BULLETIN . - . # 0f of the FLORIDA STATE MUSEUM Biological Sciences Volume 27 1981 Number 1 A REVISION OF THE FOSSIL ERETHIZONTIDAE OF NORTH AMERICA MICHAEL K. FRAZIER - UNIVERSITY OF FLORIDA GAINESVILLE Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain varying numbers of pages and are not necessarily completed in any one calendar year. OLIVER L. AUSTIN, JR., Editor RHODA J . BRYANT , Managing Editor Consultants for this issue: ROBERT WILSON CHARLES A. WOODS Communications concerning purchase or exchange of the publications and all manuscripts should be addressed to: Managing Editor, Bulletin; Florida State Museum; University of Florida; Gainesville, Florida 32611. Copyright © 1982 by the Florida State Museum of the University of Florida This public document was promulgated at an annual cost of $2028.00 or $2.028 per copy. It makes available to libraries, scholars, and all interested persons the results of researches in the natural sciences, emphasizing the circum-Caribbean region. Publication date: 5 January 1982 Price: $2.05 A REVISION OF THE FOSSIL ERETHIZONTIDAE OF NORTH AMERICA MICHAEL K. FRAZIERI SYNOPSIS: The North American fossil record of porcupines (Erethizontidae) is reviewed. Com- parisons of cranial osteology, incisor enamel, auditory ossicles, and post-cranial elements of fossil samples with Recent Coendou and Erethizon samples indicate that all of the fossil porcupines in North America are congeneric with Erethizon. The following four species of Erethizon are recognized in the fossil record of North America: (1) E. bathygnathum appeared approximately 2.5 million years ago and lived in western North America during late Blancan and early Irvingtonian times. (2) E. cascoensis is recognized only in the early Irvingtonian El Casco local fauna of California. (3) A new species of Erethizon is known only from the early-middle Irvingtonian of Florida. (4) The extant E. dorsatum first appears in the United States during middle Irvingtonian times, later dispersing throughout most of temperal North America. Erethizon apparently differentiated from an ancestral form in South America prior to its appearance in North America during late Pliocene times. Even the earliest fossils already possessed the enlarged masticatory apparatus designed to consume the bark of temperate conifers. Other functional specializations in the skeleton of Coendou and Erethizon are consid- ered. TABLE OF CONTENTS INTRODUCTION ...................................... . 3 ACKNOWLEDGEMENTS .............................. 5 MATERIALS AND METHODS .... ..... .... .. 6 NORTH AMERICAN DISTRIBUTION .......... ......... 9 CRANIAL OSTEOLOGY ....... ............ ....... 13 FOSSIL IDENTIFICATION BY CRANIAL MEASUREMENTS 25 INCISOR ENAMEL ........... ...................... . 31 AUDITORY 35 POSTCRANIAL OSTEOLOGY ................. ........... 40 SYSTEMATICS ............................................... 45 ORIGIN OF 53 APPENDIX A Pleistocene Faunas That Contain Porcupines ....... 56 APPENDIX B Graphs ofCranial Measurements ............... . 58 LITERATURE CITED .................... ......... ....... 77 iThe author is Associate Curator of Paleontology at the Mississippi Museum of Natural Science, 111 North Jefferson Street, Jackson 39202. This paper was submitted in partial fulfilment for the degree of Master of Science at the Univer·sity of Florida, Gainesville. FRAZIER, MICHAEL K. 1981. A revision of the fossil Erethizontidae of North America. Bull. Florida State Mus., Biol. Sci. 27(1):0-00. r 2 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 INTRODUCTION The oldest known erethizontids appear in the late Oligocene Deseaden sediments of Patagonia and Bolivia, along with early records of other cavio- morphs (Simpson 1950; Hoffstetter and Lavocat 1970). The origin of these early South American caviomorphs has long been a topic of controversy among rodent systematists. One school of thought favors an origin from an African phiomorph, via rafting across a narrower Atlantic Ocean during the Eocene (Hoffstetter and Lavocat 1970; Lavocat 1974; Raven and Axelrod 1975). Another school supports a North American ancestry from a primitive Eocene hystricognathous rodent, possibly a franimorph (Wood and Patter- son 1970; Wood 1974; Wahlert 1973). Presumably these ancestral popu- lations arrived in South America via island hopping across the Eocene Antillean Archipelago (Wood 1977). Wood (1975, 1977) has presented evi- dence favoring the parallel evolution of the African and South American hystricomorph rodents. Yet another school adds another dimension. Hus- sain et al. (1978) suggests the caviomorphs are derived from the Chapat- timyidae, a Southeast Asian late Eocene rodent group. Numerous fossil erethizontid genera have been described from Tertiary deposits in South America, all having the same basic tooth structure as their living representatives (Patterson 1958; Fields 1957). The erethizon- tids were restricted to the South American continent until their late Pliocene dispersal across Middle America (Simpson 1950). At that time they came north during the major faunal interchange between North and South America (Webb 1976). Fossil porcupines appear in North America during the late Pliocene, less than three million years ago. There is little doubt that they are of South American origin, where three of the four living genera still occur. This study concentrates only on the two North American genera, Coendou and Erethizon. Less is known about the other two genera, Chaetomys and Echinoprocta, which occur exclusively in South America (Anderson and Jones, 1967; Walker 1975). Chaetomys may even be an echymyid, based upon the nature of the r ((lA. Woods, pers. comm.). Erethizon lives exclusively in temperate and boreal North America, pre- ferring wooded areas of conifers, junipers, and poplars, where it feeds primarily upon the cambium layer of bark. It now ranges from northern Mexico northward into the major portion of Canada and Alaska (Woods 1974). Erethizon is the only caviomorph rodent now occupying temperate and boreal North America. Coendou inhabits lowland rain forests of the Neotropical Realm from southern Mexico (20° latitude) to northern Argen- tina, where it feeds upon leaves, tender stems, and fruit (Walker 1975). The two genera are geographically separated by a distance of about 1000 km. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 3 Coendou and Erethizon show gross morphological differences. Coendou has a quill-less, prehensile tail, a well developed prehallux in the hind foot, and a masticatory apparatus designed for the consumption of soft tropical vegetation. However some species of Coendou do not exhibit the pre- hensile tail (Karl Koopman, pers. comm.). Erethizon, on the other hand, has a short, stubby tail bearing quills, a vestigial prehallux, and a masti- eatory apparatus suited for the consumption of the much tougher tem- perate vegetation, particularly the bark of conifers and hardwoods. In a revision of the North American Erethizontidae, White (1970) presented morphological evidence for referring the fossil porcupines from Grand View (Coendou bath!/~nathum), Vallecito Creek (C. stirtoni), El Casco (C. cascoensis), Cumberland Cave (C. cumberlandicus), and other localities to the genus Coendou. He considered Coendou ancestral to Erethizon, with the middle Irvingtonian Cumberland Cave sample from Maryland being intermediate between the two genera. Discovery of an excellent sample of Erethizon-like porcupines from the early Irvingtonian Inglis IA locality in Florida prompted this further review of the fossil porcupines of North America. The principal goals of this study are (1) to decide whether the fossil porcupines of North America represent Coendou, Erethizon, or both genera, (2) to trace their stratigraphic and geographic distribution through time, and (3) to outline any evolutionary trends that may be observed in these North American erethizontids. ACKNOWLEDGEMENTS Specimens were made available for study by the following institutions: Academy of Natural Sciences at Philadelphia (ANSP), American Museum of Natural History (AMNH or F:AM), University of Arizona (UALP), Carnegie Museum of Natural History (CM), Florida State Museum (UF), Imperial Valley College Museum (IVCM), University of Kansas Museum of Natural History (KU), Los Angeles County Museum of Natural History (LACM), University of Michigan Museum of Paleontology (UMMP), National Museum of Natural History, Smith- sonian Institution (NMNH), University of Oregon Museum of Natural History (UO), and Texas Memorial Museum, University ofTexas (TMM). I am grateful to the people responsible for the loan of these materials. I express deep appreciation to S. David Webb, my supervisory chairman, for the pre- liminary review of this thesis and his helpful comments and guidance through the course of this work. I also wish to thank the other members of my supervisory committee, Pierce Brodkorb, David Nicol, and Ronald G. Wolff, for helpful criticism of the manuscript and their assistance during the last three years. Valuable fossil specimens were donated by Allen Davis, Frank Garcia, Margaret C. Tho- mas, and John S. Waldrop (Timberlane Research Organization). Mrs. Steven Kruger of.Seff- ner, Florida, loaned several porcupine mandibles from Apollo Beach and also allowed them to be cast Raymond Ciron (Sant Fe Community College Zoo) provided a Coendou for wet study. Jerry Duvendeck (Michigan Department of Natural Resources) and David Steadman provided several specimens of Erethizon. Tim Breen and Robert Smidt provided valuable assistance in the statistical analysis of the cranial measurements. John A. White and Charles 4 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 A. Woods helped with their discussions concerning the evolution and morphology of erethi- zontids. Charles A. Repenning added valuable suggestions concerning Pleistocene strati- graphy and zoogeography. Discussions with friends and fellow graduate students Jon Baskin, David Frailey, Richard Franz, Andrew Homner, Jean Klein, John Meeder, Gary Morgan, David Steadman, and John Waldrop added much to my thought processes. Additional thanks go to Dr. Wolff and Robert Allen for their photographic assistance and to Chandra Aulsbrook and Nancy Halliday for help with the figures. Special thanks go to Mr. and Mrs. Henry Danker for the donation of the scanning electron microscope to the Department of Geology. Robert Pierce provided assistance with the use of the microscope. Warm and special gratitude goes to all the people who were not mentioned, but were helpful in many ways during the course of this study. MATERIALS AND METHODS Cranial measurements of numerous Recent skulls and mandibles of Coendou and Erethizon as well as all available fossil materials were taken in millimeters using a dial caliper to the nearest tenth. On each Recent specimen 46 measurements were taken, of which the most diagnostic are presented in Figures 1 and 2. On Recent specimens where a measurement could not be taken, the sample mean of that measurement was substituted. The Recent samples (over 100 skulls per genus) included males and females, adults and subadults, and sampled the entire geographic range of each genus. Individuals with deciduous premolars were considered subadults. The skulls and skeletons used were loaned from the mammalogy collections of the American Museum of Natural History, National Museum of Natural History (Smithsonian Institution), and the Florida State Museum. A principal component analysis, analysis of variance, cluster analysis, and discriminant analysis were performed upon the cranial measurements using the Statistical Analysis System (SAS76) (Barr et al. 1976). SAS 76 also produced the graphs of the cranial measurements of Recent specimens (Appendix B). Sagittal sections of lower incisor enamel of Coendou and Erethizon and fossil specimens were studied using an ISI Super Mini-SEM II scanning elctron microscope. The preparation of the specimens followed the procedures described by Flynn and Wahlert (1978), Measure- ments taken from the enamel photomicrographs follow Escala and Gdllego (1977) and Wahlert (1968). The preparations of Recent and fossil auditory bullae were accomplished using an S. S. White Airbrasive jet machining unit. NORTH AMERICAN DISTRIBUTION The earliest porcupines in North America appear in late Pliocene (Blan- can) and early Pleistocene (Irvingtonian) faunas from Grand View, Idaho, Vallecito Creek and El Casco in southern California, and Wolf Ranch in the San Pedro Valley of Arizona. These records all occur in the western United States. Later records are known from other localities throughout North America. Stratigraphic correlation of these early records has been refned and simplified by radiometric dating and magnetic polarity stratigraphy in the Anza Borrego section of California (Opdyke et al. 1977), the San Pedro Valley of Arizona (Johnson et al. 1975; Lindsay et al. 1975), the Pearlette type ashes of the Midwest (Boellstorff 1973; Zakrzewski 1975a), and the Grand View section of Idaho (Neville et al. 1979). Other correlations in this 1981 FR A ZIE R : N .A . FO S S IL E R E T H IZ O N T ID R E V IS IO N 3 2 4 9 A B C FIGuRE 1. Key to cranial measurements. (1) width of rostrum, (2) width of frontals, (3) width of palate between P4 alveoli, (4) width of palate between M3 alveoli, (5) width at zygomatics, (6) width across auditory meatuses, (7) width across occipital condyles, (8) medial width of u rostrum, and (9) width at infraorbital foramen. 6 BULLETIN FLORIDA STATE MUSEUM Vul. 27 No. 1 11 ® 1 0 , A 17 16 B 19 20 21 C FIGURE 2. Key to cranial measurements. (10) depth of rostrum, (11) length of auditory bulla, (12) length of rostrum, (13) alveolar length of upper tooth row, (14) length from upper M3 alveolus to occipital condyle, (15) total skull length, (16) depth 6f mandible at lower P4, (17) depth of mandible at lower ME, (18) depth from condlyle to angular process, (19) alveolar length of lower tooth row, (20) length of diastema, and (21) length of symphysis. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 7 paper rely on biostratigraphic comparisons of diverse mammalian taxa (Hibbard et al. 1965; Skinner and Hibbard 1972). Harrison (1978) considered the Wolf Ranch local fauna to be late Blancan in age, based upon the fossil mammals. Lindsay et al. (1975) established a radiometric date for the Wolf Ranch sediments of approximately 2.5 my, and their paleomagnetic studies establish the time of deposition at just below the Gauss-Matuyama magnetic polarity boundary. These dates establish the Wolf Ranch record as the oldest known porcupine in North America. The Grand View local fauna, as described by Shotwell (1970) and Wilson (1933), is considered late Blancan in age by most paleontologists. A radio- metric date of 1.36 my from the Bruneau Basalt, which is stratigraphically above the Grand View deposits, provides a minimum age, and a date of 3.5 my for the Hagerman local fauna (Glenns Ferry Formation) below the Grand View provides a maximum age for the fauna (Evernden et al. 1964). Neville et al. (1979), using magnetic polarity stratigraphy, assign the Grand View deposit to the earliest part of the Matuyama polarity epoch, between 1.8 and 2.5 my old. The above information and the occurrence of Boropha- gus, Trigonictis, and Ondatra idahoensis support a late Blancan age for the fauna. Thus, the holotype of Erethizon bathygnathum Wilson (1935) is late Blancan. The Anza Borrego faunas of southern California were originally studied by Downs and White (1968), and recently the magnetic polarity stratig- raphy was correlated with the mammal-bearing section (Opdyke et al. 1977). The stratigraphic occurrence of porcupine in the section coincides with the appearance of Nothrotherium, Lepus, Microtus, Smilodon, and ?Euceratherium, which are considered early Irvingtonian indicators and occur in the middle of the Matuyama polarity epoch in the Anza Borrego section. The age of this record is approximately 1.9 or 1.6 my (Opdyke et al. 1977). The El Casco record occurs in a fauna collected in 1923 by Joseph Rak and Childs Frick (White 1970; Henry Galiano, pers. comm.). As no stratig- raphic data exist, correlation must rely on biostratigraphic methods. Canis edu>ardi, Ondatra idahoensis (advanced form), Odocoileus, and Equus (Equus) also occur in the El Casco local fauna. Nelson and Semken (1970) have shown conclusively that a chronocline exists in Ondatra during the Plio-Pleistocene, and its species are especially useful biostratigraphic indicators. Ondatra idahoensis appears in the San Pedro River Valley of Arizona approximately two million years ago (Lindsay et al. 1975). It is known from the Borchers local fauna of Kansas, dated at 1.9 my (Zakrzewski 1975a), the Grand View local fauna (Wilson 1933; Shot- well 1970), the White Rock local fauna (Eshleman 1975), the Mullen assemblage of Nebraska (Martin 1972), and the California Wash and Gidley Level local faunas from the San Pedro Valley (Johnson et al. 1975). All of 8 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 these records are considered late Blancan or early Irvingtonian. The other taxa in the El Casco local fauna also indicate an early Pleis- tocene age. Opdyke et al. (1977), in their study of the Anza Borrego sec- tion, described the appearance of Odocoileus during the Matuyama polar- ity epoch (approximately 2.1 my). Beryl Taylor (pers. comm.) considefs the wolf, Canis edwardi, to be an Irvingtonian species. Based upon the dental development of the lower Ml in Ondatra idahoensis, the presence of the post-Blancan Equus (Equus) sp. (Charles Repenning, pers. comm.), and the occurrence of other early Pleistocene taxa, the El Casco local fauna is considered early Irvingtonian in age (Frazier, MS). The earliest procupines in the eastern United States occur in the Inglis IA local fauna of Florida (Klein 1971; Webb 1974, 1976). This faunal assemblage includes Smilodon gracilis, Chasmaporthetes, Glyptotherium arizonae, Sigmo(ion curtisi, Canis edwardi, Platygonus bicalcaratus, Cap- romeryx arizonensis, and an advanced Ondatra c£ idahoensis. These mammalian taxa indicate an early Irvingtonian age. The Inglis IA local fauna compares very closely with the early Irvingtonian Curtis Ranch fauna as described by Gazin (1942). For example, G. arizonae, C. edu>ardi, Felis lacustris, S. curtisi, Ondatra, Lepus, and C. arizonensis are shared by the two faunas. Based upon these faunal similarities the Inglis IA local fauna can be considered equivalent in age with the Curtis Ranch fauna, which Johnson et al. (1975) assign to the Matuyama polarity epoch, approximately 1.9 my (near the Olduvai event). The correlations of the major North American late Pliocene and early Pleistocene faunas appear in Figure 3. Erethizontid occurrences are indi- cated by asterisks. As chronostratigraphic and paleomagnetic data are missing, the majority of the middle Irvingtonian and Rancholabrean faunas that contain porcu- pines must be correlated biostratigraphically. The Cumberland Cave, Port Kennedy, and Trout Cave local faunas contain assemblages of middle Irvingtonian arvicoline rodents (Hibbard 1955; Zakrzewski 1975b). The majority of the Rancholabrean porcupine records were collected from cave deposits throughout North America. Figure 4 presents the Recent ranges of Coendou and Erethizon in North America along with selected late Pliocene through late Pleistocene localities that have produced porcupine fossils. A complete list of localities in North America is presented in Appendix A. CRANIAL OSTEOLOGY Striking differences exist between the skulls of Erethizon and Coendou. Most of them are presumably related to the profoundly different modes of mastication in the two genera. Various cranial characters involved with the 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 9 AGE|~·S~ RADIOMETRIC - MAGNETOSTRATIGRAPHIC Biostratigraphic ~& MAMMALIAN FAUNAL 1 §;41 M ~7» Arizona California Idaho Texas Kansas ; Correlation @< DATUM PLANES PL IO C EN E ~L E IS TO C E N E | 0.69 Cudahy 06 i Irvington Bruneau 0 basalt ~ ·- Mammuthus '- Hypo/agus(extinction)1 G AU SS M A T U Y A M A Equus (Equus)Vallecito !.H Curtk; ;*Creek %*511 ,- Borophagus (ex,Encta) - 1.8, - -Ba-ncb 86rchers ,- Smifodon GrandView * Mt. Blanco Cita 2 43 WO" Canyon . Ondatra, Odocoileus * Ranch Sanders ·- Lepus ~ ·- /Vannippus (extinction) W = 5 CO 2.12 ·- Equus (Dolichohippus)2.90 Bender 3.00 Benson ReAroad ].0; Fox Canyon 3.32 Hagerman ·- Geomys3.4 r--1 reversed normal 1 1 polarity polarity * erethizontid FIGURE 3. Correlation Chart of late Pliocene and early Pleistocene local faunas. Mammalian faunal datum planes are geographically localized. LEarly Pleistocene Wellsch Valley local fauna, Saskatchewan, Canada (Harington 1978). (Chart modified from Johnson et al. 1975; Lindsay et al. 1975; Zakrzewski 1975a; Opdyke et al. 1977; Neville et al. 1979.) masticatory apparatus have been proposed previously for generic deter- mination of fossil porcupines in North America. White (1968, 1970) pre- sented the following distinctions: 1) Upper P4 in Erethizon is generally larger than upper Ml, while in Coendou the two teeth tend to be subequal in size. 2) The tooth rows are widely divergent in Erethizon, whereas they are subparallel in Coendou. 3) "The incisors of Coendou are essentially orthodont, and the upper and lower incisors occlude in such a manner as to provide an efficient cutting mechanism. This is contrasted by the occlusion of the more proodont incisors of Erethizon, which have less of a cutting and more ofa scraping function." (White 1970:12) 4) "In living Coendou the orbital width, or the greatest distance from the supraorbital ridge to the zygomatic arch, is less than 75 percent of the least interorbital constriction, whereas in Erethizon it is greater than 80 percent." (White 1970:12) 5) "The angular process of the mandible in adults of Erethizon is in- flected mediad and the posterolateral surface of the mandible is con- 10 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 T-f 7 1 62 1 -194'-f < - 7 12, 10 19 ./ 4 21 16 13 1000 KM -r.- 1~- Ill I FIGURE 4. Erthizontid distribution in North America showing Recent ranges of Erethizon (stippled) and Coendou (cross-hatched) (after Hall and Kelson 1959). Selected fossil porcupine localities below correspond with numbers on map: (1) Vallecito Creek, (2) El Casco, (3) Grand Vies, (4) Cumberland Cave, (5) Port Kennedy, (6) Conard Fissure, (7) Hartman's Cave, (8) Merritt Island, (9) Inglis IA, (10) Coleman IIA, (11) Apollo Beach, (12) Haile XVI, (13) Cedazo, (14) Cherokee Cave, (15) Durham's Cave, (16) San Josecito Cave, (17) Clamp Cave, (18) Waccasassa River, (19) Wolf Ranch, (20) Trout Cave, (21) Port Charlotte. vex, while in Coendou and in juvenile Erethizon the angular pro- cesses are subparallel and the posterolateral surface of the mandible is flattened." (White 1968:9) 6) "In adults of Erethizon the projection of a line superimposed upon the longitudinal axis of the lower tooth row and bisecting the occlusal surfaces of PI and MS, passes mediad to the incisor, while in Coen- dou it extends to the posteromedial surface or even laterad to the 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 11 incisor. This measurement makes it possible to determine if the man- dibular cheek teeth converge as in adults of Erethizon, or are sub- parallel as in Coendou and juvenile Erethizon." (White 1968:9) 7) "The fossa for the insertion of M. masseter medialis pars posterior on the side of the mandible is deeper in Coendou than in adults of Erethizon. This structure seems related to the degree of convexity seen in the posterior part of the mandible." (White 1968:9) 8) "The ascending ramus of the mandible in Coen(lou slants posteriad to a greater degree than in adults of Erethizon." (White 1968:9) 9) "The scratches on the enamel of the occlusal surfaces in both the upper and lower cheek teeth are oriented anteromediad and form an angle greater than 35 degrees with the longitudinal axis of the tooth rows in Coendou." (Landry, 1957) "In Erethizon this angle is less than 30 degrees." (White 1968:9) Martin (1974) added two other criteria to distinguish the genera: 1) The posterior border of palate at midline located opposite center or posterior border of upper M3 in Coendou, while in Erethizon the posterior border of palate is located opposite center or posterior bor- der of upper M2. 2) Upper cheek tooth row in Coendou is less than one-fourth of skull length (microdont), while in Erethizon the upper cheek tooth row is greater than one-fourth the skulllength (macrodont). A thorough comparison of the cranial osteology of Recent Coendou and Erethizon was deemed necessary to test the validity and variability of the characters presented above and to search for other differences. A large sample of skulls and mandibles of Coendou and Erethizon was measured according to the previously described procedure (N greater than 100 for each genus). The major cranial measurements of adults are presented in Table 1, along with the corresponding statistical parameters. In almost every dimension, living Coendou is much smaller than living Erethizon. The largest differences in the cranial dimensions measured in the two living genera occur in the following variables: Xl, X20 Alveolar length of upper and lower tooth rows X2, X21 Width of upper and lower P4 X3, X22 Width of upper and lower M 1 X5 Width of palate between upper M3 alvegli X6, X23 Width of upper and lower incisor X7, X24 Anteroposterior length of upper and lower incisor X8 Total skulllength X9 Length of rostrum X27 Length of diastema X10 Length of upper M3 to occipital condyle TABLE 1.-Statistical values of major cranial measurements in adult Coendou and Erethizon. Coendou Erethizon 12 B U LLE T IN F LO R ID A S TATE M U S E U M V 27 N 1 Variable' N X SD MIN MAX N X SD MIN MAX X1 117 18.0 2.138 13.5 21.8 155 26.6 1.346 23.5 30.1 X2 117 5.3 .615 4.1 6.5 154 8.7 .586 6.9 10.1 X3 116 4.8 .531 3.8 5.9 152 7.1 .450 6.0 9.3 X4 118 3.8 .916 2.0 6.3 155 3.8 .960 1.5 7.9 X5 117 6.7 1.146 3.8 9.2 155 11.0 1.153 7.2 14.4 X6 116 2.7 .351 2.0 3.8 154 4.3 .366 3.5 5.3 X7 116 3.9 .483 3.0 5.3 154 4.9 .337 4.2 6.0 X8 115 81.3 9.345 61.0 100.4 155 108.8 6.937 92.0 125.7 X9 118 26.6 4.178 17.0 36.0 155 36.9 3.434 28.7 45.3 X1O 114 35.4 4.026 26.8 47.3 153 44.9 3.036 37.7 51.1 X11 113 19.5 2.439 15.0 28.6 152 24.9 2.068 20.6 30.4 X12 116 33.6 3.096 27.3 40.7 155 45.6 2.213 41.0 51.2 X13 114 46.7 4.882 36.2 56.9 150 71.2 3.333 62.8 80.2 X14 114 37.1 5.322 27.0 48.9 148 50.4 2.996 44.4 68.2 X15 115 18.8 4.053 13.0 28.7 151 23.0 1.687 18.9 28.3 X16 118 14.7 2.378 11.0 21.0 154 18.4 1.291 14.5 217 X17 116 28.3 5.735 18.2 40.9 155 31.3 3.245 23.8 43.0 X18 116 22.0 3.700 14.2 33.7 151 29.7 2.221 24.8 34.9 X19 225 22.6 2.330 17.2 28.4 279 25.1 2.240 20.1 31.7 X20 114 19.4 2.259 15.0 23.7 154 29.1 1.583 25.3 33.1 X21 112 4.5 .599 3.5 6.0 151 7.5 .540 6.0 9.0 X22 113 4.3 .494 3.2 5.6 151 6.4 .397 5.3 7.9 X23 117 2.8 .362 2.0 3.9 153 4.5 .333 3.7 5.5 X24 117 3.7 .486 2.6 4.8 153 4.8 .312 4.2 5.7 X25 117 16.7 2.146 11.5 22.0 154 25.1 1.809 20.1 30.2 X26 117 10.5 1.452 8.0 14.1 154 16.0 1.228 12.2 18.9 X27 117 14.2 2.678 8.7 19.8 154 22.8 2.346 17.3 27.9 X28 117 22.6 3.725 13.9 30.7 154 36.5 2.750 29.6 44.4 X29 114 24.0 3.111 17.2 30.8 150 36.6 2.728 30.2 45.1 *Cranial measurements corresponding to variables are: SKULL- X1 = alveolar length of upper tooth row. X2 = width of upper P#. X3= width of upper M 1, X4 - width of palate.between P4 alveoli. X5 = width of palate between M 1 aiveoli, X6 = width of upper incisor, X7 = Interior-posterior length of upper incisor, X8 =total skull length, X9 = rostrum length, X 10 =len4th from M3 alveolus to occipital condyle, Xlli= width across occipital condyles. X12=width across auditory meatuses, X13 = width at zygomatics. X14 = width at infraorbital.foramen, 115 - width o! rostrum. X 16 = medial widih of mtrum, X17= width at frontals, X 18 = depth of rostrum. X 19 = length of auditory bulla. MANDIBLE- X20= alveolar length of lower• tooth r6w. X21 = width of lower P4. X22 = width of lower Ml, X23 = width of lower' incisor, X24 - anterior-posterior length of lower incisor, X25 = depth of mandible at P4, X26= depth of mandible at MZ, X27 = length of diastema. X28=length of symphysis, X29 = depth from condyle to angular process. 1981 FRAZIER: N.A. F()SSIL ERETHIZONTID REVISION 1.3 A C 3 - ~cm t D. -a L 4,/Al .li FIGURE 5. Comparison of fossil and Recent porcupine skulls in lateral view. (A) Recent Coendou, AMNH 190423; (B) Recent Erethizon dorsatum, NMNH 245525; (C) fossil E. dorsatum, Cumberland Cave, Maryland, NMNH 7996 (holotype, C. cumberlandicus White 1970), (D) fossil E. cascoensis, El Casco, California, F:AM 17883-1 (holotype, C. cascoensis White 1970). 14 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 0-5- n A B cm dp. r . I 4 h , 4 , D C FIGURE 6. Comparison of fossil and Recent porcupine skulls in dorsal view. A-D same as figure 5. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 15 Icm ~cm f 1, A B f 1:2 j*41 iIcm ~ t-~1 7 f i.... dja I I / L...i D C FIGURE 7. Comparison of fossil and Recent porcupine skulls in ventral view. A-D same as figure 5. 16 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 X12 Width of skull at auditory meatus X13 Width of skull at zygomatics X14 Width of skull at infraorbital foramen X25 Depth of mandible at lower P4 X26 Depth of mandible at lower M2 X28 Length of symphysis X29 Depth of mandible from condyle to angular process In order to explore the range of variation in the overall cranial dimen- sions of the two genera, a cluster analysis was performed. The computer reviewed all of the data of Recent specimens and reclassified them into two groups based upon all cranial variables. One group contained all Erethizon and some large Coendou; the other contained the rest of the Coendou sample and no Erethizon. This indicates a range of overlap in the cranial measurements of large individuals of Coendou with those of adults and subadults of Erethizon. The measurements in which there is no overlap in adults are: width of upper P4, width of upper Ml, width of skull at audi- tory meatus, width of skull at zygomatics, and alveolar length of upper and lower tooth rows. Figures in Appendix B illustrate the relationships of selected cranial measurements of subadults and adults of both genera. These morphometric differences are evidently related directly to the dif- ferent masticatory adaptations of the two genera. Other discriminatory morphological characters in the crania are also apparent (Figs. 5-10). The anterior end of the nasals is pointed in Coendou but indented in Erethizon. This feature may relate to the fact that in Coendou the soft part of the nose is much more massive than in Erethizon. In every Recent skull examined, the above character clearly distinguishes these two genera. Another character of the skull is the great inflation of the frontals in some Coendou, as noted by Ellerman (1940). This inflation may be a nasal de- velopment for more effective olfaction. In diagnosing Coendou cumberlandicus from the Cumberland Cave in Maryland, White (1970:7) stated: "Nasofrontal suture slightly curved pos- teriad," as in Coendou, "not V-shaped" as in Erethizon. This character has been shown to be quite variable in Erethizon (Anderson and Rand 1943). Indeed Ahlberg (1969) used this feature to distinguish the eastern sub- species of Erethizon, with a nearly straight transverse nasofrontal suture, from the western subspecies with a V-shaped suture. The degree of development of the sagittal crest, reflecting the size of the temporal muscle that attaches to the parietals, is relatively greater in Erethizon than Coendou. As the temporal muscles develop with age, juve- nile Erethizon do not have such a well developed sagittal crest. The rela- tive degree of divergence of the zygomatics in Erethizon is much greater than in Coendou and the dorsoventral thickness of the zygomatic is less in 1981 FRAZIER: N.A F()SSIL ERETHIZONTID REVISION 17 - - A B C D cm 1* I-®: rA, .liall.UF BM 49.4. E F lie, G H FIGURE 8. Comparison of fossil and Recent porcupine mandibles in lateral view. (A) Recent Erethizon dorsatum, N MNH 245525; (B) fossil E. dorsatum, Haile XVI, Florida, UF 21490; (C) Recent Coendou, AMNH 190423; (D) fossil E. donatum, Aguascalientes, Mexico, UMMP V47106; (E) Erethizon kleini, new species, Inglis IA, Florida, UF 21474; (F) fossil E. bathy- gnathum, Grand View, Idaho (holotype), NMNH 13684, (G) E. kleini. new species, Inglis IA, Florida (holoype), UF 21473, (H) E. bathygnathum, Vallecito Creek, California, IVCM 309. 18 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 Coendou than in £rethizon. However, some large Coendou have relatively thickened zygomatics, so that some overlap occurs in this feature. The zygomatic processes of the squamosals extend ventrolaterally in Coendou, while in Erethizon they extend laterally. All of these characters are a result of much greater masseter muscle development in Erethizon. The shape of the auditory bullae in Coendou is oval and elongate, while in Erethizon the bullae are more rounded. Internal ear structures are compared in a later section. White (1970) showed that Erethizon skulls have a greater interorbital width than those of Coendou, and suggested that this produces different visual fields. The tangent to the line of optic projection in arboreal Coen- dou makes an angle of about 65° with the horizontal plane; whereas in the more terrestrial Erethizon, the angle is approximately 45°, a more dorsal view than in Coendou. The posterior border of the palate at the skull midline was found to be quite variable in both genera and cannot be used as a distinguishing char- acteristic. Martin (1974) developed this criterion after comparing only six skulls of Erethizon and three of Coendou. Martin's use of the ratio of the upper tooth row length to the total skull length is also too variable to be statistically reliable according to the larger samples studied here. The char- acter holds for Coendou but tends to break down in the smaller Erethizon. Thus the two characters he developed have proved to be'quite variable. As noted by White (1970), the incisors of Erethizon are more extended anteriorly (proodont) than the essentially orthodont incisors of Coendou. The lower incisors of Coendou occlude with the upper incisors at a more oblique angle than Erethizon. The present study strongly supports this distinction of White, and I agree with his suggestion that the proodont incisors of Erethizon are better adapted for eating bark than the orthodont incisors of Coendou. The mandibuldr characters discussed by White (1968, 1970) are generally distinctive but are somewhat variable. The most distinctive character is the inflection of the angular process. As stated by White (1968), the angular process is inflected sharply mediad and flattened on the ventral surface in Erethizon, whereas the angle in Coendou exhibits a smaller degree of inflection and has little or no flattening of the angular process. White (1968, 1970) used the angle of divergence of the lower tooth row from the midline to distinguish Coendou and Erethizon. This angle is useful in separating the Recent genera in most cases but cannot be used with total confidence on a small fossil sample. Erethizon exhibited a lateral projection of the tooth row in relation to the lower incisor in only 2 out of 125 observations; Coendou had a medial projection in 6 of 53 cases. Among other characters that have proven to be variable are the depth of the fossa for the insertion of the M. masseter medialis pars posterior on the mandi- 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 19 1.":p.- A B m,3.K#. lig 4, f C D -cm E F I =Nill G H FIGURE 9. Comparison of fossil and Recent porcupine mandibles in medial view. A through H same as Figure 8. 20 BULLETIN FL()1111)A STATE MUSEUM VoL 27 No. 1 A B -...."h,flarziatilwl -*5¥di/'-:/9- D 2iR F G H FIGURE 10. Comparison of fossil and Recent porcupine mandibles in dorsal view, A through H same as Figure 8. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 21 ble and the orientation of the ascending ramus. The scratches on the enamel of the occlusal surface in both upper and lower cheek teeth have also been used to distinguish Coendou from Erethi- zon. Landry (1957:15-16) stated "In Coendou, ..., the scratches on the teeth indicate that the angle at which the lower teeth move across the upper is about fifty degrees from transverse. White (1968:9) described this angle as "greater than 35 degrees with the longitudinal axis of the tooth " rows in Coendou. In Erethizon this angle is less than 30 degrees. Generic distinction based solely upon the orientation of tooth scars in a small sam- ple of isolated teeth should be made with caution. Along with the other morphological characters, the orientation of the tooth scratches on the cheek teeth provides useful evidence of masticatory differences between the genera. Woods and Howland (1979) describes two types of mastication in hystri- comorphous rodents. Orthognathous mastication is propalinal (anterior- posterior) and bilateral, while plagiognathous is more oblique and unilater- al. Woods (pers. comm.) considers Coendou to be more orthognathous than the plagiognathdus £rethizon. The major osteological characters of Erethizon associated with plagiognathous mastication are the posterior di- vergence of the cheek tooth rows from the midline, the increased width of the skull and zygomatic development and the deepening of the mandible. The increase in size of the upper P4 in relation to M 1 and the enlarged upper and lower incisors are all adaptations of Erethizon for consuming coarse vegetation. FOSSIL IDENTIFICATION BY CRANIAL MEASUREMENTS Using the information supplied by statistical analysis of living Coendou and Erethizon cranial dimensions, a discriminant analysis was performed upon the cranial measurements of the fossil specimens from North Amer- ica. As some fossils are fragmentary, the analysis assigned generic prob- ability values for each specimen based only upon the available measure- ments. These values are presented in Tables 2 and 3, together with the raw measurements of the classified fossils. The fossils from Grand View, Vallecito Creek, and El Casco were classi- Bed as Erethizon wit}i a very high probability (greater than or equal to .99). The palates of the Erethizon-like porcupine from Inglis IA and Merritt Island in Florida were classified into both genera. In most cases the Inglis IA and Merritt Island fossils tend to be intermediate in size between Coendou and Erethizon. Based upon the mandibular measurements, the Inglis IA porcupines were considered Erethizon with a probability of great- er than or equal to . 89. The Cumberland Cave sample was classified as Erethizon with a probability of greater than or equal to . 98. Figures in Appendix B graphically present the measurements of most of the fossil porcupines studied, together with the values of adults and subadults of Coendou and £rethizon. 22 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 TABLE 2. -Fossil cranial measurements and discriminant analysis classifcation (d = deciduous). 4% I,ocality 3: a}Eled 4 P IM N ~e Catalogue # 4:2 GRAND VIEW UO F-16282 UO F-16284 UO F-16279 VALLECITO CREEK LACM 17633 28.1 8.9 8.4 6.7 LACM 6136 26.4 6.7 EL,CASCO F:AM 17883-1 25.0 8.7 --- 2.4* F:AM 17882 26.2 7.5 6.7 4.0* F:AM 17883-7 F:AM 18128 INGLIS IA UF 21468 ---- 6.8 6.0 3.2 UF 21469 21.2 6.7 6.2 4.7* UF 21470 23.7 6.6 5.7 UF 21471 21.1 5.2 5.8 MERRITT ISLAND UF 21465 7.5 6.1 3.7 UF 21466 6.4 5.5 CUMBERLAND CAVE NMNH 7996 26.2 8.5 7.6 6.7 NMNH 7670 25.0 7.0 6.1 4.5 NMNH 25692 26.3 8.3 6.8 4.9 PORT KENNEDY ANSP 122 7.3 6.8 COLEMAN IIA UF 11774 24.7 6.2 6.3 3.4 CONARD FISSURE AMNH 12422 29.1 9.4 8.1 5.0 HARTMAN'S CAVE ANSP 658 23.5 7.7 6.5 2.7 •Specimens either reconstructed or damaged. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 23 W id th P al at e * Classification byE E Discrim. Analysis 16 A i Coendou Erethizon 4.9 5.8 0.000 1.000 5.9 6.3 0.000 1.000 4.9 5.5 0.000 1.000 10.8 -- - 0.001 0.999 11.2 4.8 5.8 0.000 1.000 6.7* ----- 6.7* --- 0.008 0.992 5.5 6.7 0.000 1.000 4.6 5.7 0.000 1.000 9.2 0.200 0.800 6.2* 1.000 0.000 0.606 0.394 0.962 0.038d 9.6 0.038 0.962 0.998 0.002 11.2 5.2 5.6 0.000 1.000 10.0 0.000 1.000 11.6 0.000 1.000 0.000 1.000 9.0 4.0 5.1 0.017 0.983d 10.3 0.000 1.000 8.8 0.003 0.997 24 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 TABLE 3.- Fossil mandibular measurements and discriminant analysis classification (d = deciduous). -5 - Locality 2: 3 2 -1 I R A O '-1 M Catalogue # 5 H GRAND VIEW NMNH 13684 32.9 8.0 6.9 UO F-16271 33.2 7.3 7.0 -- UO F-16272 6.3 7.6 5.0 5.2 UO F-16275 5.5 6.5 VALLECITO CREEK LACM 6136 ---- LACM 61420 31.0 LACM 6210 --- IVCM 309 7.1 EL CASCO F:AM 17883-2 5.9 --- F:AM 17883 --- 5.4 5.5 F:AM 178834 6.8 --- F:AM 17883-5 4.6 5.5 F:AM 17883-6 4.9 5.1 INGLIS LA UF 21473 25.7 5.9 5.5 4.4 4.8 UF 21474 24.7 5.5 5.2 UF 21475 5.3 --- UF 21480 4.3 5.3 CUMBERLAND CAVE NMNH 8128 28.2 7.1 --- 5.4 6.0 NMNH 25689 27.8 6.3 5.9 NMNH 7672 29.7 7.0 6.7 AGUASCALIENTES UMMP V-47106 31.0 7.0 6.8 4.8 5.5 HAILE XVI UF 21490 29.7 7.1 6.7 5.5 5.7 CHEROKEE CAVE AMNH 45742 29.2 7.5 6.1 5.5 5.1 APOLLO BEACH UF 24112 30.4 8.1 6.6 4.7 4.7 UF 24113 ---- 6.5 5.9 --- --- UF 24114 27.6 6.3 5.9 4.5 4.6 HARTMAN'S CAVE ANSP 648 29.8 7.0 6.0 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 25 M an di bl e D w Classification by ~ f Discrim. Analysis m 46 % 901 C es EQ z 54 A .4 Coendou Erethizon- 30.0 19.5 --- --- 0 000 1.000 27.2 17.2 17.6 37.5 0.000 1.000 14.0 13.8 -----d 0.000 1.000 17.7 15.9 17.2 16.5 14.7 ----- 0.000 1.000 --- 0.001 0.999 0.000 1.000 22.9 14.4 18.3 31.4 0.026 0.974 20.4 14.2 13.6 28.0 0.115 0.885 15.1 ----- ----- 0.004 0.996 23.9 14.9 20.0 32.2 ----- ----- 25.3 15.9 20.3 34.4 0.000 1.000 24.2 16.0 16.6 33.3 24.1 15.4 17.7 33.2 0.000 1.000 28.4 18.9 26.7 42.1 0.000 1.000 26.1 16.0 17.2 33.9 0.000 1.000 21.8 15.2 i.- --- 21.8 16.7 25.6 16.6 14.4 26 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 The mandibles from these same localities are wholly characteristic of Erethizon rather than Coendou. The Inglis IA mandibles are referrable to Erethizon in every feature; especially diagnostic are posterior depth of mandible, ratio of mandibular depth at lower P4 and M2, and inflection of angular process. A line down the tooth row in both Inglis IA mandibles projects mediad to the incisor, a correlate of Erethizon's divergent tooth rows (White 1968, 1970). The Grand View mandibles, although incom- plete, look as though they had an angular process directed strongly mediad, as in Erethizon. This character cannot be determined in the El Casco and Vallecito Creek samples, but on all other features they were classified as £rethizon (greater than or equal to . 99) along with the Grand View specimens. Figures 5 through 11 compare fossil cranial specimens from various localities with Recent £rethizon and Coendou. Thus all fossil porcupines in North America can be referred to Erethizon using the cranial characters discussed above, although some Florida speci- mens show some Coendou dimensions in their skulls. We turn next to other sets of diagnostic characters revealed in the incisor enamel and audi- tory ossicles. INCISOR ENAMEL The value of incisor enamel microstructure in rodent taxonomy was first investigated by Tomes (1850) and later applied by Korvenkontio (1934). More recent studies involving incisor enamel microstructure in rodents include works by Wahlert (1968), Escala and Ghllego (1977), and War- shawsky (1971). Wahlert (1968) compared the incisor enamel of various fossil and Recent rodent groups as viewed in thin section. He examined the degree of variability in the enamel and discussed the three basic types of incisor enamel found in rodents. Escala and Ghllego (1971), also using thin sections, determined the variability and taxonomic usefulness of lower in- cisor enamel in several genera of Muridae. Warshawsky (1971) examined in detail the enamel microstructure of Rattus, using scanning electron micros- copy. In search of other taxonomic tools for the separation of Coendou and Erethizon, I examined the sagittal sections of lower incisor enamel in both of the living genera and in fossil specimens from various localities in North America, using scanning electron microscopy. Both genera and the fossil specimens have multiserial enamel as discussed by Wahlert (1968). Using the measurement procedures presented by Wahlert (1968) and Escala and Ghllego (1977), differences were found in the thickness of the inner enamel and the total enamel in Coendou and £rethizon. Figure 12 pictures the enamel of both genera in sagittal view and also shows the measurement methods. 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 27 436 A#*,1.USCI A B C 01\:fm D E F /1 0 1 ¥ G H ~cm i - ,=/7£'49/1..... -1-ri// - y/./ 00 000-= BU LLETIN FLO R ID A STATE M U SEU M V 27 N o. 1 = 0 .. . 9 . ~~ 00 0 . :0 I I . I " I . . 8- - I I . I . - I I . I . ...... I. . .. 0 7 - ... .. I. . * I ... ... /1 ." . I. - I 6 - . - I . -. ... I . .. * I . 5 • 0 I. I . 4 '1111,111'11 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 Ses 6.0 6.5 7.0 7.5 FIGURE B-8 PALATE WIDTH BETWEEN P4's 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 61 0 1. +3 0 61.,e 0 0 0 '@: to 1. 00 0 1. ... ..0 ® i00 0 0 . CO It O .....O 0 0 0. 0 0 18 0 0. 0 0 0.0 00 '1 1 +3 O 0.00<10.0 0 ..<3> .... 40 r 131 +3 0 0 0 0.0 0 0 0 It./Ek 1: 0 (2> .Ii. , LW X13ddn H IC IA -ril'€1 . 1... m= , . 1. 1 .liu. ... Tr. ... I. , I .3 ........ 1 10 e . ... . + *: ....... 1 .... . 1 I , ts . . 1 ... . + 0 . • T: I . 1 ..... . 1 *: : FI G UR E B -9 09:99:97:9: 10 .0 ........... rd Haddn Hial/* 62 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 0 1 11 N 0 1 60 .. 0 11 6 0 M. W O.0 00 0 0 0.00 0 .9215) EIS * n00 0.0 0 (07=\ " : 0.0.0.00 .>/L H 1 .......O ..1 o . • 1 3 0 0 00000 „liu„ 1 000 . 0 .P il. 0 , o o . 4/.) 1:00 0.00 7 0 4 T*.til , I 0 0 0 YZEk :6 0 1 : +3 : LW 213*~01 H.LCHM ,0 ... . I *. 1 . ... . . 0 ... ...... t; . . .. 1 .... . 1 . ... I .. 1. .... I . 1 ... .. /.. . 1 ..... . 1 I. . .1 .... , 1 + . 1: . 1 1. . 1 . O I-9 3Hn E Td kia/*01 HlaIM 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 63 4 : 0 .®e 10 0 00 O 000 0 0 t: 0.00000 + 9.0 000000 0 N O lilli. 0 0 o. o o 11 00,10.000 0 0 1 ,. O 0.00 0.0 0 I. 0 ...On.0 0 11e ra......„. ..,0 0000.0.41'111 1:". lIO SIO NI Pladdn H laIM O 0 0 1 00 H H 11 ... , I I i: I .. . 1. *. ...... ... .. . I. ... ..1/ ......, .. 1N ....... I , 1. I . . ,. I . 1 IN I. I. . IN . I . , . . .1 T I-9 3H nO W -/ 6 .2 + 6 .3 + 3333333231235323333333ES,2332 2 7 i r 0 . : 6 . 2 MOSIDNI 113ddn ldV 64 BULLETIN FLORIDA STATE MUSEUM Vol. 27 No. 1 @ 0 1 SO i* 9@ 0 00 10I. ..tk. 0 0 1.In00 0.4.0 :.©. ....O.0 0 +3 Go. 0e o. 1 4 0® ....O .., 40 1/I..3... 0 11 1 ®.... . 0 0 11 .. O 0 0 0 0~011 1 .~0 0 ::1/S.®8. " 1 0 0 11 0 11 0 0 1, i I I .. %1091 DNI 213/*01 H l.01/* .lak " 11'M I . ... • • • 1: .. 1.. *. 1 . . .. 0 ........ 1 . , ... I .... .. I. ..... . 11 ... I ...... .. 1 . . 1 . 43 . . I. 1 10 . I. Z I-g 3.noid ':1:19:1::9:1:t:::9:::1:19:1::9:1 . .. .. .*./................ .. .... I NOSIONI XIE1MC)1 ldV 36 1981 FR A ZIE R : N .A . FO S S IL E R E T H IZ O N T ID R E V IS IO N 65 34 0 0 00 0 032 0 0 00 @6% 0 00° o00 0-9 00 0.0 00 0 00400 030 000 000 0 0 0 00 0 00 0 000 28 - 0§DAK= o Liag =:0§ 00~: 0 0 *0 - 2*I- 7 O 00 0 0 - 000 0 0 0 00 = = 0= 0 00 26 · AL VE O LA R LE NG TH LO W ER TO O TH RO W 24 22 - * * * I I. . . .. . I I - I ** I * 20 - ... ** . I I - * *. I ** * * 18 * .** .. I . ... I. I 16 . * ... 14 * -----..----- 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 FIGURE B-13 SYMPHYSIS LENGTH 36 - 66 B U LLE T IN FLO R ID A S TATE M U S E U M Vol. 27 No. 1 34 - e 0 0 0 0 32 - 0 , 0 0 0 ® 0 00 0 00 0 00 0 0 =0 00 0 ,o - 0 oo o o 000 000 000 0 0 0 o ooo oo 00 0 0 AL VE OL AR LE NG TH LO W ER TO O TH RO W 00 B :O gi) O = O 0 00 0 0 28 - ,-O 0: oo -0 7 0. 0 00 0 00# O 00 0 0 0 00 0 0 0. 96 -00.0 0 0 0 0 24 . . .. I . . . * 22 . I . . ... . 0 - ' I . . ... I . . . . I . I I. I . 20 . * *. . * .I . * ..* I . . 18 ... **. . I . I .*. * 16 · ... I . .. I . t4 8.4 9.6 10.8 12.0 13.2 14.4 /S.6 16.8 18.0 19.2 20.4 2/.6 22.8 24.0 25.2 26.4 27.6 28.9 FIGURE B-14 DIASTEMA LENGTH 45 · 0 1981 FR A ZIE R : N .A . FO S S IL E R E T H IZ O N T ID R E V IS IO N 67 O @00 42 - 0 0 00 0 O 00 0 0· 00 0 00 0 00 0 00 0 39 O 00 0 0 o ooo ooo 0 0 0 000 0 oooo ~0~00000~%~ ~ 0 0~ 0 36 ~0 20 00 050 0 00 ®@ 10 00 0 (1© : :0~EO *Ef0 . 000 .00 00 0 00 0 0 SY M PH YS IS LE NG TH 0 0 0 33 O. 0 9(* _ 30 - - 0.. 900 -' * I I . I 27 - .-I. I = ./. ... I I I ... I .. . 24 . I. . ** I I. I . I . I . I ...... I . I 21 1 I ..... I ..* I . .. . .- I le · * i.** i .. . I . I . 15 - * I 12 8.4 9.6 10.8 12.0 13.2 14.4 15.6 16.8 18.0 69.2 20.4 2/.6 22.8 24.0 25.2 26.4 27.6 28. a FIGURE 8-15 DIASTEMA LENGTH 32 - @ 030 " e 0 0 0 28 - 0 000 00 00-0 00 0 o0 0 0 000=. 0 0 0.-0 26 - = ; 02 200* 004 Y < u 1 MA ND IB LE DE PT H AT P4 0 00 O .00.00 &'imp 000 0 0 24 - 0 0% +4@ RD o = 0. 0 r<6~ e22 - B U LLE T IN F LO R ID A S TATE M U S E U M Vol. 27 N o. 1 , * §7)61., 20 .. . I .-/ I 18 . I ... . . .. I .... I . * ... . . . I /6 .. + .... I . * 14 , I •-I .. :... 12 - 10 - '11111'1111.1,1,1,8/ &0 e.6 9.2 9.8 10.4 11.0 ll.6 /2.2 12.8 t 3.4 14.0 14.6 15.2 /5,8 l 6.4 17.0 /7.6 t 8.2 la.8 /9.4 FIGURE B-16 MANDIBLE DEPTH AT M2 32 * 1981 FR A ZIE R : N .A . FO S S IL E R E T H IZ O N T ID R E V IS IO N 69 0 30 0 0 00 0 0 00 0020 · 0 0 0 0 00 0 0 00 O 00 0 00 0 00 0 0 00 26 · .0 .0570%6&,5% M AN DI BL E DE PT H AT P4 0 0 = 00 0 0 0.0 0 00 00 0 000 0 0 0_00 = 0 00 00 24 - .. 900 000 0000 0 00 -g 00 00 0 0 = 0 00 0= 00= 00 0 0 22 - *0= 0 = 0 0 c • *0 0 = = I . 20 - I . I ...... ... . I . - I 18 - . . .. . ... I *... .-. I - . .* lilli . I . 16 4 I. I I ** I I - . I ... . 14 ****. . . 1 I . .. 12 10 + 14 16 18 20 22 24 26 28 30 32 34 36 38 .0 42 .4 46 48 FIGURE B-17 DEPTH CONDYLE TO ANGULAR PROCESS 19 - 0 0 70 B U LLE T IN FLO R ID A STATE M U S E U M Vol. 27 N o. 1 0 00 00 018 - O 000 0 0 0 00 00 0 00 0 0 0 00 000 017 - 0 0 0 00000000 0 0.0 0 0 0 0 0 0 00 0 OU 00 0¤ 0 16 · O 0 O 00 == 00~ O 0 0 00 O 0000 000 00 29 0 0 0 0 00ts - O/Tho = O . 00 0 =.. . 09~00 0=000 0 00 0 0 00 00000 0 = = = M AN DI BL E DE PT H AT M 2 0 I . 13 - = .. I . O12 · .. . I I . I . . .1 - .. I . .. . I. I I I 10 · . I ** I . I .. . . . I . I .... * 9 0-* . I. I 8 0 "* 14 16 le 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 FIGURE B-18 DEPTH CONDYLE TO ANGULAR PROCESS 1981 FRAZIER: N.A. FOSSIL ERETHIZONTID REVISION 71 LITERATURE CITED Ahlberg, H. D. 1969. Geographical variation in the North American porcupine Erethizon - (Linnaeus) Cuvier (Mammalia, Rodentia). Ph. D. 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