. fig. 17b. Durhamella floridana (Twitchell). Osborn et al., 2016. tbl. 2. Occurrence.—D. floridana is present in both the lower and upper divisions of the OLS in Florida. It occurs in the Oligopygus haldemani Zone, espe- cially northwest of Mayo (FM-IP LF001), Lafayette County. It is also found in the Oligopygus phe- lani Zone of the lower OLS, occurring along the banks of the Withlacoochee River, Levy County (FM-IP LV024) and Cross Florida Barge Canal, Cit- rus County (FM-IP CI001). The type locality for the species is Johnson’s Sink, (FM-IP 3142), 4 mi. northeast of Williston, Levy County. Kier (1968) documented the occurrence of D. floridana in the Middle Eocene Lake City Formation (now Avon Park Formation) in the subsurface of Georgia. Discussion.—Twitchell in Clark and Twitchell (1915), initially described this species as Laganum floridanum from Johnson’s Sink, Levy County, in strata that were at the time considered to ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 177 Figure 126: Durhamella floridana (UF 337983), 23 mm TL, 19.5 mm TW, 3.5 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. be early Oligocene. The stratum at the type locality is now recognized as the Upper Eocene OLS. Twitchell in Clark and Twitchell (1915) initially described this species as having four gonopores, but Cooke (1959) corrected the description to clarify that the species has five gonopores. Kier (1968) subsequently altered the species designation from floridanum to floridana. Specimens from the type locality, which has since been developed and is no longer accessible, are figured herein (Figs. 127, 128). Durhamella floridana is usually distinguished from D. ocalana by its lack of the characteristic depressed sutures of interambulacral plates on the aboral surface. These depressed sutures give the submarginal area of the aboral surface of D. ocalana a “pillowy” appearance. However, it should that noted that weakly depressed sutures may be present in D. floridana, as shown in Kier (1968: pl. 9, fig. 1). Therefore, distinguishing between D. ocalana and D. floridana can be troublesome unless one is able to examine the petal plate architecture (Fig. 106). The presence of five gonopores readily distin- guishes D. floridana from N. durhami, W. inglisensis n. sp., and D. tetrapora n. sp., with which it occurs in the lower OLS. Durhamella floridana occurs with W. cubae in the Oligopygus haldemani Zone of the OLS, from which it is easily distinguished by D. floridana’s flat oral surface (the oral surface of W. cubae is distinctly concave), its much thinner test, and the presence of five gonopores. It cannot be confused with any other element of the fauna of the OLS, within which it appears to be restricted ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 178 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 127: Figure 129: Durhamella floridana (UF 345211), 14.7 mm TL, 13.4 mm TW, 3.1 mm TH, Upper Eocene Ocala Limestone from the type locality of the species at Johnson’s Sink, Levy County, Florida (FM-IP 3142). A: aboral. B: oral. C: posterior. D: left side. E: anterior. F: tilted aboral viewpoint from posterior. G: tilted aboral viewpoint from anterior. H: oblique oral viewpoint from posterior. I: periproct. J: apical area. K: peristomal area showing oral plating. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 179 Figure 128: Durhamella floridana (UF 344747), 11.1 mm TL, 10.7 mm TW, 2.4 mm TH, Upper Eocene Ocala Limestone from the type locality of the species at Johnson’s Sink, Levy County, Florida (FM-IP 3142). A: aboral. B: oral. C: right side. D: left side. other than in the isolated Lake City Formation (now Avon Park Formation) well occurrence in the Glynn County, Georgia (Kier, 1968). As discussed further in the remarks for D. ocalana, it is curious that D. floridana and D. ocalana are usually found together in the OLS, both in the Oligopygus haldemani Zone, where both species are usually larger (Figs. 120, 121), and in the Oligopygus phelani Zone, where both species are typically much smaller. In addition, the gonopores of D. ocalana are usually, though not always, larger and it is possible that the two species are morphs of the same species: D. floridana might represent the males, and D. ocalana the females, in which the depressed sutures of the aboral surface of D. ocalana facilitate brooding. Such sutures seem to cause a reduction in interambulacra, which in turn could provide more space for the developing young. What spines remain adjacent to the depressions could form a protective screen over the juveniles. However, the fact that D. ocalana is typically much more abundant than D. floridana at all known exposures casts doubt on this hypothesis, as does the variation in gonopore size of D. ocalana, even though it does tend to have larger gonopores. The slightly depressed sutures might also conflict with the interpretation that D. floridana is the male of D. ocalana, except that as discussed below, Kier (1967) noted that males of Pentedium curator can express shallow pits in the anterior paired interambulacra ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 180 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) adjacent to the apical system. This illustrates a tendency for males of sexually dimorphic laganids to show some of the secondary sexual features expressed to a much greater degree in the females. Emended Diagnosis.—The only Durhamella with: five gonopores; absence of strong expression of depressions along the sutures in the peripheral region of the aboral surface (if these are present, they are very shallow and nearly indistinguishable); poriferous zones that are much less than half the width of the poriferous zone as measured from the perradial to abradial suture about two thirds of the length of the petal. Durhamella tetrapora n. sp. (Figs. 105, 106, 129-132; Table 8) Diagnosis.—The only Durhamella with: four rather than five gonopores; very small test with possible sexual dimorphism. Description.— Based on three specimens; UF 343902, interpreted as female, two specimens interpreted as male (UF 343903, UF 343904). Small, from 5.4 mm TL to 6.4 mm TL, 4.7 to 6.0 mm TW, slightly subpentagonal, posterior margin truncated, anterior gently rounded, greatest width through peristome; TW averages 93.6% TL in males, 88.4% in female. Test low, TH 24.6 – 26.5% TL, greatest height at apical system. Margin thick, slightly higher anteriorly. Aboral surface depressed between margin and mid-length of petals (sutures of aboral inter- ambulacral plates not depressed as in D. ocalana); depression creates shallow indentations marginally adjacent to ambulacra, strongly developed in UF 343902 (female), interpreted as brooding pouches; less depressed ridges radiate from apical area in center of all five interambulacra to margin. Oral surface flat, only slightly depressed near peristome. Apical system slightly anterior; anterior edge on average 36.9% TL from anterior ambitus, posterior edge on average 39.9% TL from posterior ambitus. Four gonopores well-developed in all three specimens; sexual dimorphism very likely. Gono- pores vary in position and size. UF 343902 (female) has large gonopores (avg. 2.3% TW), male gono- pores much smaller (avg. 1.4% TW). Gonopores of female more widely separated (anterior pair 14.8% TW apart, posterior pair 23.8 % TW apart), posi- tioned on edge of apical system. Gonopores of males well within apical area, closer together (anterior pair on average 5.4% TW apart, posterior pair on average 9.3% TW apart); males show that separation of posterior pair of gonopores is variable. Shallow, short, unbranched hydropore groove present. Petals flush, wide apically, nearly closed dis- tally. In smaller specimen (UF 343902: female) petal I and V longest, on average 18.4% TL, petal II and IV shortest, on average 14.5% TL, petal III 16.6% TL. In larger specimens (males) petal III longest, on average 22.2%, petals II and IV shortest, on average 19.3% TL; petals I and V on average 21.0 % TL. Pores round, outer pore not elongate. UF 343902 has one more pore pair per series in petal I and V than other petals, both male specimens have on average one less pore pair per series in petals II and IV than the others. Occluded plates (secondaries that extend from the perradial suture but do not touch an interambulacrum) present distally in most petals (Fig. 132), especially UF 343904 which has one occluded plate in petals I, III, and IV; two in petal II, and three in petal V. Pores more numerous in oral ambulacra; concentrated on adradial sutures; buccal pores present at edge of peristome. Basicoronal plates form subpentagonal out- line with single plate in each interambulacrum, paired plates in each ambulacrum; first post- basicoronal interambulacral plates considerably longer than first post-basicoronal ambulacral plates, extending past second post-basicoronal ambulacral plates. Number of oral interambulacral plates reduced in comparison with other neolaganids ex- cept in Weisbordella and Durhamella (Fig. 105): UF 343904 has six post-basicoronal plates per oral inter- ambulacrum; UF343902 has five post-basicoronal plates in interambulacrum 1-4 and 6 in interambu- lacrum 5. Peristome subpentagonal, anterior blunt, pos- terior pointed; nearly central (posterior edge on average 44.4 % TL from posterior margin, anterior edge on average 43.4% TL from anterior margin), peristome length on average 12.6% TL, slightly ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 181 Figure 129: Durhamella tetrapora n. sp., holotype, (UF 343902), 5.4 mm TL, 4.7 mm TW, 1.3 mm TH, a female, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, Levy County, Florida (FM-IP LV016). A: aboral. B: oral. C: posterior. D: left side. E: tilted oral viewpoint from posterior. F: petaloid area. G: periproct. H: peristome. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 182 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 130: Durhamella tetrapora n. sp., paratype, (UF 343903), 6.4 mm TL, 6.0 mm TW, 1.7 mm TH, a male, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, pit north of Hwy 40, west of Hwy 19/98 in Inglis, Levy County, Florida (FM-IP LV016). A: aboral. B: oral. C: periproct. D: oblique oral viewpoint. E: posterior. F: tilted aboral viewpoint from posterior. G: oblique aboral viewpoint from anterior. H: apical area and petaloid portion of ambulacrum III. I: left side. J: petaloid area. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 183 Figure 131: Durhamella tetrapora n. sp., paratype, (UF 343904), 6.3 mm TL, 5.9 mm TW, 1.6 mm TH, a male, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, pit north of Hwy 40, west of Hwy 19/98 in Inglis, Levy County, Florida (FM-IP LV016). A: aboral. B: oral. C: apical area. D: oblique aboral viewpoint from posterior. E: anterior. F: left side. G: peristome and periproct with some oral plating exposed. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 184 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 132: Durhamella tetrapora n. sp., comparison of aboral plate arrangement: Left: holotype, (UF 343902), a female. Right: paratype, (UF 343904), a male. Ambulacrum III is towards the top, the interambulacral plates are shaded, and pores are solid black. longer than wide: peristome width on average 87% length. Periproct ovate, wider than long (height from 74.6 to 83.0% width, average 78.4% width), first in contact with plates 5.a.2 and 5.b.2 (Fig. 105), near the margin: on average 5.8% TL from posterior margin. Interior structures such as peripheral supports, pillars and lantern not recorded. Zoobank Nomenclatural Act.—6C5CC4EA- 6074-41B1-ADAE-03629917B860 Discussion.—Muriel Hunter collected the holotype (UF 343902) and two paratypes (UF 343903, UF 343904) of D. tetrapora n. sp. west of Inglis in Levy County (FM-IP LV016). The spec- imens are small, from 5.4 to 6.4 mm TL, but likely mature or nearly so, as evidenced by their fully de- veloped gonopores. This species is very similar to the other known species of Durhamella; D. ocalana and D. floridana. However, it is readily differentiated by the presence of four gonopores, instead of the five seen in its two congeners. Kier (1968) stated the presence of five gono- pores, fewer pseudocompound plates in the petals, and the outer petaloid pore not being in a pronounced slit distinguishes Durhamella from the other genera of Neolaganidae: as noted in the description, D. tetra- pora n. sp., shares all these characters. Other than the presence of four rather than five gonopores, the new species shares all characteristics of Durhamella, so we place this taxon within Durhamella rather than erect yet another new genus. As discussed in the remarks for Durhamella, the genus was previously understood only to include taxa with five gonopores. Therefore, assigning D. tetrapora n. sp., which has four gonopores, to Durhamella required analysis of additional characteristics that diagnose the genus itself. Durhamella is further characterized by a mar- gin that is usually slightly thickened, but with a depressed aboral submarginal region, and D. tetra- pora n. sp., also displays this feature. Furthermore, analysis of oral plate structure of D. floridana, D. ocalana, and D. tetrapora n. sp. (Fig. 105), shows they share very similar oral plate arrangement, with a reduced number of plates and greatly elongated first post-basicoronals. Two of the three specimens of D. tetrapora n. sp. (UF 343903, UF 343904) has gonopores situated well within the apical system and are likely males. However, UF 343902 has much larger gonopores ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 185 Figure 133: Weisbordella cubae (UF 337960), 29 mm TL, 25 mm TW, 6 mm TH, Oligopygus haldemani Zone of Upper Eocene Ocala Limestone, Sumter County, Florida (FM-IP SM010). A: aboral. B: oral. C: left side. D: right side. that are more widely separated and positioned just outside of the apical area, and we interpret this spec- imen to be a female. The gonopore arrangement is very similar to the sexual dimorphism seen in Pente- dium curator Kier, 1967. However, as noted by Kier (1967), the males of P. curator do not have adapical depressions except for one specimen that has two slight pits in the anterior paired interambulacra ad- jacent to the apical system, suggesting a tendency for males of some neolaganid taxa to show some of the marsupium-like features seen in the females. Kier (1967) noted that males of P. curator did not grow as large as the females, whereas the female D. tetrapora n. sp., is 5.4 mm TL, and smaller than the two males which are 6.3 mm and 6.4 mm TL. Etymology.—The species name highlights the four gonopores, unique to Durhamella, seen in each of the known specimens. Materials and Occurrence.—Holotype (UF 343902) and two paratypes (UF 343903, UF 343904) from the Oligopygus phelani Zone of the Upper Eocene OLS, pit on north side of Rt. 40, west of Inglis, Levy County, Florida (FM-IP LV016). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 186 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 134: Weisbordella cubae (UF 337971), 13.5 mm TL, 12 mm TW, 3 mm TH, Oligopygus wetherbyi Zone of Upper Eocene Ocala Limestone, Dixie County, Florida (FM-IP DI001). A: aboral. B: oral. C: left side. D: right side. Genus Weisbordella Durham, 1954 Durham (1954) designated Peronella caribbeana Weisbord, 1934 as the type species of Weisbordella, stating that Peronella quinquen- odulata Weisbord, 1934; P. caribbeana; Peronella cubae Weisbord, 1934; L. dalli, and Peronella mirabilis Jackson, 1922 also likely belong in the genus. We agree with Cooke (1959) and recognize L. dalli in Neolaganum. Cooke (1942, 1959) also considered P. quinquenodulata and P. caribbeana to be subjective junior synonyms of P. cubae. Durham (1954) did not recognize Laganum johnsoni Twitchell in Clark and Twitchell, 1915 as a Weisbordella, leaving it unassigned to genus because he was unable to examine any specimens of the species. Cooke (1959) subsequently placed L. johnsoni in Weisbordella and we concur. Therefore, our concept of Weisbordella follows previous treat- ments in containing W. cubae and W. johnsoni from our regional faunas, further adding W. inglisensis n. sp. and W. libum n. sp. to the genus. The status of P. mirabilis, from Trinidad, is uncertain but in any case, this species falls outside the scope of the present work. Durham (1954) did not publish a viable diag- nosis for Weisbordella (or any of his new neolaganid genera) as there are no features in his descriptions that we cannot also find in other genera. He dis- tinguished Weisbordella from Neolaganum by the lack of a hydropore groove, but well-developed hy- dropore grooves are clearly evident in both W. cubae ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 187 Figure 135: Weisbordella cubae (UF 337972), 12 mm TL, 10 mm TW, 2.5 mm TH, Oligopygus wetherbyi Zone of Upper Eocene Ocala Limestone, Dixie County, Florida (FM-IP DI001). A: aboral. B: oral. C: left side. D: right side. and W. johnsoni. Smith and Kroh (2011) acknowledged the presence of hydropore grooves in Weisbordella, but stated that the slightly sunken oral surface, short, strongly lanceolate petals, and the sparsity of tu- bercles surrounding the peristome in Weisbordella are more significant in distinguishing it from Ne- olaganum. The relatively dense concentration of tubercles around the peristome used by Smith and Kroh (2011) must be based on examination of N. durhami, because in N. archerensis, the type species of the genus, there are no more tubercles in the peristomial area than is typical of W. cubae, and perhaps even less than is typical for W. johnsoni. This feature is not considered reliable to distinguish the two genera. Cooke (1959) stated that Weisbordella can be distinguished from Neolaganum by its concave oral surface and absence of food grooves. However, we are unaware of food grooves in any neolaganids. Although some specimens of W. johnsoni have al- most planar oral surfaces, a concavity is usually expressed to some degree. The concavity is strongly expressed in W. cubae, so the concave oral surface, as variable as it is, is a reliable feature of Weis- bordella. However, a concave oral surface is not ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 188 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 136: Weisbordella cubae (UF 337973), 10.5 mm TL, 9 mm TW, 2.5 mm TH, Oligopygus wetherbyi Zone of Upper Eocene Ocala Limestone, Dixie County, Florida (FM-IP DI001). A: aboral. B: oral. C: left side. D: right side. unique to this genus among laganiforms, and in addition, we herein describe, based on other char- acters, W. inglisensis n. sp., which has a planar oral surface. Durham (1954: 681) seemed to question the validity of his new genera Weisbordella and Neo- laganum when he stated: “In the present studies the groove in the madreporite seems to be a more significant difference. If these differences should be found to be of lesser value, Neolaganum should have priority over Weisbordella.” Weisbordella is more reliably distinguished from Neolaganum by the fact that it has only four gonopores, and reduced number of post-basicoronal plates (Fig. 105; Tables 8, 9). The oral plate ar- rangement of Weisbordella is much more like that of Durhamella. However, Weisbordella expresses many more demiplates in the petals, has four gono- pores instead of the five seen in all Durhamella except D. tetrapora n. sp., and lacks the depressed sutures at the aboral margins of Durhamella. Emended Diagnosis.—Neolaganids that com- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 189 Figure 137: Weisbordella johnsoni (UF 329697), 44 mm TL, 41 mm TW, 8 mm TH, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA086). A: aboral. B: oral. C: left side. D: right side. bine the presence of: four gonopores; a long, sinuous hydropore groove; usually concave oral surface with between 54 and 59 ambulacral oral plates and 41 to 46 interambulacral plates; poriferous zones slightly to much less than half the width of the poriferous zone as measured from the perradial to abradial suture at about two thirds of the length of the petal; and many demiplates in the petals. Weisbordella cubae (Weisbord, 1934) (Figs. 105, 106, 133-136; Tables 8, 9) Peronella cubae Weisbord, 1934. p. 217 (53). pl. 24 (5), figs. 4-6. Peronella cubae (Weisbord). Cooke, 1942. p. 25. pl. 2, figs. 15-21. Peronella cubae (Weisbord). Cooke, 1945. fig. 6, no. 6. Peronella cubae (Weisbord). Cooke, 1948b. p. 91. Peronella cubae (Weisbord). Sánchez-Roig, 1949. p. 97. Weisbordella cubae (Weisbord). Durham, 1954. p. 682. Weisbordella cubae (Weisbord). Cooke, 1959. pp. 53-54. pl. 20. figs. 1-4. Weisbordella cubae (Weisbord). Scolaro and Ross, 1963. pp. 304-307. Weisbordella cubae (Weisbord). Toulmin, 1977. p. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 190 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 138: Weisbordella johnsoni (UF 329698), 52.5 mm TL, 45.5 mm TW, Oligopygus haldemani Zone, Upper Eocene upper Ocala Limestone, Jackson County, Florida (FM-IP JA031). A: aboral. B: oral. C: left side. D: right side. 347. pl. 72, fig. 3-4. Weisbordella cubae (Weisbord). Osborn et al., 2016. tbl. 2. Occurrence.—Weisbordella cubae occurs throughout the OLS, but it is most abundant in the upper part of the unit just below the Wythella el- dridgei Zone, and the underlying Oligopygus wether- byi Zone. This species is also documented in the Eocene of Georgia (Cooke, 1942). The type locality of W. cubae is in Cuba (Weisbord, 1934). Discussion.—This is the common “sand dol- lar” of most exposures of the Oligopygus wetherbyi Zone of the upper OLS in peninsular Florida. In northern Florida, especially Jackson County, W. johnsoni replaces W. cubae as the abundant neola- ganid in the OLS. Weisbordella cubae has shorter, broader, and less lanceolate petals than W. johnsoni. Its test is also smaller and more ovate, whereas W. johnsoni tends towards a more subpentagonal outline, this being more pronounced in larger specimens. The concave oral surface readily differentiates W. cubae from W. inglisensis n. sp., as well as any species of Neolaganum or Durhamella with which it may be found. Although Fischer (1951: table 1) had W. cubae in his species list for the Inglis Formation (= lower OLS; Oligopygus phelani Zone), it is exceptionally rare in the lowermost OLS. Emended Diagnosis.—Weisbordella with: a deeply concave oral surface; primary plates lacking ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 191 Figure 139: Weisbordella johnsoni (UF 329699), 41 mm TL, 35 mm TW, Oligopygus haldemani Zone, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA031). A: aboral. B: oral. C: left side. D: right side. in the petals. Weisbordella johnsoni (Twitchell in Clark and Twitchell, 1915) (Figs. 105, 106, 137-139; Tables 8, 9) Laganum crustuloides (Morton). Clark and Twitchell, 1915. p. 122. figs. 5a-d, 6a-d. Laganum johnsoni Twitchell, in Clark and Twitchell, 1915. p. 162. pl. 75, figs. 2a-d. Echinodiscus crustuloides (Morton). Lambert and Thiéry, 1925. p. 581. Echinodiscus johnsoni (Twitchell). Lambert and Thiéry, 1925. p. 581. Laganum johnsoni (Twitchell). Cooke, 1926. pl. 96, fig. 2. Peronella crustuloides (Morton). Cooke, 1942. p. 24. Weisbordella johnsoni (Twitchell). Cooke, 1959. pp. 54-55. pl. 20, figs. 5-7. Weisbordella johnsoni (Twitchell). Kier, 1970. txt. fig. 1G. Weisbordella johnsoni (Twitchell). Toulmin, 1977. p. 347. pl. 71, figs. 7-8. Weisbordella johnsoni (Twitchell). Osborn et al., 2016. tbl. 2. Occurrence.—Weisbordella johnsoni is very abundant in the OLS in Jackson County, especially in the Sills Pit (FM-IP JA086) and Brooks Quarries ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 192 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 140: Weisbordella inglisensis n. sp., holotype (UF 342099), 21.8 mm TL, 19.1 mm TW, 4.1 mm TH, Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, mouth of Withlacoochee River, west of Yankeetown, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: posterior. D: tilted aboral viewpoint from posterior. E: tilted aboral viewpoint from anterior. F: anterior. G: oblique oral viewpoint from posterior. H: peristome. I: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 193 Figure 141: Weisbordella inglisensis n. sp., paratype (UF 342100), 23.8 mm TL, 20.9 mm TW, 4.0 mm TH, Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, mouth of Withlacoochee River, west of Yankeetown, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: oblique oral viewpoint from posterior. E: posterior. F: anterior. G: apical area and petaloid portion of ambulacrum III. H: tilted aboral viewpoint from anterior. I: peristome. J: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 194 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) (FM-IP JA009, (FM-IP JA018, FM-IP JA027, FM- IP JA031, FM-IP JA039), northwest of Marianna. This species also occurs in the OLS of Alabama (Cooke, 1959; Toulmin, 1977) and Georgia (Cooke, 1959). The type locality is Turks Cave, in Conecuh County, Alabama. Turks Cave is located about 3 mi. west of Brooklyn and is now commonly referred to as Sanders Cave. Discussion.—Twitchell in Clark and Twitchell (1915), described Laganum johnsoni from strata considered to be Lower Oligocene at the time, but now recognized as the Upper Eocene OLS, at Turks (Sanders) Cave in Conecuh County, Alabama. Twitchell in Clark and Twitchell (1915) named the species in honor of Mr. L. C. Johnson, the collector of the holotype, and noted that his new species is very similar to Laganum crustuloides, which Morton (1833) described as Scutella crustuloides from the Eocene of South Carolina and is now recognized as Protoscutella plana (Conrad, 1865). Twitchell in Clark and Twitchell (1915) noted that the specimen he described as L. johnsoni from Alabama had a higher test, thicker margin, and significantly concave oral surface. Cooke (1942) was puzzled by the fact that he was able to document L. crustuloides (with L. johnsoni as a junior synonym) from the Gulf coastal plain but not from South Carolina (the type area). He therefore suspected that Morton’s type of S. crustuloides came from Alabama and not from South Carolina. Cooke (1942) asserted that Morton’s (1833) original figure did not match his description of S. crustuloides, which Morton described as having a thick margin. Cooke presumed that the specimen Morton illustrated is in fact a specimen of P. plana. Therefore, Cooke (1942) asserted that Morton de- scribed a Gulf Coast specimen as S. crustuloides (the type of this species has been lost), and that Morton (1833) inadvertently figured a different specimen, likely P. plana. To attempt to bring some clarity to this epic of misidentifications, Cooke (1959) asserted that Morton’s type material of S. crustuloides was lost and that the figure included by Morton (1833) did not match the description of the species. Cooke placed the Gulf Coast form recognized as L. crustu- loides (Clark and Twitchell, 1915; Cooke, 1942) into synonymy with Twitchell’s (in Clark and Twitchell, 1915) L. johnsoni. Therefore, the first documenta- tion of what we now refer to as W. johnsoni is in Clark and Twitchell (1915) as both L. crustuloides and L. johnsoni, as recorded in the synonymy above. When Durham (1954) erected Weisbordella, he did not list W. johnsoni as a member. He stated that he did not move L. johnsoni into any other genus because he was unable to examine material of this species. Cooke (1959), with more material at hand, moved the species to Weisbordella and clarified that W. johnsoni had not been documented from South Carolina. Therefore, W. johnsoni appears to be restricted to the northern depositional area of the OLS, north of the Suwannee Strait, in Jackson County, Florida, and northward into Georgia and Alabama. Weisbordella johnsoni, as stated by Cooke (1942, 1959), is highly variable, with great varia- tion in test thickness, outline, and marginal thick- ness. Some specimens are thick and wafer-like with very swollen margins and a depressed area just inward from the margin. Others are thin- ner and have a much less swollen margin, with many specimens having intermediate characteris- tics within the same population. This variation has not helped with clarification of the concept of the species. Fischer (1951: 57) documented P. crustu- loides (= W. johnsoni) in the lower OLS near Inglis, Florida. As we discussed in the remarks for W. in- glisensis n. sp., these specimens (UF 317144) are inseparable from W. inglisensis n. sp., and we there- fore do not recognize W. johnsoni as occurring below the Oligopygus haldemani Zone of the OLS in the region. The concave oral surface readily differentiates W. johnsoni from any of the species that occur in the lower OLS. However, specimens of W. johnsoni from the type area near Brooklyn, Conecuh County, Alabama demonstrate some variation in oral surface concavity, with rare specimens having a nearly flat oral surface. Nevertheless, the typical concave oral ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 195 surface is a dominant trait in this species. Weisbordella johnsoni has proportionately longer and more lanceolate petals than W. cubae. Weisbordella cubae is also smaller and more ovate, whereas W. johnsoni tends towards a more subpen- tagonal outline, especially the posterior portion of the test. This trait is more pronounced in larger specimens. We figure specimens of varying sizes to illustrate this variation (Figs. 137-139). The largest of these (UF 329698) is 52.7 mm in length and 45.4 mm wide. Emended Diagnosis.—Weisbordella with a combination of: proportionately longer, more lance- olate petals than its congeners; a predominance of demiplates in the petals with only rare primary plates; highly variable degree of concavity on the oral surface. Weisbordella inglisensis n. sp. (Figs. 105, 106, 140-146; Tables 8, 9) not Peronella crustuloides (Morton). Fischer, 1951. p. 57, tbl.1. Diagnosis.—Weisbordella with a combina- tion of: a planar oral surface; petals with an almost regular pattern of a demiplate for every primary plate. Description.— Description based on holotype (UF 342099) and eight paratypes (UF 342092, UF 342100-342105, and CASG 103249). Moderately sized neolaganid, largest specimen (UF342100) 23.8 mm TL, 20.9 mm TW, 4.0 mm TH; holotype: 21.8 mm TL, 19.1 mm TW, 4.1 mm TH; small specimens (less than 10 mm TL) nearly circular outline, subo- vate above 12 mm TL, TW on average 88.5% TL; posterior margin usually truncated at ambulacrum I and V, giving outline subpentagonal aspect; mar- gin at anterior ambulacra less frequently slightly truncated. Test low, largely flattened, apical area slightly raised, highest point anterior of center; TH on average 18.9% TL, slightly wedge-shaped pro- file, lowest posteriorly; posterior ambitus on average 57.8% height of anterior ambitus; upper surface often very slightly depressed inside margin at end of petals. Apical area slightly anterior; center of apical system on average 54.1% TL from posterior margin; four gonopores, hydropores in sinuate groove. Petals flush, lanceolate, nearly closed distally, broad; width of widest point of petal I on average 53.4% length of petal; petal II: 57.1%, petal III: 53.4%, petal IV: 58.6%, petal V: 53.7%. Petals extend more than two- thirds of distance from apical system to ambitus; petal III longest, petal II and IV shortest: petal I on average 27.9% TL, petal II: 24% TL, petal III: 26.4% TL, petal IV: 23.7% TL, petal V: 27.3% TL; pore pairs conjugate, outer pore slightly elongate. Petals with one demiplate for each primary plate (Fig. 105). Oral surface nearly planar. Periproct on aver- age 11.4% TL from posterior margin, first in contact with plates 5a2 and 5b2 (Fig. 105); slightly ovate, wider than long: length on average 83.6% width. Basicoronal plates form subpentagonal outline, with single plate in each interambulacrum, paired plates in each ambulacrum, ambulacral plates about one- half as wide as adjacent interambulacral plates; first pair of postbasicoronal interambulacral plates elongate, usually three shorter plates between them and ambitus. Total of approximately 54 ambulacral and 41 interambulacral plates on oral surface in adult specimens. Peristome subpentagonal to nearly round, nearly central; posterior edge of peristome on average 49.1% TL from posterior margin of test; peristome diameter on average 7.8% TL. Zoobank Nomenclatural Act.—40BCA279- F8B9-426E-A1F7-3B028C77BE6D Discussion.—The Oligopygus phelani Zone of the lower OLS is rich in neolaganids, with most well-weathered exposures of the limestone portion of the strata formerly known as the Inglis Formation containing D. ocalana. Durhamella ocalana has unmistakable depressions along the sutures in the area just proximal to margin on the aboral surface. Neolaganum durhami, which also occurs in the lower OLS, is distinguished primarily by its much great number of oral ambulacral and interambulacral plates and the fact that it has five gonopores, not four as in all species of Weisbordella (including W. inglisensis n. sp.). Members of Neolaganum also tend to have a branched hydropore groove, whereas W. inglisensis n. sp. usually has an unbranched, sinuous groove. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 196 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) While identification of these two species is rel- atively simple and straightforward, additional small neolaganids that have previously been misidentified also occur in this horizon. In addition to D. ocalana and N. durhami, Fischer (1951) documented Per- onella dalli (= N. dalli) from the Inglis Formation (= lower OLS). He admitted that this species had not been found in outcrops of the unit, and we concur. Fischer (1951: table 1) also listed Peronella cubae (= W. cubae) in the unit, but it is very rare in the lowermost OLS. Fischer (1951) also documented Peronella crustuloides (= W. johnsoni) in the In- glis Formation. This assertion was based on two poorly preserved specimens measuring 12 mm and 13 mm in length. Fischer did not provide catalog numbers for his specimens of P. crustuloides (= W. johnsoni). However, we were able to locate them in the UF collections (UF 317144). These two speci- mens were collected 1 mi. east of the Withlacoochee River bridge at Inglis, south of the river (FM-IP CI053), and are poorly preserved but indistinguish- able from W. inglisensis n. sp. We are not aware of any occurrence of W. johnsoni in the Oligopygus phelani Zone (that portion of the lowermost OLS previously referred to as the Inglis Formation). We therefore do not recognize W. johnsoni as occurring in Florida below the Oligopygus haldemani Zone of the OLS. Fischer’s misidentification of the speci- mens is understandable because, as discussed below, the petals of W. johnsoni are very similar to those of W. inglisensis n. sp. The specimens we describe as W. inglisensis n. sp. were collected from the limestone portion of the Oligopygus phelani Zone of the lower OLS (strata previously included in the Inglis Formation) along the Withlacoochee River at its mouth west of Yankeetown, Levy County (FM-IP LV024), where it occurs with D. ocalana, D. floridana, N. durhami, P. floridana, E. mooreanus, O. phelani, and other, rarer species. Weisbordella inglisensis n. sp. occurs more rarely in the same horizon along the banks of the Cross Florida Barge Canal south of Inglis (e.g., FM-IP CI001) in Citrus County, at the type locality of the Inglis Formation behind the powerplant on the Withlacoochee River east of Inglis, Levy County, and in Briar Cave, Marion County (FM-IP MR018). Weisbordella inglisensis n. sp. shares char- acteristics of both Neolaganum and Weisbordella, further discussed in the remarks on the genus, above. However, the oral plate architecture of W. inglisensis n. sp. (Fig. 105; Table 9) clearly shows a signifi- cantly reduced number of plates that is not typical for Neolaganum. In addition, differs in having four gonopores and a sinuous, rather than branched hy- dropore groove, and more demiplates in the petal plate pattern. Thus, even though W. inglisensis n. sp. lacks the concave oral surface typical of the type species of Weisbordella, W. cubae, it is clearly more closely related to other taxa in Weisbordella than to any member of Neolaganum. Weisbordella inglisensis n. sp. is unlike any of the known Weisbordella. In the regional fauna, W. inglisensis n. sp. is very similar to W. johnsoni from an aboral viewpoint alone (similar petaloid and apical structure). However, the consistently planar oral surface of W. inglisensis n. sp. distinguishes it from W. johnsoni, as does the more frequent occurrence of demiplates in the petals of the latter. The petals of W. cubae have even more demiplates, so along with its very concave oral surface, it is easily distinguished from W. inglisensis n. sp. Etymology.—Named after the community of Inglis, in Levy County, Florida, near the type locality of W. inglisensis n. sp. Material and Occurrence.—Holotype (UF 342099) and paratypes (UF 342100, UF 342101, UF 342102, UF 342103, UF 342104, UF 342105, CASG 103249) from the lower OLS, Oligopygus phelani Zone, at the mouth of the Withlacoochee River at the Gulf of Mexico (FM-IP LV024), east of Yankeetown, Levy County, Florida. Also, a paratype UF 342092 from the lower OLS, Oligopygus phelani Zone, Cross Florida Barge Canal, south of Inglis, west of Hwy 19/98, Citrus County, Florida. This species has also been found in the Oligopygus phe- lani Zone of the lower OLS in Briar Cave, Marion County (FM-IP MR018). Weisbordella libum n. sp. (Figs. 106, 147; Table 8) not Clypeaster oxybaphon (Jackson). Oyen, 2001. pp. 97-98. figs. 3-13 a, b. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 197 Figure 142: Weisbordella inglisensis n. sp., paratype (UF 342103), 12.7 mm TL, 11.4 mm TW, 2.5 mm TH, Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, mouth of Withlacoochee River, west of Yankeetown, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: tilted aboral viewpoint from posterior. D: tilted oral viewpoint from posterior. E: apical area. F: periproct. G: peristome. H: anterior. I: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 198 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 143: Weisbordella inglisensis n. sp., paratype (UF 342104), 11.2 mm TL, 10.2 mm TW, 2.1 mm TH, Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, mouth of Withlacoochee River, west of Yankeetown, Levy County, Florida (FM-IP LV024). A: aboral. B: oral. C: left side. D: tilted oral viewpoint from posterior. E: petaloid area. F: periproct. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 199 Figure 144: Weisbordella inglisensis n. sp., paratype (UF 342092), 24.8 mm TL, 22.2 mm TW, 4.9 mm TH, Upper Eocene, Oligopygus phelani Zone of lower Ocala Limestone, Cross Florida Barge Canal south of Inglis, west of Hwy. 19/98, Citrus County, Florida (FM-IP CI001). A: aboral. B: oral. C: posterior. D: anterior. E: tilted aboral viewpoint from posterior. F: tilted oral viewpoint from posterior. G: petaloid portion of ambulacrum III. H: tilted aboral viewpoint from anterior. I: periproct. J: left side. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 200 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 145: Weisbordella inglisensis n. sp., (UF 343023), 18.1 mm TL, 16.9 mm TW, 3.55 mm TH, Oligopygus phelani Zone, Upper Eocene, lower Ocala Limestone, Citrus County, Florida (FM-IP CI012). A: aboral. B: oral. C: left side. D: right side. Diagnosis.—Very large Weisbordella with very thick test, TH estimated to be 20-25% TL; highly complex system of hydropore grooves; petals with very long, slit-like outer pores and two to three demiplates alternating with a single primary. Description.—Description based on the holo- type (UF 4926) and only known specimen of W. libum n. sp. Specimen is incomplete (missing poste- rior portion of test); large, 64.4 mm TL (incomplete) and 69.2 mm TW (estimated dimensions of com- plete test 85 mm long and 70 mm wide: Fig. 147), estimated ratio of TW to TL 82%; thick, 17.8 mm TH (only measurement that is complete), between 20-25% TL. Test high at the margin and not abo- rally depressed just proximal to that. Aboral surface with even dispersal of small primary tubercles and fewer, scattered larger tubercles; oral surface evenly covered in tubercles of approximately equal size. Posterior portion of test missing, but test likely elongate; ovate to subpentagonal (Fig. 147). Anterior edge of apical area 37 mm from anterior margin; four gonopores, average .25 mm in diameter; anterior pair 2.3 mm apart, posterior pair 4 mm apart: nearly twice as far apart as anterior pair. highly complex system of hydropore grooves (Fig. 147). Petals flush, lanceolate, nearly closed distally, broad; width of widest point of petal II 55% length of petal; petal III: 48%, petal IV: 54. Petals extend more than two-thirds of distance from apical system to ambitus; petal III longest (22.7 mm), petal II (21.8 mm) and IV (21.1 mm) shortest. Poriferous zones wide, but still less than half as wide as distance from perradial suture to abradial suture, pore pairs conjugate, outer pore elongate, slit-like. Petals with two to three demiplates alternating with a single ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 201 Figure 146: Weisbordella inglisensis n. sp., (UF 344347), 18.5 mm TL, 16.5 mm TW, 4.0 mm TH, Oligopygus phelani Zone, Upper Eocene lower portion of Ocala Limestone, Briar Cave, Marion County, Florida (FM-IP MR018). A: aboral. B: oral. C: left side. D: right side. primary (Fig. 105). Oral surface with slight concavity that starts from just proximal to ambitus; peristome subpentagonal; width equals 89% peristome height; peristome 39.5 mm from anterior margin. Periproct not preserved. Food grooves absent. Basicoronal plates form subpentagonal outline, with a single plate in each interambulacrum, paired plates in each ambulacrum, ambulacral plates about one-half as wide as adjacent interambulacral plates; first pair of post-basicoronal interambulacral plates elongate, extending to second or third ambulacra post- basicoronals (Fig. 147). Peristome subpentagonal to nearly round, nearly central; posterior edge of peri- stome on average 49.1% TL from posterior margin of test; peristome diameter on average 7.8% TL. Internal structure partially visible along bro- ken edge of test; massive peripheral buttress system, extremely thick test, extremely high lantern pre- served and in living position within test, spanning the body cavity from aboral to oral interior surfaces of the test, auricle in interambulacrum 5 massive. Zoobank Nomenclatural Act.—9082B149- 35D3-4ED1-890B-8C0CE6D4E6E9 Discussion.—This specimen consists of a large, incomplete test (UF 4926) collected 50 years ago in the upper portion of the Upper Eocene, OLS in the now inactive Mill Creek Quarry (FM-IP LF002), west of Dowling Park in Lafayette County. The precise placement of the specimen within the OLS is not possible, as these data are not indicated on the labels with the specimen, but this quarry exposed both the Oligopygus wetherbyi Zone and overlying Wythella eldridgei Zone of the upper OLS. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 202 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 147: Weisbordella libum n. sp., holotype (UF 4926), 64.4 mm TL and 69.2 mm TW (not complete), 17.8 mm TH, Upper Eocene, upper portion of Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: detail of apical system. D: plate detail of oral surface; ambulacrum III towards the top, the interambulacral plates are shaded, and peristome and periproct are solid black. Reconstructed dotted outline is an extrapolation based on the dimensions of other neolaganids. E: detail of peristome. F: detail of aboral interambulacrum 2 showing presence of large and small primary tubercles, as well as portions of the poriferous zones in petals II and III. G: left side. H: anterior (directly ambital view of ambulacrum III). I: slightly upward-tilted posterior view showing the massive peripheral buttress system, extremely thick test, posterior view of the Aristotle’s lantern in place, and lantern support (auricle) in interambulacrum 5. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 203 Field notes for this site made by Muriel Hunter during the time the specimen was collected indicate a significant presence of the uppermost OLS (Wythella eldridgei Zone) and it is likely the spec- imen originated from this horizon. However, this taxon is not documented elsewhere, and this cannot be confirmed until additional specimens are found. The specimen was first recorded by Oyen (2001) as Clypeaster oxybaphon. However, the pres- ence of four gonopores, rather than the five typical for Clypeaster, and other characters discussed below readily distinguish the specimen from any member of the Clypeasteroida sensu lato. As shown in Fig. 147, the broken test shows the interior of the test, revealing a massive peripheral buttress system and a posterior view of an extremely robust Aristotle’s lantern in place, with a large, robust auricle visi- ble in interambulacrum 5. This further indicates that this is not a clypeasteroid, but a scutelloid, as clypeasterines do not have interambulacral auricles. The large test (estimated to have been 85 mm TL and 70 mm TW) with an exceptionally thick test (17.8 mm TH) and highly complex system of hydropore grooves is unique among neolaganids from Florida. We compared the specimen with neola- ganids from elsewhere in the eastern Americas and Caribbean region to ensure it was not synonymous with an already described species, and to support the genus designation. Cubanaster torrei (Lambert in Sánchez Roig, 1926) the type species of Cubanaster, and C. acunai (Lambert in Sánchez Roig, 1926), the two earliest described Cubanaster species, differ from W. libum n. sp. in having a much lower test, especially at the margin, and depressed just proximal to that, a greater number of plates on the oral sur- face (even though Weisbordella libum n. sp. is much larger), and much narrower interporiferous zones. In addition, C. torrei has a more equilateral (i.e., almost circular) test, whereas C. acunai appears to have a more elongate test than Weisbordella libum n. sp. Laganum cubensis Lambert, 1925 has been confused in past literature. Lambert (1925: 582) lists the species under the name Jacksonaster, stating that it was listed by Egozcue (in Cotteau, 1897: 26, pl. 4) under the name ”Laganum elongatum”. Since that time, it has appeared in works by Sánchez Roig (1924, 1926, 1949) using ”Laganum-elongatum” in synonymies (which is odd as Egozcue did not use the hyphen in his version of the name), but always unfig- ured. In Sánchez Roig (1924), the taxon was placed in Clypeaster as C. elongatum, but subsequently moved to Jacksonaster as a new species, J. cubensis, in Sánchez Roig (1926), but attributed to Lambert, 1925. It seems that Lambert’s coining was a nomen nudum, and that Sánchez Roig (1926) should be the author of record. In his entry on Clypeaster oxy- baphon, Cooke (1942: 13) used the name Laganum elongatum (Egozcue in Cotteau, 1897), citing the illustrations in Egozcue’s work while stating, ”It [meaning C. oxybaphon] bears considerable resem- blance to Laganum elongatum Egozcue (1897, pl. 4), which appears to be a Clypeaster.” Apparently, Cooke missed Sánchez Roig’s (1924) earlier referral to that genus but seemed to be advocating a return to the name ”elongatum”. Jackson (1922: 47) indicated that he had never seen an example of L. elongatum, and reiterated Egozcue’s description (in Cotteau, 1897). The description, dimensions, and appearance of the specimen in the figures provided by Egozcue (1897: pl. 4) for L. elongatum and cited by Lambert (1925: 582) for his Jacksonaster cubensis, back up the supposition that the echinoid is most likely a Clypeaster. Laganum elongatum has a deep infundibu- lum. Although W. libum n. sp. has a highly localized depression surrounding the peristome, this is com- pletely unlike what is depicted for L. elongatum. The petals, though like those of W. libum n. sp., are also consistent in shape with those of other Clypeaster. However poorly known, and whatever its correct name might be, ”L. elongatum/cubensis” does not seem to be related to W. libum n. sp. Jacksonaster depressus [sic], as discussed in Sánchez Roig, 1949, is large, but not so large as W. libum n. sp., has a distinctly inflated edge, and longer petals with parallel poriferous zones. Jacksonaster depressus has a stronger resemblance to C. torrei or C. acunai than to W. libum n. sp. It is presently uncertain what J. depressus represents, but it is highly unlikely to be conspecific with Jacksonaster ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 204 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) depressum (L. Agassiz, 1841) Jacksonaster remediensis Sánchez Roig, 1949 is very much like Sanchezella sanchezi except that the petals are not deeply depressed, as Sánchez Roig (1949) himself notes. It is also a small species, and is not comparable to W. libum n. sp., which is not only larger, but much wider relative to its TL, and with much narrower interporiferous zones. Jacksonaster sandiegensis Sánchez Roig, 1949 is a small species relative to others described by Sánchez Roig, and although the specimen illustrated by him seems to be crushed laterally, it still appears to be elongated, much as for other species he later considered to be members of Cubanaster, such as S. sanchezi. The apical system of J. sandiegensis is unknown due to the weathering of the upper surface, but Sánchez Roig (1949) reports that the petals are very large, occupying almost the entire upper surface of the test. It is therefore very unlike W. libum n. sp. Laganum lamberti Sánchez Roig, 1949 is very similar to L. santanae (see below, and not to be confused with Cubanaster santanae Sánchez Roig, 1952e, also see below), including in petal shape, gonopore number and test shape, and is likely simply a smaller specimen of L. santanae Sánchez Roig, 1949), with all the same objections to its identity with W. libum n. sp. Laganum santanae Sánchez Roig, 1949 is a very large species, even larger than the specimen of W. libum n. sp., and is relatively thick-edged, and appears, at first glance, to be the closest to W. libum n. sp. of all the species considered here. The petals of this species are also remarkably similar to those of W. libum n. sp., being somewhat sinuous with wide poriferous zones. However, Sánchez Roig (1949) reports that L. santanae has 5 gonopores (presumably part of the reason that he placed it in Laganum). In addition, the peristome of L. santanae is transversely elongated, and if this is a good charac- ter for the species, then it is quite different in shape from the peristome of W. libum n. sp. Cubanaster acunai gigas Sánchez Roig, 1952e is the only form described in Sánchez Roig (1952e) that rivals W. libum n. sp. in size, other than L. santanae. However, the estimated ratio of TW to TL for W. libum is 82%, which is considerably greater than for C. acunai gigas. The latter also has a conspicuously sunken aboral region just inside the ambitus (i.e., a very ”inflated” margin), and the petals are also much narrower, with a very narrow interporiferous zone. Cubanaster camagueyensis Sánchez Roig, 1952e is, like many of the new species described in Sánchez Roig (1952e), elongate and thick edged, but much more like Sanchezella than W. libum n. sp. in having depressed, long, narrow petals and, if the illustrations can be believed, very distinct food grooves. Gonopore number is not mentioned in the description by Sánchez Roig (1952e), but a specimen of C. camagueyensis (UF 216680), clearly shows that there are four. Cubanaster herrerai Sánchez Roig, 1952e has a conspicuously sunken aboral surface, and long, nar- row petals with very narrow interporiferous zones quite unlike those of W. libum n. sp. Cubanaster pla- nipetalum Sánchez Roig, 1952e seems again to be a more elongate form than W. libum n. sp., with short but distinct food grooves which are lacking in the latter, and a slightly depressed aboral surface show- ing relatively wide petals with narrow poriferous zones. Cubanaster santanae Sánchez Roig, 1952e is like W. libum n. sp. in that it has 4 gonopores and a thickened edge, but it is very elongate, similar in test shape to Sanchezella, and has somewhat depressed petals. Examination of oral plate arrangement (Figs. 105, 147) revealed this taxon is more closely aligned to Weisbordella than any other neolaganid. Neolaganum and Cubanaster have high plate num- bers on the oral surface and the plate numbers of W. libum n. sp. are fairly low, in spite of its large size. The large, thick test with a highly complex system of hydropore grooves readily distinguishes this species from its congeners. It is hoped that additional col- lecting will provide more complete specimens of this species to enable a better understanding of its characters (such as true test dimensions, periproct position, and oral surface plate architecture). Etymology.—The species is named libum (Latin, neuter) for its resemblance to a consecrated cake or pancake made in Roman times. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 205 Figure 148: Wythella eldridgei (UF 337964), 55 mm TL, 44 mm TW, uppermost portion of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF023). A: aboral. B: oral. C: left side. D: right side. Material and Occurrence.—This species is known only from the holotype (UF 4926) which was collected in upper portion of the Upper Eocene OLS in the Mill Creek Quarry (FM-IP LF002) west of Dowling Park, Lafayette County, Florida. Genus Wythella Durham, 1954 When Durham (1954) described Wythella it contained Wythella eldridgei as its type and only species. Durham (1954) also designated Rumphia elegans Sánchez-Roig, 1949, from the Oligocene of Cuba, as the type species of Neorumphia. Durham (1954) distinguished Wythella from the very similar Neorumphia by noting that the former had fewer plates on the oral surface, a thinner margin, and narrower interambulacral areas at the ambitus. Smith and Kroh (2011) noted that although Neorumphia has somewhat wider interambulacral zones at the ambitus, the otherwise very similar plating and appearance render Durham’s distinction between the two genera suspect. Smith and Kroh (2011) therefore treated Neorumphia as a subjective junior synonym of Wythella, with which we agree. Durham did not provide a diagnosis for the genus, but it is readily distinguished from other neolaganids by its large, very thin test (W. eldridgei has the thinnest test relative to TL of all known ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 206 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 149: Wythella eldridgei (UF 338005), 36 mm TL, 32 mm TW, 5 mm TH, uppermost portion of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF023). A: aboral. B: oral. C: left side. D: right side. neolaganids), and flat oral surface. In addition, anal- ysis of oral plate architecture of W. eldridgei (Fig. 105) reveals that it has a greater number of plates than Weisbordella or Durhamella and is more sim- ilar to Neolaganum in this aspect. Wythella has at least one plate more in each of the anterior paired interambulacral regions than in the posterior pair. This is also like the condition found in many Neolaganum. Emended Diagnosis.—Large Neolaganidae with a combination of: extremely thin test, espe- cially near the ambitus; short, unbranched hydropore groove; long, slit-like outer pore in petal pore pairs; two or three demiplates alternating with single pri- mary plate; poriferous zone approximately half as wide as the distance from the perradial suture to the abradial suture at a point about 2/3 the length of the petal. Wythella eldridgei (Twitchell in Clark and Twitchell, 1915) (Figs. 105, 106, 148, 149; Table 8) Laganum eldridgei Twitchell in Clark and Twitchell, 1915. p. 160. pl. 74, figs. 2a-d. Rumphia eldridgei (Twitchell). Stefanini, 1924. pp. 831, 843. Rumphia eldridgei (Twitchell). Cooke and Mossom, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 207 Figure 150: Porpitella micra (USNM 462807), Ohio Oil Co Well, 5520-5530 ft depth, Late Cretaceous Atkinson Formation (Applin and Applin, 1965), Hernando County, Florida. A: aboral. B: oral. 1929. pl. 3, fig. 7. Rumphia eldridgei (Twitchell). Cooke, 1942. p. 27. pl. 2, fig. 14. Rumphia eldridgei (Twitchell). Cooke, 1945. fig. 5, no. 1. Wythella eldridgei (Twitchell). Durham, 1954. p. 682. txt fig. 3D. Wythella eldridgei (Twitchell). Durham, 1955. txt fig. 27d. Wythella eldridgei (Twitchell). Cooke, 1959. p. 53. pl. 21, figs. 8-10. Wythella eldridgei (Twitchell). Durham, 1966. p. 475. fig. 365, 5a, 5b. Wythella eldridgei (Twitchell). Toulmin, 1977. p. 348, pl. 72, figs. 1-2. Wythella eldridgei (Twitchell). Mooi, 1989. figs. 10a, 16b, 18a. Wythella eldridgei (Twitchell). Osborn et al., 2016. tbl. 2. Occurrence.—Wythella eldridgei is most com- monly found in abundance in a horizon resting just above the Oligopygus wetherbyi Zone in the up- per OLS; this is the Wythella eldridgei Zone, and best exposed in Lafayette, Suwannee, and northern Dixie Counties, Florida, especially west of Dowl- ing Park, Lafayette County (FM-IP LF002), quar- ries northwest of Mayo, Lafayette County (FM- IP LF001), and O’Brien Quarry (FM-IP SU002) north of Branford, Suwannee County. The type locality of the species is along the Suwannee River, 24 km below Ellaville. Carter and McK- inney (1992) documented this species in the OLS of Georgia. Discussion.—Twitchell in Clark and Twitchell (1915), initially described this very distinctive species as Laganum eldridgei from strata referred to the Lower Oligocene from the Suwannee River at Ellaville, Florida. Subsequently, Durham ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 208 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 151: Porpitella micra (USNM 462806), 3 mm TL, 2.5 mm TW, 1 mm TH, Ohio Oil Co Well, 5509-5519 ft. depth, late Cretaceous Atkinson Formation (Applin and Applin, 1965), Hernando County, Florida. A: aboral. B: oral. (1954) erected Wythella, with W. eldridgei as the type species. Wythella eldridgei is thinner, larger, and more elongate than any other sand dollar occurring in the upper OLS. It is found in abundance in a lens of pale soft limestone upwards 1 m thick that rests atop the Oligopygus wetherbyi Zone of the up- per OLS and below the overlying Rhyncholampas gouldii-bearing Oligocene sediments of the Suwan- nee Limestone in the northern Florida Peninsula. Puri (1957) named this horizon the Spirolaea (now Rotularia) vernoni Zone. Here, W. eldridgei oc- curs with W. cubae, R. trojana, and a diverse as- semblage of spatangoids that includes P. dixie, E. ocalanus, P. curvus, S. armiger, B. steinhatchee, and O. beckeri. Emended Diagnosis.—As for the genus. Family SCUTELLINIDAE Pomel, 1888 Genus Porpitella Pomel, 1883 Porpitella micra H. L. Clark, 1937 (Figs. 150, 151) Porpitella micra H. L. Clark, 1937. p. 248. Porpitella micra (H. L. Clark). Cooke, 1942. p. 30. Porpitella micra (H. L. Clark). Cooke, 1959. p. 33. pl. 7, figs. 5-7. Occurrence.—The type locality for this minis- cule species is at 3800-4000 ft. depth in the Oakley Estate well 2 in sec. 9, T. 3 N., R. 29 E., Houston County, Alabama. Cooke (1959) stated that the spec- imens were associated with the oyster Cubitostrea sellaeformis (Conrad, 1832), which would indicate the material is from the Middle Eocene Lisbon Formation. However, this Eocene age was debated by Applin and Applin (1965), who also documented specimens of P. micra from a well of the Ohio Oil Company, Hernasco Corp., in sec. 19, T. 23 S., R. 18 E., in Hernando County, Florida. These P. micra ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 209 were from 5509–5519 ft. (1679–1682 m) depth (USNM 462806) and 5520–5530 ft. (1682–1686 m) depth (USNM 462807) in what Applin and Applin assert is the Late Cretaceous Atkinson Formation. If this proves correct, this remarkable finding will make P. micra the earliest known of all the scutelloids (see below). Porpitella micra is not known from surface exposures in Florida, or anywhere else, being documented only from well cores. Discussion.—H. L. Clark (1937) described this species from well cuttings in Houston County, Alabama, at a depth of 3800-4000 ft. (1158–1219 m) below the surface, from strata Clark (1937) surmised to be the Middle Eocene Lisbon Formation. As noted above, Cooke (1959) associated the echinoids with C. sellaeformis, the characteristic oyster of the upper Lisbon Formation, and reiterated the Eocene age asserted by Clark (1937). Clark (1937) stated that there were 26 paratypes from the same locality that show great diversity in form and preservation and that few were as well preserved as the holotype, and in most cases, the petals cannot be distinguished. The smallest specimen he recorded is 5 mm TL x 4 mm TW and the largest is 9.5 mm TL x 8.5 mm TW. Clark noted that while most individuals are distinctly longer than wide, with the anterior end narrowed and the poste- rior margin nearly straight, some are almost circular, one notable example 6 mm TW and less than 7 mm TL. The apical system is nearly central in these more circular specimens, so the anterior position of the apical system is not a constant character. Clark (1937) also mentioned there is great diversity in the concavity of the oral surface. In some cases, the con- cavity is pronounced and extends from the posterior margin nearly to the anterior end of the test. At the other extreme are individuals in which the entire oral surface is almost flat except for a localized depres- sion around the peristome. Clark (1937) stated there is no American echinoid with which P. micra can be confused because the supramarginal periproct and the character of the petals are distinctive. The Florida specimens (Figs. 150, 151) are very small: 3.2 mm TL, 2.8 mm TW, and 1.1 mm TH; and 3.3 mm TL, 2.9 mm TW, 1.2 mm TH, and not well preserved. Although the aboral surface is well preserved on USNM 462807, and the characteristic aboral periproct is readily discernable, the petals are not. Applin and Applin (1965) questioned the Mid- dle Eocene age Clark (1937) and Cooke (1942, 1959) attributed to the species at its type locality. The age determination was in part based on the associa- tion of the specimens with Cubitostrea sellaeformis noted by Cooke (1959). This is a characteristic, saddle-shaped oyster diagnostic for the NP16 zone of the Claibornian Middle Eocene of the Gulf Coast (Baum and Vail, 1988; Toulmin, 1977) and the up- per portion of the Lisbon Formation in Alabama. Applin and Applin (1965) reported that Cooke had examined the Florida specimens and identified them as P. micra. Our own examination indicated that they are certainly most similar to P. micra, but they are not well preserved. As noted by Clark (1937), and discussed above, this is a highly variable species, and the Florida specimens fall within the ranges of variation seen in the type material from Alabama. As already noted, Applin and Applin (1965) strongly suggested that the Florida specimens were collected in Upper Cretaceous strata within the well core, the exceptionally deep origin of which does imply they come from below the Eocene. Applin and Applin provided the well logs that support this de- termination. In addition, Applin and Applin (1965) stated that the Hernando County specimens were collected with Exogyra woolmani Richards, 1947, (a diagnostic Late Cretaceous species), supporting their claim of a Late Cretaceous age for the speci- mens. To reconcile this Late Cretaceous occurrence of a previously reported Middle Eocene popula- tion of P. micra, they studied the cores from the Alabama well (type locality for the species). Ap- plin and Applin (1965: 69) stated the Alabama Geological Survey had also questioned the age of the material from the Alabama well and that their subsurface investigations in the region indi- cated that the specimens of Porpitella from the Houston County well are, like the Florida spec- imens, of early Atkinson (early Gulf) age (Late Cretaceous). As mentioned, Clark (1937) and Cooke (1959) based their age determination of P. micra on the ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 210 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 152: Periarchus floridanus (UF 12797), 110 mm TL, 110 mm TW, external mold with RTV cast, dolomitic beds of Upper Eocene lowermost Ocala Limestone, Citrus County, Florida (FM-IP CI009). A: external mold. B: RTV cast. association of C. sellaeformis with the type speci- mens in the Alabama core. Cubitostrea sellaeformis is typically a large species of oyster. An example surviving sufficiently intact for full identification in well cuttings would be remarkable, so it is possible this identification was made from incomplete oyster valves. A Late Cretaceous occurrence of Porpitella is remarkable, as the earliest known scutelliforms discussed in recent literature are suggested to ap- pear in the Early Eocene (Smith and Kroh, 2011). However, Mongiardino Koch et al. (2022) provided robust molecular and phylogenetic dating results that strongly suggest an origin of scutelliforms before the Paleocene, making the data concerning the oc- currence of Porpitella of significance in supporting these findings. Strictly speaking, a Cretaceous occurrence of a scutelliform does not fit the Paleogene focus of this work. However, the significance of P. micra in Upper Cretaceous strata compels us to include the taxon as the earliest documented occurrence of a species of Scutelloida. Accordingly, we have not included this species in the Eocene distribution table of Florida taxa (Table 1). Infraorder SCUTELLIFORMES Haeckel, 1896 Family PROTOSCUTELLIDAE Durham, 1955 Genus Periarchus Conrad, 1866 Periarchus floridanus Fischer, 1951 (Figs. 152, 153) Periarchus lyelli (Conrad). Cooke, 1942. p. 14. (in part, references to this species in Florida are P. floridanus). Periarchus lyelli floridanus Fischer, 1951. p. 60. pl. 1, figs. 1-4; txt figs. 4, 5. Periarchus lyelli floridanus (Fischer). Cooke, 1959. p. 42. pl. 13, fig. 4. Periarchus lyelli floridanus (Fischer). Toulmin, 1977. p. 344. pl. 70, fig.4. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 211 Figure 153: Periarchus floridanus (UF 12882), 64.5 mm TL, 65 mm TW, external mold with RTV cast, dolomitic beds of Upper Eocene lowermost Ocala Limestone, Citrus County, Florida (FM-IP CI009). A: external mold. B: RTV cast. Periarchus floridanus (Fischer). Oyen and Portell, 2001. pp. 193-218. pl. I, fig. 1. Periarchus lyelli floridanus (Fischer). Osborn et al., 2016. tbl. 2. Occurrence.—This is the common large sand dollar of the Oligopygus phelani Zone of the lower OLS (formerly Inglis Formation) of Citrus and Levy Counties and is present in varying concentrations (usually as test fragments) at nearly all exposures of the unit. A few localities include: Cross Florida Barge Canal (e.g., FM-IP CI001) south of Inglis, Citrus and Levy Counties; mouth of Withlacoochee River west of Yankeetown, Levy County (type lo- cality: FM-IP LV024); and Cemex (formerly Inde- pendent Aggregates) Quarry, southwest of Inglis, Citrus County (FM-IP CI017). Toulmin (1977) documented the occurrence of P. floridanus in the Moodys Branch Formation along the east bank of the Chattahoochee River at mile 36.4 above the confluence with the Flint River in Early County, Georgia. Discussion.—Although Cooke (1942) docu- mented the presence of Periarchus lyelli within the Eocene strata of Florida, Fischer (1951) was first to recognize the differences between this west central Florida population and typical P. lyelli when he described the form as P. lyelli floridanus. Fischer (1951) documented his new subspecies at numerous localities near Inglis in Citrus and Levy Counties in strata that were at the time recognized as the Moodys Branch Formation. This Upper Eocene de- posit would subsequently be designated as the Inglis Member of the Moodys Branch Formation, Inglis Formation, and later as the lower OLS; it is the Oligopygus phelani Zone of the lower OLS. Al- though this species would later be documented from strata outside the Inglis area, the type area is its area of greatest concentration. Like Carter et al. (1989: tables 1, 4) and Oyen ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 212 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 154: Periarchus quinquefarius (UF 5275), 49 mm TL, 51 mm TW, 9.5 mm TH, upper portion of Upper Eocene Ocala Limestone, Suwannee County, Florida (FM-IP SU004). A: aboral. B: oral. C: left side. D: right side. and Portell (2001), we consider the very thin mar- gin, flat test with a gentle bevel leading adapically from the margin, and proportionately shorter petals distinct enough to warrant considering P. floridanus at the species level, rather than as a subspecies of P. lyelli. These features are very stable and consistent in mature specimens. Though P. floridanus is exceptionally abun- dant in the lower OLS, it is typically represented as a dense accumulation of fragments. Complete individ- uals are rare, due to the very large, thin, and fragile test of the species. In this horizon, P. floridanus oc- curs with an assemblage of much smaller scutelloids represented by the neolaganids W. inglisensis n. sp., D. ocalana, N. durhami, E. mooreanus, O. phelani, and many other, rarer species. Periarchus floridanus also occurs in the dolomite beds of the basal OLS, which form the lowest beds of the OLS and are well exposed along the Cross Florida Barge Canal south of Inglis. Here, P. floridanus is preserved as internal and external molds along with E. clevei. We figure a few of these molds (Figs. 152, 153) Periarchus quinquefarius (Say, 1825) (Figs. 154, 155) Scutella 5-faria Say, 1825. p. 228 (read to the academy in 1825, published in 1827). not Scutella rogersi (Morton). Agassiz, 1841. p. 85. pl. 19a, figs. 1-4. Mortonia rogersi (Morton). Desor, 1858. p. 231. Mortonia quinquefaria (Say). Conrad, 1866. p. 32. not Mortonella rogersi (Morton). Pomel, 1883. p. 70. not Scutella (Mortonia) rogersi (Morton). Gregorio, 1890. p. 250. pl. 43, figs. 18-19 (not fig. 16). ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 213 Figure 155: Periarchus quinquefarius (UF 338250), 53 mm TL, 54 mm TW, 8 mm TH, upper portion of Upper Eocene Ocala Limestone, Suwannee County, Florida (FM-IP SU003). A: aboral. B: oral. C: left side. D: right side. Mortonia quinquefaria (Say). Grabau and Shimer, 1910. p. 593. not Mortonella rogersi (Agassiz not Morton). Ste- fanini, 1911. p. 685. pl. 22, figs. 2, 3. Mortonella quinquefaria (Say). Clark and Twitchell, 1915. p. 128. pl. 60, figs. 2a-f; pl. 61, figs. la-b (includes additional synonymy). Periarchus quinquefarius (Say). Cooke, 1942. p. 15. Periarchus kewi Cooke, 1942. p. 16. pl. 1, figs. 12-14. Mortonella quinquefaria (Say). Mortensen, 1948. p. 391, txt figs. 228a-c. Mortonella quinquefaria (Say). Durham, 1955. p. 155. txt figs. 1L, 18c. Periarchus quinquefarius (Say). Cooke, 1959. pp. 43-44. pl. 14, figs. 6-8. Periarchus quinquefarius kewi (Cooke). Cooke, 1959. p. 44. pl. 14, figs. 4-5. Mortonella quinquefaria (Say). Durham, 1966. p. U477. figs. 368, 4. Periarchus quinquefarius kewi (Cooke). Pickering, 1970. p. 20. p. 28, p. 62. Periarchus quinquefarius (Say). Pickering, 1970. pp. 20, 33, 61. Periarchus quinquefarius (Say). Huddlestun and Hetrick, 1986. p. 15. Mortonella quinquefaria (Say). Mooi, 1989. fig. 27a. Mortonella quinquefarius kewi (Cooke). Oyen, 2001. pp. 41, 41. figs. 3-5, c-d. Mortonella quinquefarius (Say). Osborn et al., 2016. tbl. 2. Occurrence.—This species is the characteris- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 214 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) tic sand dollar of the Sandersville Limestone Mem- ber of the Tobacco Road Sand (sensu Huddlestun and Hetrick, 1986) in Georgia, especially near Sander- sville, Washington County. The species is exception- ally rare in Florida. Carter (1987a) and Oyen (2001) documented this species in the upper OLS of Suwan- nee County. We confirmed this after examining two specimens in the FM-IP collections, UF 5275 from the Watermelon Pit (FM-IP SU004) and UF 338250 from the greatly enlarged quarry operation currently known as the Branford 01A Quarry (FM-IP SU003), north of Branford, Suwannee County. Both speci- mens were collected ex situ, but the strata exposed in this quarry represent the Oligopygus wetherbyi Zone and overlying Wythella eldridgei Zone. There- fore, the specimens came from no lower than the Oligopygus wetherbyi Zone of the upper OLS. These are the only documented occurrences of this species in Florida. Discussion.—This species has the distinction of being the first echinoid described from North America. Thomas Say described the species in 1825 but did not figure it. Clark and Twitchell (1915) and Cooke (1959) attributed the species to Say’s date of reading (November 8, 1825) at the Academy of Natural Sciences in Philadelphia. However, Say’s paper: “On the species of the Linnaean genus Echinus, inhabiting the coast of the United States”, within which he described P. quinquefarius, was published in 1827 as clearly indicated on the title page. This led to some confusion and caused difficulty researching the initial description of P. quinquefarius. Agassiz (1841) was the first to figure and describe what was Say’s Scutella quinquefarius, al- though he called it Scutella rogersi Morton, 1834. Subsequently, Desor (1858) founded his genus Mor- tonia using the thick margin for this species as di- agnostic, but he replicated the error of Agassiz, and placed the species under the name Mortonia rogersi. The confusion between M. quinquefarius and C. rogersi was first noted by Gregory (1892) but it persisted until Clark and Twitchell (1915) clar- ified the synonymies of both species, which are very distinct. These latter authors properly appor- tioned which species were being referred to by which name in the previous literature. These cor- rections are provided in our synonymies for each species. Pomel (1883) changed the genus for this species from Mortonia to Mortonella because Gray (1851) had already used the name Mortonia for a subgenus of Echinocyamus (now considered a genus). Durham (1955) appealed to the Interna- tional Commission on Zoological Nomenclature to formalize this designation and asserted that Desor (1858) intended to designate the form recognized as Scutella quinquefarius Say, 1825, as the type species of his new genus Mortonia, and not Scutella rogersi Morton, 1834, and that misidentification of the two species created the confusion. The ICZN approved Durham’s requests in opinion 358 (1955). Although this species was the type of Mor- tonella Pomel (1883: 70), the differences between Mortonella and Periarchus were too insignificant for Cooke (1959) to justify the retention of Mortonella. The most conspicuous difference between the gen- era is in the thickness of the margin, but margin thickness is variable in both genera, and as clarified by Cooke, some individuals of P. quinquefarius are as thin as some of P. lyelli. Cooke (1959) asserted that if Mortonella is to be retained, it should not rank higher than subgenus. Durham (1966) retained Mortonella with- out commenting on Cooke’s (1959) assertion. Some subsequent workers have continued usage of Mortonella (Mooi, 1989; Carter, 1987a). Like Cooke (1959), we do not see sufficient differences between Mortonella and Periarchus to warrant retaining Mor- tonella as a distinct genus. Periarchus quinquefarius can be distinguished from the other species of Periarchus by its propor- tionately longer petals, its usually thicker margin, and by its deeper and more conspicuous food grooves (Cooke, 1959). Carter (1987a) was the first to document the occurrence of this species outside of Georgia when he listed it as occurring near Suwannee, Florida. Oyen (2001) subsequently documented numerous specimens in the Florida Museum collections from Florida [UF 2202, UF 2203, and UF 2204 (each lot consists of numerous specimens)]. However, associ- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 215 ated data attributes them to Georgia. Oyen (2001) also figured and described a specimen he referred to Mortonella quinquefarius kewi (UF 5275) from Florida, and this specimen is attributed to the upper OLS of Suwannee County. We have examined this specimen in the FM-IP collections, and it is indeed P. quinquefarius from the Watermelon Pit (FM-IP SU004), expanded and now known as the Branford 01A Quarry, (FM-IP SU003) north of Branford, Suwannee County. A second specimen attributable to this species was found recently in the Denali Quarry. These are the only confirmed occurrences of the species in Florida or anywhere south of the Suwannee Strait. The specimens (Figs. 154, 155) have a distinctly beveled margin characteristic of Cooke’s (1942) P. kewi. Cooke (1942) described Periarchus kewi as a distinct species. However, in 1959 he changed that opinion and reassigned it as a subspecies of P. quin- quefarius, mostly because it is associated with the typical form, whose margin often shows a tendency to become beveled. The distinguishing features of the variety are the strongly tumid central region, the swollen submargin, and the plainly beveled edge (Cooke, 1959). Cooke (1959) described his subspecies P. quinquefarius kewi as follows: hori- zontal outline circular; upper surface tumid medially, swollen beyond the petals, sloping steeply to the margin; margin thin; oral side flat; other features as in the typical variety. Because the typical form also shows a tendency for a beveled margin, as stated by Cooke (1959), and both forms occur together in Georgia, we cannot justify differentiating them at either the species or subspecies level. Genus Protoscutella Stefanini, 1924 Protoscutella pentagonium Cooke, 1942 (Fig. 156) Protoscutella pentagonium Cooke, 1942. p. 18, pl. 2, figs. 4-6. Protoscutella pentagonium (Cooke). Mortensen, 1948. p. 390. Protoscutella pentagonium (Cooke). Cooke, 1959. p. 39. pl. 15, figs. 6-8. not Protoscutella tuomeyi (Twitchell). Zachos and Molineux, 2003. pp. 495, 497. figs. 3.10-3.12, 3.14, 3.15 (in part, they referred specimens of P. pentagonium from the Weches Formation to P. tuomeyi). Protoscutella sp. (Cooke). Osborn et al., 2013. fig. 2. Occurrence.—The type locality of this species is a well of the Chipley Oil Company at Falling Waters, about 6.4 km south of Chipley, Washington County: depth unknown; Cooke (1942) stated that it was probably about 183 m (about 600 ft.), embedded in white sand (USGS 13145). It is not otherwise known from Florida. Cooke (1959) documented P. pentagonium in the Eocene of Texas, but Zachos and Molineux (2003) did not. They provided localities for P. tuomeyi in the Middle Eocene Weches Formation near Nacogdoches and San Augustine Texas, which we believe are likely P. pentagonium. As stated in Osborn et al. (2013), they are not P. tuomeyi. Discussion.—The type locality of P. pentago- nium is inconveniently located approximately 183 m below the surface from a well core in Washington County in the panhandle of Florida (Cooke, 1942; 1959). Cooke (1942) stated that the holotype was collected in strata likely attributable to the Lisbon Formation (Middle Eocene), though he was not certain of this. Bryan (2018) provided an updated log of the Chipley Oil Company Well (FGS W-1) with updated stratigraphic nomenclature that states strata at the 183 m level (about 600 ft.: depth of P. pentagonium provided by Cooke) is Claibornian and notes the presence of a small scutellid at 650 ft. They also note the presence of another scutellid fragment at ˜975 ft. in a clay bed, likely of the Lisbon Formation. This species has not otherwise been docu- mented from the fossil record of Florida. Subterclass ATELOSTOMATA von Zittel, 1879 Order SPATANGOIDA L. Agassiz, 1840 Suborder MICRASTERINA Fischer, 1966 incertae sedis Genus Gillechinus Fell, 1964 Gillechinus alabamensis (Cooke, 1942) (Fig. 157) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 216 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 156: Protoscutella pentagonium holotype (USNM 498992), 30.0 mm TL, 31.4 mm TW, 6.1 mm TH, well of the Chipley Oil Company at Falling Water, 4 miles south of Chipley, Washington County, Florida. A: aboral. B: oral. C: left side. D: right side. Eupatagus (Brissopatagus) alabamensis Cooke, 1942. p. 58. pl. 4, figs. 7, 8. Eupatagus (Brissopatagus) georgianus Cooke, 1942. p. 58. pl. 7, figs. 8-11. Eupatagus (Brissopatagus) alabamensis (Cooke). Cooke, 1959. p. 92. pl. 43, figs. 5-11. Gillechinus alabamensis (Cooke). Henderson and Fell, 1969. pp. 2-29. Brissopatagus alabamensis (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—Within Florida, this species is known from one documented specimen (UF 337963), from the uppermost portion of the OLS exposed in Brooks Quarry, near Marianna (FM- IP JA009), Jackson County. The species is more widespread in the Upper Eocene Shubuta Clay of Alabama and OLS of Georgia. The type locality of the species is St. Stephens Bluff, Washington County, Alabama. Discussion.—Cooke (1942) described Eu- patagus (Brissopatagus) georgianus from the OLS of Georgia and Eupatagus (Brissopatagus) alaba- mensis from the Shubuta Clay of Alabama. Cooke (1959) later placed E. georgianus in synonymy with E. alabamensis. We agree with his assessment. Henderson and Fell (1969) referred this species to Gillechinus. This species is rare throughout its distribution, and prior to Osborn et al. (2016), was undocumented in Florida deposits. Within Brooks Quarry (FM- IP JA009), a single, small (32 mm TL, 29 mm TW) specimen (UF 337963) of G. alabamensis was found with M. mortoni, O. haldemani, and W. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 217 Figure 157: Gillechinus alabamensis (UF 337963), 32 mm TL, 29.5 mm TW, uppermost portion of Upper Ocala Limestone in the Marianna Lime Quarry, Jackson County, Florida (FM-IP JA009). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 218 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) johnsoni, in the upper portion of the OLS. The reduced number of very large tubercles confined to the region aboral to the peripetalous fasciole in the paired interambulacra readily distinguishes this species from any other spatangoid echinoid in the Cenozoic fauna of the region. Suborder PALEOPLEUSTINA Markov and Solovjev, 2001 Family SCHIZASTERIDAE Lambert, 1905 Genus Ova Gray, 1825 Ova beckeri (Cooke, 1942) (Figs. 158-160) Schizaster beckeri Cooke, 1942. p. 40. pl. 3, figs. 5-8. Ditremaster beckeri (Cooke). Cooke, 1959. p. 77. pl. 30, figs. 9-12. Ditremaster beckeri (Cooke). Toulmin, 1977. pp. 339-340. pl. 65, figs. 11-13. Schizaster beckeri (Cooke). Kier, 1984. p. 57. Ditremaster beckeri (Cooke). Neraudeau, 1994. tbl 3. Schizaster beckeri (Cooke). Osborn et al., 2016. tbl 2. Occurrence.—Ova beckeri is most commonly found in the Wythella eldridgei Zone near the top of the upper OLS, especially south of Tennille (FM- IP DI001), west of Dowling Park (FM-IP LF002), and west of Center Hill in Sumter County (FM-IP SM010). The holotype (USNM 499008) was col- lected in the Gainesville Rock Pit near Arredondo, and the paratype (USNM 499009) is from the Suwan- nee River west of Dowling Park, Lafayette County. This species was documented in the Eocene of Al- abama by Toulmin (1977). Discussion.— Cooke (1942) described Schiza- ster beckeri from the OLS in Florida. Cooke (1942) stated that it differs from the other members of Schizaster in the region by its higher, more ro- tund test, and presence of only two gonopores. In this same work, he described Schizaster (Linthia) ocalanus, which also has only two gonopores, but he referred it to the subgenus Linthia. The traits Cooke (1942) listed do not serve to distinguish between O. ocalanus and O. beckeri. However, they are distinct, with O. beckeri being proportionately longer, with a shallower anterior sulcus, more posterior apical system, and anterior paired petals that are angled more anteriorly Cooke (1959) placed S. beckeri in Ditremas- ter, stating that Ditremaster differs from Schizaster by its lack of a deep, continuous, anterior sulcus and its apparent lack of lateral fascioles. Kier (1984: 57) subsequently stated that Ditremaster beckeri should be referred to Schizaster, as its latero-anal fasciole is distinct, whereas no latero-anal fasciole is present in Ditremaster. The two gonopores of Ova beckeri presently exclude it from placement in Schizaster (Smith and Kroh, 2011), as the membership in the latter is restricted to those forms with four gonopores. There- fore, we place S. beckeri Cooke, 1942 in Ova. Smith and Kroh (2011) list the very similar Schizaster subcylindricus (Cotteau, 1875) from the Eocene of St. Bartholomew, Cuba, and Jamaica as belonging to Ova but do not discuss O. beckeri. Cooke (1959) noted the latter’s similarity with Ova subcylindri- cus, indicating that it is very closely related but has somewhat shorter anterior petals. This affinity with O. subcylindricus is unde- niable and the two species are obviously closely related. Kier (1984) stated that these two species differ only in that the anterior petals in O. subcylin- dricus are slightly shorter and the apical system is more anterior. Kier indicated the anterior petals in O. beckeri have a length 48% TL as opposed to 34- 41% in O. subcylindricus. Ova beckeri is likewise similar in general form to Schizaster camagueyensis (Weisbord, 1934), from the Late Eocene of Cuba. However, the number of gonopores of this species is not provided in the original description of Weisbord (1934) or Kier (1984). In Florida, O. beckeri most commonly occurs near the top of the OLS with W. eldridgei, R. trojana, S. armiger, and many other, rarer species. The pres- ence of two gonopores and ovate, proportionately elongate test that lacks an anterior sulcus distin- guishes O. beckeri from any species with which it occurs. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 219 Figure 158: Ova beckeri (UF 5819), 33 mm TL, 31 mm TW, 24 mm TH, upper beds of Ocala Limestone, Lafayette County, Florida (FM-IP LF001). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 220 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 159: Ova beckeri (UF 322676), 27 mm TL, 25 mm TW, 19 mm TH, Upper Eocene upper Ocala Limestone, Taylor County, Florida (FM-IP 5649). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 221 Figure 160: Ova beckeri (UF 341764), 19 mm TL, 17 mm TW, 13 mm TH, upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA033). A: aboral. B: oral. C: left side. D: right side. E: anterior. F: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 222 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Ova ocalanus (Cooke, 1942) (Fig. 161) Schizaster (Linthia) ocalanus Cooke, 1942. p. 42. pl. 5, figs. 18-22. Schizaster (Brachybrissus) ocalanus (Cooke). Cooke, 1959. p. 73. pl. 29, figs. 13-17. Schizaster ocalanus (Cooke). Toulmin, 1977. p. 347. pl. 71, figs. 4-6. Schizaster ocalanus (Cooke). Osborn et al., 2016. tbl 2. Occurrence.—This species is most commonly found in the Oligopygus wetherbyi Zone of the OLS, where it is especially abundant near Kendrick, Mar- ion County (type locality), in the Cemex Quarry (FM-IP SM010) west of Center Hill, Sumter County, and the O’Brien Quarry in Suwannee County (FM- IP SU002); see Cooke (1959) for additional locali- ties. Discussion.—This species is more rotund than other American species of schizasterid, and al- though it has historically been referred to Schizaster (Cooke, 1942; 1959), clarification of the characters of the genus by Smith and Kroh (2011) indicate that this assignment is incorrect in the present con- cepts of these schizasterid genera. Cooke (1959) referred it to the subgenus Brachybrissus. However, Smith and Kroh (2011) considered Brachybrissus a synonym of Schizaster and referred schizasterids with a well-developed latero-anal portion of the marginal fasciole and two gonopores to Ova. We concur, placing S. ocalanus in Ova. The presence of two gonopores readily distinguishes this species from other schizasterids of the region except for O. beckeri, which is proportionately longer, with a shallower anterior sulcus, more posterior apical system, and anterior paired petals that are angled more anteriorly. Genus Schizaster L. Agassiz, 1836 Schizaster americanus Clark in Clark and Twitchell, 1915 (Figs. 162-168) Schizaster americanus Clark in Clark and Twitchell, 1915. p. 176. pl. 82, figs. 2a-d. Schizaster americanus (Clark). Lambert and Thiéry, 1925. p. 526. Schizaster americanus (Clark). Cooke, 1942. p. 40. Paraster americanus (Clark). Cooke, 1959. p. 72. pl. 30, figs. 5-8. Paraster americanus (Clark). Pickering, 1970. pp. 20, 28, 34. Schizaster (Paraster) americanus (Clark). Dockery, 1980. p. 193. pl. 82, fig. 1. Schizaster americanus (Clark). Oyen and Portell, 2001. p. 202. Schizaster americanus (Clark). Osborn and Ciampaglio 2014. p. 142. Schizaster sp. cf. S. americanus (Clark). Donovan et al., 2015. pp. 5-6. fig. 3 (not S. americanus). Occurrence.—Within Florida, this species is most abundant in the Lower Oligocene Marianna Limestone where it occurs in the Brooks Quarries (FM-IP JA013, FM-IP JA019, FM-IP JA026, FM-IP JA029) near Marianna, Jackson County; Dry Creek (FM-IP JA010), south of Marianna, Jackson County; and in a dolomite quarry north of Altha, Jackson County (FM-IP JA014). Schizaster americanus is much more rarely found in the Suwannee Limestone in the Terramar Quarry (FM-IP PO017) in Polk County, and the Bridgeboro Limestone in the Dun- can Church Quarry (FM-IP WG002) in Washington County. Schizaster americanus also occurs in Oligocene strata of Alabama, Georgia, and Missis- sippi. The type locality is the Marianna Limestone near Brandon, in Rankin County, Mississippi. Discussion.—Schizaster americanus is the only spatangoid present in the typical Marianna Limestone in Jackson County, Florida. However, in the dolomitic facies of the Marianna Limestone, ex- posed along the banks of Dry Creek (FM-IP JA010) south of Marianna, S. americanus occurs with P. cassadyi n. sp. and C. rogersi. A few kilometers eastward in quarries north of Altha (FM-IP JA014), also in the dolomitic facies, S. americanus occurs with E. aldrichi and C. rogersi. See the discussion for the Marianna Limestone in the stratigraphic overview section for more on these dolomitic facies ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 223 Figure 161: Ova ocalanus (UF 338006), 23 mm TL, 24 mm TW, 19 mm TH, Oligopygus wetherbyi zone of Upper Eocene upper Ocala Limestone, Sumter County, Florida (FM-IP SM010). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 224 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 162: Schizaster americanus (UF 337961), 35.5 mm TL, 33.5 mm TW, 24.5 mm TH, Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA029). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 225 Figure 163: Schizaster cf. S. americanus (UF 112437), 57 mm TL, 52.5 mm TW, 36.5 mm TH, dolomitic portion of Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA014). A: aboral. B: oral. C: left side. D: posterior. in southern Jackson County. Schizaster americanus is much more rarely found in the Suwannee Limestone and is repre- sented by one specimen (UF 27345) from the Terramar Quarry (FM-IP PO017) in Polk County. It is also rarely found in the Bridgeboro Lime- stone and is represented by three specimens [UF 55006 (2 specimens), UF 61440] from the Duncan Church Quarry in Washington County (FM-IP WG002). Cooke (1959) noted that S. americanus is proportionately shorter than S. armiger, and that its posterior petals are somewhat shorter. However, S. americanus demonstrates considerable variability in the degree of divergence of the anterior petals and position of its apical system (often more posterior in specimens from the Chickasawhay Limestone of Alabama). A series of specimens preserved as molds from the dolomitic facies of the Marianna Limestone north of Altha (FM-IP JA014) show significant vari- ation in apical system placement (more posterior in some specimens), degree of divergence of ambu- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 226 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 164: Schizaster cf. S. americanus (UF 112449), 58 mm TL, 56 mm TW, 37 mm TH, dolomitic portion of Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA014). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 227 Figure 165: Schizaster cf. S. americanus (UF 112348), 52.5 mm TL, 49.5 mm TW, 33.5 mm TH, dolomitic portion of Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA014). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 228 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 166: Schizaster cf. S. americanus (UF 112344), 34.5 mm TL, 33 mm TW, 22.5 mm TH, dolomitic portion of Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA014). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 229 Figure 167: Schizaster americanus (UF 55006), 31 mm TL, 29.5 mm TW, 23.5 mm TH, Oligocene Bridgeboro Limestone, Washington County, Florida (FM-IP WG002). A: aboral. B: oral. C: left side. D: right side. lacra II and IV, and height of the anterior portion of the test, with some being much more wedge-shaped in profile than is typical for the species (Fig. 165). Schizaster armiger Clark in Clark and Twitchell, 1915 (Figs. 169, 170) A sea urchin, Harris, 1894. p. 172. pl. 6, fig. 11. Schizaster armiger Clark in Clark and Twitchell, 1915. p. 152. pl. 70, figs. 1a-d. Schizaster floridanus Clark in Clark and Twitchell, 1915. p. 175. pl. 82, figs. 1a-c. Schizaster armiger (Clark). Lambert and Thiéry, 1925. p. 524. Schizaster armiger (Clark). Cooke, 1942. p. 39. Schizaster armiger (Clark). Cooke, 1948b. p. 92. pl. 22, figs. 1-4. Paraster armiger (Clark). Cooke, 1959. pp. 71-72. pl. 30, figs. 13-15. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 230 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 168: Schizaster americanus (UF 27345), 20 mm TL, 21 mm TW, 15 mm TH, Lower Oligocene Suwannee Limestone, Polk County, Florida (FM-IP PO017). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 231 Figure 169: Schizaster armiger (UF 114429), 74.5 mm TL, 72 mm TW, 37.5 mm TH (specimen is slightly compressed), upper portion of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF002). A: aboral. B: oral. C: left side. D: posterior. Schizaster (Paraster) armiger (Clark). Toulmin, 1977. p. 346. pl. 70, figs. 5-7. Schizaster armiger (Clark). Osborn et al., 2016. tbl 2. Occurrence.—This species is present in the upper OLS where it is most commonly encountered in association with W. eldridgei in the uppermost por- tion of the unit. A few localities include: Johnson’s Sink, Levy County (type of Schizaster floridanus Clark in Clark and Twitchell, 1915); northwest of Mayo (FM-IP LF001), Lafayette County; north of Branford (FM-IP SU002), Suwannee County; and south of Tennille (FM-IP DI001), Dixie County. Schizaster armiger is also found in Upper Eocene strata of Alabama, Arkansas, Georgia, Mis- sissippi, and Panama (Cooke, 1948b; 1959). The holotype of S. armiger (USNM 141104) was col- lected in the Upper Eocene Yazoo Formation near the Cocoa Post Office, Choctaw County, Alabama. Discussion.—Gilbert Harris first documented this species in the Annual Report of the Geologi- cal Survey of Arkansas for 1892 (1894: 172), in which he identified it as “A Sea Urchin” from Cornish Ferry on the Sabine River. The figure he provided was good enough to permit its identifi- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 232 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 170: Schizaster armiger (UF 114448), 50 mm TL, 47 mm TW, 36.5 mm TH, Upper Eocene Ocala Limestone, Alachua County, Florida (FM-IP AL004). A: aboral. B: oral. C: left side. D: posterior. cation as S. armiger, which Clark, in Clark and Twitchell (1915), subsequently described from a specimen collected in the Upper Eocene strata of Choctaw County, Alabama. Cooke (1942) acknowl- edged the identification of Harris’ specimen when he documented the occurrence of S. armiger at the east bank of the Sabine River at Cornish Ferry (Caveness Landing) about 8 km east-northwest of Warren, Bradley County, Arkansas, and listed G. B. Harris as the collector of the specimen. Clark in Clark and Twitchell (1915), described S. floridanus from Johnson’s Sink in Levy County, Florida, in the same work in which he described S. armiger, though Cooke (1942) considered S. flori- danus a synonym of S. armiger. Subsequent authors have followed this synonymy, as do we. Cooke (1959) stated that S. armiger is propor- tionately longer, and its posterior petals are some- what longer than in S. americanus. Cooke (1959) also stated that the type and other specimens from ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 233 western Alabama have wider petals than those from Florida and that there is some variation in the rel- ative length of the posterior petals, an observation we have also made. Schizaster armiger is commonly encountered in the Wythella eldridgei Zone in the upper OLS of the northern Florida peninsula where it occurs with W. eldridgei and a diverse assemblage of spatan- goids, including P. dixie, E. ocalanus, P. curvus, B. steinhatchee, and O. beckeri. Schizaster carlsoni n. sp. (Figs. 171-173) Diagnosis.— Schizaster with high (TH on average 70% TL), narrow test (TW on average 90.8% TL) that is not sharply wedge-shaped in lateral view, with a nearly central apical system (center of apical system on average 47.3% TL from posterior margin), petals of ambulacra I and V short (length on average 21.9% TL) and diverging on average at 47° from each other, petals of ambulacra II and IV long, extending nearly to margin (length on average 40.4% TL), slightly sinuous, and divergent on average 95° from each other, curving towards the anterior until their ends, at which point they curve slightly posteriorly. Description.—Description based on the holo- type (UF 342111), three paratypes (UF 342112, UF 342113, UF 342114), and non-type material, all from the type locality (FM-IP HE038). Moderately sized Schizaster, largest com- plete specimen (UF 342114) 50.2 mm TL, small- est (UF 342111) 38.0 mm TL; test ovate, narrow, narrowing posteriorly, TW on average 90.8% TL; test high, TH on average 70% TL, greatest height posterior, test slopes gently anteriorly from api- cal; only slightly wedge-shaped in lateral view- point; greatest width nearly central, adjacent to apical area; test narrows laterally above ambitus towards aboral highest point of test. Apical system ethmolytic; nearly central, center of apical system on average 47.3% TL from posterior margin; four gonopores, posterior pair slightly larger and fur- ther apart than anterior pair. Ambulacrum III not petaloid, in very deep, steep-sided depression ex- tending from apical system to anterior notch, and continuing to the peristome, depression deepest and widest midway between apical system and anterior margin; furrow on average 12% TW at widest point, sides of furrow nearly vertical, often overhung, en- larged pore pairs present from apical system to about two-thirds distance to margin. Pore-pairs of paired petals strongly conjugate; interporiferous zones slightly wider than single poriferous zone. Petals of ambulacra I and V short, extending less than halfway from apical system to margin (length on average 21.9% TL); narrow (width on average 34% petal length); bent sharply posteriorly, in deep, ditch-like depression; on average divergent from each other by 47°, range 37-55°; widest point me- dially; tapered but open distally. Petals of ambu- lacra II and IV slightly sinuous, divergent from each other on average 95°; curved towards anterior until end where they often curve slightly posteri- orly; long, extending more than two-thirds distance from apical system to margin; length on average 40.4% TL; widest point medially, tapered but open distally. Periproct small (periproct height on aver- age 26.5% TH), ovate, taller than wide (width on average 49.7% periproct height), high on near- vertical, slightly overhung truncate face, lower edge of periproct on average 39.6% TH above posterior margin. Peristome anterior, posterior edge on aver- age 63% TL from posterior margin, kidney shaped, open towards anterior; wider than high, height on average 43.4% peristome width. Labrum short and wide, plastron wide posteriorly, widest point on average 14.5% TW, tapers anteriorly. Tubercles on aboral surface minute, uni- form, evenly and densely distributed. Oral tuber- culation fine, dense and uniform. Well-developed peripetalous and lateral fascioles present, but not completely discernable on any specimen (largely due to air abrasive preparation required to remove specimens from stubborn matrix). Zoobank Nomenclatural Act.—451038E1- 8AED-4A66-9C4A-0699500242E9 Discussion.— Schizaster carlsoni n. sp. has not been documented outside of the basal portion of the Lower Oligocene Suwannee Limestone in ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 234 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 171: Schizaster carlsoni n. sp., holotype (UF 342111), 38.0 mm TL, 34.1 mm TW, 25.2 mm TH, lower bed of the Lower Oligocene Suwannee Limestone, Vulcan Quarry northwest of Brooksville, Hernando County, Florida (FM-IP HE038). A: aboral. B: apical area. C: oral. D: anterior. E: oblique lateral viewpoint from anterior. F: posterior. G: left side. H: peristome. I: oblique lateral viewpoint from posterior. J: petaloid portion of ambulacrum I. K: petaloid portion of ambulacrum IV. L: ambulacrum III and apical area. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 235 Figure 172: Schizaster carlsoni n. sp., paratype (UF 342112), 50.9 mm TL (TL not complete due to damaged posterior), 46.8 mm TW, 34.0 mm TH, lower bed of the Lower Oligocene Suwannee Limestone, Vulcan Quarry northwest of Brooksville, Hernando County, Florida (FM-IP HE038). A: aboral. B: oral. C: anterior. D, G: oblique aboral viewpoints from anterior. E: left side. F: posterior. H: apical and ambulacrum III. I: apical area. J: tilted viewpoint of petaloid area from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 236 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 173: Schizaster carlsoni n. sp., paratype (UF 342113), 41.0 mm TL, 36.9 mm TW, 27.0 mm TH (complete TW and TH obscured by matrix), lower bed of the Lower Oligocene Suwannee Limestone, Vulcan Quarry northwest of Brooksville, Hernando County, Florida (FM-IP HE038). A: aboral. B: anterior. C: oblique aboral viewpoint. D: tilted viewpoint of petaloid area from posterior. E: left side. F: apical area. G: peristome. H: petaloid portion of ambulacrum II. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 237 the Vulcan Quarry, roughly 5 mi. northwest of Brooksville, Hernando County, Florida (FM-IP HE038). The horizon bearing S. carlsoni n. sp. is be- low the general floor of the quarry and usually only exposed during the excavation of drainage basins and canals. Schizaster carlsoni n. sp. is rarely found with more abundant Clypeaster sp. B, R. gouldii, which are typically larger in this horizon than in the overlying beds of the Suwannee Limestone, E. dumonti n. sp., and rare specimens of Phyllacanthus cf. P. mortoni. Schizaster carlsoni n. sp. joins S. americanus, from which it is readily distinguished by its narrower test and longer and more sinuous ambulacra II and IV, as the only other Schizaster documented from the Oligocene of North America. Schizaster carlsoni n. sp. is not similar to the other five species of Schizaster that occur in the Cenozoic deposits of the extended region, which in- cludes: S. alabamensis Clark in Clark and Twitchell, 1915 (Lower Paleocene of the Gulf Coast), S. cad- doensis Zachos in Zachos and Molineux, 2003 (Mid- dle Eocene of Texas), S. armiger (Upper Eocene of Florida and the Gulf Coast), S. susana Zachos in Zachos and Molineux, 2003 (Upper Eocene of Texas and Louisiana), and S. kieri (Pliocene of Florida); not S. ocalanus, S. beckeri, or S. stenzeli Zachos in Zachos and Molineux, 2003, which have only two gonopores and are placed in Ova. The modern Schizaster of the region includes S. doederleini (Chesher, 1972) and S. floridiensis (Kier and Grant, 1965), but not S. orbignyanus Agassiz, 1880, which only has two gonopores and therefore belongs in Ova. Schizaster is abundantly represented in the fos- sil faunas of the remainder of the eastern Americas and the Caribbean region. We compare S. carlsoni n. sp. to all of them, below. Kier (1984) recognized 16 distinct species of Schizaster in Eocene to Miocene strata of Cuba. These 16 species are reduced from 37 species previ- ously documented in the Cuban faunas, largely in the works of Sánchez-Roig. Of the 21 species Kier (1984) did not recognize, he noted the types of ten are lost or too poorly preserved to permit recognition of distinguishing characters, one is transferred to a different genus, and the remainder are considered subjective junior synonyms of the 16 species he retained. Among the Cuban species, 13 of the 16 species recognized by Kier (1984) are: S. bathy- petalus Clark, in Arnold and Clark, 1927, Eocene (this species was initially described from Jamaica); S. cartagensis (Sánchez-Roig, 1949), Oligocene- Miocene; S. delgadoi (Sánchez-Roig, 1953b), Late Oligocene; S. egozcuei Lambert, 1925, Oligocene- Miocene; S. fernandezi Sánchez-Roig, 1952c, Miocene; S. formelli Kier, 1984, Eocene; S. gerthi Pijpers, 1933, Eocene, initially described from Bonaire; S. llagunoi Lambert and Sánchez-Roig in Sánchez-Roig, 1949, Eocene-Oligocene; S. munozi Sánchez-Roig, 1949, Oligocene-Miocene; S. neuvi- tasensis (Weisbord, 1934), Late Eocene; S. rojasi Sánchez-Roig, 1952c, Oligocene-Miocene; S. sanc- tamariae Sánchez-Roig, 1949, Oligocene, and S. santanae Sánchez-Roig, 1949, Late Eocene. Agassiz (1872) placed S. cubensis in Meoma. Kier (1984) omitted Schizaster cubensis D’Orbigny, 1847 from his catalog of Cuban Schizaster because he considered the form unrecognizable beyond the genus level. However, the taxon is notable because it seems to be the member of the genus attributed to the Pliocene of Cuba. The remaining three species Kier (1984) rec- ognized are S. subcylindricus Cotteau, 1875, initially described from the Eocene of St. Bartholomew, which has two gonopores and therefore belongs in Ova, and two species which are very similar to it: Schizaster cubitabellae (Weisbord, 1934) and S. camagueyensis (Weisbord, 1934), both from the Late Eocene of Cuba. These two species are very similar to O. subcylindricus as well as O. beckeri. However, the apical systems appear to be damaged in the types, thus the number of gonopores is not apparent in their figures and is also not provided in the descriptions of Weisbord (1934) and Kier (1984). Both both species could be conspecific with O. subcylindricus and therefore a member of Ova (having two gonopores). In addition, the type and only known specimen of S. cubitabellae is only 50% complete, making determination of its features difficult. Specimens with better preserved apical ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 238 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) systems would be required to establish their genus placement with certainty. Of these Cuban taxa, S. carlsoni n. sp. is most like S. bathypetalus Clark, 1927, and the similarities of these two taxa are discussed below. However, S. carlsoni n. sp. is also like S. cartagensis. The holotype of this species is well-figured by Kier (1984: pl. 16, figs. 4-6), and has a similar globose form that lacks a sharply wedge-shaped profile, a more anterior apical system, and long petals. How- ever, ambulacrum II and IV of S. cartagensis are more divergent and less sinuous than S. carlsoni n. sp. The fossil record of Jamaica shares S. bathy- petalus and O. subcylindricus with Cuba, and also includes the following species: S. altissimus Clark, in Arnold and Clark, 1927, from the Eocene; S. dum- blei Israelsky, 1924, which was initially described from Mexico; however, Kier (1984) thought the specimens Arnold and Clark (1927) documented from Jamaica are likely O. subcylindricus, though Donovan (2003) did not entirely agree, and S. hexag- onalis Clark, 1927, from the Eocene. Specimens described from Jamaica, but no longer recognized in Schizaster, are S. dyscritus Clark, in Arnold and Clark, 1927, which was moved to Caribbaster by Kier (1984), and S. brachypetalus Clark, in Arnold and Clark, 1927, which Kier (1984) demonstrated is a synonym of O. subcylindricus. Of these taxa, S. carlsoni n. sp. can only be confused with S. bathy- petalus, which as mentioned above, also occurs in the Eocene of Cuba. Schizaster bathypetalus is figured by Arnold and Clark (1927: pl. 12, figs. 1-4), and Kier (1984: pl. 15, fig. 2) persuasively figured the holotype (MCZ 3294). Kier (1984) reasonably placed Schizaster gigas Sánchez-Roig, 1953c and Schizaster pentag- onalis Sánchez-Roig, 1953c in synonymy with S. bathypetalus. Schizaster carlsoni n. sp. can be dis- tinguished from the type of S. bathypetalus by its proportionately shorter posterior petals. Kier noted the holotype of S. bathypetalus has posterior petals 34% TL, whereas in S. carlsoni n. sp., they range from 20-25% TL (21.9 % TL average). Furthermore, Clark (in Arnold and Clark, 1927), stated TW is practically equal to TL on S. bathypetalus, but in S. carlsoni n. sp., TW is on average 90% TL. Cotteau (1875) described the previously dis- cussed O. subcylindricus from the Eocene of St. Bartholomew, but he also named S. antillarum Cot- teau, 1875, from St. Barts, as well as S. clevei Cotteau, 1875, from the Oligocene of Antigua and Anguilla, and S. loveni Cotteau, 1875, from An- guilla, which was also documented in Puerto Rico (Jackson, 1922; Gordon, 1963). Schizaster carlsoni n. sp. cannot be confused with any of Cotteau’s species. The fossil faunas of eastern Mexico contain the previously mentioned S. dumblei, which also questionably occurs in Jamaica; S. cristatus Jackson, 1917, from the Oligocene (S. cristatus was initially described from the Miocene of Panama; see below); S. tampicoensis Israelsky, 1924, from the Miocene, and S. scherzeri Gabb, 1881, originally described from the Miocene of Costa Rica (see also Dickerson and Kew, 1917). Schizaster carlsoni n. sp. cannot be confused with any of these Mexican species. In addition, S. eustatii Engle, 1961 was de- scribed from the Pleistocene of St. Eustatius, based on a poorly preserved specimen that clearly has shorter posterior ambulacra than S. carlsoni n. sp., among other distinguishing features. Jackson (1917) described S. panamensis and S. cristatus from the Miocene of Panama. However, Kier (1984: 10) stated both Panamanian species are too poorly preserved for comparisons. Durham (1961) described S. costari- censis Durham, 1961, from the Miocene of Costa Rica. However, as well documented by Durham (1961) and Fischer (1985) this species has only two gonopores and should be reassigned to Ova. Therefore, with the addition of S. carlsoni n. sp., Schizaster includes no fewer than 34 species in the Cenozoic fossil faunas of the eastern Americas and Caribbean region. However, as discussed above, many of these species are inadequately described and figured, but all readily distinguished from S. carlsoni n. sp. Etymology.—Named in honor of Robert Carl- son of St. Petersburg Florida, collector of the type material. Material and Occurrence.—Holotype (UF 342111) and paratypes (UF 342112, UF 342113, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 239 UF 342114) from the lower beds of the Suwan- nee Limestone in the Vulcan Quarry, northwest of Brooksville, Hernando County (FM-IP HE038). Family PRENASTERIDAE Lambert, 1905 Genus Agassizia Valenciennes, 1846 Agassizia clevei Cotteau, 1875 (Figs. 174, 175) Agassizia clevei Cotteau, 1875. p. 33. pl. 6, figs. 2-8 (in part, not figs. 9, 10). Agassizia floridana de Loriol, 1887. p. 398. pl. 17, figs. 9-9f. Agassizia conradi (Bouvé). Clark and Twitchell, 1915. p. 174. pl. 81, figs. 3a-d. Agassizia clevei (Cotteau). Jackson, 1922. p. 71. pl. 12, figs. 5-7. Agassizia inflata Jackson. Arnold and Clark, 1927. p. 56. Agassizia caribbeana Weisbord, 1934. p. 238. pl. 27, figs. 1-6. Agassizia camagueyana Weisbord, 1934. p. 247. pl. 9, figs. 5,6. Agassizia floridana (de Loriol). Cooke, 1942. p. 44. pl. 3, figs. 1-4. Agassizia clevei (Cotteau). Sánchez-Roig, 1949. p. 256. Agassizia avilensis Sánchez-Roig, 1949. p. 260. pl. 8, figs. 1-6. Agassizia camagueyana (Weisbord). Sánchez-Roig, 1949. p. 253. Agassizia floridana (de Loriol). Fischer, 1951. p. 73. pl. 6, figs. 3, 4. Agassizia floridana (de Loriol). Cooke, 1959. p. 75. pl. 32, figs. 1-4. Agassizia clevei (Cotteau). Gordon, 1963. p. 640. fig. 1. pl. 80, figs. 1-3. Agassizia floridana (de Loriol). Toulmin, 1977. p. 338. pl. 64, fig. 1-3. Agassizia clevei (Cotteau). Kier, 1984. pp. 62-65. fig. 23. pls. 29-31. Agassizia clevei (Cotteau). Osborn et al., 2016. tbl 2. Occurrence.—Agassizia clevei occurs throughout the OLS. It is commonly encountered in the upper OLS in the Brooks Quarries (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031), northwest of Marianna, Jackson County; quarry northwest of Mayo (FM-IP LF001), Lafayette County; and numerous other localities. This species is also widespread in the Oligopygus phelani Zone of the lower OLS, especially along the Withlacoochee River west of Yankeetown (FM-IP LV024), Levy County and along the Cross Florida Barge Canal (FM-IP CI001), south of Inglis, Citrus County. The type locality of A. floridana is near Gainesville (de Loriol, 1887). This species is also found in the OLS of Geor- gia (Cooke, 1959), and throughout the Caribbean re- gion. In Cuba [(type of A. caribbeana, Late Eocene; type of A. camagueyana, Oligo-Miocene; and type of A. avilensis, Oligo-Miocene (Kier, 1984)]. In Anguilla [(type of A. clevei, Miocene Anguilla For- mation (Cotteau, 1875; Kier, 1984)]; Puerto Rico (Gordon, 1963); and Trinidad [Eocene (Jeannet, 1928)]. Discussion.—Bouvé (1851) described Hemi- aster conradi from Paleogene strata of Georgia. However, his brief description and sketched outlines of the specimen are not diagnostic. The test is most likely an Agassizia and, as it was found with E. patel- liformis (the other species Bouvé described from the deposit), it is perhaps what is now referred to A. clevei. However, this cannot be determined with certainty. De Loriol (1887) described Agassizia flori- dana from the Eocene of Florida. However, Clark and Twitchell (1915) believed A. floridana was a subjective junior synonym of Bouvé’s H. conradi, which they placed in Agassizia. Cooke (1942: 45) invited H. L. Clark to examine Bouvé’s type of H. conradi in the Museum of Comparative Zoology, and he notes Clark stated that specimen is “abso- lutely unidentifiable even to the family, let alone genus or species.” Therefore, Cooke (1942) recog- nized de Loriol’s A. floridana for the specimens from Florida and Georgia that he discussed. However, if Bouvé’s H. conradi were recognizable, and verified as conspecific, it would have precedence over A. clevei, as well as de Loriol’s (1887) A. floridana. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 240 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 174: Agassizia clevei (UF 329693), 24.5 mm TL, 23 mm TW, 19 mm TH, Oligopygus haldemani Zone of Upper Eocene upper Ocala Limestone, Jackson County, Florida (FM-IP JA009). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 241 Figure 175: Agassizia clevei (UF 337959), 25.5 mm TL, 21.5 mm TW, 19 mm TH, Oligopygus haldemani Zone Upper Eocene Ocala Limestone, Sumter County, Florida (FM-IP SM010). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 242 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Kier (1984) compared the Cuban species of Agassizia with their North American and Caribbean region counterparts. Kier (1984) studied many spec- imens of A. floridana from the Late Eocene of Florida and could not distinguish them from the types of A. clevei from Anguilla. He searched for differences with particular care because the Florida specimens are Eocene and those from Anguilla are Miocene, and he acknowledged that echinoids do not commonly have such a wide stratigraphic range. However, he could find no substantive differences. He therefore considered A. floridana a subjective junior synonym of A. clevei. Cooke (1942, 1959) considered A. inflata Jackson, 1922 and A. egozcuei Lambert, 1925, from the Eocene of St. Bartholomew, to be synonyms of A. floridana. Kier (1984) asserted that A. egozcuei is a synonym of A. inflata, also stating that A. inflata can be distinguished from A. clevei by its more anterior apical system and possession of a few enlarged pore pairs in the anterior poriferous zone. He stated that in none of the 17 specimens of A. floridana (= A. clevei) he examined were large pore pairs present in the anterior poriferous zones. Within the eastern United States, A. clevei is most found in the Upper Eocene OLS of Florida where it is a consistent presence in the Oligopygus haldemani Zone, but also occurs more rarely in the overlying Oligopygus wetherbyi Zone, as well as in the underlying Oligopygus phelani Zone in Citrus and Levy Counties. It is much more rarely represented in the equivalent strata in southwestern Georgia. The species is typically small but can reach significant size, the largest specimen in the USNM, from the Steinhatchee River in Florida, being 36 mm TL, 34 mm TW, and 28 mm TH (Cooke, 1942). Agassizia mossomi Cooke, 1942 (Figs. 176, 177) Agassizia (Anisaster) mossomi Cooke, 1942. p. 46. pl. 5, figs. 14-17. Agassizia (Anisaster) mossomi (Cooke). Cooke, 1959. p. 76. pl. 32, figs. 5-9. Agassizia mossomi (Cooke). Kier, 1997. pp. 10, 11. fig. 5. pl. 7, figs. 1-8; pl. 8, figs. 1-8. Agassizia mossomi (Cooke). Zachos and Molineux, 2007. pp. 79-91. fig. 5e. Agassizia mossomi (Cooke). Osborn and Ciampaglio, 2014. p. 142. Occurrence.—Within Florida, A. mossomi oc- curs in the Lower Oligocene Suwannee Limestone at numerous localities, including quarries west of Brooksville (FM-IP HE019), Hernando County and in the Bridgeboro Limestone south of Chipley (FM- IP WG002), Washington County. The type locality is in the Suwannee Limestone at the Florida Rock Prod- ucts Quarry southwest of Brooksville, Hernando County. It is perhaps most abundant in the Suwannee Limestone at the Terramar Quarry in Polk County (FM-IP PO017). This species is more commonly found in the Oligocene of Alabama (Cooke, 1959; Osborn and Ciampaglio, 2014), Georgia (Cooke, 1959), Missis- sippi (Cooke, 1959), North Carolina (Kier, 1997), and Texas (Zachos and Molineux, 2007). Discussion.—Although Cooke (1942) doc- umented this species from the Lower Oligocene Suwannee Limestone of Florida, it has since been proven to be more widely distributed in the region. Agassizia mossomi occurs throughout nearly the entire extent (both geographically and stratigraph- ically) of the Oligocene strata of the region, from the Lower Oligocene Marianna Limestone of the Gulf Coast to the Upper Oligocene Chickasawhay Limestone of Alabama and River Bend Formation of North Carolina. Cooke (1959) stated that A. mossomi is higher and more nearly spherical, its apex is farther forward, its petals are more deeply sunken, and the anterior pores of the front pair of ambulacra are more fully developed than in A. floridana (= A. clevei) or A. wilmingtonica (= A. inflata). The presence of these pore pairs in the distal half of the anterior column of ambulacra II and IV has caused this species to be placed in the subgenus Anisaster by some authors (e.g., Cooke, 1959). This species cannot be confused with any echinoid documented from Oligocene deposits in the region. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 243 Figure 176: Agassizia mossomi (UF 344754), 21 mm TL, 20 mm TW, 19 mm TH, Lower Oligocene Suwannee Limestone, Polk County, Florida (FM-IP PO017). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 244 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 177: Agassizia mossomi (UF 344755), 15 mm TL, 14 mm TW, 13 mm TH, Lower Oligocene Suwannee Limestone, Polk County, Florida (FM-IP PO017). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 245 Figure 178: cf. Prenaster sp. (UF 337992), 64.2 mm TL, 57.4 mm TW, 49.5 mm TH, collected from the Haimea brooksi Zone, Upper Eocene Ocala Limestone, dredged ∼80 ft. below top of unit and overlying Bumpnose Limestone, Brooks Quarry, northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: posterior. C: oblique lateral viewpoint from posterior. D: oral. E: left side. F: spines near lateral ambitus near posterior. G: oblique aboral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 246 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 179: Brissopsis steinhatchee (UF 322462), 52 mm TL, 42.5 mm TW, 24 mm TH, Uppermost portion of Upper Eocene Ocala Limestone, Suwannee County, Florida (FM-IP SU002). A: aboral. B: oral. C: left side. D: right side. E: posterior. Genus Prenaster Desor, 1853 cf. Prenaster sp. (Fig. 178) Discussion.—This specimen in unlike any species currently documented from the North Amer- ican echinoid faunas. It was collected in the OLS, in the Haimea brooksi Zone, dredged 29-30 m below the top of the Eocene exposures within the Brooks Quarry northwest of Marianna, Jackson County (FM-IP JA039). Some test is preserved, but the specimen largely consists of an internal mold of a rather large echinoid. The specimen is 64 mm TL, 57 mm TW, and 49 mm TH, although none of these measurements indicate actual dimensions of the complete test. The specimen is rotund, with a nearly com- pletely preserved flat posterior surface with the periproct situated high on the posterior face. The apical system is missing, but based on the pre- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 247 Figure 180: Brissopsis cf. B. steinhatchee (UF 2122), 96.5 mm TL, 72 mm TW, 34.5 mm TH, an exceptionally large specimen, Oligopygus phelani Zone of Upper Eocene lower Ocala Limestone, Levy County, Florida (FM-IP 2409). A: aboral. B: oral. C: left side. D: posterior. served portions of the ambulacra, would have been positioned far anteriorly on the aboral surface. A few spines are preserved on the posterior and oral surfaces. Faint traces of a lateroanal fasciole are discernable low on the posterior margin. We are reluctant to do any additional preparation to further expose any fascioles given the incompleteness of the test, particularly considering that even if fascioles were further exposed, they would not be sufficient to determine either genus or species of the specimen. We likewise prefer not to risk removal of any of the preserved spines. What is preserved of the specimen is not dis- tinguishable from Prenaster, especially the general test shape and very anterior apical system. The spec- imen is most like Prenaster jeanneti Pijpers, 1933, from the Eocene of Bonaire, which was well figured by Kier (1984: pl. 41, figs. 6-8). The Florida speci- men is slightly larger than the maximum size of 52 mm TL, 45 mm TW of P. jeanneti. If additional material is collected that enables the genus identification of Prenaster to be confirmed, this is the first documentation of Prenaster on main- land North America. However, it is represented in the Eocene of Bonaire, as discussed above, and by five species in the Cuban Eocene faunas: P. clarcki Sánchez Roig, 1949 [which Kier (1984) corrected to P. clarki], P. elongatus Sánchez Roig, 1949, and P. sanchezi Lambert, in Sánchez Roig, 1949 [Kier (1984) asserted these three species were unrecogniz- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 248 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) able]; P. nuevitasensis Sánchez Roig, 1949 (which was transferred to Aguayoaster by Zitt, 1981), and P. parvus Palmer, in Sánchez Roig, 1949, which Kier (1984) redescribed and is readily distinguish- able from this specimen. Given the presence of Prenaster in the Cuban faunas, its presence in the Florida Eocene would not be surprising. Carolinaster varnami Osborn et al., 2016 is comparable in general prenasterid features to the Florida specimen. However, without further infor- mation concerning genus features of cf. Prenaster sp., notably the number of gonopores, it is not cer- tain that the two forms are congeneric, let alone conspecific. Chesher (1968) described Saviniaster enoda- tus, which has subsequently been placed in Pre- naster, from the modern faunas off the Bahama Islands. Thus, the genus is known from regional waters today. Suborder BRISSIDINA Kroh and Smith, 2010 Family BRISSIDAE Gray, 1855 Genus Brissopsis L. Agassiz, 1840 Brissopsis steinhatchee Cooke, 1942 (Figs. 179, 180) Brissopsis steinhatchee Cooke, 1942. p. 49. pl. 5, figs. 29-32. Brissopsis steinhatchee (Cooke) Cooke, 1945. p. 61, fig. 6, no. 4 (after Cooke, 1942). not Brissopsis biarritzensis (Cotteau). Cooke, 1959. p. 85. pl. 38, figs. 14-18 (in part, specimen figured is B. steinhatchee). Brissopsis steinhatchee (Cooke). Cooke, 1959. p. 85. pl. 38, figs. 9-13. Brissopsis steinhatchee (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—This species occurs throughout the OLS of Florida but is most commonly found in the upper portion of the unit in association with W. eldridgei. Localities include that of the type, in a pit near the Steinhatchee River east of Clara, Dixie County; north of Branford, Suwannee County (FM- IP SU002); northwest of Mayo (FM-IP LF001), Lafayette County; and south of Tennille (FM-IP DI001), Dixie County. Brissopsis steinhatchee is much rarer in the lower OLS where it occurs along the Cross Florida Barge Canal south of Inglis (FM-IP CI001), Levy County. This species rarely occurs in the OLS of Al- abama (Cooke, 1959). Discussion.—Brissopsis steinhatchee is most abundant near the top of the OLS, where it occurs in the Wythella eldridgei Zone with S. armiger, P. dixie, O. beckeri, R. trojana, and other, rarer species. In Jackson County, the species rarely occurs in the Oligopygus haldemani Zone, and it is even rarer near the base of the OLS in the Oligopygus phelani Zone. Cooke (1959) discussed and figured a speci- men of Brissopsis from the “OLS”, likely strata now referred to the Shubuta Member of the Yazoo Clay of Monroe County, Alabama that he referred to Bris- sopsis biarritzensis Cotteau, 1884. The latter was described from the Eocene Lou Cout, Villa Eugenie near Biarritz, France. The specimens Cooke (1959) described (USNM 562454a, b) have anterior petals that were less divergent than in the two specimens of B. steinhatchee he had available for study (the holotype and paratype). Carter et al. (1989) stated that the figures of the specimen provided by Cooke (1959) display a specimen no less similar to B. stein- hatchee than to the figures of B. biarritzensis in Cahuzac and Roman (1984) and that the specimen may simply be a variant of B. steinhatchee. We agree, as additional specimens of B. steinhatchee reveal that the specimen figured by Cooke (1959: pl. 38, figs. 14-18) falls within the variability of B. steinhatchee displayed in specimens from the type area of Dixie County, Florida. The species is now known to show a greater degree of variability in divergence of ambulacra I and V, as well as ambu- lacra II and IV than was understood from the two specimens Cooke had available for study. The largest specimen available (UF 2122) is from the Oligopygus phelani Zone at the Cross Florida Barge Canal (FM-IP 2409) and measures 96.5 mm TL, 72.1 mm TW, 34.4 mm TH (Fig. 180). cf. Brissopsis sp. (Fig. 181) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 249 Figure 181: cf. Brissopsis sp. (UF 337993) portion of ambulacra II and III of a large spatangoid that cannot be attributed to any known taxa from the region. The specimen has a peripetalous fasciole that very closely follows the ambulacra and is very similar to the genus Brissopsis; thus, the generic attribution questionably assigned herein. Specimen measures: 72.9 mm TL, 52.2 mm TW, 46.4 mm TH, though none of the measurements are complete, Wythella eldridgei Zone at the top of the Upper Eocene, upper Ocala Limestone, quarry west of Dowling Park, Lafayette County, Florida (FM-IP LF002) A: aboral. B, C: petaloid portion of ambulacrum II, note how tightly the peripetalous fasciole follows the ambulacrum. D: oblique lateral viewpoint. E: oblique posterior viewpoint. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 250 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Discussion.—This specimen consists of ap- proximately a posterior quarter of the test, with portions of ambulacra II and III, of what would have been a very large spatangoid, measuring 73 mm TL, 52 mm TW, and 46 mm TH. Given its incomplete- ness, none of these measurements reflects its true dimensions. The specimen was collected with W. eldridgei from the uppermost bed of the OLS (Rotu- laria vernoni Zone) west of Dowling Park, Lafayette County (FM-IP LF002). Although the specimen is incomplete, enough is preserved to determine that it does not belong to any species currently known from the Paleogene faunas of the region. However, it remains impossible to attribute it to any named species from the fossil faunas of the remainder of the eastern Americas. The specimen is very similar to Brissopsis in that it has a peripetalous fasciole that closely follows the ambulacra. Therefore, it is conditionally assigned to Brissopsis. If this is a Brissopsis, it is significantly larger than any documented specimens of B. steinhatchee, the only Eocene representative of the genus in the eastern United States. The largest specimen of B. steinhatchee (UF 2122: Fig. 180) is 96 mm in length. Given similar proportions, and conservatively estimating that this specimen shows less than 50% of its TL, it would have exceeded 145 mm in length. Genus Brissus Gray, 1825 Brissus bridgeboroensis Carter, 1987b (Fig. 182) Brissus bridgeboroensis Carter, 1987b. pp. 1043- 1046. Occurrence.—Suwannee Limestone, Sunwest Mine, southwest of Aripeka, Pasco County. This species is also documented from the Bridgeboro Limestone near Florala, Covington County, Al- abama, and southwest of Bridgeboro, Mitchell County, Georgia (the type locality). Discussion.—Carter (1987b) described B. bridgeboroensis from a single specimen collected in the Lower Oligocene Bridgeboro Limestone of Figure 182: Brissus cf. B. bridgeboroensis (UF 342093), 19.9 mm TL, Lower Oligocene Suwannee Limestone, Sunwest Mine, west of Aripeka, Florida (FM-IP PA010). A: specimen in matrix. B: aboral. C: left side. D: tilted aboral viewpoint from anterior. E: petaloid area with apical system. F: tilted aboral viewpoint from posterior. G: posterior. H: anterior. Mitchell County, Georgia. He differentiated it from the extant B. unicolor by its longer posterior petals, fewer pore pairs in the paired petals, its apical system closer to the anterior margin, and complete lack of a dorsal keel in interambulacrum 5 (Carter, 1987b). Heller and Bryan (1992) subsequently doc- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 251 umented the occurrence of B. bridgeboroensis at Huddlestun’s (1993) type locality for the Florala Member of the Bridgeboro Limestone in Covington County, Alabama. This is the first record of the species both outside of the Bridgeboro Limestone, and within Florida. The occurrence of this single, small (20 mm TL) specimen from Aripeka expands the dis- tribution of the species geographically southward and stratigraphically upward into the Suwannee Limestone. Zachos and Molineux (2007) documented imperfect specimens of a Brissus from Oligocene strata at Damon Mound, Brazoria County, Texas which they attributed to Brissus exiguus Cotteau, 1875 [which Cooke (1961) synonymized with B. uni- color]. They stated that B. bridgeboroensis differed from the specimens from Damon Mound in having a much more anteriorly eccentric apical system and a greater divergence of the anterior paired ambulacra (these differences are not evident in the specimens figured). However, they did not further describe, provide measurements of, or further discuss the specimens and conspecificity cannot be ruled out. Until additional specimens of B. bridgeboroensis are available for comparison, the variability within the species remains largely unknown. Brissus jonesi n. sp. (Figs. 183-187) Diagnosis.—Brissus with large ovate periproct; periproct width on average 60.8% periproct height; periproct height on average 35.1% TL and 54.5% TH and center of apical system on average 70.2 % TL from posterior margin; 77.1% TL in holotype. Description.—Description based on the holo- type (UF 342106) and four paratypes (UF 342107, UF 341761, UF 341762, UF 341765). Test small to moderately sized, largest complete specimen desig- nated the holotype (37.7 mm TL, 30.7 mm TW, 24.1 mm TH); largest specimen is paratype UF 341765: approximately 45 mm TL (posterior missing so the specimen could be much larger), 43.3 mm TW, 33.7 mm TH; smallest is paratype UF 341761 (12.0 mm TL, 9.6 mm TW, 7.8 mm TH); TW 81.4% TL in holotype, average 82.3%; TH 63.9% TL in holotype, average 66.2%; greatest TW posterior of apical system, greatest TH posterior. Test ovate, narrows posteriorly in holotype, less so in smaller specimens. Anterior abrupt and nearly vertical in holotype, more gently rounded in smaller specimens. Posterior ambitus above periproct in holotype, an- gles sharply anteriorly towards oral surface, less angled in smaller specimens. Aboral surface gently rounded in smaller specimens, raised medially in holotype. Test covered with relatively evenly dis- persed, small tubercles, more pronounced in smaller specimens. Apical system anterior, center of apical sys- tem 77.1% TL from posterior margin in holotype, more posterior in smaller specimens: 69.8 % TL in specimen 20.9 mm TL, 63.7% TL in specimen 12.0 mm TL; center of apical system on average 70.2 % TL from posterior margin; apical system ethmolytic; 4 gonopores. Ambulacrum III not petaloid, in very shal- low depression from apical system to peristome, pores minute; ambulacra II and IV transverse, nearly forming 180° angle to each other; ambulacra I, II, IV, and V in deep grooves in holotype, shal- lower depressions in smaller paratypes; petals of ambulacrum I and V longest: on average 43.7% TL in holotype, proportionately shorter in smaller specimens: average 37.3% TL, extending two-thirds distance to ambitus; petals of ambulacrum II and IV on average 29.3% TL in holotype, average 25.3% TL; extending nearly to ambitus. Petals narrow, am- bulacrals I and V width on average 20% petal length; petal II and IV width on average 25.7% petal length; interporiferous zones moderately narrow; inner pore circular, outer pore elongate, pores conjugate; petals closed distally. Periproct very large, inframarginal, high on overhanging posterior truncation; higher than wide; periproct height on average 35.1% TL and 54.5% TH; periproct width on average 60.8% periproct height. Peristome anterior, posterior edge of peris- tome on average 66.9% TL from posterior margin; peristome large, wider than high; peristome height on average 42% peristome width; labrum very short; ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 252 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 183: Brissus jonesi n. sp., holotype (UF 342106), 37.7 mm TL, 30.7 mm TW, 24.1 mm TH, Upper Eocene, Haimea brooksi Zone of the Ocala Limestone, Brooks Quarry northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: apical area. C: oral. D: posterior. E: anterior. F: oblique lateral viewpoint from posterior. G: peristome. H: oblique lateral viewpoint from posterior. I: peripetalous fasciole and distal end of petal of ambulacrum II. J: left side. K: periproct. L: oblique lateral viewpoint from anterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 253 Figure 184: Brissus jonesi n. sp., paratype (UF 342107), 20.9 mm TL, 17.9 mm TW, 14.6 mm TH, a very corroded specimen lacking most surface details, Upper Eocene, Haimea brooksi Zone of the Ocala Limestone, Brooks Quarry northwest of Marianna, Jackson County, Florida (FM-IP JA039). A: aboral. B: tilted aboral viewpoint from posterior. C: oblique lateral viewpoint from posterior. D: left side. E: oral. F: tilted aboral viewpoint from anterior. G: posterior. H: anterior. plastron long and wide, bowed outward medially. Due to weathering, only short traces of fasci- oles preserved; peripetalous and subanal fascioles present. Peripetalous fasciole moderately indented in interambulacrum 5; not otherwise preserved but presumably indented in all 5 interambulacra. Sub- anal fasciole not completely preserved; consistent with being bilobed. Zoobank Nomenclatural Act.—BDC07B8D- 897C-45DD-B552-371BA875A472 Discussion.—Brissus jonesi n. sp. is the first species of Brissus described from the Eocene of the eastern United States. It is unknown outside of Jackson County, Florida, where it occurs in the OLS dredged from 24 m below the top of the Eocene in the Brooks Quarry (FM-IP JA039) west of Mari- anna, in a horizon of Asterocyclina-rich limestone with a diverse echinoid assemblage that includes H. brooksi, O. haldemani, O. rotundus, W. johnsoni, R. georgiensis, R. mariannaensis n. sp., and other, rarer species (Table 1; Fig. 6). Brooks Quarry is the type locality of the species. However, paratypes were also obtained from the same horizon (with H. brooksi) in Jackson Blue Spring east of Marianna, at a depth of 26-27 m (FM-IP JA033). The holotype (UF 342106) and paratype (UF 342107) from Brooks Quarry (FM-IP JA039) are complete. However, only one (UF 341761) of the three paratypes from Jackson Blue Spring (FM-IP JA039) retains the posterior portion of the test. The posterior portion of the two complete paratypes is hollow and exceptionally fragile because the sed- iment that infilled, and provided support for, the test has weathered out through the periproct. This is problematic because the holotype (37.7 mm TL) has a much more tapered posterior outline, has a steeper anterior, less rounded (steeper laterally) ab- oral surface, and more anterior apical area than the two smaller paratypes, which are 12.0 and 20.9 mm TL. Lacking additional complete, larger specimens, it is not possible to verify if this represents typical ontogenetic narrowing of the posterior end of the test, movement of the apical area anteriorly, and development of a steeper anterior end or if these features are unique to the holotype. Though Brissus is well documented in the global fossil record from the Eocene to Pleistocene strata, it is relatively rare in both the fossil record and modern fauna in the eastern United States, being rep- resented only by the Oligocene B. bridgeboroensis, Pliocene B. glenni, and extant B. unicolor. The fossil faunas of the Caribbean region also include B. exiguus, from the Miocene of Anguilla, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 254 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 185: Brissus jonesi n. sp., paratype (UF 341761), 12.0 mm TL, 9.6 mm TW, 7.8 mm TH, Upper Eocene, Haimea brooksi Zone of the Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033). A: aboral. B: anterior. C: oblique lateral viewpoint from posterior. D: left side. E: posterior. F: peristome. G: apical area and petaloid portion of ambulacra I and V. H: oral. I: oblique oral viewpoint from posterior. J: oblique posterior viewpoint from oral surface; no trace of subanal fasciole is preserved in this corroded and very weathered specimen. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 255 Figure 186: Brissus jonesi n. sp., paratype (UF 341762), 34 mm TL (incomplete), 29.3 mm TW, 21,4 mm TH, Upper Eocene, Haimea brooksi Zone of the Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033). A: aboral. B: oblique lateral viewpoint from anterior. C: oblique lateral viewpoint from posterior. D: anterior. E: tilted aboral viewpoint from anterior. F: aboral surface with strong cross lighting to enhance peripetalous fasciole detail; indicated with stippled line where discernable. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 256 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 187: Brissus jonesi n. sp., paratype (UF 341765), 45.5 mm TL (incomplete), 43.3 mm TW, 33.7 mm TH (incomplete), Upper Eocene Haimea brooksi Zone of the Ocala Limestone, Jackson Blue Spring, Jackson County, Florida (FM-IP JA033. A: aboral. B: tilted aboral viewpoint from anterior. C: left side. D: posterior. E: apical area. F, G: oral; posterior end is largely missing but anterior portion of subanal fasciole is preserved, indicated by stippling. H: petaloid portion of ambulacrum IV. I: petaloid portion of ambulacrum II. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 257 which, as noted by Kier (1984: 81), is based on two deformed specimens that make it difficult to be certain of diagnostic characters. Cooke (1961) considered B. exiguus a subjective junior synonym of B. unicolor. Brissus jonesi n. sp. cannot be confused with B. exiguus. Kier (1984) recognized, redescribed, and fig- ured five species of Brissus in the Cuban Ceno- zoic faunas: B. cabrerai (Sánchez-Roig, 1953c); B. camagueyensis Weisbord, 1934; B. caobaense Sánchez-Roig, 1953b; B. durhami (Sánchez-Roig, 1952a), and B. minutus Sánchez-Roig, 1949. In ad- dition, Kier (1984) noted that Sánchez-Roig (1949) reported a specimen of B. unicolor from the Pleis- tocene of Cuba, but the specimen is no longer avail- able for study. Of these, Brissus jonesi n. sp. is most similar to B. caobaense from the Cuban Eocene, but the two species are readily distinguished by the greater height of B. jonesi n. sp. (63.9-69.8% TL vs 49% TL in B. caobaense), and the much larger and proportionately narrower periproct of B. jonesi n. sp. (periproct height on average 35.1% TL in B. jonesi n. sp. and 20% in B. caobaense; periproct width 60% periproct height in B. jonesi n. sp. and 50% in B. caobaense). Donovan et al. (2015) documented an assem- blage of internal molds that are readily identifiable as belonging to Brissus from the Miocene of the Cay- man Islands. They recognized the form as Brissus cf. B. oblongus Wright, 1855, noting its similari- ties with the form from Malta. The specimens may represent a new species, but Donovan et al. (2015) considered it best not to erect a new species based on internal molds. These specimens are not comparable to B. jonesi n. sp. Etymology.—Named in honor of Douglas S. Jones, Florida Museum of Natural History Director and Invertebrate Paleontology Curator, for his many years of contributions and service. Material and Occurrence.—Holotype UF 342106 and paratype UF 342107 from the type locality in the OLS, where it occurs with H. brooksi and O. haldemani in material dredged from 24 m below the top of the Eocene stratum in the north- westernmost pit in the quarry complex of Leon Brooks, northwest of Marianna, Jackson County, Florida (FM-IP JA039). Additional paratypes (UF 341761, UF 341762, UF 341765) are from Jackson Blue Spring east of Marianna at a depth of 26-27 m (FM-IP JA033). Genus Plagiobrissus Pomel, 1883 Plagiobrissus curvus (Cooke, 1942) (Fig. 188) Eupatagus (Plagiobrissus) curvus Cooke, 1942. p. 56. pl. 7, figs. 5-7. Eupatagus sp. Fischer, 1951. pp. 84, 85. pl. 5, fig. 5. Plagiobrissus curvus (Cooke). Cooke, 1959. pp. 87-88. pl. 39, figs. 3-6. Plagiobrissus curvus (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—This species is rarely encoun- tered in the OLS of Florida. The type locality is along the Chipola River above the bridge at Mari- anna, Jackson County. Plagiobrissus curvus is also found at the nearby Brooks Quarries (FM-IP JA009, FM-IP JA018, FM-IP JA027, FM-IP JA031, FM-IP JA039) northwest of Marianna, Jackson County, in the Wythella eldridgei Zone west of Dowling Park, Lafayette County (FM-IP LF002), and northwest of Mayo (FM-IP LF001), Lafayette County, among other localities. The species is more rarely encoun- tered in the Oligopygus phelani Zone of the lower OLS where Fischer (1951) documented it along the Withlacoochee River west of Yankeetown, Levy County (FM-IP LV024). It also occurs in the OLS of Georgia (Cooke, 1959; Carter and McKinney, 1992). Discussion.—Plagiobrissus curvus is typically rare but widespread in the OLS. Near Marianna in Jackson County Florida, it occurs in the Oligopygus haldemani Zone, and in Lafayette County it occurs in the Oligopygus wetherbyi Zone of the upper OLS. The species is much rarer in the Oligopygus phelani Zone of the lower OLS (Fischer, 1951). The prominent large tubercles and curving ambulacra readily distinguish this species from other members of the genus in the region, as well as from E. ocalanus, with which it often occurs. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 258 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 188: Plagiobrissus curvus (UF 322381), 93.5 mm TL, 76 mm TW, uppermost portion of Upper Eocene Ocala Limestone, Lafayette County, Florida (FM-IP LF015) A: aboral. B: oral. C: left side. D: right side. Plagiobrissus dixie (Cooke, 1942) (Fig. 189) ?Brissoides (Koilospatangus) floridanus (Clark). Lambert and Thiéry, 1924. p. 454. Eupatagus (Plagiobrissus) dixie Cooke, 1942. p. 55. pl. 6, figs. 1-3. Plagiobrissus? dixie (Cooke). Cooke, 1959. p. 87. pl. 40, figs. 1-5. Plagiobrissus? dixie (Cooke). Toulmin, 1977. p. 345. pl.70, figs. 1-3. Plagiobrissus dixie (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—Within Florida, P. dixie occurs in the upper portion of the OLS, especially the uppermost portion of the unit where it occurs in the Wythella eldridgei Zone. The type locality is north of Mayo, Lafayette County. The species also occurs north of Branford (FM-IP SU002), Suwannee County; northwest of Mayo (FM-IP LF001), Lafayette County; the Brooks Quarries near Mar- ianna (FM-IP JA009, (FM-IP JA018, FM-IP JA027, FM-IP JA031), and in the Shubuta Clay of Alabama (Toulmin, 1977) and OLS of Georgia (Cooke, 1959). Discussion.—Plagiobrissus dixie commonly can be found at the very top of the OLS, in the Wythella eldridgei Zone where it occurs with R. trojana and an assemblage of spatangoids that are otherwise rare in the unit, including B. steinhatchee, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 259 Figure 189: Plagiobrissus dixie (UF 343599), 75 mm TL, 62 mm TW, 32 mm TH, uppermost portion of Upper Eocene Ocala Limestone, Dixie County, Florida (FM-IP DI013). A: aboral. B: oral. C: left side. D: right side. E: posterior. F: tilted oral viewpoint from posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 260 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 190: Plagiobrissus cassadyi n. sp., holotype (UF 97921), 82 mm TL, 70.5 mm TW, 30.5 mm TH (TW and TH incomplete), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: aboral. B: apical area. C: oral. D: tilted aboral viewpoint from posterior. E: oblique aboral viewpoint from posterior. F: left side. G: tilted aboral viewpoint from anterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 261 Figure 191: Plagiobrissus cassadyi n. sp., paratype (UF 105223), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). External mold of specimen showing large tubercles confined within peripetalous fasciole, which is stippled where visible. E. ocalanus, O. beckeri, S. armiger, and other uncom- mon taxa. The species is more rarely encountered in the Oligopygus haldemani Zone in Jackson County, Florida where it is found with W. johnsoni and other, rarer species. Cooke (1959) reassigned Eupatagus dixie to Plagiobrissus because of the narrowness of the am- bulacra and the length of the plastron. However, he only tentatively placed this species in that genus, because of the lack of evidence of an anal fasciole in specimens he had available for study. Examination of additional well-preserved specimens (e.g., UF 343599) does reveal traces of anal branches on the subanal fasciole (Fig. 189), so assignment to Pla- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 262 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) giobrissus is confirmed. In addition, P. dixie shares other traits of that distinguish it from Eupatagus, such as the long, narrow, somewhat curving ambu- lacra, rather than straight, lanceolate ambulacra as in typical Eupatagus. Cooke (1959) also indicated that Plagiobrissus has a long plastron connected with a short labrum, and this contrasts with the V-shaped plastron and long, narrow labrum of Eupatagus. In addition to the presence of the anal fascioles, the narrow, slightly curved petals readily differentiate P. dixie from E. ocalanus, which is the only species that it could possibly be confused with in the OLS. Plagiobrissus cassadyi n. sp. (Figs. 190-195) Brissopatagus sp. Oyen 2001. pp. 136-138. figs. 3-18, A, B. Diagnosis.—Plagiobrissus with broad test (TW on average 89.7% TL); ambulacra II and IV di- vergent from each other at 148° (on average), flexed distally towards anterior; distinct sulcus where am- bulacrum III crosses anterior margin; large tubercles confined aboral to peripetalous fasciole of interam- bulacra 1, 2, 3 and 4 only, tubercles in interambu- lacra 1 and 4 arranged in rows, tubercles in anterior interambulacra haphazardly distributed. Description.—Based on the holotype (UF 97921), five paratypes (UF 40441, UF 105223, UF 105225, UF 105226, UF 105227), and non-type material in the FM-IP collections. Specimens con- sist of both internal and external molds. Therefore, measurements of TL, TW, and TH are slightly less than if specimens still had test preserved. Test subovate, broad anteriorly, tapering pos- teriorly, with shallow anterior sulcus; moderately sized, largest specimen 83.9 mm TL; holotype 82 mm TL, 70.5 mm TW, 30.5 mm TH (TW and TH incomplete), TW on average 89.7% TL; test de- pressed overall, TH on average 36.9% TL; highest point posterior, widest point just posterior of apical system. Apical system anterior, on average 67% TL from posterior margin; four gonopores, no further details discernable, but appears to be ethmolytic. Petals narrow, weakly sunken, flexed outward dis- tally; ambulacrum III not petaloid, in shallow notch at anterior margin. Petals of ambulacra I and V longest; petal I on average 37% TL, petal V on average 38.6% TL; on average divergent from each other by 51°, narrow (width on average 20.2 petal length); petals II and IV shortest, with petal II on average 33.4% TL, petal IV on average 33.6 TL; narrow (width on average 22.3% petal length), on average divergent from each other by 148°; distally curved towards anterior. Large tubercles confined aboral to peripetalous fasciole in interambulacra 1, 2, 3, and 4, but absent in interambulacrum 5. Tubercles in interambulacra 1 and 4 arranged in slightly crooked rows, tubercles in anterior paired interambulacra haphazardly arranged, not in distinct rows. Peripetalous fasciole prominent, running close to tips of paired petals, not indented between them. Subanal fasciole present, wide (width on average 44.1% TW); anal branches not discernable. Periproct on truncate, overhanging posterior; ovate, height on average 94% periproct width; height on average 38% TH. Peristome anterior, posterior edge on average 82% TL from posterior margin; wider than high, kidney-shaped, with short, wide labrum. Plastron broad, medially convex. Oral plating and tuberculation unknown. Zoobank Nomenclatural Act.—20D5AF6C- 6090-408F-837F-8E1BA04992F7 Discussion.— All known specimens of this taxon are preserved as molds (both external and internal) from a dolomitic bed in the upper portion of the Lower Oligocene Marianna Limestone in the bed and banks of Dry Creek, a tributary of the Chipola River, south of Marianna in southern Jackson County (FM-IP JA010). Here, P. cassadyi n. sp. occurs with moldic C. rogersi and rarer S. americanus and Phyllacanthus cf. P. mortoni. Although the moldic preservation is not optimal, sufficient detail is preserved, especially on the external molds, to describe the species and recognize it as a new taxon. Although measurements of TL, TW, and TH are slightly less than what the specimens would be with preserved test, ratios recorded above remain useful for comparison with other taxa. This is the first known occurrence of Plagio- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 263 Figure 192: Plagiobrissus cassadyi n. sp., paratype (UF 105227), 76.7 mm TL, 65.8 mm TW, 25.9 mm TH (no dimensions are complete), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: aboral. B: apical area. C: tilted aboral viewpoint from posterior. D: oblique aboral viewpoint from posterior. E: close-up of pore-pairs at widest point of petal in ambulacrum V. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 264 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 193: Plagiobrissus cassadyi n. sp., paratype (UF105225), 77.5 mm TL, 73.6 mm TW, 28.0 mm TH (no dimensions are complete), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: oral. B: peristome. C: periproct and indications of subanal fasciole. D: oblique lateral viewpoint from posterior. E: tilted oral viewpoint from posterior. F: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 265 Figure 194: Plagiobrissus cassadyi n. sp., paratype (UF40441), 83.9 mm TL, 76.6 mm TW (TH incomplete; aboral surface is completely corroded), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: oral. B: peristome. C: tilted oral viewpoint from posterior. D: tilted oral viewpoint from anterior. E: tilted aboral viewpoint from posterior. F: posterior end of oral surface showing indications of subanal fasciole. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 266 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 195: Plagiobrissus cassadyi n. sp., paratype (UF105226), 73.4 mm TL, 67.7 mm TW (TH incomplete), dolomitic facies in upper part of the Lower Oligocene Marianna Limestone, Jackson County, Florida (FM-IP JA010). A: aboral. B: ambulacrum I. C: tilted aboral viewpoint from posterior. D: tilted aboral viewpoint from anterior. E: pore-pairs at widest point of petal in ambulacrum I. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 267 brissus in the Oligocene of North America, which, together with E. dumonti n. sp., adds data to an otherwise relatively depauperate regional Oligocene spatangoid fauna (Table 4). Oyen (2001) first documented this species and placed it in Brissopatagus. However, he recognized that it likely represented a new species, but did not name it. Brissopatagus was reviewed by Henderson and Fell (1969), and P. cassadyi n. sp. is readily separated from members of Brissopatagus since the anterior interambulacra are not depressed. In Brissopatagus, the paired interambulacral regions immediately posterior to the paired anterior petals is distinctly concave, forming a depression that is confluent with the petals. This species cannot be assigned to Eupatagus due to the presence of slightly depressed petals and distinct anterior notch in P. cassadyi n. sp. Although details of the apical system are largely lacking, P. cas- sadyi n. sp. has a distinct subanal and peripetalous fasciole, and narrow, slightly sunken petals, all fea- tures suggesting the family Brissidae. The low test, large aboral, interambulacral primary tubercles that are confined to the region aboral to the peripetalous fasciole, and narrow petals with the posterior pair flexed laterally at their tips suggests Plagiobrissus. Although the currently available specimens do not provide sufficient detail of the posterior region to detect anal branches on the subanal fasciole, the specimens are otherwise more like Plagiobrissus than any other genus. Plagiobrissus occurs as old as the Eocene, where it is represented in eastern North Ameri- can faunas by P. curvus and P. dixie in the Upper Eocene deposits of the southeastern United States. Prior to the recognition of P. cassadyi n. sp., the genus appeared to have a lengthy regional gap in the fossil record between Late Eocene occurrences noted in this paper, and Late Pliocene occurrence of Plagiobrissus sarae Ciampaglio et al., 2009, and the Recent P. grandis (Gmelin, 1791). The latter is only tentatively documented in the fossil record of the region (Osborn et al., 2020). Plagiobrissus cassadyi n. sp. cannot be confused with any of these species. It is perhaps nearest in overall morphology to P. dixie, from which it is readily distinguished by its possession of more numerous large primary tubercles within the peripetalous fasciole, broader, more cordate test, and deeper anterior notch. The tuberculation of P. cassadyi n. sp. is more like that of P. curvus, but the petals of these two species are very different. Although there are other occurrences of Pla- giobrissus in the fossil record of the eastern Ameri- cas, it is relatively rare. We compare P. cassadyi n. sp. to all the known taxa below. Kier (1980: 11) documented Plagiobrissus sp. from the Castle Hayne Limestone of North Car- olina but did not figure or describe the material. Plagiobrissus is present in sporadic concentrations in the fossil record of the Caribbean region, largely concentrated in Eocene deposits. Kier (1984) did not find the genus in Cuban faunas. Plagiobrissus loveni (Cotteau, 1875) is com- mon in the Eocene of Jamaica (Arnold and Clark, 1927; Donovan, 1993) as well as at its type locality in St. Bartholomew (Cotteau, 1875; Jackson, 1922). The abundant large tubercles on the aboral surface of this species are unique. Arnold and Clark (1927) stated that the plastron of P. loveni is distinctly differ- ent than that of P. grandis and suggested the species may not be a Plagiobrissus. However, subsequent authors have not reassigned the species, and we con- sider P. loveni to be correctly assigned. The aboral tuberculation of P. loveni readily distinguishes it from P. cassadyi n. sp. In addition, P. cassadyi n. sp. has a much more tapered posterior region than P. loveni, and its peripetalous fasciole does not closely follow the ambitus, as it does in P. loveni. Arnold and Clark (1927) also described P. abruptus, P. elevatus, P. latus, P. perplexus, and P. robustus from Eocene deposits of Jamaica. These five species, most of which were described from incomplete holotypes, were further discussed by Donovan (1993) and P. cassadyi n. sp. is not con- specific with any of them. Jeannet (1928) described Plagiobrissus lam- berti using a very incomplete test from Venezuela, which Cooke (1961) did not figure in his review of the fauna, perhaps due to the poor quality of the fossil. Anisgard (1954) figured an unnamed Eocene species of Plagiobrissus from a deep well in ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 268 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Venezuela, but the specimen consists of the anterior portion of the test and lacks features that would allow comparison to other heart urchins. Sánchez-Roig (1949) described Brissolampas santanae, which he subsequently assigned into his new genus Moronaster (Sánchez-Roig, 1952a). This species shares similar gross morphology to P. cas- sadyi n. sp. However, as noted by Kier (1984), no types of this species or the other species Sánchez- Roig placed in Moronaster can be located for com- parison. The existing figures are poor and show badly weathered specimens, making assignment even to genus uncertain. Kier (1984) presumed the species belong to either Pericosmus or Antillaster. Cooke (1961) documented fragmentary ma- terial from the Miocene? of Venezuela that he at- tributed to P. grandis. The fragment he discussed and figured (Cooke, 1961: pl. 13, fig. 1) likely belongs to Plagiobrissus, but its assignment to P. grandis is questionable due to the lack of diagnostic features on the incomplete specimen. Conspecificity with P. sarae, a stratigraphically more logical comparison, could not be ruled out. Kew, in Dickerson and Kew, 1917 described Metalia cumminsi from the Cenozoic of Mexico, but this was referred to Plagiobrissus by Israelsky (1924). However, Durham (1961) clarified that this species belongs to Lajanaster Lambert and Sánchez Roig in Sánchez Roig, 1926. Durham (1961) described the very elongate Plagiobrissus costaricensis as well as P. malavas- sii, which occur together in the Miocene of Costa Rica. Neither of these species could be confused with P. cassadyi n. sp. The narrow test of P. costari- censis readily differentiates it from P. cassadyi n. sp., and petals II and IV of P. malavassii are not flexed distally towards the anterior as in P. cassadyi n. sp. Macropneustes mexicanus Kew, in Dickerson and Kew, 1917, was described from the Oligocene to Miocene strata of Topila, Mexico, and seems to be the most similar taxon to P. cassadyi n. sp. in the Americas. Macropneustes mexicanus was sub- sequently transferred to Brissopatagus by Israelsky (1924) and later to Gillechinus by Henderson and Fell (1969). Although G. mexicanus and P. cas- sadyi n. sp. are very similar in petaloid structure and test shape, the primary tubercles are arranged differently. In Gillechinus, the primary tubercles are arranged in rows expressed within the peripetalous fasciole only on the posterior columns of plates in the paired interambulacra. In external molds of P. cassadyi n. sp. (Fig. 191), primary tubercles are arranged within the peripetalous fasciole throughout the paired interambulacra. With the addition of P. cassadyi n. sp., Pla- giobrissus now includes 18 species from the eastern Americas. However, as mentioned above, several of these species remain poorly known. Etymology.—Named for Tim Cassady, who not only brought the locality to the attention of RWP, but donated most of the specimens collected from the site, including the holotype and paratypes. Material and Occurrence.—This species is represented by the holotype (UF 97921) and paratypes (UF 40441, UF 105223, UF 105225, UF 105226, UF 105227), all internal and external molds, from the dolomitic facies of the Lower Oligocene Marianna Limestone along Dry Creek, south of Mar- ianna, in southern Jackson County (FM-IP JA010). Superfamily SPATANGOIDEA Gray, 1825 Family MACROPNEUSTIDAE Lambert, 1905 Genus Macropneustes L. Agassiz in Agassiz and Desor, 1847 Macropneustes mortoni (Conrad, 1850) (Fig. 196) Holaster mortoni Conrad, 1850. p. 40. pl. 1, fig. 10. Holaster mortoni (Conrad). Boyle, 1893. p. 150. Macropneustes mortoni (Conrad). Stefanini, 1911. p. 700. Macropneustes mortoni (Conrad). Clark and Twitchell, 1915. p. 155. pl. 72, figs. 1a-d. Macropneustes mortoni (Conrad). Cooke, 1926. pl. 96, figs. 1a-b. Macropneustes mortoni (Conrad). Cooke, 1942. p. 51. Macropneustes mortoni (Conrad). Cooke, 1959. p. 82. pl. 37, figs. 5-9. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 269 Figure 196: Macropneustes mortoni (UF 278706), 106 mm TL, 97.5 mm TW, 61.5 mm TH, Upper Eocene Ocala Limestone, Jackson County, Florida (FM-IP JA085). A: aboral. B: oral. C: right side. D: posterior. Macropneustes mortoni (Conrad). Pickering, 1970. pp. 20, 27, 64. Macropneustes mortoni (Conrad). Toulmin, 1977. p. 342. pl. 67, figs.7, 8. Macropneustes mortoni (Conrad). Heller and Bryan, 1992. pp. 345-346. Macropneustes mortoni (Conrad). Osborn et al., 2016. tbl. 2. Occurrence.—Within Florida, this species appears to be restricted to the OLS of Jackson County (FM-IP JA009, FM-IP JA018, FM- IP JA027, FM-IP JA031) where it occurs with Oligopygus haldemani. It is also found in the Upper Eocene of Alabama (Toulmin, 1977) and Georgia (Cooke, 1959). The type locality is near Palmyra in Lee County, Georgia (Conrad, 1850). Discussion.—Macropneustes mortoni has not been documented south of the Suwannee Strait in peninsular Florida. In Jackson County, especially in the Brooks Quarries (FM-IP JA009, (FM-IP JA018, (FM-IP JA027, FM-IP JA031, FM-IP JA039), it occurs with O. haldemani, W. johnsoni, R. conradi, and numerous other, less common species in the ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 270 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 197: Eupatagus clevei (UF 12808), 101.5 mm TL, 82.5 mm TW, 40.5 mm TH, internal mold from dolomitic beds of lowermost portion of the Upper Eocene Ocala Limestone, Citrus County, Florida (FM-IP CI009). A: aboral. B: oral. C: left side. D: posterior. Asterocyclina Zone just below the contact with the Bumpnose Limestone. Heller and Bryan (1992) documented the pres- ence of a large echinoid they identified as M. mor- toni in the Lower Oligocene (Vicksburgian) Florala Member of the Bridgeboro Limestone near Florala, Covington County Alabama. They discussed the significance of the species occurring in Oligocene strata, as M. mortoni had previously been only doc- umented from the Upper Eocene. We have collected specimens of M. mortoni from the OLS in this quarry, with other, typically Upper Eocene species such as R. conradi and W. johnsoni. Unless the Heller and Bryan (1992) specimen was extracted in situ from the Bridgeboro limestone, it is possible it came from Eocene strata in the quarry. We do not consider the species to occur in the Oligocene (Fig. 4). The specimen is incomplete with damage on the posterior margin. If additional specimens are collected from the definitive Bridgeboro Limestone strata in the Florala Quarry, they should be com- pared with Eupatagus dumonti n. sp. The low test, flush petals and very large periproct of E. dumonti n. sp. would be distinctive. The large test with thin walls of M. mortoni cannot be confused with that of any other member of the Eocene echinoid fauna of the region. Family EUPATAGIDAE Lambert, 1905 Genus Eupatagus L. Agassiz in Agassiz and Desor, 1847 Eupatagus clevei (Cotteau, 1875) (Figs. 197-200) Euspatangus clevei Cotteau, 1875. p. 44. pl. 8, figs. 1-4. Euspatangus grandiflorus Cotteau, 1875. p. 45. pl. 8, figs. 5, 6. Eupatagus clevei (Cotteau). Guppy, 1882. p. 199. Eupatagus grandiflorus (Cotteau). Jackson, 1922. p. 89. pl. 15, figs. 5, 6. Eupatagus clevei (Cotteau). Jackson, 1922. p. 90. pl. 16, figs. 1, 2. Eupatagus hildae Hawkins, in Arnold and Clark, 1927. p. 81-82. pl. 22, figs. 9, 10. txt. fig. 1. Eupatagus grandiflorus (Cotteau). Molengraaff, 1929. p. 72. pl. 24, figs. 1, 2; pl. 25, fig. 1. Eupatagus grandiflorus (Cotteau). Arnold and Clark, 1934. p. 156. Eupatagus clevei (Cotteau). Cooke, 1948b. p. 92. pl. 22, fig. 9. Brissoides grandiflorus (Cotteau). Sánchez-Roig, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 271 Figure 198: Holotype of Eupatagus ingens (= E. clevei), (UF 1008), 128 mm TL, 105 mm TW, 54 mm TH (test height is obscured by squishing of the test and missing portion of oral surface), Upper Eocene lower Ocala Limestone, Citrus County, Florida (FM-IP CI010). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 272 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 199: Eupatagus clevei (UF 341014) rare specimen from the Ocala Limestone with preserved test: 124 mm TL, 105 mm TW, 60.5 mm TH, Upper Eocene Ocala Limestone (precise zone within OLS is unknown), Hernando County, Florida (FM-IP HE006). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 273 Figure 200: Eupatagus hildae Hawkins, in Arnold and Clark, 1927 (= Eupatagus clevei) (UF 4496) cast of holotype (Natural History Museum E17664), 105 mm TL, 81 mm TW, 63 mm TH, Oligocene, White Limestone. St. Hilda’s School, Browns Town, Jamaica (FM-IP 2822). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 274 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) 1949. p. 208. Eupatagus clevei (Cotteau). Fischer, 1951. p. 83. pl. 7, figs. 1-3; txt fig. 18. ?Eupatagus (Gymnopatagus) venturillae Sánchez- Roig, 1951. p. 43. pl. 26, figs. 1, 2. ?Eupatagus (Gymnopatagus) roajasi Sánchez-Roig, 1951. p. 42. pl. 34, fig. 3. ?Eupatagus (Gymnopatagus) zanoletti Sánchez- Roig, 1951. p. 43. pl. 32, fig. 3. Lajanaster guevarai Sánchez-Roig, 1951. p. 53. pl. 24, figs. 2-4. ?Eupatagus (Plagiobrissus) herrerae Sánchez-Roig, 1951. p. 46. pl. 25, figs. 1, 2. ?Lajanaster venturillae Sánchez-Roig, 1951. p. 54. pl. 25, figs. 3, 4. ?Zanolettiaster herrerae Sánchez-Roig, 1952c. p. 15. pl. 8, figs. 1, 2; pl. 9, fig. 4. ?Megapatagus franciscanus Sánchez-Roig, 1953c. p. 59. pl. 11, 2 figs. Eupatagus clevei (Cotteau). Cooke, 1959. p. 89. pl. 41, figs. 6-8. Eupatagus clevei (Cotteau). Cooke, 1961. pp. 26-27. pl. 10, figs. 2-5. Eupatagus ingens Zachos, 1968. pp. 161-164. Eupatagus clevei (Cotteau). Kier, 1974. txt. fig. 26B. Eupatagus clevei (Cotteau). Toulmin, 1977. p. 341. pl. 66, figs. 4, 5. Eupatagus ingens (Zachos). Zachos and Shaak, 1978. pp. 921-927. Eupatagus clevei (Cotteau). Kier, 1984. pp. 98-100. pl. 61, figs. 3-6. pls. 62-66. Eupatagus clevei (Cotteau) Donovan, 1993. p. 402. Eupatagus clevei (Cotteau). Osborn et al. 2016. tbl. 2. Occurrence.—This species is very common in sporadic concentrations along the Withlacoochee River, Cross Florida Barge Canal, Dolime Quarry (CI009), and Cemex Quarry (FM-IP CI017) south of Inglis, Citrus County, where it occurs as internal and external molds in the dolomitic lowermost portion of the OLS (formerly Inglis Formation). It occurs more rarely in the overlying limestone beds of the Oligopygus phelani Zone with E. mooreanus at these same locations. It is much rarer in the Oligopygus wetherbyi Zone of the upper OLS in a quarry west of Dowling Park (FM-IP LF002), Lafayette County and south of Tennille (FM-IP DI001), Taylor County. Eupatagus clevei is widespread in the Caribbean region, including St. Bartholomew (Eocene; type locality), Panama (Eocene), Jamaica (Eocene), and Venezuela (Eocene). In Cuba, based on the synonymies and information in Kier (1984), it ranges from the Oligocene to Miocene, though the certainty of these stratigraphic designations is questionable. Discussion.—Cotteau (1875) described E. clevei and E. grandiflorus from the Eocene of St. Bartholomew. Guppy (1882) later asserted that E. grandiflorus was a junior synonym of E. clevei, an opinion subsequent authors have largely upheld. Zachos (1968) described Eupatagus ingens (Fig. 198) from the OLS of Florida. However, Kier (1984) completed an extensive review of E. clevei and considered E. ingens to be a subjective junior synonym. Subsequent workers (Carter, 1987a; Oyen, 2001) have not recognized E. ingens as a distinct species and neither do we, so it remains in synonymy with E. clevei. Eupatagus clevei is most abundant in the lower dolomitic beds of the lower OLS near Inglis where it is preserved as internal and external molds with P. floridanus. In the overlying limestone beds of the Oligopygus phelani Zone of the lower OLS, E. clevei is much rarer, but test detail is typically better preserved, as the material is not moldic (Fig. 199). In these limestone beds, E. clevei is associated with extremely abundant E. mooreanus, as well as numerous other echinoid taxa. Eupatagus clevei is much rarer in the upper OLS, where it is associated with the typical fauna of the Oligopygus wetherbyi Zone of the unit. Eupatagus mooreanus Pilsbry, 1914 (Figs. 201-207) Eupatagus mooreanus Pilsbry, 1914. p. 206. pl. 8. Eupatagus floridanus (part) Clark in Clark and ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 275 Figure 201: Comparison of Eupatagus antillarum and E. mooreanus. A: Eupatagus antillarum holotype (specimen designated as USNM 115371 by Jackson [1922] but since redesignated USNM 207225), Eocene St. Bartholomew Limestone, St. Bartholomew (image from Smithsonian NMNH online database). B: Eupatagus antillarum (UF 245010), Oligopygus phelani Zone of lower Ocala Limestone, Citrus County, Florida (FM-IP CI012). Twitchell, 1915. p. 176. pl. 83, figs. 1a-2b (not pl. 82, figs. 2a-d). Maretia twitchelli Lambert in Sánchez-Roig, 1926. p. 112. Maretia clarki Lambert in Sánchez-Roig, 1926. p. 112. Eupatagus (Gymnopatagus) mooreanus (Pilsbry). Cooke, 1942. p. 54. pl. 1, fig. 16. Eupatagus (Gymnopatagus) mooreanus (Pilsbry). Cooke, 1945. p. 60, fig. 6, no. 1. Eupatagus (Gymnopatagus) mooreanus (Pilsbry). Fischer, 1951. p. 74. pls. 2, 5, 6; txt figs. 12-17. Eupatagus (Gymnopatagus) antillarum (Cotteau). Cooke, 1959. p. 90. pl. 41, figs. 1-5 (in part, Cooke considered E. mooreanus a synonym of E. antillarum). Eupatagus antillarum (Cotteau). Toulmin, 1977. pp. 340-341. pl. 66, figs. 1-3 (in part, Toulmin followed Cooke 1959). Eupatagus mooreanus (Pilsbry). Buitrón and Sánchez, 1979. pp. 123-125. figs. 3, A-E. Eupatagus cf. antillarum (Cotteau). Donovan and Rowe, 2000. p. 659. fig. 4 (species likely referable to Eupatagus mooreanus). Eupatagus sp. cf. E. antillarum. (Cotteau). Donovan, 2004. fig.1, #8. p. 144 (species likely referable to Eupatagus mooreanus). Eupatagus antillarum (Cotteau). Osborn et al., 2016. tbl. 2. Occurrence.—This ubiquitous echinoid of the Oligopygus phelani Zone of the OLS (formerly Inglis Formation) is especially common near Inglis, in Levy and Citrus Counties. A few notable localities include the banks of the Withlacoochee River west of Inglis, especially at its mouth west of Yankeetown (type locality FM-IP LV024), as well as along the banks of the Cross Florida Barge Canal (FM-IP ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 276 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 202: Eupatagus mooreanus (UF 111613), 27.5 mm TL, 23.5 mm TW, 12.5 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Levy County, Florida (FM-IP LV035). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 277 Figure 203: Eupatagus mooreanus (UF 182819), 34 mm TL, 29 mm TW, 14.5 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Levy County, Florida (FM-IP LV035). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 278 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 204: Eupatagus mooreanus (UF 111609), 44 mm TL, 38 mm TW, 21 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Levy County, Florida (FM-IP LV035). A: aboral. B: oral. C: left side. D: right side. E: posterior. CI001). See Fischer (1951) and Cooke (1959) for additional localities. This species also occurs in the Moodys Branch Formation in Covington and Conecuh Counties, Alabama. Eupatagus mooreanus also occurs in the Eocene of Jamaica (Donovan, 2004) and Mexico (Buitrón and Sánchez, 1979). Discussion.—Eupatagus mooreanus is the unofficial fossil of the state of Florida (although most often erroneously called E. antillarum). It is ubiquitous wherever the Upper Eocene lower OLS (formerly Inglis Formation) is exposed. Fischer (1951: 74) stated that next to Periarchus floridanus, the robust E. mooreanus is the most conspicuous macrofossil in the middle portion of the “Inglis Formation” in Citrus and Levy Counties, Florida. Pilsbry (1914) described E. mooreanus from specimens collected in material dredged at the mouth of the Withlacoochee River. He referred the species to the Pliocene, but this stratum is now firmly established as Upper Eocene. Pils- bry stated that the species is about the size of E. clevei. However, additional material of both species shows this is not always the case, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 279 Figure 205: Eupatagus mooreanus (UF 338554), 59 mm TL, 51 mm TW, 32 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Citrus County, Florida (FM-IP CI012). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 280 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 206: Eupatagus mooreanus (UF 245010), 68.5 mm TL, 57.5 mm TW, 33.5 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Citrus County, Florida (FM-IP CI012). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 281 Figure 207: Eupatagus mooreanus (UF 344272), deformed specimen with merely three petals, 32.5 mm TL, 47 mm TW, 27 mm TH, Upper Eocene, Oligopygus phelani Zone, lower Ocala Limestone, Citrus County, Florida (FM-IP CI004). A: aboral. B: oral. C: left side. D: right side. E: posterior. as E. clevei can attain much greater size than E. mooreanus. Part of the early confusion with the species name for this taxon stems from Pilsbry (1914). Subsequently, Clark in Clark and Twitchell (1915), described Eupatagus floridanus from Levy County. However, of the three specimens Clark used for his description, the holotype (Clark and Twitchell, 1915: pl. 82, figs. 2a-d) from Johnson’s Sink, Levy County, is an internal mold that, as pointed out by Cooke ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 282 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) (1942), represents a different species, having much shorter and narrower ambulacra. That specimen is likely E. ocalanus, although remains uncertain due to the moldic preservation of the specimen. The other two specimens of E. floridanus that Clark in Clark and Twitchell (1915: pl. 83, figs. 1, 2) figured are what Pilsbry described a year earlier as E. mooreanus. Cooke (1942, 1959) stated that Lambert in Sánchez Roig (1926) chose the two specimens of E. mooreanus figured by Clark and Twitchell (1915) as the types of his Maretia clarki (Clark in Clark and Twitchell, 1915: pl. 83, fig. 1) and Maretia twitchelli (Clark in Clark and Twitchell, 1915: pl. 83, fig. 2), which Cooke (1942) then placed into synonymy with E. mooreanus. Mortensen (1951: 462) noted that Lambert’s taxa clearly did not belong in Maretia but was not prepared to sup- port the suggestion that they were different than E. floridanus. Fischer (1951) completed a very thorough description of E. mooreanus. Cooke, via personal communication, informed Fischer that E. mooreanus is very closely related to, and likely identical with E. antillarum, from the Eocene of St. Bartholomew. However, Fischer (1951) retained the name E. moore- anus. Cooke (1959) declared E. mooreanus a sub- jective junior synonym of E. antillarum. Cotteau’s figure of the holotype of E. antillarum is some- what stylized. However, the specimen was figured by Jackson (1922) and these two taxa are clearly not conspecific (Fig. 201). Concerning E. antillarum, Jackson (1922) stated that the small size, ovate or subovate form, and almost perfectly transverse anterior pair of am- bulacra distinguish this species from other West Indian fossil species of Eupatagus. Some of these same traits also serve to distinguish it from E. moore- anus (Fig. 201). As stated by Jackson, the anterior paired petals of E. antillarum are nearly straight, whereas those of E. mooreanus are arched anteri- orly and diverge from each other at 145° per Cooke (1959). Furthermore, the primary tubercles of E. antil- larum are retained within the peripetalous fasciole, which does not extend outward to nearly the ambitus as viewed from above, as clearly shown in the holotype of E. mooreanus. The aboral surface of E. mooreanus is much more heavily tuberculated, with the zigzagging rows of tubercles extending nearly to the ambitus, anteriorly and laterally. The structure of the primary tubercles and the anterior petals alone serve readily to distinguish E. mooreanus from E. antillarum. We do not consider E. mooreanus a subjective junior synonym of E. antillarum and in addition, do not recognize E. antillarum in the Eocene faunas of the eastern United States. Doubts concerning Cooke’s (1959) synonymy of E. mooreanus with E. antillarum are not new. Kier (1984: 84), in comparing Eupatagus alatus from the Cuban Eocene stated: “it differs from the Late Eocene Floridian specimens that Cooke referred to E. antillarum (which belong to a dif- ferent species, Eupatagus mooreanus Pilsbry) in having shorter posterior petals, a less-pointed poste- rior, and blunter anterior surface.” The documented occurrences of E. antillarum in the Caribbean re- gion need to be reconsidered, given Cooke’s (1959) synonymy, because we cannot be certain whether subsequent authors were referencing the Floridian E. antillarum (= Eupatagus mooreanus) or the typical E. antillarum from St. Bartholomew. For example, Donovan (1993: 402) discussed, but did not figure, a very worn specimen from the Eocene of Jamaica that he questionably attributed to E. antillarum. He compared it to the figures of Jackson (1922), which show E. antillarum from St. Bartholomew, so it is likely that Donovan was not comparing his material to the Florida E. antillarum (= E. mooreanus). Donovan and Rowe (2000) documented Eupatagus cf. E. antillarum from the Eocene Clare- mont Formation of Jamaica. The angle of the petals and tuberculation of the species is more reminis- cent of E. mooreanus than it is of E. antillarum (as evident in their fig. 4, #1: BMNH EE 6341). How- ever, as Donovan (1993) noted, the specimen is damaged and incomplete. Curiously, Donovan (2004: figs. 1, 8) also discussed and figured a specimen he tentatively referred to E. antillarum from the Eocene Troy ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 283 Formation of the White Limestone Group of Jamaica. However, the specimen shown is the same as that figured in Donovan and Rowe (2000: fig. 4, #1: BMNH EE 6341), from the Claremont Formation, that we concluded is E. antillarum. Buitrón and Sánchez (1979: fig. 3) docu- mented E. mooreanus from the Upper Eocene Tan Toyuca Formation in the northeastern portion of the state of Veracruz, Mexico. The figured specimens are clearly conspecific with E. mooreanus, and not E. antillarum. Eupatagus mooreanus is most abundant in its type area, where it occurs in the Oligopygus phelani Zone of the lower OLS, in the vicinity of Inglis in Citrus and Levy Counties, where it occurs with P. floridana, R. georgiensis, N. durhami, and other, rarer species. Eupatagus ocalanus Cooke, 1942 (Fig. 208) ?Eupatagus floridanus (part) Clark in Clark and Twitchell, 1915. p. 176. pl. 82, figs. 2a-d (not pl. 83, figs. 1a-c, 2a-d). Eupatagus (Plagiobrissus) ocalanus Cooke, 1942. p. 57. pl. 6, figs. 4-8. Eupatagus (Gymnopatagus) ocalanus (Cooke). Cooke, 1959. p. 91. pl. 42, figs. 8-12. Eupatagus ocalanus (Cooke). Osborn et al., 2016. tbl. 2. Occurrence.—This species occurs within the upper OLS of Florida, where it is most abundant near the top of the Wythella eldridgei Zone, es- pecially north of Mayo, Lafayette County (type locality); north of Branford (FM-IP SU002), Suwan- nee County; south of Tennille (FM-IP DI001), Dixie County; and west of Center Hill, Sumter County (FM-IP SM010). It is much more rarely found in the Oligopygus haldemani Zone where it occurs in the Brooks Quarries (FM-IP JA009, FM-IP JA018, FM- IP JA027, FM-IP JA031, FM-IP JA039), northwest of Marianna, Jackson County. Eupatagus ocalanus also occurs in the Upper Eocene of Alabama and Georgia (Cooke, 1959). Discussion.—This species occurs with P. dixie at most known localities, and can be read- ily differentiated from the much wider poriferous zones, flattened anterior ambulacral area, and dis- tinct zigzag arrangement of the primary tubercles in E. ocalanus. It is not known to occur with E. carolinensis Clark in Clark and Twitchell, 1915, but it could be confused with it. It is differentiated from E. carolinensis by its proportionately narrower and longer petals. In addition, E. carolinensis appears not to have attained as great a size as E. ocalanus (Cooke, 1942). Although E. ocalanus has not been documented in the Oligopygus phelani Zone in the lower portion of the OLS, it is readily differenti- ated from E. mooreanus, the dominant spatangoid species of the horizon, by the much less heavily tuberculated aboral surface of E. ocalanus. Eupatagus dumonti n. sp. (Figs. 209-214) Diagnosis.—Eupatagus with broad (TW on average 92.9 % TL), low (TH on average 49.1% TL) test, large periproct that encompasses nearly the entire height of the vertical part of the posterior surface (periproct width on average 71% periproct height; periproct height on average 49.6% TH), and broad, long, heavily tuberculated plastron. Description.—Description based on the holo- type (UF 238275), paratypes (UF 342109, UF 342110), and non-type specimens from the Suwan- nee Limestone in Hernando County, Florida (FM-IP HE013, FM-IP HE038). Test moderate in size (five specimens range 59.2 mm to 74.5 mm TL), broad (TW on aver- age 92.9 % TL), low (TH on average 49.1% TL), widest point posterior of apical system, highest point posterior, truncated anterior margin, lacking ante- rior sulcus. Apical system ethmolytic, madreporic plate projects well posterior of other apical system plates, anterior edge of apical system on average 64% TL from posterior margin; 4 gonopores; pos- terior gonopores further apart than anterior pair. Aboral tubercles small, not prominent, randomly arranged, largely confined aboral to peripetalous fasciole, densest in anterior paired interambulacra. Ambulacrum III not petaloid, pores small, non- ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 284 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 208: Eupatagus ocalanus (UF 129328), 46.5 mm TL, 40.5 mm TW, 21 mm TH, Upper Eocene, upper portion of Ocala Limestone, Sumter County, Florida (FM-IP SM010). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 285 Figure 209: Eupatagus dumonti n. sp., holotype, (UF 238275), 64 mm TL, 60 mm TW, 33 mm TH, Lower Oligocene, basal beds Suwannee Limestone, Hernando County, Florida (FM-IP HE013). A: aboral. B: oral. C: left side. D: right side. E: posterior. conjugate. Paired petals flush, not sunken; tapered but open distally; pores conjugate, outer pore more elongate than inner; petals I and V longest, extending almost to margin, on average 42.1% TL, petals II and IV shortest, extending two-thirds dis- tance to margin, on average 33.4% TL; petals narrow, greatest width of petal I and V on average 17.3% petal length; width of petals II and IV on average 20.9% petal length. Peripetalous and subanal fascioles present; peripetalous fasciole thin, extending to margin an- teriorly, just beyond petals I and V posteriorly, not indented aborally between petals. Subanal fasciole broadly bilobed, aboral tract passes just below lower ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 286 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 210: Eupatagus dumonti n. sp., paratype (UF 342110), 66 mm TL, 61 mm TW, 31 mm TH, Lower Oligocene, basal beds of the Suwannee Limestone, Hernando County, Florida (FM-IP HE038). A: aboral. B: posterior. C: oblique aboral viewpoint from posterior. D: posterior. E: oral. F: tilted oral viewpoint from anterior. G: tilted oral viewpoint from posterior. H: tilted aboral viewpoint from posterior. I: subanal fasciole. J: left side. K: peristome. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 287 Figure 211: Eupatagus dumonti n. sp., paratype (UF 342109), 68 mm TL, 61.5 mm TW, 32 mm TH, Lower Oligocene, basal beds of the Suwannee Limestone, Hernando County, Florida (FM-IP HE038). A: aboral. B: oblique aboral viewpoint from anterior. C: left side. D: oral. E: anterior. F: posterior. G: apical area. H: peristome. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 288 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 212: Eupatagus cf. E. dumonti n. sp., (UF 342108), 93 mm TL, 87 mm TW, 46 mm TH, Oligocene Bridgeboro Limestone, Duncan Church Beds, Washington County, Florida (FM-IP WG002). A: aboral. B: oblique lateral viewpoints, posterior to right. D: left side. E: oral. F: apical area. G: oblique aboral viewpoint. H: posterior. I: tilted aboral viewpoint from posterior. J: posterior portion of oral surface showing subanal fasciole. K: portion of subanal fasciole at top, tubercles within plastron below. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 289 Figure 213: Eupatagus dumonti n. sp., paratype (UF 341412), 74.5 mm TL, 72 mm TW, 39 mm TH, Lower Oligocene, basal beds of the Suwannee Limestone, Hernando County, Florida (FM-IP HE038). A: aboral. B: oral. C: left side. D: right side. E: posterior. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 290 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 214: Top: Comparison of test height vs test length in Eupatagus dumonti n. sp. and Macropneustes mortoni. Bottom: Comparison of periproct height vs test height in Eupatagus dumonti n. sp. and Macropneustes mortoni. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 291 edge of periproct; wider than tall, height on average 66% fasciole width; width on average 41.8% TW. Periproct on nearly vertical to slightly overhung posterior truncation, ovate, taller than wide, periproct width on average 71% periproct height; periproct very large, height on average 49.6% TH; periproct encompasses nearly entire height of posterior. Oral surface more heavily tuberculated than aboral. Peristome anterior, distance of poste- rior edge (i.e., anterior edge of labrum) to posterior margin on average 68.6% TL; kidney-shaped; wider than high, peristome height on average 49% peris- tome width; labrum elongate, narrow; plastron fully tuberculated, long, extends anteriorly nearly to peri- stome (i.e., labrum); wide, greatest width posterior and on average 36.4% TW; narrows anteriorly. Zoobank Nomenclatural Act.—2EBF3767- FC5D-4D80-A4BC-8A82F6ECF16B Discussion.—Eupatagus dumonti n. sp. represents the first known occurrence of Eupatagus in the Oligocene of North America. The species is most found in the lowermost bed of the Suwannee Limestone, which is usually exposed below the typical floor of the Vulcan Quarry northwest of Brooksville, Hernando County, where E. dumonti n. sp. occurs with Clypeaster sp., S. carlsoni n. sp., and R. gouldii. This horizon is just above the contact with the underlying OLS, and although E. dumonti n. sp. is relatively common in this bed, it is typically represented by incomplete tests. The complete specimens figured (Figs. 209-211, 213) are rare. A specimen (UF 342108) from the Oligocene Duncan Church beds of the Bridgeboro Limestone in the Florida panhandle appears to represent this species (Fig. 212). Although it is much larger than the Suwannee Limestone specimens from the type locality (93 mm TL, 87 mm TW, 46 mm TH), and the overall dimensions are slightly obscured by slight compression of the specimen, it is otherwise similar to E. dumonti n. sp. It has a TH 49% TL, and a periproct roughly 38% TH. We refer this specimen to Eupatagus cf. E. dumonti. Eupatagus dumonti n. sp., has affinities with both Macropneustes (broad, rotund test) and Eupata- gus (flush petals, lack of anterior notch). Fischer (1966) stated Macropneustes (Agassiz, 1847) differs from Eupatagus (Agassiz, 1847) chiefly in having depressed petals, a broad test, and distinct anterior notch. Duncan (1889) noted the similarities of Eu- patagus and Macropneustes and made Macrop- neustes a subgenus of Eupatagus. However, Mortensen (1951) asserted that although Macrop- neustes clearly has some relation to Eupatagus, it should be treated as a distinct genus. Subsequently, echinologists have continued to recognize Macrop- neustes as distinct from Eupatagus. Cooke (1959) noted that Macropneustes shows significant variation in the depth of the am- bulacral depressions, with Middle Eocene species usually having a conspicuous anterior notch and moderately depressed petals, but the Late Eocene species tend to have almost flush petals. Kier (1984) stated Macropneustes is difficult to distinguish from Eupatagus, but clarified the petals are generally nar- rower and that a distinct anterior ”groove” is present in Macropneustes. But his use of the term groove is unfortunate because this anterior sinus (to use Fischer’s word) is often merely a slight indentation in the posterior margin, as seen in M. mortoni, and could not be recognized as a groove. We prefer the term ”notch”, which does not connote an impression of considerable depth, and distinguishes this feature from the sinus associated with the periproct in some cassiduloids. We also reserve the term groove for the narrow depression in which hydropores occur, or for food grooves of clypeasteroids and scutelloids. The flush petals and lack of a discernable anterior notch place this species in Eupatagus. We compare Eupatagus dumonti n. sp., to known species of both Macropneustes and Eupatagus from the eastern Americas. In the North American faunas, Macropneustes is represented only by M. mortoni from the Upper Eocene OLS of the eastern Gulf Coast. Eupatagus dumonti n. sp. is similar in general morphology to M. mortoni but has a lower test and significantly larger periproct (Fig. 214). The average periproct height of M. mortoni is 26.9 % TH. The average periproct height in E. dumonti n. sp. is 49.6% TH (Fig. 214). In addition, on average, the TH of M. mortoni is 62.1% TL, whereas in E. dumonti n. sp. it is 49.1%. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 292 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Eupatagus dumonti n. sp. is further distinguished from M. mortoni by its shorter petals, which are flush and not at all depressed. Macropneustes is sparsely represented in the fossil faunas of the Caribbean region and remain- der of the eastern Americas. Kier (1984) stated that in the Cuban faunas, only the Late Eocene Macropneustes (Deakia) armadilloensis Sánchez- Roig, 1953b belongs to Macropneustes. This is the only species within the subgenus Deakia in the Western Hemisphere (Kier, 1984). Kier (1984) ten- tatively placed Macropneustes cubensis Cotteau, 1875 and M. palmeri (Sánchez-Roig, 1953b), from the Oligocene-Miocene of Cuba, within this genus. Eupatagus dumonti n. sp., is not at all like any of these species, which were all figured and described well by Kier (1984). Arnold and Clark (1927, 1934) named six species of Macropneustes from the Eocene of Ja- maica: M. altus Arnold and Clark, 1927; M. au- gustus Arnold and Clark, 1927; M. parvus Arnold and Clark, 1927; M. dyscritus Arnold and Clark, 1934; M. sinuosus Arnold and Clark, 1934, and M. stenopetalus Arnold and Clark, 1934. As noted by Mortensen (1951), these species are generally very poorly preserved, and as noted by Arnold and Clark (1927), their actual assignment to genus is ques- tionable. Mortensen (1951: 451) stated: “it might perhaps have been better to leave these poor speci- mens undescribed.” The figures of Arnold and Clark (1927, 1934) are sufficient to permit comparison with E. dumonti n. sp. which does not resemble any of them closely enough to warrant further consid- eration. However, even the most poorly preserved of these specimens shows distinctly sunken petals, among other features that distinguish them from E. dumonti n. sp. Macropneustes dubius Israelsky, 1924 was described from the Miocene of the Tampico region of Mexico, but among other features, its deeply sunken petals are not at all similar to E. dumonti n. sp. Kew (1917) described Macropneustes mexicanus from the Oligocene to Miocene of Mexico, but this species has been referred to Gillechinus (Henderson and Fell, 1969). Peripneustes antillarum Cotteau, 1875, was described from the Eocene of St. Bartholomew, and subsequently recorded from Cuba (Cottteau, 1897; Kier, 1980), and potentially Veracruz, México (Is- raelsky, 1924). This species was subsequently placed in Macropneustes (Guppy, 1882; Jackson, 1922), but then in Meoma by Chesher (1970) and Kier (1984). It is clearly distinct from E. dumonti n. sp. Eupatagus is commonly found in the Eocene strata of the region, and less so in overlying Oligocene or Neogene strata of the region. Dis- tinguishing between these myriad fossil forms is not easy. As noted by Mortensen (1951: 463) when he stated the following concerning the genus: “going over the numerous fossil species I have the impres- sion that it must be an exceedingly difficult task to dis- tinguish them with certainty.” We try to do so below. Eupatagus was previously represented by eight species in the fossil record of the eastern United States. All are Eocene: E. carolinensis Clark in Clark and Twitchell, 1915; E. clevei; E. gardnerae; E. lawsonae Kier, 1980; E. mooreanus; E. ocalanus; E. texanus Zachos in Zachos and Molineux, 2003, and E. wilsoni Kier, 1980. The broad test and dis- tinctly larger periproct separate E. dumonti n. sp. from any of these species. Kier (1984: 95), recognized the following Cuban species as distinct: E. alatus Arnold and Clark, 1927 (Middle to Late Eocene); E. clevei (Eocene to Oligocene); E. cubensis (Cotteau, 1875) (Eocene to Miocene); E. sanchezi (Lambert in Sánchez-Roig, 1949) (Miocene); E. santanae Sánchez-Roig, 1951 (Oligocene to Miocene); E. turibacoensis Sánchez-Roig, 1953b (Late Eocene), and E. siboneyensis Weisbord, 1934 (Late Eocene). These seven species are reduced from 41 species documented from the Cuban faunas that Kier (1984) examined, eight of which were unrecognizable and had to be set aside, but the remainder were placed in synonymy of the preceding seven species. Many of these were described in the works of Sánchez-Roig (1949, 1951, 1953, inter alia). Although poorly fig- ured by the latter, the taxa recognized by Kier (1984) are not conspecific with Eupatagus dumonti n. sp. Arnold and Clark (1927, 1934) and Hawkins in Arnold and Clark (1927) documented seven species of Eupatagus from Jamaica. All are Eocene ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 293 with the exception of the Oligocene occurrence of E. hildae Hawkins, 1927 (= E. clevei), including: E. grandiflorus Cotteau, 1875 (= E. clevei); E. alatus, which Kier (1984) identified in the Cuban faunas; E. attenuatus (Arnold and Clark, 1927) which Kier (1984: 7) stated is likely synonymous with E. alatus; E. longipetalus Arnold and Clark, 1927, which Kier (1984: 7) placed in Antillaster; E. defectus Arnold and Clark, 1927; E. hildae Hawkins, 1927, from the Oligocene which Kier (1984) considered synony- mous with E. clevei, and Eupatagus cf. E. antillarum (Cotteau, 1875). These Jamaican occurrences were further discussed by Dixon and Donovan (1994), Donovan (1994, 2003, 2004), and Donovan and Rowe (2000). Based on comparison with all these records, E. dumonti n. sp. cannot be confused with any of them. E. clevei and E. antillarum are documented from their type localities in the Eocene of St. Bartholomew and elsewhere (Cotteau, 1875; Jack- son, 1922; Cooke, 1959; Cooke, 1961; Kier, 1984), and Pijpers (1933) named Brissoides aloysii from the Eocene of Bonaire, which is clearly a Eupatagus. Jackson (1922) documented the occurrence of E. depressus Jackson, 1922, in the Oligocene of Puerto Rico, which was considered by Kier (1984) to be a synonym of E. cubensis (Cotteau, 1875). Jack- son (1922) also named E. elegans and E. vaughani, which Kier (1984) suggested belonged to Antillaster. Eupatagus dumonti n. sp. cannot be confused with any of these species. Some Caribbean species of Antillaster (Lam- bert, 1909) are very similar in general morphology to E. dumonti n. sp., such as Eupatagus mexicanus Jackson, 1937, from the Oligocene of Mexico. How- ever, Kier (1984) placed E. mexicanus in Antillaster and then synonymized it with A. vaughani (Jackson 1922), first described from the Oligocene-Miocene of Antigua. Antillaster is a wide-spread genus in Cenozoic strata of the Caribbean region and north- eastern South America. However, it has not been documented from North America. Antillaster is readily distinguished from Macropneustes and Eu- patagus by the absence of fascioles, whereas E. dumonti n. sp. has both peripetalous and subanal fascioles. Eupatagus dumonti n. sp., is similar to Eu- patagus longipetalus Clark, 1927, from the Eocene of Jamaica. Although the petals and general mor- phology (shape and size) are similar, Kier (1984) placed this species in Antillaster, presumably due to the lack of discernable fascioles in the type and only known specimen. Eupatagus dumonti n. sp. represents the first documented occurrence of Eupatagus in the Oligocene of North America. Including the 17 species of Eupatagus previously recognized in the fossil record of the eastern Americas and Caribbean region reviewed above, 15 are documented from the Eocene. Peak of diversity of Eupatagus in the Eocene is not restricted to the Americas, as this seems to be true globally as well: Lambert and Thiéry (1925) listed 33 species of Eupatagus from the Eocene, Kier and Lawson (1978) listed an additional 27 species, and Kroh (2010) documented another 13 species from the global faunas, of which five are Eocene. Therefore, no fewer than 65 species of Eupatagus have been documented from Eocene strata worldwide. Etymology.—Named in honor of Joe Dumont, collector and donor of many of the specimens used to describe this species. Material and Occurrence.—Eupatagus du- monti n. sp. is represented by the holotype (UF 238275) and paratypes (UF 342109, UF 342110, UF 341412) from the type locality in the lower beds of the Suwannee Limestone in the Vulcan Quarry, west of Brooksville, Hernando County (FM-IP 6360, FM-IP HE013, FM-IP HE038). It is also represented by a single specimen UF 342108 we tentatively as- sign to this species from the Duncan Church beds of the Bridgeboro Limestone in the Duncan Church Quarry, south of Chipley, Washington County (FM-IP WG002). Eupatagus sp. A (Fig. 215) Occurrence.—Upper Eocene OLS, (occurs with H. brooksi), Jackson Blue Spring (FM-IP JA033), east of Marianna, Jackson County. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 294 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Figure 215: Eupatagus sp. A, (UF 341760), 22.1 mm TL, 16.3 mm TW, 9.5 mm TH (TL and TH are incomplete) Upper Eocene Ocala Limestone (Haimea brooksi Zone) Jackson Blue Spring (FM-IP JA033), Jackson County, Florida A: aboral. B: anterior. C: left side. D: tilted aboral viewpoint from posterior. E: posterior (largely missing). F: oblique aboral viewpoint from anterior. G: tilted aboral viewpoint from anterior. H: petaloid area with cross lighting to highlight seemingly randomly arranged, sparse, large tubercles. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 295 Figure 216: Eupatagus sp. B, (UF 344742), 108 mm TL, 137 mm TW, 59 mm TH (no dimensions are complete), Lower Oligocene Suwannee Limestone, Washington County, Florida (FM-IP WG034). A: aboral. B: oral. C: left side. D: right side. E: posterior. Discussion.—This species is represented by a single specimen collected at Jackson Blue Spring. The incomplete specimen was associated with H. brooksi, R. georgiensis, and other species, and is the only representative of this form found within these deposits. The specimen is corroded and missing much of the posterior and oral surfaces but does show traces of a peripetalous fasciole. This fragment (UF 341760) is small (22.1 mm TL, 16.3 mm TW, 9.5 mm TH) but TL and TH are not accurate assessments due to the missing portions of the test. The test has sparse, seemingly random, large, perforate tubercles, flush petals, and a rounded anterior margin that lacks a notch. It is not like any other spatangoid, including any documented species of Eupatagus from the North American or Caribbean faunas, but remains is too incomplete to describe. We document it as a taxon in Eupatagus ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 296 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) facies of the Bridgeboro Limestone, which occurs near the surface in the Duncan Church Quarry (FM- IP WG002) roughly 30 km northeastward. However, R. gouldii is not present in the Duncan Church beds of the Bridgeboro Limestone. The stratigraphic deter- mination is therefore questionable, but its Oligocene age is certain. None of the dimensions of this specimen is complete (108 mm TL, 137 mm TW, 59 mm TH) but enough of the test is preserved to determine that it was very large, almost undoubtedly the largest documented from the Oligocene strata of North America. The presence of a peripetalous fasciole rules out assignment to Antillaster (a genus also containing some very large representatives in the Eocene and Oligocene of the Caribbean region), and likely belongs in Eupatagus. As for Eupatagus sp. A, above, we leave this taxon in open nomenclature pending discovery of better specimens. CLASSIFICATION OF SPECIES TREATED Class ECHINOIDEA Schumacher, 1817 Subclass CIDAROIDEA Smith, 1984 Order CIDAROIDA Claus, 1880 Family CIDARIDAE Gray, 1825 Genus Phyllacanthus Brandt, 1835 Phyllacanthus mortoni (Conrad, 1850) Genus Prionocidaris A. Agassiz, 1863 Prionocidaris robertsi n. sp. Subclass EUECHINOIDEA Bronn, 1860 Infraclass AULODONTA Jackson, 1912 Aulodonta gen. et. sp. indet. Infraclass CARINACEA Kroh and Smith, 2010 Order PHYMOSOMATOIDA Mortensen, 1904 Family PHYMOSOMATIDAE Pomel, 1883 Genus Acanthechinus Duncan and Sladen, 1882 Acanthechinus dixie (Cooke, 1941a) Order STOMOPNEUSTOIDA Kroh and Smith, 2010 Family STOMOPNEUSTIDAE Mortensen, 1903 Genus Phymotaxis Lambert and Thiéry, 1914 Phymotaxis mansfieldi Cooke, 1941a Order CAMARODONTA Jackson, 1912 ”Triplacidiids” temporary family (Kroh and Mooi, 2022) Genus Gagaria Duncan, 1889 Gagaria mossomi (Cooke, 1941a) Family TRIGONOCIDARIDAE Mortensen, 1903 Genus Brochopleurus Fourtau, 1920 Brochopleurus pretiosus (Clark, 1915) Family TOXOPNEUSTIDAE Troschel, 1872 Genus Lytechinus A. Agassiz, 1863 Lytechinus floralanus (Cooke, 1941a) Euechinoidea gen. et. sp. indet. Infraclass IRREGULARIA Latreille, 1825 Subterclass NEOGNATHOSTOMATA Smith, 1981 Order ECHINONEOIDA H.L. Clark, 1925 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 297 Family ECHINONEIDAE Agassiz and Desor, 1847 Genus Amblypygus L. Agassiz, 1840 Amblypygus americanus Michelin, 1856 Superorder LUMINACEA Mongiardino Koch, Thompson, Hiley, McCowin, Armstrong, Coppard, Aguilera, Bronstein, Kroh, Mooi, & Rouse, 2022 Order CLYPEASTEROIDA A. Agassiz, 1872 Family OLIGOPYGIDAE Duncan, 1889 Genus Haimea Michelin, 1851 Haimea brooksi Osborn et al., 2016 Genus Oligopygus de Loriol, 1887 Oligopygus haldemani (Conrad, 1850) Oligopygus phelani Kier, 1967 Oligopygus rotundus Cooke, 1942 Oligopygus wetherbyi de Loriol, 1887 Suborder CLYPEASTERINA L. Agassiz, 1835 Family CLYPEASTERIDAE L. Agassiz, 1835 Genus Clypeaster Lamarck, 1801 Clypeaster cotteaui Egozcue in Cotteau, 1897 Clypeaster marinanus Jackson, 1937 Clypeaster oxybaphon Jackson, 1922 Clypeaster rogersi (Morton, 1834) Clypeaster sp. A Clypeaster sp. B Clypeaster sp. C Order ECHINOLAMPADACEA Mongiardino Koch et al., 2018 Suborder CASSIDULOIDA Agassiz and Desor, 1847 Family EURHODIIDAE Souto et al., 2019 Genus Eurhodia Haime in d’Archiac and Haime, 1853 Eurhodia patelliformis (Bouvé, 1851) Family CASSIDULIDAE L. Agassiz and Desor, 1847 Genus Rhyncholampas A. Agassiz, 1869 Rhyncholampas conradi (Conrad, 1850) Rhyncholampas ericsoni (Fischer, 1951) Rhyncholampas fontis (Cooke, 1942) Rhyncholampas georgiensis (Twitchell, 1915) Rhyncholampas gouldii (Bouvé, 1846) Rhyncholampas trojanus (Cooke, 1942) Rhyncholampas mariannaensis n. sp. Rhyncholampas bao n. sp. Suborder ECHINOLAMPADOIDA Kroh and Smith, 2010 Family ECHINOLAMPADIDAE Gray, 1851 Genus Echinolampas Gray, 1825 Echinolampas aldrichi Twitchell, 1915 Echinolampas tanypetalis Harper and Shaak, 1974 Suborder SCUTELLOIDA Mongiardino Koch et al., 2018 Infraorder LAGANIFORMES Desor in L. Agassiz and Desor, 1847 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 298 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Family FIBULARIIDAE Gray, 1855 Genus Echinocyamus van Phelsum, 1774 Echinocyamus macneili Cooke, 1959 Genus Fibularia Lamarck, 1816 Fibularia vaughani (Twitchell, 1915) Family NEOLAGANIDAE Durham, 1954 Genus Neolaganum Durham, 1954 Neolaganum archerensis (Twitchell, 1915) Neolaganum dalli (Twitchell, 1915) Neolaganum durhami Cooke, 1959 Genus Durhamella Kier, 1968 Durhamella ocalana (Cooke, 1942) Durhamella floridana (Twitchell, 1915) Durhamella tetrapora n. sp. Genus Weisbordella Durham, 1954 Weisbordella cubae (Weisbord, 1934) Weisbordella johnsoni (Twitchell, 1915) Weisbordella inglisensis n. sp. Weisbordella libum n. sp. Genus Wythella Durham, 1954 Wythella eldridgei (Twitchell, 1915) Family SCUTELLINIDAE Pomel, 1888 (Scutelliformes stem group) Genus Porpitella Pomel, 1883 Porpitella micra H. L. Clark, 1937 Infraorder SCUTELLIFORMES Haeckel, 1896 Family PROTOSCUTELLIDAE Durham, 1955 Genus Periarchus Conrad, 1866 Periarchus floridanus Fischer, 1951 Periarchus quinquefarius (Say, 1825) Genus Protoscutella Stefanini, 1924 Protoscutella pentagonium Cooke, 1942 Subterclass ATELOSTOMATA von Zittel, 1879 Order SPATANGOIDA L. Agassiz, 1840 Suborder MICRASTERINA Fischer, 1966 incertae sedis Genus Gillechinus Fell, 1964 Gillechinus alabamensis (Cooke, 1942) Suborder PALEOPLEUSTINA Markov and Solovjev, 2001 Family SCHIZASTERIDAE Lambert, 1905 Genus Ova Gray, 1825 Ova beckeri (Cooke, 1942) Ova ocalanus (Cooke, 1942) Genus Schizaster L. Agassiz, 1836 Schizaster americanus Clark, 1915 Schizaster armiger Clark, 1915 Schizaster carlsoni n. sp. Family PRENASTERIDAE Lambert, 1905 ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida 299 Genus Agassizia Valenciennes, 1846 Agassizia mossomi Cooke, 1942 Genus Prenaster Desor, 1853 cf. Prenaster sp. Suborder BRISSIDINA Kroh and Smith, 2010 Family BRISSIDAE Gray, 1855 Genus Brissopsis L. Agassiz, 1840 Brissopsis steinhatchee Cooke, 1942 cf. Brissopsis sp. Genus Brissus Gray, 1825 Brissus bridgeboroensis Carter, 1987b Brissus jonesi n. sp. Genus Plagiobrissus Pomel, 1883 Plagiobrissus curvus (Cooke, 1942) Plagiobrissus dixie (Cooke, 1942) Plagiobrissus cassadyi n. sp. Superfamily SPATANGOIDEA Gray, 1825 Family MACROPNEUSTIDAE Lambert, 1905 Genus Macropneustes L. Agassiz, in Agassiz and Desor, 1847 Macropneustes mortoni (Conrad, 1850) Family EUPATAGIDAE Lambert, 1905 Genus Eupatagus L. Agassiz, in Agassiz and Desor, 1847 Eupatagus clevei (Cotteau, 1875) Eupatagus mooreanus Pilsbry, 1914 Eupatagus ocalanus Cooke, 1942 Eupatagus dumonti n. sp. Eupatagus sp. A Eupatagus sp. B CONCLUSIONS This work documents occurrences of all echinoids currently known from Florida’s Paleogene strata. Our work is based on decades of extensive col- lecting, thorough examination of the FM-IP collec- tions, the collections of regional institutions, and the collections of numerous regional avocational collectors. These collectors have contributed much through their diligence, careful documentation, and willingness to share important specimens. Despite intense efforts of professional pale- ontologists and avocational collectors alike, there undoubtedly remain large numbers of undocumented echinoid species in Florida. It is hoped that a mono- graphic treatment such as this will inspire and aid those who will have further interest in the echinoids of these remarkable fossil-bearing strata spanning crucial time intervals in the evolutionary history of the Echinoidea. The present work is not intended to be the final word, but a progress report for a better understanding of the evolution and diversity of these animals within the sedimentary sequences of Florida. ACKNOWLEDGMENTS We thank the following individuals for contributions that led directly to the completion of this work, either through specimen donation, time spent col- lecting in the field, or discussions about echinoids that helped improve the manuscript. Leon Brooks, ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 300 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) owner of the Marianna Lime Quarry in Marianna was beyond generous with his time and access to his quarry. Without his kindness, Rhyncholampas mariannaensis n sp., Rhyncholampas bao n. sp., and Brissus jonesi n. sp. would remain undocumented. Larry Rogers and Sandy Owens (Limestone Products, Inc., FM-IP AL004) and Wayne Beaver (Denali Quarry, FM-IP SU003) allowed RWP decades of unfettered access for research and education. Sean and Rebecca Roberts collected, by SCUBA, most of the specimens from Jackson Blue Spring that are figured within this work, and Sean took many of the digital images, herein. Don Clements and Don Rideout guided AO on numerous trips to the type area of Echinocyamus parvus in Craven County, North Carolina in an effort to collect specimens to enable a better understanding of this poorly known species, specimens of which are figured as comparisons to Echinocyamus from Florida. John DuBois facilitated access to numerous Carolina quarries for comparative specimens, some of which we have figured. George Phillips guided AO and RWP to Paleogene exposures in the coastal plain of Mississippi and Alabama, and specimens of C. rogersi from these outings are herein figured. We are also indebted to the efforts of nu- merous collectors that either provided specimens or better-preserved exemplars necessary for this work. These individuals include Ron Bear, Tina Bell, Bill Birdsall, Mary Jo Bopp, Jon Bryan, Tabitha Cale, Robert Carlson, Burt Carter, Rick Carter, Tim Cassady, Don Clements, Wendy Conway, Steve Denahan, Joseph Dumont, Wiley Dykes, Barbara Fite, Dan Frederick, Muriel Hunter, Linda Ivany, David Kendrick, Curtis Klug, Carl Mayers, Mar- shal McCullough, Guy Harley Means, Jack Meeder, Gary Morgan, Craig Oyen, Pete Paradise, Carol and Bernie Peterson, Al Rieck, Sean and Rebecca Roberts, Larry Rooks, Craig and Laurie Samuel, Kevin Schindler, Thomas Scott, Steve and Michael Sharpe, Byron and Cindy Shumaker, John Waldrop, Charles West, and Louis Zachos. Curation assistance at the FLMNH was pro- vided by Sean Roberts, Samantha Zbiden, and Tin Yan (Jessie) So. Specimen loans by Terry Harrell (Geological Survey of Alabama) and Jessica Nakano and Gene Hunt (National Museum of Natural His- tory, Smithsonian Institution) are gratefully acknowl- edged. Funding for fieldwork and equipment related to this project was provided to RWP by Barbara Toomey and son James Toomey and the FLMNH Invertebrate Paleontology Endowment. The Florida Geological Survey is also thanked for transferring both their micro- and macro- invertebrate collections to the Florida Museum of Natural History. Some specimens used in this project were curated and available for study due to National Science Founda- tion collection improvement grants received by RWP (DBI 0645865, DBI 1055588, and DBI 1458151) and a National Science Foundation Collaborative Research grant to RM (DBI 2036298). In addition, we are indebted to James H. Nebelsick (University of Tübingen) and Jonathan Bryan (Northwest Florida State College) for their thoughtful reviews, though any errors or omissions are solely the responsibility of the authors. This is University of Florida Contribution to Paleobiology 886. LITERATURE CITED Agassiz, A. 1863. 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Casopis pro mineralogii a geologii 26:273–284, pls 1–4. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-1 LOCALITY APPENDIX FM-IP FLORIDA LOCALITIES 2409 Levy County, 1.0 mi. west of US19 along Cross Florida Barge Canal, Upper Eocene lower OLS. 2432 Alachua County, 5.0 mi. southwest of Gainesville at Gainesville Limestone Pit on Archer Road, Upper Eocene OLS. 2434 Alachua County, Upper Eocene OLS. 2613 Alachua County, in and around Archer, Upper Eocene OLS. 3142 Levy County, 4.0 mi. north of Williston at Johnsons Sink on southwest end of Johnson’s Lake, Upper Eocene OLS. 3573 Polk County, Lake Wales from well at 200 ft. depth, Middle Eocene Avon Park Formation. 5649 Taylor County, Upper Eocene upper OLS. 6340 Hernando County, Lower Oligocene Suwannee Limestone. 6341 Citrus County, Upper Eocene lower OLS. 6360 Hernando County, Upper Eocene OLS. AL002 (FT. CLARK QUARRY) Alachua County, quarry southwest of Gainesville on Ft. Clark Church Road (29.631667°, -82.440807°) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ A1-2 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) (WGS84), Upper Eocene OLS. AL004 (DICKERSON LIMEROCK MINES (HAILE COMPLEX) Alachua County, 5.0 mi. northeast of Newberry (29.678299°, -82.568823°) (WGS84), Upper Eocene OLS. AL030 (OLD SANTE FE RIVER BRIDGE 01) Alachua County, 1.0 mi. north of High Springs along riverbank just west of US441 bridge over Sante Fe River (29.851999°, -82.611524°) (WGS84), Lower Oligocene Suwannee Limestone. CI001 (INGLIS 01A) Citrus County, 1.5 mi. southwest of Inglis along Florida Cross State Barge Canal (29.007775°, -82.688094°) (WGS84), Upper Eocene lower OLS. CI003 (FLORIDA BARGE CANAL, NORTH BANK) Citrus County, 1.0 mi. south of Inglis along north bank of Florida Cross State Barge Canal, east of US98 bridge (29.014996°, -82.658378°) (WGS84), Upper Eocene lower OLS. CI004 (FLORIDA CROSS STATE CANAL 02) Citrus County, approximately 6.0 mi. west of Inglis, spoil banks on an island approximately 2.0 mi. offshore from the west end of Florida Cross State Barge Canal (28.981799°, -82.765421°) (WGS84), Upper Eocene lower OLS. CI009 (DOLIME QUARRY 01) Citrus County, 2.5 km south of Inglis; southeast side of Florida Cross State Barge Canal at US19/98 Bridge (29.010862°, -82.653961°) (WGS84), Upper Eocene lower OLS. CI010 (MAPLE SUGAR CAVE) Citrus County, about 10 mi. south of Lecanto in cave (28.7017°, -82.4629°) (NAD27), Upper Eocene lower OLS. CI012 (FLORIDA CROSS STATE CANAL 03) Citrus County, approximately 2.5 mi. southwest of Inglis on spoil pile approximately 2 mi. west of US19 Bridge (29.0060870°, -82.6940940°) (WGS84), Upper Eocene lower OLS. CI014 (RED LEVEL DOLOMITE MINE) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-3 Citrus County, Red Level Quarry complex on west side of US19/US98 (28.965858°, -82.640819°) (NAD27), Upper Eocene lower OLS. CI015 (FLORIDA CROSS STATE CANAL 04) Citrus County, approximately 1.5 mi. southwest of Inglis, 1.25 mi. southwest of US19 Bridge on north bank of canal (29.009092°, -82.683842°) (WGS84), Upper Eocene lower OLS. CI017 (INDEPENDENT AGGREGATES 01) Citrus County, quarry approximately 2 mi. southwest of Inglis (29.014478°, -82.702498°) (WGS84), Upper Eocene lower OLS. CI020 (FLORIDA CROSS STATE CANAL 06) Citrus County, 3.6 mi. southwest of Inglis, Florida Cross State Barge Canal 3.5 mi. west of US19 (29.001074°, -82.715253°) (WGS84), Upper Eocene lower OLS. CI021 (FLORIDA CROSS STATE CANAL 07) Citrus County, 4.6 mi. west southwest of Inglis along Barge Canal (28.9956°, -82.7353°) (NAD27), Upper Eocene lower OLS. CI022 (FLORIDA CROSS STATE CANAL 08) Citrus County, southwest of Inglis along Barge Canal (29.0055°, -82.6918°) (NAD27), Upper Eocene lower OLS. CI024 (FLORIDA CROSS STATE CANAL 10) Citrus County, 3-4 mi. west southwest of Barge Canal in Gulf of Mexico (28.9726°, -82.7922°) (NAD27), Upper Eocene lower OLS. CI038 (INGLIS 05) Citrus County, 4.23 mi. west southwest of Inglis, U.S.E.D. Well on south side of the Florida Cross State Barge Canal (28.997472°, -82.727882°) (WGS84), Upper Eocene lower OLS. CI052 (CRACKERTOWN 03) Citrus County, south side of Crackertown on south bank of the Withlacoochee River, 0.25 mi. up and down river from Faris Landing (29.032574°, -82.688149°) (WGS84), Upper Eocene lower OLS. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ A1-4 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) CR013 (BELLE MEADE 01) Collier County, about 3 mi. northwest of Golden Gate (26.2443°, -81.6827°) (NAD27), Middle Eocene Avon Park Formation. DI001 (STEINHATCHEE 02) Dixie County, near Tennille at quarry east of Steinhatchee River (29.769017°, -83.317604°) (WGS84), Upper Eocene OLS. DI013 (STEINHATCHEE 05) Dixie County, near Tennille at quarry east of Steinhatchee River (29.769017°, -83.317604) (WGS84), Upper Eocene upper OLS. HE006 (RIDGE MANOR 01) Hernndo County, Ridge Manor in limerock quarry northeast of corner of US301 and SR50 (28.519215°, -82.160042°) (NAD27), Upper Eocene OLS. HE007 (BROOKSVILLE ROCK QUARRY 01) Hernando County, northwest of Brooksville at Cargil Vulcan Quarry, 4.25 mi. south of county line and 2.0 mi. southwest of US98 (28.6316°, -82.4806°) (NAD27), Lower Oligocene Suwannee Limestone. HE012 (BROOKSVILLE ROCK QUARRY 02) Hernando County, 9.01 mi. northwest of Brooksville at Cargil Vulcan Quarry, mine 2.05 mi. south southeast of intersection of SR589 and US98 (28.6562260°, -82.4851720°) (NAD27), Lower Oligocene Suwannee Limestone. HE013 (BROOKSVILLE ROCK QUARRY [GENERAL 01]) Hernando County, Brooksville Rock Company Quarry 4.0 mi. south of county line and 1.5 mi. southeast of US98 (28.636318°, -82.476333°) (NAD27), Lower Oligocene Suwannee Limestone. HE019 (BROOKSVILLE ROCK QUARRY 06) Hernando County, Brooksville Rock Quarry 1.57 mi. northeast of the intersection of SR589 and Centralia Road (28.629629°, -82.474583°) (NAD27), Lower Oligocene Suwannee Limestone. HE026 (BROOKSVILLE ROCK QUARRY 12) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-5 Hernando County, 9.0 mi. northwest of Brooksville at Cargil Vulcan Quarry, pit 2.09 mi. south southeast of intersection of SR589 and US98 (28.65453°, -82.483983°) (NAD27), Lower Oligocene Suwannee Limestone. HE028 (BROOKSVILLE ROCK QUARRY 14) Hernando County, Brooksville Rock Quarry 1.33 mi. northeast of the intersection of SR589 and Centralia Road (28.628452°, -82.478524°) (NAD27), Lower Oligocene Suwannee Limestone. HE034 (BROOKSVILLE ROCK QUARRY 17) Hernando County, Brooksville Rock Quarry 1.72 mi. northeast of the intersection of SR589 and Centralia Road (28.630768°, -82.472073°) (WGS84), Lower Oligocene Suwannee Limestone. HE038 (BROOKSVILLE ROCK QUARRY 20) Hernando County, Brooksville Rock Quarry 1.5 mi. northeast of the intersection of SR589 and Centralia Road (28.629297°, -82.475314°) (WGS84), Lower Oligocene Suwannee Limestone. JA009 (MARJAX PIT) Jackson County, 3.38 mi. northwest of Marianna, 1.5 mi. northwest of the junction of US90 and SR73 (30.801477°, -85.274942°) (WGS84), Upper Eocene upper OLS. JA010 (DRY CREEK 01) Jackson County, 2.5 mi. southwest of Oakdale at Dry Creek (30.673458°, -85.207758°) (WGS84), Lower Oligocene Marianna Limestone. JA013 (MARJAX PIT A) Jackson County, 3.38 mi. northwest of Marianna (30.801477°, -85.274942°) (WGS84), Lower Oligocene Marianna Limestone. JA014 (OAKDALE 01) Jackson County, quarry north of Altha (30.639833°, -85.173137°) (WGS84), Lower Oligocene Marianna Limestone. JA018 (MARIANNA LIME PRODUCTS) Jackson County, quarry northwest of Marianna (30.807°, -85.2606°) (NAD27), Upper Eocene upper OLS. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ A1-6 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) JA019 (MARIANNA LIME PRODUCTS 01) Jackson County, quarry northwest of Marianna (30.807°, -85.2606°) (NAD27), Lower Oligocene Marianna Limestone. JA020 (MARIANNA LIME PRODUCTS 02) Jackson County, quarry northwest of Marianna (30.807°, -85.2606°) (NAD27), ex situ, Upper Eocene upper OLS, Upper Eocene to Lower Oligocene Bumpnose Limestone, and Lower Oligocene Marianna Limestone. JA021 (MARIANNA LIME PRODUCTS 03) Jackson County, quarry northwest of Marianna (30.807°, -85.2606°) (NAD27), Upper Eocene to Lower Oligocene Bumpnose Limestone. JA025 (MARIANNA LIME PRODUCTS 05) Jackson County, quarry northwest of Marianna (30.8052°, -85.2634°) (NAD27), Upper Eocene to Lower Oligocene Bumpnose Limestone. JA026 (MARIANNA LIME PRODUCTS 06) Jackson County, quarry northwest of Marianna (30.8052°, -85.2634°) (NAD27), Lower Oligocene Marianna Limestone. JA027 (MARIANNA LIME PRODUCTS 07) Jackson County, quarry northwest of Marianna (30.8052°, -85.2634°) (NAD27), Upper Eocene upper OLS. JA029 (MARIANNA LIME PRODUCTS 09) Jackson County, quarry northwest of Marianna (30.8068°, -85.2567°) (NAD27), Lower Oligocene Marianna Limestone. JA030 (MARIANNA LIME PRODUCTS 10) Jackson County, quarry northwest of Marianna (30.8068°, -85.2567°) (NAD27), Upper Eocene to Lower Oligocene Bumpnose Limestone. JA031 (MARIANNA LIME PRODUCTS 11) Jackson County, quarry northwest of Marianna (30.8068°, -85.2567°) (NAD27), Upper Eocene upper OLS. JA033 (JACKSON BLUE SPRING 01) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-7 Jackson County, 5.8 km east northeast of Marianna at head of Merritt’s Millpond (30.7903°, -85.1402°) (NAD27), Upper Eocene OLS. JA039 (MARIANNA LIME PRODUCTS 13) Jackson County, northwest of Marianna (30.811°, -85.2689°) (NAD27), Upper Eocene upper OLS. JA084 (MARIANNA 12 [TYPE MARIANNA LIMESTONE]) Jackson County, about 1 mi. east of Marianna (30.773°, -85.2165°) (NAD27), Lower Oligocene Marianna Limestone. JA085 (CHIPOLA 29) Jackson County, east side of Marianna along Chipola River between US90 Bridge and SR166 Bridge (30.784072°, -85.213872°) (WGS84), Upper Eocene upper OLS. JA086 (SILLS PIT 06) Jackson County, approximately 7.0 mi. north northeast of Cottondale, east of Union Road and northwest of Waddell’s Millpond (30.88°, -85.3489°) (WGS84), Upper Eocene upper OLS. LF001 (DELL LIMEROCK MINE) Lafayette County, 4.5 mi. northwest of Mayo (30.0894°, -83.2345°) (NAD27), Upper Eocene upper OLS. LF002 (MILL CREEK QUARRY 01) Lafayette County, 2.0 mi. west of Dowling Park at quarry complex (30.2529°, -83.2578°) (NAD27), Upper Eocene OLS. LF010 (MILL CREEK QUARRY 03) Lafayette County, 2.0 mi. west of Dowling Park at quarry complex (30.2529°, -83.2578°) (NAD27), Lower Oligocene Suwannacoochee Dolostone. LF015 (DOWLING PARK 04) Lafayette County, 1.04 mi. west of Dwling Park at quarry just west of Suwannee River (30.244454°, -83.253597°) (WGS84), Upper Eocene upper OLS. LF023 (SUWANNEE RIVER 12) Lafayette County, 0.75 mi. west of Dowling Park on west bank of Suwannee River (30.245852°, -83.249011°) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ A1-8 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) (WGS84), Upper Eocene upper OLS. LV004 (GULF HAMMOCK QUARRY 01) Levy County, 1.0 mi. southeast of Gulf Hammock (29.237046°, -82.700545°) (NAD27), Middle Eocene Avon Park Formation. LV005 (DEVIL’S DEN) Levy County, 2.0 mi. north and 1.0 mi. west of Williston (29.407548°, -82.47642°) (WGS84), Upper Eocene OLS. LV014 (GULF HAMMOCK 02) Levy County, quarry 2.4 mi. south of Gulf Hammock (29.21619°, -82.703966°) (WGS84). Upper Eocene lower OLS. LV016 (INGLIS 02 [COTYPE LOCALITY INGLIS MEMBER]) Levy County, quarry 1.0 mi. west of Inglis north of SR40 (29.037226°, -82.681656°) (WGS84), Upper Eocene lower OLS. LV021 (INGLIS 03 [COTYPE LOCALITY INGLIS MEMBER]) Levy County, 0.125 mi. south of Inglis at Florida Power Corporation Plant, north bank of the Withlacoochee (29.0308°, -82.6817°) (NAD27), Upper Eocene lower OLS. LV024 (WITHLACOOCHEE BAY 01) Levy County, dredged from the mouth of the Withlacoochee River (29.00131°, -82.759994°) (WGS84), Upper Eocene lower OLS. LV035 (WITHLACOOCHEE BAY 02) Levy County, approximately 2.5 mi. southwest of Yankeetown, first and second spoil islands west of boat ramp at end of SR40 (29.001221°, -82.766074°) (WGS84), Upper Eocene lower OLS. LV039 (GULF HAMMOCK QUARRY 04) Levy County, 2.25 mi. southeast of Gulf Hammock, 1.16 mi. northeast of SR19 (29.238256°, -82.692481°) (NAD27), Middle Eocene Avon Park Formation. LV101 (WITHLACOOCHEE RIVER 09) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-9 Levy County, southeast of Crackertown, dredging along the Withlacoochee River between Inglis and Crackertown (29.030179°, -82.678837°) (WGS84), Upper Eocene lower OLS. LV104 (WITHLACOOCHEE RIVER 07) Levy County, east of Yankeetown, dredging on both sides of the Withlacoochee River between Inglis and Yankeetown (29.032912°, -82.694589°) (WGS84), Upper Eocene lower OLS. LV106 (WITHLACOOCHEE RIVER 11) Levy County, 1.46 mi. west of Inglis, dredging on north bank of the Withlacoochee River 1.0 mi. below Power Plant at Inglis (29.03273°, -82.692578°) (WGS84), Upper Eocene lower OLS. LV114 (ROYAL ROAD PIT 01) Levy County, 6.15 mi. south of Otter Creek, pit 0.57 mi. south of the intersection of Royal Road and Buck Island Road (29.235935°, -82.778488°) (WGS84), Upper Eocene lower OLS. MR018 (BRIAR CAVE) Marion County, 3.5 mi. southwest of Ocala, Upper Eocene OLS. OK005 (FRED GANNON ROCKY BAYOU STATE PARK 01) Okaloosa County, 0.96 mi. northwest of Seminole, well core in Fred Gannon Rocky Bayou State Park (30.4912247°, -86.4194612°) (WGS84), Upper Eocene OLS. PA002 (MORRELL PIT 06) Pasco County, limerock mine approximately 2.5 mi. northeast of Branchborough (28.286469°, -82.066235°) (WGS84), Lower Oligocene Suwannee Limestone. PO017 (TERRAMAR 01 [WEST COAST MINE]) Polk County, limerock mine approximately 6 mi. northwest of Socrum (28.232729°, -82.098447°) (NAD27), Lower Oligocene Suwannee Limestone. PO050 (PALMETTO PHOSPHATE COMPANY 02) Polk County, phosphate mine 2.63 mi. west northwest of Ft. Meade (27.761748°, -81.843375°) (WGS84), Middle Eocene Avon Park Formation. SM010 (CEMEX CENTER HILL QUARRY 01) ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ A1-10 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) Sumter County, 1.63 mi. northwest of Center Hill (28.6641°, -82.01397°) (WGS84), Upper Eocene OLS. SU002 (O’BRIEN QUARRY) Suwannee County, 1.0 mi. northwest of O’Brien at quarry off SR349 (30.047505°, -82.952708°) (WGS84), Upper Eocene upper OLS and Lower Oligocene Suwannee Limestone. SU003 (BRANFORD 01A) Suwannee County, Denali Quarry north of Branford (30.0041°, -82.9388°) (NAD27), Upper Eocene upper OLS. SU004 (WATERMELON PIT 01) Suwannee County, 10.0 mi. northeast of Branford along SR247, near Suwannee/Columbia County line (30.047505°, -82.952708°) (WGS84), Upper Eocene upper OLS. TA001 (CABBAGE GROVE 01) Taylor County, road metal pit approximately 3 mi. west of Cabbage Grove (30.208337°, -83.920824°) (WGS84), Lower Oligocene Suwannee Limestone. TA004 (TENNILLE 01 [LIMEROCK INDUSTRIES INC.]) Taylor County, southwest of Tennille (29.776211°, -83.330453°) (WGS84), Upper Eocene upper OLS. VO001 (BLUE SPRINGS 01) Volusia County, approximately 2.5 mi. west of Orange City (28.947472°, -81.339528°) (WGS84), Middle Eocene Avon Park Formation. WG002 (DUNCAN CHURCH 01) Washington County, 4.5 mi. north of Wausau (30.705215°, -85.590275°) (WGS84), Lower Oligocene Bridgeboro Limestone, Duncan Church beds. WG033 (DUNCAN CHURCH 02) Washington County, quarry 4.5 mi. north of Wausau (30.705215°, -85.590275°) (WGS84), Lower Oligocene Marianna Limestone. WG034 (EBRO BLUE SPRING 01) Washington County, 5.0 mi. north northeast of Ebro (30.5134°, -85.84652°) (WGS84), Lower Oligocene ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ OSBORN ET AL.: Paleogene echinoids of Florida A1-11 Suwannee Limestone. FM-IP LOCALITIES OUTSIDE OF FLORIDA 2822 Jamaica, St. Ann Parish, Browns Town at St. Hilda’s School, Upper Oligocene Moneague Formation. ZA023 (STOVALL QUARRY 01) Alabama, Covington County, approximately 7.5 mi. northeast of Florala northwest of SR54 (31.035°, -86.2315°) (NAD27), Lower Oligocene Florala Limestone. ZA026 (PERDUE HILL) Alabama, Monroe County, Perdue Hill, Claiborne Lime Plant Pit, Eocene. ZA145 (CONECUH RIVER 08) Alabama, Covington County, 5.51 mi. west southwest of Andalusia (31.279206°, -86.569924°) (WGS84), Upper Eocene Moodys Branch Formation. ZA147 (BROOKLYN QUARRY 01) Alabama, Conecuh County, 3.0 mi. south southeast of Brooklyn (31.248186°, -86.723239) (WGS84), Lower Oligocene Marianna Limestone. ZA181 (BROOKLYN 01) Alabama, Conecuh County, 1.68 mi. southeast of Brooklyn (31.246556°, -86.747938°) (WGS84), Lower Oligocene Marianna Limestone. ZA192 (TOMBIGBEE RIVER 02) Alabama, Washington County, 1.94 mi. northeast of St. Stephens, bluff on the west bank of the Tombigbee River northwest of Cowans Bluff (31.557915°, -88.028804°) (WGS84), Lower Oligocene Marianna Limestone. ZN039 (LANIER PIT 01) North Carolina, Pender County, 17.5 km northwest of Holly Ridge, Lanier Pit 1.1 km south southwest of intersection of SR50 and Lee Road, just west of Williams Road (34.6268°, -77.6763°) (NAD27), Middle Eocene Castle Hayne Formation. ✐ ✐ ✐ ✐ ✐ ✐ ✐ ✐ 12 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 61(1) ZN125 (TRENT RIVER 02) North Carolina, Jones County, 0.75 mi. east of Trenton, west bank of Trent River at boat ramp (35.062529°, -77.341431°) (WGS84), Middle Eocene Castle Hayne Formation. ZN126 (MARTIN MARIETTA RICHLAND QUARRY 01) North Carolina, Onslow County, northwest of Jacksonville, quarry northwest of intersection of Union Chapel Church Road and Duffy Field Road (34.841991°, -77.542205°) (WGS84), Middle Eocene Castle Hayne Formation. ZZ104 (SMITH COUNTY LIME PLANT 02) Mississippi, Smith County, 1.92 mi. southeast of Sylvarena, Smith County Lime Plant at the end of CR18-8 (31.988756°, -89.361082°) (WGS84), Lower Oligocene Marianna Limestone. =Introduction =Materials and Methods =The Eocene =The Oligocene =Systematic Paleontology =Classification of Species Treated =Conclusions =Acknowledgments =Literature Cited