- ORIDA MUSEUM BULLETIN OF NATURAL HISTORY LANDSNAILS OF THE GENUS HUMBOLDTIANA FROM NORTHERN MEXICO (GASTROPODA, PULMONATA, HELICOIDEA, HUMBOLDTIANIDAE) Fred G. Thompson and Greg R Brewer Vol. 43, No. 2, pp. 49-77 2000 UNIVERSITY OF FLORIDA GAINESVILLE 1 .IGU ' 1"'.:pit. 'r' L~14 4 2 ,?1- : , ''* '" .1 Rhjd b ft I ' ,1 '' Numbers dfihe BULLETIN OF THE FLORIDA MUSEUMfOF NATURAL HISTORY' -·: I are published at'irregular intervals. Voiumes contain about 300 pages andare not ndcessarily completed in any one calendar year. FRED G. THOMPSON, Editor MARGARET E.B. JOYNER, Managing Editor Send communications concerning purchase or exchange of the publication and manuscripts queries to: Managing Editor of the BULLETIN Florida Museum ofNatural History University of Florida, P0 Box 117800 Gainesville, FL 326}11-7800, U.S.A. Phbne: 352-392-6724 Fax: 352-846-0287 e-mail: mjoyner @flmnh.ufl.edu ISSN: 0071-6154 CODEN: BF 5BA5 Publication date: November 15, 2000 Price: $5.00 - LANDSNAILS CiF'THE GENUS HmlfBOLD*1*t FROM NORTHERN MEXICO e «GASTROPODA, PULM0NATA, HELICOIDEA; mjMBOLDTIANIDAE) Fred G. Thompson? Florida Museum of Namral'Histoty Universifytof Florid* Gainesville, F-16fida, USA 32611 and Greg R Brewer 6071 Ranch Rd' 965 Fredericksburg,fTexas, USA 78624 ABSTRACT Seven new species are described fr6m the following Mexican states: NuevoLe6n: Humboldtiana porterae n. sp. and H edesma n. sp.; Zacatecas: H. bicincta n. sp.; Durango: # latizona n. sp. and H gra*i n. sp.; Querttaro:«H. pinicola n. sp.; Coahuila: H. oreina n. sp. The description of H plana Metcalf and Ridtind 1976 is amended, and the holotype is refigured. 'Ikvo new subgenera are propased: Okades n. subgen. (type species: Humboldtiona porterae n. sp.) and Polyomphalus n. subgen. (type species: H. oreina n. sp.). The generic definition of Humboldtiana s. s. is amended. Key words: Landsnail, Gastropoda, Pulmonata, Humboldtianidae, Humboldtiana, Oreades, Polymophala, Mexico. RESUMEN Se describen siete especies nuevas para los siguientes estados de Mexico: Nuevo Le6n: Humboldtiana porterae n. sp. y H. edesma n. sp.; Zacatecas: H. bicincta n. sp.; Durango: H. latizona n. sp. y H gra*i n. sp.; Quer6taro: H. pinicola n. sp: Coahuila: H. omina n. sp. Se corrige la description de H. plana Metmify Rielrind 1976 y se re,lpfine el holotipo. Se proponen dos nuevos subgeneros: Onades n. subgen. (especie tipica: HumboWtiana porterae n. sp.) y Polyomphalus n. subgen. (especie tipica: H. oreina n. sp.) y se corrige la definicion generica de Humboldtiana s. s. TABLE OF CONTENTS~ti~.f~~.g..········,·,······,················--.-.-.-- -. - ---.-.------- ---.--- .. 55~ Genus Humboldtiana Ihering, 1892 Subgenus Humboldtiana s. s. The Humboldtiana bulronian, species-group . ....._........................................... 51 Humboldbana gradyi n. ,p 51 Humboldtiana pinicola n. er 55 The Humboldtiand bicincm species-group...............,„............................................ 59 Humboldtiana. bicincta n. sp..............................„............................................ 59 Humboldliana, edemna n. *f - 62 The·H:imboW#a-*a texana species-group - 63 Humboldtiana latizona n. sp. 66 Oreades n.:suben 66 Hui4914ti*:d,parterae & ap 68 Pobumpha/a n. subgen. R+Arati**0*na n. sf 71 Huinbolatiana plana Metcalf and Risking, 1976 75 Acknowledgments.....„..................................................................-..„...............v_............ 77 Lit¢*4*re Cited....--------------------------------------. 77 "Correspondi~g 9**gEEISid~#ih~«9fNatur*1'History.,Wniversity of Florida, PO'Bo~-11~99k_98***116. FL 51611-7860 iii*i*©f.6., and F.P. Sewer. 2600. 1:andsnails'ofthe genus'Humboldlioilo-#diii n6rd~*rii®[exico'(dashpoda;'PulmonataiHeli@idea, H*S@Iii@idae) . Bull.:F16*6 Mus. NaL jHist: 43(2):49-77. 50 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) cannot be made. Solem (1974) attempted to determine INTRODUCTION species boundaries within Humboldtiana and to look for patterns of morphological specialization over broader The purpose ofthis paper is to describe seven new species geographic regions. He arbitrarily concluded that and two new subgenera of Humboldtiana found in the populations that differed anatomically from other Mexican states of Nuevo Le6ni Quer6taro, Coahuila, populations represented distinct species, and populations Zacatecas, and Durango. Another previously known that differed only in shell characters might or might not species from Coahuila is redescribed. Prior to this work be different species. He found no geographically the genus Humboldtiana contained thirty species and two associated patterns of anatomical differentiation. Our subspecies. Up to the present twenty-two species and observations are nearly consistent with his, except that one subspecies have been described from the Mexican we recognize two new subgenera which have state ofM6xico. Humboldtiana is distributed from Texas geographically restricted distributions. south to the states of M6xico and Veracruz. Populations Burch and Thompson (1957) recognized two occur in isolated colonies and no colony is known to species groups. Group4 was proposed for those species contain more than a single species. The intrinsic ability that have an enlarged epiphallic chamber, have a short, for geographic dispersal is limited, as evidenced by the stout verge, and have the dart-glands juxtaposed to the high degree of local endemism. Dispersal probably is dart-sacs. Presumably this group includes the type the result of long-term passive events associated with species, H. humboldtiana (Pfdiffbr, 1847), which remains tectonic and climactic changes and other adventitious anatomically unknown. Group-II was proposed for those means. Speciation appears to have been a result ofgenetic species that have a vestigial epiphallic chamber, a long, isolation from other colonies because of ecological slender verge, and have dart-glands separated from the barriers coupled with local processes of natural selection dart-sacs by a distance equal to or greater than the length and genetic drift. This produced a large number of ofthe glands. The division of Humboldtiana into these morphologically and geographically isolated species of two groups is no longer tenable. All the species we discuss which interspecific competition seems not to have been below have dart-glands juxtaposed to the dart-sacs and a factor in their evolution. none has an enlarged epiphallic chamber, whereas the Most species ofHumboldtiana are conservative in verge may be short and stout, orelongate andtubular, or the basic plan of the reproductive anatomy (Fig. 7). The even absent. For purposes of this paper we continue to genus is characterized by having four dart-sacs use group designations for species within the subgenus surrounding the vagina. Two dart-bulbs are associated Humboldtiana, but the designations are somewhat with each ofthe dart sacs. Four dart-glands form a ring arbitrary because of the lack ofanatomical information around the vagina abovethe dart-sacs and below the point on some previously described species. Also, it is apparent where the spermathecal duct diverges from the uterus. that many species remain to be discovered. Group The spermathecal duct bears an appendix near its distal boundaries and geographic patterns ofspeciation cannot end. The interior of the penis contains a verge. A great be resolved until then. Papers dealing with the deal of itinerant local variations from this plan may take phylogenetic relationships of the family Humboldtiana place. Reduction or modification of any of these are presumptive and contribute nothingto our Imowledge structures occurs in various species. Normally the dart- of its relationships (Cuezo, 1997). bulbs are imbedded in the wall of the vagina at the base of the dart-sacs and are not visible superficially. In a new subgenus from Coahuila the dart-bulbs are METHODS AND TERMINOLOGY conspicuous at the base of each dart-sac, as they are in the related genus Bunnya (Baker, 1942). A most striking The shell descriptions of new species are based departure from the general anatomical plan is found in primarily on the holotype. Meristic data, as well as another new subgenus that lacks dart-glands, dart-sacs, significant variations pertaining to the paratypes, are also and dart-bulbs on the vagina lacks an appendix on the included in the description. Morphological features spermathecal duct, and lacks a verge within the penis. among the paratypes that differ significantly from the Most Humboldtiana are known only from shells. character-states in the holotype are indicated in Anatomical data for the others are incomplete so that in parenthesis. Individual measurements are listed for the some instances critical comparisons between species holotype and the paratypes. THOMPSON & BREWER Landsnails of the genus Humboldtiana from northern Mexico 51 In this paper we use the term lateral expansion which encompass a great deal ofmorphological variation. rate as a ratio of the transverse distance from the apex Additional subgeneric divisions may be warranted as new to the outer lip divided by the width of the shell. The material becomes available for study. ratio is useful for comparisons between species. In the Etymology.-The genus is named forthe Prussian female reproductive system the dart-sacs and dart-glands naturalist Alexander von Humboldt (1769-1859). may be reduced in size or be absent in particular species. Two species may have non-homologous but equal The Humbo/dtiana bu#oniana species-group numbers of reduced dart-sacs. For comparative purposes we assign a number (dst, ds2, dsi, ds4) for each of the This group ofspecies is discussed in Pilsbry (1927, dart-sacs in a clockwise sequence beginning with the dart- 1948), and Burch and Thompson (1958). The group is sac that faces the atrium and penis (see Fig. 37). typical of the genus. It is characterized by having dull, Abbreviations for other structures in anatomical figures lusterless rough shells that are covered for the most part are as follows: with a dense mesh ofgranules on and between the rugose incremental striations and wrinkles. The reproductive agl albumen gland system has a ring of four dart-sacs on the vagina with app spennathecal appendix the four dart-glands discharging into the vagina between atr genital atrium the openings of the dart-sacs. The following two new db dart-bulb species belong here. dgl dart-gland epi epiphallus Humboldtiana gradyi new species fig flagellum (Figures 1-6,7-9)pen penis pr penis retractor muscle Diagnosis.-A medium-sized helicoid species of spt spermatheca the buSoniana species-group. Shell about 35 mm wide sptd spermathecal duct and with 3.9-4.2 whorls; lateral expansion rate 0.59-0.65. utr uterus The dull-colored three-banded shell is rugosely sculptured vag vagina with rough incremental striations and wrinkles. vd vas deferens Superimposed upon the wrinkles is a dense mesh of vrg verge minute granules that cover the last two whorls. The growth wrinkles and the granules are white, and obscure the tan background and the three bands. The translucentGenus Humboldtiana Ihering, 1892 interior of the aperture has a light brown wash. TheSubgenus Humboldtiana s. s. umbilicus is partially open. The aperture lies at an angle Type species: Helix humboldtiana Pfeiffer, 1847 of22-30° to the shell axis. Vagina with four equal-sized dart-sacs. Spermathecal appendix less than length of The shell is helicoid or depressed-helicoid in shape; distal segment of spennathecal duct. Penis bulbous at usually solid; color pattern three-banded. Sculpture apical end, with a large tri-lobed verge internally variable, smooth, and glossy to coarsely wrinkled; formed from a single fold of tissue pendent from the granules when present aligned parallel to axial growth apex and open along its side. Epiphallus longer than striations. Spermathecal duct slender, about as long as penis. Flagellum 1.2-1.4 times combined length of or longer than the combined length ofthe uterus+vagina; penis+epiphallus. spermathecal duct with an appendix. ~gina bearing dan- Among known species, the shell ofHumboldtiana glands, dart-sacs, and two dart-bulbs on each side ofthe gra*i is most similar to H. durangoensis Solem, 1954. dart-sacs; dart-bulbs imbedded in vaginal wall and not The latter is larger, has better defined color bands on a apparent externally; dart-glands forming a ring or a lighter background, is not as strongly streaked with white, partial ring around the vagina above the dart-sacs. Penis the suture is more deeply impressed, and the umbilicus bearing a verge internally. Flagellum moderately long. is reduced to a rimate perforation. The anatomy of H. Distribution.-Widely distributed from the durangoensis remains unknown. Guadalupe Mountains in Texas south to Puebla and Shell (Figs. 1-6).-Medium-sized for the genus, Veracruz. Humbold#anas. s. contains numerous species, 33-37 mm in diameter; helicoid in shape, being 0.89- 52 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) 1 2c 4 1 5 6 Figs. 1 -6 . Humboldtiana gradyi new species. Figs. 1 -3 , 6 : holotype (UF 267738). Figs. 4-5 : paratypes (UF 267720). THOMPSON & BREWER: Landsnails of the genus Humboldtiona from northern Mexico 53 0.98 times as high as wide. Ground color lusterless light Reproductive Anatomy (Figs. 7-9, Table 1).- tan with three brown bands and numerous close white Typical for the subgroup of Humboldtiana bu«foniana. axial streaks; the upper band is about halfway between The following description is based on five paratypes (UF the suture and the periphery; the second band is just above 267720). Genital atrium very short, 1-3 mm long. Penis the periphery; the lower band is about halfway between slender at base, upper half bulbous; interior (Fig. 9) with the periphery and the base of the body whorl; bands six longitudinal folds and containing a short, stout verge partially obscured by white axial streaks and a dense that is formed from a vertical fold of tissue that is tri- mesh of white granules; bands become decreasingly lobed at its lower end and is open along one edge to the distinct on last halfwhorl; apex cream-colored; peristome roof of the penis. Penis retractor muscle about equal to white; interior of aperture with a slight brown flush; or slightly longerthan the penis; originating on the middle exterior bands visible inside of shell. Spire conical and ofthe inner wall of the lung about one fourth of distance straight-sided with amoderately impressed suture. Shell posterior to the mantle collar; inserting on and enveloping consisting of 3.9-4.2 rapidly expanding whorls; lateral the base ofthe epiphallus. Epiphallus stocky, about half expansion rate 0.59-0.65; embryonic shell smooth (Fig. again as long as penis, lined internally with four 2), 4.9 mm in diameter and consisting of 1.5 whorls; longitudinal columns of tissue (Fig. 9a). Vas deferens first quarter of embryonic shell depressed so that apex nearly uniform in diameter between prostate and appears flat-topped in frontal view; subsequent whorls epiphallus. Flagellum about 1.2-1.4 times the combined regularly descending along the periphery; last whorl length of the penis+epiphallus. Vagina relatively short expanding laterally, and last quarter abruptly descending with four equal-sized dart-sacs. Dart-glands juxtaposed to the lower band; umbilicus narrowly perforate; about to the dart-sacs and coalesced into a ring around the 75 percent covered by the columellar reflection. Sculpture vagina. Spermathecal duct long (Fig. 7), about 1.5-1.7 consisting of irregular, closely spaced axial growth times the length ofthe uterus; spermathecal sac elongate- threads and wrinkles; superimposed on and between the elliptical in shape. Spermathecal appendix short, about growth threads are numerous minute axially-elongated 0.2-0.3 times the distal segment ofthe spermathecal duct; granules that are about 0.1 mm in length and tend to branching from duct at about 0.7-0.8 ofthe distance from coalesce (Fig. 6); striations and granules continuous over its base. Albumen gland large and elongate. Talon and surface of post embryonic shell. Aperture prosocline at carrefour imbedded in albumen gland. 22-30° to vertical axis of shell; aperture oblong-ovate in Type Locality.-DURANGO: area near the Dos shape; width equal to or slightly greater than height; with de Diciembre Microwave Tower, 2 km southeast, 5 lan athin, opaque, tan parietal callus; peristome very slightly east of Dos de Diciembre (24°44.6' N, 103°32.7' W); reflected along base, and more so along the columellar 2080 m alt. The area is on the edge of a knoll in a small reflection; upper end of outer lip inserted just above the range of low hills consisting of metamorphic rocks edge ofthe lower brown band. intruding through limestone and capped with caliche. The vegetation consisted of numerous Agave americana, Dasylirion sp. (sotal), Yucca spp., and occasional mesquite. Snails were found aestivating under Agave, dead Yucca, and sotal. HOLOTYPE: UF 267738; collec- Measurements in mm are given for the Humboldtiana ted 15 June, 1997, by Grady B. Taylor, Greg R Brewer, gra*i n. sp. holotype (UF 267738) and four paratypes and Fred G. Thompson. PARAIYPES: UF 267720 (10), (UF 267720) selected to show size variations in mature UF 267721 (10, animals of267720), ITCVZ 8012 (10); shells. same data as the holotype. Other Localities.-DURANGO: a limestone ridge Specimen Height Width ApH ApW Whorls 2 km SE, and 6.5 km ESE ofDos de Diciembre (24°23.1' holotype 31.3 34.5 23 24 3.9 N, 103°30.8' W); 2010 m alt. (UF 267685, ITCVZ). Remarks.-The two localities where this species paratype 30.5 34.0 22 23 4.1 was collected are about 3 km apart. We observe no paratype 31.0 35.0 22 23 4.0 significant differences between the two populations in paratype 32.5 33.0 22 22 4.1 the shell or the reproductive anatomy. paratype 34.0 37.0 23 24 4.2 The verge ofHumboldtiana gra*i is unique among 54 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) pr fig spt lt- vd app wl &1~ ds 3 vd -... vag pen atr8 10 mm agl - fig 9a epi epi Y a «»ng pen i.. »»/ ds 3 u «" vrg atratr 7 9 10 mm 2 mm Figs. 7-9. Humbola'anagra*i new species. Reproductive anatomy oftwo paratypes (UF 267721). Fig. 7: complete reproductive system. Fig. 8: terminal genital structures of a second specimen. Fig. 9: interior ofpenis fromFig. 8. Fig. 9a: transverse section ofepiphallus in Fig. 9. THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 55 known species in the H. bu#oniana species-group. It on the growth wrinkles and generally are the same color consists ofa pendent fold oftissue that is open to the apex as their background; granules become weaker near the along one side. Most other species have a tubular verge, umbilicus. Aperture broadly ovate, 0.80-1.06 times as although it may be very short, as in the following species. high as wide, and about 0.58-0.69 times the height ofthe Etymology.-We take pleasure in naming this shell; plane ofaperture prosocline at an angle of32-39° species after Grady B. Taylor (San Antonio, Texas) for to the shell axis. Outer lip sharp but slightly reflected; his companionship in the field and in recognition of his basal lip thicker; columellar lip more broadly reflected lifelong dedication to and love for shells. and partially obstructing the narrow umbilicus; parietal wall with a thin transparent callus. Reproductive anatomy (Figs.16-18, Table 1).- Humbo/dtiana pinicola new species (Figs. 10-15, 16-18) The following description is based on five paratypes (UF 268440). Genital atrium long and relatively slender, about Diagnosis.-A large helicoid shell that is about 40 mm equal to the length of the penis. Penis short and stocky, wide and consists of 4.4-4.8 whorls. Lateral expansion shorterthan length ofvagina; slightly compressed antero- rate 0.59-0.64. The lusterless shell is relatively rugose posteriorly. Interior of penis with a long verge that has with rough white incremental wrinkles and a dense mesh three or four twisted terminal lobes (Fig. 18). Verge of granules on the last 2.5 whorls. The light brown shell extending to opening ofvagina. Inner wall ofdistal half bears three black bands intermittently interrupted by the of penis with several large, thick transverse fleshy folds, white growth wrinkle. The aperture is prosocline at an and about 6-8 smaller longitudinal ridges below. Penis angle of 32-39° to the axis of the shell. retractor muscle long, originating on middle of lung wall Superficially Humboldtiana pinicola is most just behind mantle collar and inserting on base of similar in shell features to H. fasciata Burch and epiphallus . Epiphallus short and stout; enlarged at base; Thompson, 1957 from Hidalgo, which has 4.24.3 whorls entering penis eccentrically due to longitudinal and an aperture that lies at a steeper angle of about 45°. compression ofpenis (Fig. 18);interior ofepiphallus with However, the reproductive anatomy ofH. fasciata isvery four longitudinal folds (Figs . 17a, 17b). Flagellum different in that the penis is elliptical-elongate and the relatively short for the genus (Fig. 17), about 1.7-2.6 mucus glands on the vagina are widely separated from times the combined length of the penis+epiphallus. The the dart-sacs. Humboldtiana jasciata is placed in the high ratio is due to very short length of penis and te=ina species-group. epiphallus . Vas deferens stout, but constricted at Shell (Figs. 10-15).-Shell conical-globose in epiphallus and again above atrium (Fig. 16). Vagina shape; medium- to large-sized for the genus, 36.5-42.6 capacious, almost as long as combined length of mm wide; 0.85-1.061imes as high as wide. Spire straight- penis+atrium; bearing four unequal sized dart-sacs; dsl sided, 0.33-0.44 times height of shell. Shell lusterless, largest; ds# second largest; ds2 smallest. Dart-glands ground color light brown with three broad, bold, black forming a thick ring around vagina juxtaposed to dart- spiral bands, of which the middle band is usually the sacs. Spermathecal duct (Fig. 16) moderately long, stout narrowest; interior of aperture with a white flush, but about 1.04-1.26 times length ofthe uterus; spermathecal clearly showing the external banded pattern through the sac small, bulbous; appendix shorter than distal segment thin shell. Suture moderately impressed. Whorls 4.4-4.8; of spermathecal duct; branching from duct at about 0.5- body whorl rapidly expanding, lateral expansion rate 0.7 the distance from its base. Albumen gland large, 0.59-0.64; last quarter whorl descending below the reniform in shape; carrefour imbedded in albumen gland. periphery to the lower band. Embryonic shell (Fig. 11) Talon elongate, exposed. caramel in color, with 1.6-1.7 whorls; raised and pitted; Type Locality.-QUERMARO: southeast side of first whorl 4.2 mm wide and planar and then descending Cerro Puerto El Pino, limestone outcrop in an oak-pine along the suture. Following whorls sculptured with coarse forest ca. 6 km by road southwest of Pinal de Amoles growth wrinkles that tend to be white and which (21°07.5'N, 99°38.2' W); 2650 malt. The area consisted intermittently disrupt the black bands. The last two whorls of sparse forest and open grassy pastures with stone ofthe shell are covered with a dense mesh ofsmall oblong- fences and walls along their borders. Snails were found shaped granules, which are about 0.3 mm in size and aestivating in crevices in limestone outcrops and stone tend to be aligned axially;'~granules largest and densest walls and on pine trees. HOLOTYPE: UF 268441; 56 BULLETIN FLORIDA MUSEUM NArURAL HISTORY VOL. 43(2) Table 1.- Humboldtiana. Comparative measurements of the reproductive anatomy of: 1-5 H. pinicola n. sp. (UF 268440); 6-9) H. gradyi n. sp. (UF 267720); and 10) H. gradyi n. sp. (UF267685). Legend: PR = penis retractor muscle, PN = penis, EP = epiphallus, FL = flagellum, AT = atrium, VG = vagina. FV = free vagina, UT = uterus, SD = spermathecal duct, SB = base of sper-mathecal duct, SP = end of spermathecal duct, AP = appendix. The length of the spermathecal duct includes the spermatheca. The end of the spennathecal duct includes the spermathecal bulb and that segment of the duct above the appendix. The base of the spermathecal duct includes that segment below the appendix. Spec. no. PR PN EP FL AT VG FV UT SD SB Sp Ap H. pinicola n. sp. (UF 268440) 1 17 12 7 38 4 13 0 61 69 40 29 19 2 18 9 7 36 4 16 0 63 67 34 33 20 3 18 8 6 37 3 15 0 57 59 40 19 14 4 18 8 7 38 4 14 0 49 62 36 26 14 5 13 11 8 33 3 14 0 48 50 31 19 16 H. gradyi n. sp. (UF 267720) 6 11 11 14 33 3 11 2 47 71 54 17 5 7 5 11 22 41 2 14 2 44 71 59 12 - 8 22 9 15 33 2 10 2 46 70 55 15 3 9 16 9 16 30 1 15 3 43 74 54 20 6 H. gradyi n. sp. (UF 267685) 10 20 9 9 32 3 16 7 44 69 49 20 8 collected February 1, 1998, by Steven R Christman. and Other records of specimens referred to this species are: Fred G. Thompson. PARATYPES: UF 268440 (7), UF Pinal de Amoles, 2,600 m alt. (UF 251482); 2 km S of 268276 (7), ITCVZ 8013 (6); same data as the holotype. Pinal de Amoles, 2470 m alt. (UF 218724); 2.5 km NE Distribution.-This species is 1mown only from of Pinal de Amoles, 2170 m alt. (21°18.5 ' N, 99°37.1' W) the immediate vicinity of the lype locality where it is (UF 200692, UF 268248). found in pine forests and mixed hardwood-pine forests. Remarks.-There is little variation among the population samples we have examined. Shells from 2.5 km NE of Pinal de Amoles have darker bands and the Measurements in mm are given for the Humboldtiana ground color is brighter orange-yellow. pinicola n. sp. holotype (UF 268441) and six paratypes This species occurs in the same region as (UF 268440) selected to show variation. Humboldtiana quentaroana (Dall, 1897). The latter has a higher spire, the umbilicus is nearly occluded by the Specimen Height Width ApH ApW Whorls reflected columellar lip, the band4ess adult shell has prominent white axial streaks, and the interior of the holatype 39.0 42.5 22.5 28.0 4.5 aperture is dark reddish brown (Solem, 1955: 41). We have examined two juvenile specimens of H. paratype 38.5 42.0 24.3 27.7 4.7 queretaroana from Pinal de Amoles (UF 267270), the paratype 41.0 40.0 25.5 25.5 4.7 type locality. We do not have exact geographic paratype 40.4 42.6 29.0 28.0 4.7 coordinates for the locality from which the specimens paratype 39.5 37.0 24.2 27.0 4.6 were collected or the proximity of H. queretaroana to paratype 37.0 42.0 24.8 26.5 4.5 known populations of H. pinicola. The shells of H. paratype 35.0 36.4 24.0 24.0 4.4 quentamana have faint indications of narrow bands. THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 57 10 11 12 13 S , \Y'4 j ''i f 14 ; :C.* k - 15 Figs. 10- 15 . Humboldtianapinicola new species. Figs. 10- 13 , 15 : holotype (UF 268441). Fig. 14 : paratype (UF 268440). 58 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) utr17a 17b p 1F X ! 1, dsl -W, dgi a spt .(.3 -- I .rnsapp .fl 41 / -,L atr vd y fig L-,4& 17 1 Ililli 10 mm epi ds4 4r- \42# / / pen --4 epi ds2 vrg pen vag atr atr16 18 1 111,11 10 mm 5 mm Figs. 16-18. Humboldliana pinicola new species. Reproductive anatomy of a paratype (UF 268440). Fig. 16: complete system. Fig. 17: mesad side of lower genital anatomy; Figs. 174 17b transverse sections of the epiphallus at points a and b in Fig. 17. Fig. 18: interior ofpenis. THOMPSON & BREWER: I.andsnails of the genus Humboldtiana from northern Mexico 59 The sculpture is similar to that of H pinicola, except Measurements in mm are given of the Humboldtiana that the granules are more numerous and more densely bicincta n. sp. holotype (UF 268291) and three paratypes packed together. (UF 268289, 268290). Etymology.-The species name pinicola is from the Latin pinus, a pine tree, and cola, a dweller, and Specimen Height Width ApH ApW Whorls alludes to its habitat ofpine forests. holotype 25.1 33.2 20.1 20.5 4.0 The Humboldtiana bicincta species-group paratype 24.0 33.0 20.5 21.3 3.9 paratype 23.5 29.8 19.1 19.5 3.9 The following two new species differ from other north paratype 25.6 32.0 21.0 21.2 3.9 Mexican Humboldtiana bytheir depressed, shiny, nearly smooth light-colored shells, and by the tendency for reduction of the dart-sac apparatus in the female reproductive system. expansion rate 0.68-0.71. Embryonic shell tan colored, consists of 1.4-1.5 whorls which are 4.4-4.6 mm wide Humbo/dtiana bicincm new species and have weak radial striation along the suture of the (Figs. 19-24, 25-29) last halfwhorl. Subsequent whorls with close incremental growth striations; granular sculpture absent. ApertureDiagnosis.-A depressed-helicoid shell that is about 30 mm wide, and 0.73-0.80 times as high as wide with an obovate in shape; 0.96-0.99 times as high as wide; prosocline at an angle of 40-44° to shell axis; parietalinflated body whorl. The lateral expansion rate is 0.68- callus thin and transparent; slightly convex in outline;0.71. The shell has two prominent peripheral dark brown bands on a lavender-white background. A very faint peristome white or flesh-colors; slightly reflected along upper, outer, and basal margins, more so along columellarobsolete third dorsal band is also present. The semi-glossy margin which nearly covers the umbilicus.shell is sculptured with fine but distinct incremental Reproductive system (Figs. 25-29; Table 2).-growth striations. Granular sculpture is absent. The The following description is based on the holotype andvagina has a single functional dart-sac (ds,) and one vestigial dart-sac (ds~· The degree of reduction of the one paratype (UF 268290). Genital opening below right eye-stalk just above margin offoot. Genital atrium short.dart-sacs and dart-glands is unique within the subgenus Penis retractor muscle short and stout; originating onHumboldnana, as is also the presence of only three nght side of lung just behind mantle collar, inserting onlongitudinal folds lining the epiphallic lumen. and encompassing lower quarter of epiphallus. PenisHumboldtiana bicincta also differs from other known elongate; slender below and club-shaped distally; slenderspecies of the subgenus by lacking an appendix on the base lined internally with five longitudinal folds; interiorspermathecal duct, or by having only a vestigial appendix. of distal club-shaped portion with two transverse fleshyShell (Figs. 19-24).-Shell medium-sized, 23.5- folds at the bottom of the cavity; an epiphallic papilla25.6 mm wide; depressed-helicoid in shape with a rotund (verge) extends into the cavity almost to its base. Thebody whorl, 0.73-0.80 times as high as wide. Spire obtuse, forming an angle of 118-126°. Moderately thick- end ofthe verge has three short lobes. Epiphallus stout and muscular; about three times as long as penis; linedshelled semi-glossy, smooth. Suture paralleling upper internally with two stout longitudinal folds and oneband until near the last quarter ofthe body whorl where smaller fold (Fig. 25a). Flagellum stout and relativelyit abruptly descends to the aperture. Color lavender-white long, about three times the combined length ofwith numerous darker growth streaks and three brown bands. The uppermost band on the dorsal side of the penis+epiphallus (Fig. 25). Vagina about as long as penis; basal half of vagina slender, and with six sub-equalbody whorl is faint but persistent. Two prominent bands longitudinal folds internally; upper portion of vaginaparallel the periphery, one above and one below, and greatly enlarged; with only two dart-sacs, ds, ontheatrialsuperficially give the shell a two-banded appearance. side is large and functional, ds3 on the opposite side isInterior ofaperture light tan, approaching opaque white small and vestigial (Figs. 25,26); dart-sac dsl with twotoward the peristome; external bands barely dart-bulbs imbedded in vaginal wall at base of sac (Fig.distinguishable internally. Suture weakly impressed. 26); dart-sac ds3 without darts or dart-bulbs. TwoUmbilicus rimate. Shell with 3.9-4.1 whorls; lateral 60 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) 19 20 21 22 23 24 Figs. 19-24. Humboldtiana bicincta new species. Figs. 19-23 : holotype (UF 268291). Fig. 24 : paratype (UF 268290). TH6*SoN *BREWER: Landsdik bithe,*ehus Humb.6/dtia#a from northem Mexico ,61 . f * _agl /-fig ~ pro 25a epi * db vas i 9 utrd53 Vf 1126 spt ds3 atr , 9,w i --vd Spt» ~ & C ..23625 - pen 10 mm 29 lili ll , I11IIIf 10 mmspd 1 ,rspd dgi pen f dsl -, 3 / dgI Vag 1 vag 27 28 lilli 11- 1 1- 1 5 mm10 mm Figs. 25-29. Humbohffiaha bicineta new species. Fig. 25: reproductive system of holotype. Fig. 258: transverse secti6n through,epiphallus'bat·point a hi~fig: *3. Eig. 166 transverse section *ugkdart-gl~*andvaginajat point V in Fig. 23. Fig. 27: mes*tdiae,gf lower genitalia in Fig. 25. Fig. 28: dart apparatus infig. 27. Fi*. 29: pallidlj Rmale system of paratype showing-vestigialappendixon spermathecal duct. 62 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) unequal-sized dart-glands nearly surround the vagina sized and ds2 and ds3 are vestigial. Thus, the functional above andjuxtaposed to the dart-sacs (Figs. 25,26,29). dart-sacs in H bicincta, H. nuevoleonis, and H. fortis The larger gland lies above dart-sac ds, Spermathecal are not homologous. Dart-gland reductions do not occur duct diverging from vagina immediately above dart- in H. nuevoleonis or H. fortis. Rather, eachretahis four glands; spermatheca small and bulbous; appressed to nearly equal-sized dart-glands that form a ring around mesad side of uterus just below albumen gland; the vagina. spermathecal duct very long, about twice or more the The shell of Humboldtiana bicincta is similar to length of the uterus+vagina, spermathecal duct without H tomi Pilsbry, 1935 from"Chihuahua"and H eulaliae an appendix gig. 25), or with only a small vestigial Metcalf, 1984 fromthe Chihuahua-Coahuilaborderarea appendix (Fig . 29). Uterus strongly folded and near Guimbalete, Chihuahua, 1250 m alt . (27° 12 . 0 ' N, convoluted. Albumen gland very large and solid, about 103°47.6' W). Both these species also lack granular half as long as uterus. Hermaphroditic duct enlarged and sculpture on the shell. Both are three-banded, and the strongly convoluted. Ovotestis imbedded in lower two- bands are narrower than they are in H. bicincm. thirds of digestive gland; consisting of about three to Humboldtiana torrei and H. ealaliae remam unknown six, usually four, clusters ofmultilobed, follicular alveoli. anatomically. Type Locality.-ZACATECAS: Sierra de Etymology.-The species name bicincta is from Mascaron, Cuesta de Judas, 4 lan south by road from the Latin bi-, two, and cinctum, a belt, and refers to the Las Crucitas, 2500 m alt. (24°41.7' N, 101°39.0' W). two bold spiral bands that distinguish this species. HOLOTYPE: UF 268291; collected February 5, 1998, by Fred G. Thompson and Steven P. Christman. Humboldttana edesma new species PARAIYPES: UF 268290 (1), UF 268289 (2), ITCVZ (Figs. 30-34,36-41) 8014 (2); same data as the holotype. The Cuesta de Judas forms a narrow east-west oriented ridge which bears a Diagnosis.-The shell is 30-35 mm wide and has 3.8- sparse forest oflow pines and oaks. Snails were collected 4.9 whorls with a lateral expansion rate of 0.66-0.71. at the top ofthe Cafion de Ju(las, on the south side of the The thin shell is depressed, medium-sized, glossy whitish ridge below the forest zone among dense stands ofAgave with three uninterrupted bold brown bands. The sculpture sp. and some Yucca sp. Las Crucitas is on the north side consists ofnumerous growth striations and wrinkles; the of the Sierra de Mascar6n on the road to Terminal de dorsal surface of the last whorl has obsolete granular Providencia, and is about 37 km west of Hwy 54. The sculpture between the wrinkles. The umbilicus is only type locality is reached south of the mine Mineral de partially obstructed by the columellar reflection. The Nochebuena at I..as Crucitas by a very steep mining road peristome is prosocline at about 42-48°. It is most similar with numerous narrow switch-backs. to and geographically closest to the following species. Distribution.-Known only from the type locality. Differences are discussed below The reproductive system Remarks .-The systematic relationship of is distinct from other known Humboldtiana because of Humboldtiana bicincta is uncertain. It differs from other the short bulbous penis and the very short penis retractor known species by the combination of its depressed muscle that inserts on the lower right side of the mantle smooth glossy white shell that is devoid of granular collar. sculpture, by the very much reduced dart-sac complex Shell(Figs. 30-34).-Shell small- tomedium-sized to one functional dart-sac and one vestigial dart-sac, by for the genus; depressed-helicoid in shape; adults 30-35 reduction of the dart-gland complex surrounding the mm in diameter, and 0.77-0.87 times as high as wide. upper vagina, and by the vestigial appendix on the Spire low and obtuse, forming anangle of 110-122°. Shell spermathecal duct. Dart-sac reduction also occurs in such smooth, glossy, cream-buff with three uninterrupted other species as H. nuevoleonis Pilsbry, 1927 (see strong brown bands . Interior of aperture with a light Pilsbry, 1948) and H. fbrns Pilsbry, 1948. Each has only brown tinge; translucent and showing external dark two functional dart-sacs. Pilsbry (1948) does not identify bands. Suture moderately impressed, descending abruptly which of the dart-sacs are lost, but interpretations can just behind aperture (Fig. 33). Umbilicus narrow and be made from his illustrations based on the relationship partially obstructed by reflected columellar lip. Whorls ofthe dart-sacs to the origin ofthe spermathecal duct. In 3.8-4.0, rapidly expand laterally and separated by a H. nuevoleonis, dart sacs dsl and ds2 are lost; ds3 and ds4 moderately weakly impressed suture; last quarter whorl are normal-sized. In H. fortis, dsl and ds4 are normal- descending to the lower band. Lateral expansion rate THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 63 0.66-0.71. Embryonic shell (Fig. 34) consisting of 1.3- gland stout, reniform; talon and carrefour imbedded in 1.4 buff- or cream-colored shiny whorls; 4.1-4.5 mm albumen gland. wide; very weak radial threads along outer edge of the Type locality.-NUEVO LEON: Sierra San last quarter whorl. The post-embryonic whorls have close Francisco de Desmontes, east slope of Cerro La Mota, growth striations and wrinkles that are strongest and Cueva Ahumada5 lansouthwest ofLos Fierros (25°41.2' recurved along the suture and become diminished below N, 100°42.9' W); 1,200 m alt. HOLOTYPE: UF 244468; the periphery. Dorsal side of last whorl has small, weak, collected by Greg P. Brewer, 13 August, 1995. granular sculpture aligned radially between the wrinkles. PARAIYPES: UF 271007 (2), UF 271403 (3), ITCVZ The granules become indistinguishable below the lower 8015 (2); same data as the holotype. Specimens were band. Aperture obovate in shape and 1.01-1.15 times as found crawling on a damp, iron oxide-colored sandstone high as wide. Upper corners of peristome converging and under sotal (Dasylirion sp.) skirts in a zone densely closely. Parietal callus thin and transparent. Plane of vegetamd with the ubiquitous Agave lechuguilla and other aperture prosocline at 42-48° to shell axis. Peristome low-growing vegetation and grasses. Numerous rodent- white or cream-colored, weakly expanded along outer gnawed snail shells were scattered about the area. and basal margins, more so along columellar reflection, Distribution.-This species is found in the Sierra which covers about half'ofumbilicus. San Francisco de Desmontes. The sierra is a low, dry Reproductive system (Figs. 36-41, Table 2).- limestone range that is aligned NE-SW across theNuevo The following description is based on the holotype and Le6n-Coahuila border east of Saltillo. The type locality two paratypes. Atrium very short and stout, hardly is at the east end of the range. The following population distinguishable. Penis short and bulbous. Interior with a is referred to this species: COAHUILA: Cuesta de Boey, pendant fold oftissue that extends the length ofthe cavity 5 km NE of Mes6n del Norte, 1450 m (25°37.7' N, (Figs. 38,39,40) and surrounds a much smaller bi-lobed 100°55.2' W) (UF 268317, UF 268318, UF 271008). epiphallic papilla that extends downward as a verge (Fig. Remarks.-No significant shell differences exist 40). Epiphallus relatively stout and about as long as, or among the specimens from the two localities where this longer than, the penis, with four longitudinal folds species was collected. However, a specimen from internally (Fig. 408). Penis retractor muscle originating northeast of Mes6n del Norte differs in the reproductive on the lower right margin of the lung at the edge of the anatomy from the type population in several conspicuous mantle collar and inserting onthe base ofthe epiphallus; features (Fig. 41). The penis and vagina complex is larger, retractormuscle very thick and short abouthalfthe length the penis has a short neck above the genital atrium, the of the penis. Flagellum long, 3-6 times the combined long slenderpenis retractor muscle originates on the mid- length ofthe penis+epiphallus. Vagina bearing four equal- dorsal margin of the inner lung wall, the epiphallus is sized dart-sacs and dart-glands. Dart-glands separated proportionally longer and not as stocky, andthe flagellum from dart-sacs by a distance almost as great as the base is longer. Otherwise it is similar to typical Humboldtimm ofthe vagina. Spermathecal duct long; spermathecal sac edes/na in development ofthe bulbous penis and in the elliptical; spermathecal appendix about equal to orgreater configuration of the dart-sacs and dart-glands. We than the length of spermathecal duct (Fig. 36); entering tentatively consider the two populations to be conspecific. duct at 0.7-0.8 ofthe distance from its base. Albumen Humboldtiana edesma is similar to H. bicincta because both have relatively smooth, white shells. They differ from each other in many features of the reproductive Measurements in mm are of Humboldtiana edesma n. systems, as are described above. sp. holotype (UF 244468) and three paratypes (UF Etymology.-The species name edesma is from 271403). the Greek 66&apa, meaning food or edibles such as delectable snacks, and alludes tothe factthatthis species, Specimen Height Width ApH ApW Whorls like many other land snails, is devoured by rodents. holotype 24.7 35.0 21.1 20.8 4.0 The Humboldtiana texana species-group paratype 28.0 32.0 23.0 20.0 4.0 paratype 24.3 30.5 20.5 18.2 3.8 The following species tentatively is placed in this group. paratype 31.0 24.7 21.0 20.4 3.8 Its anatomy is unknown. The shells of H. hogeana (Martens, 1890), H tomi Pilsbry, 1935, andH eulaliae 64 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) 30 31 32 33 Figs. 30-35. Humboldtiana. Figs. 30-34: H. edesma new species, holotype (UF 244468). Fig. 35: H. latizona new species, holotype (UF 271009). THOMPSON.&,BREWER: Lahds,iails,of the *enta Humbblatiana.from northern *exico '65 app 5 is app 41 I 1 Y app spd E 852 - , vd 3 36 4=f pen 1 :11 11,1 10 mm ds2 (Creproductive system ofspetimen from Cuesta'~de Biley (UF 268817). 66 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) Metcalf, 1984 are geographically proximal. They too forming a broad sub-sutural nist-colored zone. Middle remain unknown anatomically. band variable in width on different specimens, separated from other bands by a light tan zone above and below. Lower band irregular in width along lower margin, and Humboldtiana latizona new species tending to expand and form blotches as it nears the (Figs. 42-47,35) peristome. Base grayish white. Parietal wall and interior Diagnosis.-This is assumed to be in the Humboldtiana of aperture with a transparent wash, clearly showing texana species-group because of its semi-glossy external color pattern . Umbilicus narrow, obscured by depressed-helicoid shell, which is a common feature reflected columellar lip. Whorls 3.9-4.2;lateral expansion among species ofthis group, although not diagnostic. It rate O.60-0.68. Suture moderately impressed, last quarter is similar to H tomi and H. eulaliae because of its whorl descending to the lower band (Fig. 45). Embryonic smooth, glossy, light-colored shell. Its medium-sized shell shell (Fig. 35) with 1.3-1.4 whorls; light tan in color, is 31-34 mm wide, with 3.9-4.2 whorls, and is 0.77-0.85 smooth with fine wavy radial striations along the suture times as high as wide. Its lateral expansion rate is 0.60- of the last quarter whorl. Lower whorls with fine 0.68. The shell is sculptured with close incremental incremental striations and growth wrinkles, and covered striations and weak growth wrinkles over which are very with very fine elongate granules that are axially aligned; fine granules on the dorsal and lateral surfaces of the granules becoming smaller and more widely spaced on whorls. The granules continue onto the base but are the base of the shell. Aperture obovate in shape, 0.99- diminished in size and frequency. This species' most 1.08 times as high as wide; prosocline at an angle of 40- distinctive feature is the color pattern. The shell has three 46° to axis of shell; upper ends of peristome closely reddish-brown bands on a light tan background. A wide, converging. Upper and outer lip ofperistome not reflec- diffuse rusty zone extends from the suture to the upper ted, or reflected very slightly; basallip weakly reflected; band and merges with it. columellar margin of peristome narrowly reflected. Shell (Figs. 42-47, 35).-Shell medium-sized, 31- Type Locality.-DURANGO: Sierra San Lorenzo, 35 mm wide; depressed helicoid in shape, 0.77-0.85 times Mesitas Coloradas, 7.5 km east of Pedricefia, 1850 m as high as wide. Shiny, moderately thick. Spire broadly alt. (25°04.4' N, 103°41.5' W). HOLOTYPE: UF conical, forming an angle of 117-124°. Shell with three 271009; collected 27 December, 1969, by Fred G. reddish-brown bands on a whitish-tan background. The Thompson. PARAIYPES: UF 130585 (5), ITCVZ 8017 zone between the suture and the upper band is light rust (5); same data as the holotype. colored and blends with the band, causing the inner Distribution.-Known only from the type locality. margin of the upper band to be poorly defined, and Remarks.-The color pattern of this species is unique within Humboldtiana. The broad rusty zohelying between the suture and the upper band, as well as the poorly defined rusty lower band, contrast strongly with Measurements in mm of the Humboldtiana latizona n. other species from adjacent areas with similar shells. In sp. holotype (UF 271009) and nine paratypes are as H. tomi the three bands are sharply defined, uneven, follows. narrow, and woody-brown (Pilsbry, 1935). The shell is Specimen Height Width ApH ApW Whorls devoid of granular sculpture. The shell of H eulahae has three very narrow dark brown bands, and it also lacks holotype 26.8 34.8 22.7 22.1 4.0 granular sculpture (Metcalf, 1984). Elymology.-The name latizona is from the Latin paratype 26.0 35.0 22.1 23.0 3.9 lams, broad, and zona, a belt or girdle. It alludes to the paratype 27.0 34.2 23.0 21.5 3.9 wide reddish zone on the last whorl between the suture paratype 30.0 34.5 22.8 22.6 4.3 and the upper band. paratype 27.3 33.0 23.2 23.0 4.1 paratype 27.0 33.0 22.5 21.2 4.0 paratype 26 .0 32 . 0 21 . 8 21 .0 3 .9 Oreades new subgenus paratype 26.0 31.0 20.5 20.5 4.2 7ype species.-Humboldrianaporferae new species paratype 25.5 31.2 21.0 20.5 4.0 The shell is helicoid in shape; thin and fragile compared paratype 29.8 35.0 24.1 23.0 4.2 to other Humboldtiana; color pattern three-banded, as THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 67 A A - 4 Figs. 42-47 . Humboldtiana latizona new species. Figs . 42-45 : holotype (UF 271009). Figs. 46-47 : paratypes (UF 130585). 68 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) is typical for the genus; sculpture consisting of coarse whorl descending below the periphery tothe lower band. incremental growth striation and elongate granules Embryonic shell 5.0-5.3 mm wide, consisting of 1.3 aligned parallel to growth striations. Spermathecal duct smooth whorls with radial striation along the suture. short and stout; duct lacking an appendix. Vagina lacking Postembryonic whorls with rugose transverse striations dart sacs, dartbulbs, or dartglands. Penis slender, lacking and wrinkles. The latter bear numerous weak elongate a verge internally. Flagellum very short about one-third granules that are axially aligned and occur on the last the combined length ofthe penis+epiphallus. 2.5 whorls; they tend to diminish on the fourth whorl The reproductive anatomy ofOreades differs from above and below the periphery, but they may occur that of all other Humboldtianidae by its highly derived sporadically into the umbilical area in occasional apomorphic character states. These include the complete specimens. Aperture prosocline, lying at an angle of40- loss ofdart sacs, dartglands, and dartbulbs onthe vagina: 48° to shell axis. Aperture obovate, 0.97-1.04 times as having a short spermathecal duct that is about one-half high as wide. Parietal callus thin, transparent. Peristome the combined length of the vagina-uterus; lacking an uniformly reflected along outer and b=,Al lip; columellar appendix on the spermathecal stalk; and lacking a verge lip strongly reflected, appressed, and nearly occluding in the penis. There is a tendency for reduction or loss of unnbilicus. various of these structures among particular species Reproductive system (Figs. 52-54, Table 2).- within the subgenus Humboldtiana, butnone approaches The following description is based on two topotypes (UF the reduced character state that distinguishes Oreades. 271399). The shells of both were shattered while Distribution.-Oreades is known only from the extracting the bodies. Atrium moderately short and Caaon de Garcia, Nueyo Le6n. narrow. The penis retractormuscle originates onthe right Etymology.-The subgeneric name Oreades (f.) side ofthe inner wall ofthe lung immediately behind the is fromthe Greek opeaS< meaning nymphs who inhabit mantle collar and inserts on the apex of the penis. The hills, and alludes to the snail's graceful, delicate shell penis is slender, nearly cylindrical, slightly narrowednear and its natural habitat. base; undifferentiated externally from epiphallus, except for insertion of the penis retractor muscle. Interior of penis lined with several longitudinal fleshy pilasters (Fig. Humboldtiana porterae new species 53). A verge is absent. The opening from the epiphallus (Figs. 48-51,52-54) is surrounded by four triangular fleshy papillae that Diagnosis.-This species has a large depressed-helicoid protrude into the penis lumen (Fig. 53). The papillae are shell. It is readily recognized by its color pattern, which short extensions of four longitudinal folds lining the consists of three uninterrupted bold chocolate-brown interior of the epiphallus (Fig. 52a). The epiphallus is bands overlaying a tan background streaked with relatively long, stout, and nearly uniform in diameter numerous transverse white growth wrinkles. The sharply throughout its length. It is 2.2-2.5 times the length ofthe defined spiral bands on the lighter streaked background penis. Itbears a very short flagellum that ends in a slender give this species a very elegant appearance. fhunent. Thevery short flagellum is about one-third the Shell (Figs. 48-51).-Adults 38-41 mm in width, combined length of the penis+epiphallus. The vas depressed-helicoid in shape, 0.78-0.83 times as high as deferens is slightly enlarged near the epiphallus, but wide. Thin shiny. Color pattern consisting of three otherwise is nearly uniformly narrow throughout its sharply defined chocolate-brown bands on a chalky-white length. The vagina is elongate and slender. It lacks dart ground color which becomes increasingly tan toward sacs, dart glands, and dart bulbs. The apex ofthe vagina aperture. Ground color interrupted by numerous is slightly enlarged below the uterus. The spermathecal transverse, narrow, white-streaked growth wrinkles th,t duct is short and stout, and lacks an appendix. It is donotinterrupt spiral bands. Spiral bands arenotuniform appressed against the middle of the uterus and is 0.48- in width; middle band usually the widest. Peristome light 0.82 times the length of the uterus. The spermathecal tan. Interior of aperture with a light brown translucent bulb is elongate-elliptical in shape. wash showing external bands. Apex low, obtuse, f6rming Type Locality.-NUEVO LEON, Cafion de an angle of 104-115°. Suture moderately impressed, Garcia, 7 lan northeast of Villa de Garcia (25°50.5' N, descending below theperiphery onlhe last quarter whorl. 100°31.5' W), 950 m alt. HOLOTYPE: UF 39938; Umbilicus rimate in adults. Whorls 4.1-4.3, rapidly collected June 21, 1982, by Fred G. Thompson and expanding lateral expansion rate 0.65-0.67. Last quarter Charlotte M. Porter. THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 69 \. *i 48 49 - ¥4 7 -y'lli~.:. \ 63- 4 .,5 'r * Figs. 48-51 . Humboldtiana porterae new species, holotype (UF 39938). 70 BULLEnN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) agi agl utr52a pro spt C spt spd~ fig -*.2 utr utr a vd fig epi epi pen vag 1 : " epi b -J atr vd 1 vag 10 mm f pen b --# 53 54 1,1 10 mm 5 mm Figs. 52-54. Humboldtianaporterae new species. Reproductive anatomy of two topotypes (UF 271399). Fig. 52: complete reproductive system. Fig. 52a: transverse section through epiphallus at point a in Fig. 52. Fig. 53: interior of penis and epiphallus between points a and b in Fig. 52. Fig. 54: reproductive system of a second topotype. THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 71 Measurements in mm ofthe Humboldtiana porterae n. aligned transverse to the incremental growthstriations. sp. holotype (UF 39938) and four mature paratypes (UF Inner wall of aperture rippled by external sculpture. 251528, UF 271005) are as follows. Spermathecal duct slender, longer than the combined length of the uterus+vagina; spermathecal duct with an Specimen Height Width ApH ApW Whorls appendix. Vagina bearing four dart glands, four dart sacs, and two dart bulbs on each side of the dart sacs; dartholotype 32.0 39.8 26.5 27.2 4.3 bulbs exposed. Penis asymmetrical; bearing a large verge paratype 32.0 41.0 27.7 28.1 4.1 internally. Flagellum very long, about eight times the paratype 31.0 38.1 27.2 26.1 4.2 combined length ofthe penis+epiphallus. paratype 33.5 41.0 27.0 27.2 4.3 The depressed shell, the pustular sculpture arranged paratype 32.0 38.6 26.1 34.5 4.1 into short segments transverse to the growth striations, the asymmetrical bulbous penis, the length of the flagellum, and the exposed dart-bulbs at the base of the dart sacs are unique features among known Humboldtiana. In the subgenus Humboldtiana dart-bulbs PARATYPES: UF 271005 (2 mature and 4 are deeply imbedded in the vaginal wall and are not immature shells); UF 271008 (1 immature), ITCVZ 8016 superficially visible. Bunnya Baker, 1942 isthe only other (2 mature shells), same data as the holotype; UF 251528 genus besides Humboldnana in the Humboldtianidae. It (7), collected December15, 1995, by Fred G. Thompson; is a slug-like snail with a vestigial shell imbedded in the UF 251528(3), collected February 1, 1999, by Greg R mantle. Like Polyomphala, it too haq two superficially Brewer. Non-paratypic topotypes: UF 271399 (2, visible dart-bulbs associated with each dart-sac. anatomical specimens, shell crushed), collected February Distribution.-Known only from the Sierra 1, 1999, by G. R Brewer. Encantada and the adjacent Sierra Santa Rosa in northern The type locality is alongthe north face ofthe south Coahuila. rim of the canyon, on a nearly vertical limestone cliff Etymology.-The subgeneric name Polyomphala and is on the west side ofthe pass into the canyon. The (f.) is from the Greek *olu, meaning many, and xeric vegetation consists ofclusters ofAgave lechugilla, owl,al< a knob or boss, and alludes to the coarse, A. bractiosa, Opuntia sp., Dasylyrion sp., Sedum sp., nodular shell sculpture characteristic ofthis subgenus Acacia sp., and a bunch-grass. Live snails appear to be highly dependent on Agave bractiosa for coverage. This Humboldtiana oreina new speciessmall clumping Agave was growing out of the north- (Figs. 55-59, 61-64, 68)facing vertical limestone cliffs. Live specimens were attached to limestone under A. bractiosa or in rock Diagnosis.-A moderately large, nearly planular shell crevices. We did not find Humboldtiana porterae with rugose sculpture that is arranged into elongate associated with other vegetation. pustular ridge-segments along the periphery. The Distribution.-This species is known only from segments are aligned transverse to the incremental growth the type locality. striations. The sculpture becomes beaded near the suture Etymology.-We take pleasure in naming this (Fig. 68), and below the periphery. The base ofthe shell species after Charlotte M. Porter, Curator of History of is smooth. The umbilical perforation is only partially Science, Florida Museum of Natural History, who covered by the reflected columellar lip (Fig. 58). The assisted in collecting the type specimens. shell has three brown bands that are disrupted and obscured by the white pustular sculpture and by white growth wrinkles. The reproductive system isPo/yomphala new subgenus characterized in the diagnosis for the subgenus~pe species.-Humboldtiana oreina new species Polyomphala. The shell is discoidal in shape; moderately thin; color Shell (Figs. 55-59).-Shell moderately large, about pattern three-banded, obscured by a diaphanous wash in 36-41 mm in width; nearly planular, 0.53-0.60 times as thepedostracum. Sculptum consisting of sparse large high as wide; apex very low and forming an obtuse angle nodular tubercles that are arranged in short segments of 136-148°. Shell with 3 . 9-4 . 3 rapidly expanding whorls; 72 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) «> 55 57 58 V 59 60 Figs. 55-60. Shells of Humboldtiana. Figs. 55-59: Humboldtiana oreina new species. Figs. 55-58. holotype (UF 271518) Fig. 59: paratype (UF 244910). Fig. 60: Humboldtianaplana Metcalf and Riskind, 1976, holotype (DMNH 106681). THOMPSON & BREWER: Landsnails of the genus Humboldtiana from northern Mexico 73 lateral expansion rate 0.59-0.63; last whorl flattened epiphallus. Penis very short and bulbous; inner wall of above and forming an obtuse angle at the periphery. penis with three thick longitudinal feshy pilasters, one Embryonic whorls 1.3, protruding. Last quarter of body or more of which is enlarged, causing the penis to be whorl descending below lower band (Fig. 55). Suture acentric on one side (Fig, 63). Interior of penis with a weakly impressed. Color pattern consisting of three large verge that extends nearly to base of penis lumen diffuse brown bands on a light gray background above (Fig. 62). End of verge with a complexly twisted collar and laterally; base with streaks and patches ofdiaphanous and lobes. Epiphallus short and stout enlarged near penis, light brown near umbilicus and peristome. Bands lined internally with four longitudinal folds (Fig. 62b). obscured by the white pustular ridge-segments and Flagellum very long and slender, about eight times incremental white growth wrinkles. Dorsal band about combined length of penis+epiphallus; Ragellum lined as wide as its distance from suture. Middle band lying internally with four feshy columns (Fig. 62a). Vas just above the periphery, and is the least clearly defined. deferens uniformly slender. Vagina relatively long and Lower band located just below periphery, narrow and conical compared to most other Humboldtiana, bearing tending to be the most sharply defined of the bands. four equal-sized dart-sacs, each ofwhich has two distinct Peristome whitish. Interior of aperture light magenta- dart-bulbs at its base (Fig. 64). Four dart-glands brown vaguely showing exterior bands, and may appear coalesced to form a ring around vagina immediately reticulated due to external pustular sculpture; parietal above dart-sacs. Spermatheca small, elongate-bulbous; callus opaque white. Embryonic whorls smooth but with impressed against upper end of prostate-uterus at base fine radial striations along the suture ofthe last quarter ofalbumen gland (Fig. 61). Spermathecal duct very long, whorl (Fig. 68). Following whorls sculptured with coarse diverging from uterus just above dart-glands; bearing a white granular pustules, which on last whorl form short appendix which originat6s from spermathecal duct numerous parallel pustular ridge-segments that are about 0.8 of distance from base of duct (Fig. 63); aligned transverse to the incremental growth striations appendix about as long as distal end of the (Fig. 59); pustular segments strongest along periphery duct+spermathecal bulb. Albumen gland with a long Dorsally they become smaller scattered beads near the groove on its ventral surface where it overlays intestine. suture. Below the periphery they are reduced in size to Talon exposed along axial edge of groove. Carrefour scattered granules; base ofshell smooth. Aperture oblong- partially exposed at base of hermaphroditic duct. elliptical in shape; 0.86-0.88 times as high as wide. Ovotestis forming a large tri-lobed cluster of acini in Peristome reflected along outer lip, becoming narrow third whorl of digestive gland (Fig. 61). along base, and thenexpanding along columellar margin topartially occlude umbilicus. Plane ofaperture lying at Measurements in mm of the Humboldtiana oreina n.49-59° to shell axis. sp. holotype (UF 271518) and of five adult paratypesAnatomy (Figs. 61-64).-The following descrip- (UF 244190) selected to show variation are as follows.tion is based on two topotypic specimens (UF 244911). Measurements for the holotype of H. plana (DMNHThe shells of both were crushed while extracting the 106681) are also included. Exact measurements for H.bodies. External features. The head-foot is brown, plana are estimated because the last (quarter) whorl isbecoming lighter-colored on the nape and darker fractured.posteriorly to the end of the tail. The mantle collar is bright orange. The mantle is dark gray. The tail is dorso- ventrally flattened and is covered by very fine granular Specimen Height Width ApH ApW Whorls pustules, which become enlarged along the sides of the holotype 21.0 39.8 23.0 26.0 3.9 foot, and change into large feshy pebbles on the upper sides and nape. The kidney is elongate and tapers to a paratype 24.0 40.0 22.5 26.0 4.2 narrow rounded point anteriorly. The aorta branches paratype 24.2 40.5 23.0 26.0 4.3 profuselytothe mantle collar. Reproductive system (Figs. paratype 20.2 38.1 21.8 25.0 4.0 61-64, Table 1). Genital atrium very short, nearly paratype 21.0 37.0 20.1 22.8 4.2 indistinct. Penis retractor muscle very slender, originating paratype 20.0 36.0 21.0 24.0 4.0 on center of inner wall of lung at edge of mantle collar; H. plana 23 . 8 42. 5 20. 1 26 .0 4 .0 inserting on and forming a sheath surrounding base of 74 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 43(2) fig talon ~ =Ur=- utr spt ds)- 1 *--~---C|54 Pr ds2 - :A ft. ap- dsi ovotestis XY utr T /1 r vag/~ · I i d52 :~jie- 64 1 1,1,11 5 mmvd vag 1 mm 61 agl spd 10 mm .f 1 , 62a -. -fig 11 f-- epi apputr Ye fig vrg ) spt atr ds2 -b vag ) uk vd 62 at r \ Sr-*44 5 mm 63 , , ,,,,, 10 mm Figs. 61-64. Humboldtiana oreina new species.. Fig. 61: reproductive anatomy of a topotypic specimen (UF 244911). Fig. 62: Lower male genital system of Fig. 61 showing interior of penis. Fig. 62a: transverse section through flagellum at point a in Fig. 62. Fig. 62b: transverse section through epiphallus at point b in Fig. 62. Fig. 63: reproductive system of a second topotypic specimen. Fig. 64: enlargement ofvagina in Fig. 63 showing details of dart apparatus. THOMPSON & BREWER: Landsnails Of the ge#us H#»!boldtigna from nortliern Mexico 75 Tablet.- Humboldtiana. Comparative,measurements of the teproduative,ajElatamy of. -1-2) H. bicideta n. sp. (UP 268291, UF 268290),(1, holot*e b 2, a +Wntracted paratype);, 3-4)H. edesmatn. sp. NF'27,1007, 2 ~ar*typts); 5).H. eddsina Ii. *. (UP 268317.);,T*ky,j'H. potfer#e n,,sp. (UF'*71399),jand 8) H.*reina n: [sp: (UFL2449.jl). Iiegeud: ER = p~istrefrRefpr mgcle;'PN='penis,.EP = epiphallus, FL=flagellum, AT = atrium, VG = vagina, FV = free vaginbtlr= uterus, SD = *6rmathd.diicti SB.9,base Ofs#ar-matlitcal'duteSP -re#dof**rinath*al,duct, AP = appendix. The leng®of tlk¢ sperm*b*gal ductipol»des -Be sp~smat~~ Thtend ofithe,spermathecal duct includes the spermathecal bulb and that segment of the duet abbve,thed#pe~idi*. Thb ba,66£ thei*ermath-e~ duettincludds that segment below the appadi*. Spec. no. PR PN EP FL AT VG FV UT SD SB SP AP H. bicincta n. sp. (UF 26891, UF 268290) 1 4 4 12 45 3 13 8 31 82 - 4 - 2561342---42775232 H. edesma n. sp. (UF 271007) 3 1.5 4 7 34 1 7 2 33 43 34 9 42 4 2 4 4 48 1 6 3 23 36 26 12 43 H. edesma n. sp. (UF 268317) 5 12 7 11 61 3 12 5 36 31 21 10 39 H. porterae n. sp. (UF271399) 6 10 12 27 13 4 14 41 34 7 7 12 29 13 3 10 36 16 H. oreina n. sp. (UF 244911) 8 15 5 4 72 1 11 0 36 87 71 16 17 Distribution:-Known only fromthetype locality. Elymology.-The species name oreina is fromthe Type locality.-COAHUILA: Sierra La Greek opsiva, meaning a mountaineer and alludes to Encantada, La Ventana Cahon de Boquillg 271an WNW the snails habitat. ofHacien(la La Babia, 91 km NW ofMelchor Musquiz , (28°39.1' N, 102°19.5' W), 1150 m. HOLOTYPE: UF Humboldtianaplana Metcalf and Riskind, 1976271,518:collected by F. G. Thompson~ Scott Ogden, Val (Figs. 60,65-67)Roessling, and Grady B. Taylor 12 eSeptember, 1995. Humboldtianaplana Metcalf'and Riskind, 1976; PARATYPES: UF 244910 (10), UF'268907/ (6)6 ITCVZ Nautilus 90:99-100; Figs. 1-3 (shell) 8018 (5). Ot-her specimeos: UF 244911 0 topotypic preserved specimens). This species is 1(nownonly from the holotype shell which Remarks.-Ift Humbdldtiana oretha the 16ng is broken along the last,quarter whorl (Figs. 60,65-67). spermathedal duct, the long flagellum, and the bulbous Its measurements are, given with' those of the previous penis' with its highly modified elongate verge comprise species. The shell is Slightly larger than that of differences that are unique within Humboldtiana. Humboldtiana oreina, it is mote depressed (Figs. 65, However, these peduliaritids of the spermathecal duct, 56, respectively) and the umbilical perforation is more the flagellum, and tlie penis are diff6rences of degree narrowly rimate because of the reflected columellar lip and, may be significant. only as :species critpria. (Fig. 66). As is typical.for the subgenus Polyomphala, Humbold#ana plano Metcalf and Riskind, 1976 isthe the sculpture is dominated by lai* n6dular tybercles only other species assigned:to Polyomph