identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03AA87B0FFDAFFA9FF581751FB92FDB9.text	03AA87B0FFDAFFA9FF581751FB92FDB9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Nesophontidae Anthony 1916	<div><p>Family Nesophontidae Anthony, 1916</p> <p>REMARKS: Nesophontidae is a family of solenodonotan lipotyphlans, the only known representatives of which are species of the late Quaternary Antillean genus Nesophontes (Whidden and Asher, 2001; Brace et al., 2016). The affinities of Nesophontes have been diversely treated in the century since the discovery of the first member of the genus (Anthony, 1916). Most authors have favored a close relationship to soricoids, usually on the basis of an apparently shared pattern of molar dilambdodonty (e.g., Saban, 1954; Van Valen, 1967), but others have discussed whether Nesophontes might instead be the sister taxon of Solenodon, another lipotyphlan restricted to the Caribbean but exhibiting molars that are morphologically zalambdodont (e.g., Gregory, 1920; McDowell, 1958; Asher and Sánchez-Villagra, 2005). This dispute was recently settled in favor of the latter hypothesis, on the basis of ancient DNA evidence collected from a 750 year old N. paramicrus specimen from Hispaniola (Brace et al., 2016). The closest known relatives of Nesophontes and Solenodon are probably Eocene and Oligocene insectivores in the family Geolabididae from North America (Simpson, 1956; McDowell, 1958; MacFadden, 1980; Lillegraven et al., 1981; Morgan and Woods, 1986). Molecular dating indicates that the split between Solenodontidae and Nesophontidae occurred around 40 Ma. In view of their distinctiveness and antiquity within Lipotyphla, these families were recently placed in their own suborder, Solenodonota, by Brace et al. (2016). Their mutual divergence may have occurred on the North American mainland, in which case two invasions of the Greater Antilles by solenodonotans must be inferred. Alternatively, divergence may have occurred after initial colonization of the island arc by a joint ancestor, requiring only a single invasion (Simpson, 1956; McDowell, 1958; MacFadden, 1980; Lillegraven et al., 1981; Morgan and Woods, 1986; Iturralde- Vinent and MacPhee, 1999). Nesophontes and Solenodon are the only two endemic genera of land mammals in the West Indies that are clearly of Nearctic origin. All other Antillean terrestrial mammals have explicitly South American or more general Neotropical affinities (Morgan and Woods, 1986).</p> <p>The type species and largest member of the genus, Nesophontes edithae, was originally described from Puerto Rican cave deposits (Anthony, 1916). It is also known from Amerindian archaeological sites on Vieques and the Virgin Islands (Morgan and Woods, 1986; Quitmyer, 2003). Five species of Nesophontes have been described from Cuba: N. micrus Allen, 1917, N. longirostris Anthony, 1919, N. major Arredondo, 1970, N. submicrus Arredondo, 1970, and N. superstes Fischer, 1977. Three species of Nesophontes have also been described from Hispaniola by Miller (1929): N. hypomicrus, N. paramicrus, and N. zamicrus. Nesophontes is unknown from Jamaica, Bahamas, or Lesser Antilles.</p> <p>Considerable size variation exists within Puerto Rican samples of N. edithae, which was initially interpreted by Anthony (1916) as representing sexual dimorphism. As large-scale sexual dimorphism is unknown in other recent lipotyphlans, this variation was used as part of the original justification for erecting a new family to accommodate the species (Anthony, 1916). However, size variation in N. edithae has been subsequently interpreted as more likely to reflect allochronic plasticity, with specimens of varying sizes probably originating from different depositional horizons across the Late Quaternary (Choate and Birney, 1968; McFarlane, 1999). Condis Fernández et al. (2005) analyzed metrical and morphological variation in Cuban Nesophontes, and concluded that only two species could be supported, N. major and N. micrus, the latter incorporating all of the other nominal taxa. Importantly, these authors showed that size is a particularly labile character within Cuban Nesophontes and is therefore not a dependable basis for erecting species boundaries, especially when considered in isolation. Silva Taboada et al. (2007) went even further, suggesting that all previously diagnosed Cuban species of Nesophontes are in fact morphs of N. micrus. However, their subordination of all nominal taxa within a single species was made without any commentary, and they did not review the diagnostic value of the characters cited by Condis Fernández et al. (2005) for distinguishing N. micrus and N. major. The Hispaniolan species have not been subjected to revision in recent years, and the scale of intraspecific variation in Cuban Nesophontes raises the question whether these three species are truly distinct, or instead just size morphs of one or two variable species. Since this issue cannot be settled here, for the purposes of this study we retain Miller’s taxa as valid.</p> <p>The previous existence of Nesophontes on Cayman Brac and Grand Cayman has been repeatedly noted in the literature (e.g., Patton, 1966; Varona, 1974; Morgan et al., 1980; Steadman and Morgan, 1985; Morgan and Woods, 1986; Morgan, 1994a), but no formal systematic determination of its status has ever been undertaken. Regrettably, the Cayman material is too fragmentary or otherwise damaged to make a morphometric treatment worthwhile.</p> </div>	https://treatment.plazi.org/id/03AA87B0FFDAFFA9FF581751FB92FDB9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Morgan, Gary S.;Macphee, Ross D. E.;Woods, Roseina;Turvey, Samuel T.	Morgan, Gary S., Macphee, Ross D. E., Woods, Roseina, Turvey, Samuel T. (2019): Late Quaternary Fossil Mammals From The Cayman Islands, West Indies. Bulletin of the American Museum of Natural History 2019 (428): 1-81, DOI: 10.1206/0003-0090.428.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2019/issue-428/0003-0090.428.1.1/Late-Quaternary-Fossil-Mammals-from-the-Cayman-Islands-West-Indies/10.1206/0003-0090.428.1.1.full
03AA87B0FFDBFFAAFD2410D7FCA5FD3D.text	03AA87B0FFDBFFAAFD2410D7FCA5FD3D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Nesophontes hemicingulus Morgan & Macphee & Woods & Turvey 2019	<div><p>Nesophontes hemicingulus, new species</p> <p>Figures 3–9</p> <p>HOLOTYPE: UF 23295, partial skull lacking neurocranium (fig. 3); C1–M3 present on right side, C1–M2 on left side, with partial alveoli for I2–I3. Unfortunately, the skull broke along its long axis after collection, although little bone has been lost from complementary edges. All the cranial specimens from the Cayman Islands are damaged to a greater or lesser degree, and dentitions tend to be incomplete and much worn. UF 23295 was chosen as the holotype because its teeth are on average less worn than those in comparable specimens and all cheekteeth are preserved (albeit on one side only).</p> <p>TYPE LOCALITY: Patton’s Fissure, near Spot Bay on the northern coast of Cayman Brac. The holotype was recovered from layer 5 (80–100 cm below the surface), which has been radiocarbon dated on the basis of land snail shell carbonate to 11,180 ± 105 14 C yr BP (see Radiocarbon Dating).</p> <p>ETYMOLOGY: Frp, Latin hemi-, “half,” and cingulus, “belt,” in reference to absence of the precingulum on all upper molars.</p> <p>AGE: Late Pleistocene-Holocene (see Radiocarbon Dating).</p> <p>DISTRIBUTION: Known only from Cayman Brac and Grand Cayman. This species is the only member of the genus known outside the Greater Antilles and their satellite islands.</p> <p>REFERRED SPECIMENS: Cayman Brac: Patton’s Fissure: partial skulls, UF 23258, 23264, 23277, 23279, 23293–23301, 23332–23337, 23360- 23363, 23393, 23394, 23407; mandibles, UF 23241, 23242, 23245, 23251–23256, 23259, 23265–23269, 23280–23285, 23311–23313, 23324–23326, 23343, 23347–23359, 23364, 23365, 23367–23378, 23383, 23389, 23397, 23398, 23400, 23404, 23408–23417, 23421, 23426–23432, 23436, 23437, 23448–23450 (also includes a large sample of postcranial material not listed here). Pollard Bay Cave, Shearwater Cave 2. Grand Cayman: Dolphin Cave: UF 172845, right mandible with c1–m3; UF 172846, right mandible with c1–m3; UF 172863, right mandible with p2–m3; UF 172908, left mandible with p2, p4–m3; UF 172909, left mandible with c1, p2, m1–m3; UF 172910, left mandible with m1–m3; UF 172926, right mandible with m1– m2; UF 172939, partial skull lacking braincase, with right C1–M2 and left P2-M2; UF 172940, left mandible with p2, p4–m3; UF 172950, left mandible m1–m2. Barn Owl Cave: UF 23242, right mandible with p2–m3. Bodden Cave: UF 23241, right mandible with p4–m3. Furtherland Farms: UF 172803, left mandible with p2, p4– m3. Old Man Cave: UF 23245, right mandible with c1, p2, m1; UF 23246, right mandible with p2, p4–m3; UF 23247, edentulous right mandible; UF 23248, partial edentulous right mandible; UF 23249, right mandible with p2, p 4m 3; UF 23250, left mandible with p2, p4–m3; UF 23251, left mandible with p4–m3.</p> <p>DIAGNOSIS: Within Nesophontes, N. hemicingulus expresses unique reductions in cingulum/ ectocingulid features on upper/lower molars. It can be distinguished in lacking precingula on all molars, attenuation of postcingula on M1 and M2, and uniform de-emphasis of ectocingulids on lower cheekteeth. Closest morphological similarities are to Cuban N. micrus in regard to tooth shape, dimensions, and discrete characters.</p> <p>DESCRIPTION: In the absence of good cranial remains of Cayman nesophontids, teeth are the main source of characters (figs. 3–9). In the following set of differential diagnoses, we compare N. hemicingulus to species from the major islands on which Nesophontes formerly occurred: Puerto Rico (N. edithae), Cuba (N. major, N. micrus), and Hispaniola (N. hypomicrus, N. paramicrus, N. zamicrus). Hispaniolan N. paramicrus and Cuban N. micrus are very similar, and were in fact synonymized by Varona (1974) on the basis of his comparisons and those of Patterson (1962). However, these two species differ in some characters, such as the absence of a constricted infraorbital foramen in N. micrus, and for this reason are distinguished here. Cingulum/cingulid characters, the only reliably diagnostic characters for N. hemicingulus in the current hypodigm, are treated separately.</p> </div>	https://treatment.plazi.org/id/03AA87B0FFDBFFAAFD2410D7FCA5FD3D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Morgan, Gary S.;Macphee, Ross D. E.;Woods, Roseina;Turvey, Samuel T.	Morgan, Gary S., Macphee, Ross D. E., Woods, Roseina, Turvey, Samuel T. (2019): Late Quaternary Fossil Mammals From The Cayman Islands, West Indies. Bulletin of the American Museum of Natural History 2019 (428): 1-81, DOI: 10.1206/0003-0090.428.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2019/issue-428/0003-0090.428.1.1/Late-Quaternary-Fossil-Mammals-from-the-Cayman-Islands-West-Indies/10.1206/0003-0090.428.1.1.full
03AA87B0FFCCFFB8FF561731FBB0FE7B.text	03AA87B0FFCCFFB8FF561731FBB0FE7B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Capromyinae Smith 1842	<div><p>Subfamily Capromyinae Smith, 1842</p> <p>REMARKS: The Capromyidae is a diverse group of rodents of fairly large body size (~0.5– 4 kg) endemic to the West Indies. Capromyids are most closely related to the Echimyidae (spiny rats) and Myocastoridae (coypu) among mainland Neotropical hystricognath rodents. These three families have been referred to the hystricognath superfamily Octodontoidea, which also includes the southern South American families Abrocomidae, Ctenomyidae, and Octodontidae (Woods and Kilpatrick, 2005). Molecular analysis by Leite and Patton (2002) suggested placement of the Capromyidae as a subfamily (Capromyinae) within the Echimyidae, although these authors sampled only a single species of capromyid, Capromys pilorides. More recent molecular phylogenies, which have included all extant genera of capromyids, have reached different conclusions over whether the Capromyidae could be distinguished as a family separate from the Echimyidae and Myocastoridae (Kilpatrick et al., 2012; Fabre et al., 2014, 2017). Pending further molecular analyses, we tentatively recognize the Capromyidae as an endemic West Indian family, with close phylogenetic relationships to both the Echimyidae and Myocastoridae.</p> <p>Current taxonomy recognizes five extant genera in the Capromyidae, separated into two subfamilies: Plagiodontia (Hispaniola) in the subfamily Plagiodontinae; and Capromys, Mesocapromys, and Mysateles (Cuba) and Geo- capromys (Jamaica, Bahamas) in the subfamily Capromyinae (Morgan, 1985; Woods and Kilpatrick, 2005; Silva Taboada et al., 2007; Borroto-Páez et al., 2012; Dávalos and Turvey, 2012). A recently extinct species of Geocapromys is known from Little Swan Island in the western Caribbean, and the Capromyidae also has an extensive late Quaternary history in the Greater Antilles and Bahamas as well as the Cayman Islands. In addition to material from the Cayman Islands described here, the capromyine Quaternary fossil record includes Capromys, Geocapromys, Mesocapromys, and Mysateles from Cuba, and Geocapromys from Jamaica and many Bahamian islands where this genus no longer occurs (Morgan, 1985, 1989b, 1994a; Silva Taboada et al., 2007; Borroto- Páez et al., 2012; Dávalos and Turvey, 2012). The Quaternary record also documents several extinct capromyids from Hispaniola, divided among three subfamilies: two additional species in the extant genus Plagiodontia, and additional species in the extinct genera Hyperplagiodontia and Rhizoplagiodontia in the Plagiodontinae; and two other extinct genera, each referred to an extinct subfamily, Hexolobodon (Hexolobodontinae) and Isolobodon (Isolobodontinae) (Woods and Kilpatrick, 2005; Borroto-Páez et al., 2012; Dávalos and Turvey, 2012; Hansford et al., 2012). Isolobodon was originally described from Puerto Rico (I. portoricensis) but occurs there and on Vieques and the Virgin Islands only in archaeological sites, apparently because it was translocated from Hispaniola by Amerindian peoples. Quaternary fossil sites from Puerto Rico lack native capromyids but instead contain the large hystricognath Elasmodontomys, which has been referred to the Heptaxodontidae (“giant hutias”). Other heptaxodontids are also known in the Greater Antilles from Hispaniola and Jamaica, where they cooccurred with capromyids (Woods and Kilpatrick, 2005; MacPhee, 2011). The presence of an early Miocene capromyid from Cuba, the extinct genus Zazamys from the Domo de Zaza fauna, establishes a long history of this family in the Greater Antilles (MacPhee and Iturralde- Vinent, 1995).</p> <p>TABLE 2</p> <p>Comparative measurements (in mm) of the mandible and lower dentition of Nesophontes hemicingulus from Cayman Brac and Grand Cayman, and selected species of Nesophontes from Cuba, Hispaniola, and Puerto Rico. Abbreviations: M, mean; R, range; and N, sample size.</p> <p>The genus Capromys has received considerable attention in the past few decades, much of which has focused on the description of new species, both living and extinct. Until the mid 1970s, almost all Cuban species in the family Capromyidae were referred to Capromys (Varona, 1974). During the 1970s, a proliferation of new generic and subgeneric names was proposed for Cuban capromyids formerly included in Capromys (e.g., Kratochvil et al., 1978; Varona and Arredondo, 1979). The systematic review by Silva Taboada et al. (2007) synonymized many of the previously described genera, subgenera, and species of Cuban capromyids, and recognized five genera of Capromyidae in Cuba, Capromys, Geocapromys, Macrocapromys, Mesocapromys, and Mysateles, one of which (the extinct genus Macrocapromys) has since been synonymized with Capromys (Borroto-Páez et al., 2012). The Quaternary and extant Cuban Capromyidae includes four genera and 11 described species (although see further discussion below): three species of Capromys (one living, C. pilorides, and two extinct, C. acevedo and C. latus); one extinct species of Geocapromys (G. columbianus); six species of Mesocapromys (five living, M. angelcabrerai, M. auritus, M. melanurus, M. nanus, and M. sanfelipensis, and one extinct, M. kraglievichi); and one living species of Mysateles (M. prehensilis).</p> </div>	https://treatment.plazi.org/id/03AA87B0FFCCFFB8FF561731FBB0FE7B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Morgan, Gary S.;Macphee, Ross D. E.;Woods, Roseina;Turvey, Samuel T.	Morgan, Gary S., Macphee, Ross D. E., Woods, Roseina, Turvey, Samuel T. (2019): Late Quaternary Fossil Mammals From The Cayman Islands, West Indies. Bulletin of the American Museum of Natural History 2019 (428): 1-81, DOI: 10.1206/0003-0090.428.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2019/issue-428/0003-0090.428.1.1/Late-Quaternary-Fossil-Mammals-from-the-Cayman-Islands-West-Indies/10.1206/0003-0090.428.1.1.full
03AA87B0FFCAFF8FFD3410CAFEAFFBAE.text	03AA87B0FFCAFF8FFD3410CAFEAFFBAE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Capromys pilorides subsp. lewisi Morgan & Macphee & Woods & Turvey 2019	<div><p>Capromys pilorides lewisi, new subspecies</p> <p>Figures 10–12, 14–17</p> <p>HOLOTYPE: NHM (Mammalogy) 71.1558/ M15705, complete skull with left M1 and right PM4 –M3 (figs. 10A, 12A).</p> <p>TYPE LOCALITY: Stake Bay Cave, 0.25 miles west of Stake Bay, Cayman Brac.</p> <p>ETYMOLOGY: Named in honor of the late C. Bernard Lewis, former curator and director of the Institute of Jamaica and a member of the 1938 Oxford University Cayman Islands Biological Expedition, who collected the holotype skull of this new subspecies.</p> <p>AGE: Late Pleistocene-Holocene (see Radiocarbon Dating).</p> <p>DISTRIBUTION: Known only from Cayman Brac, Little Cayman, and Grand Cayman. The only representative of the genus Capromys known outside Cuba and its satellite islands.</p> <p>REFERRED SPECIMENS: Cayman Brac: Stake Bay Cave: NHM (Mammalogy) 71.1558/ M15704, NHM (Palaeontology) M15733, M15734, M42027, M42028, skulls; UF 61291, nearly complete skull lacking nasals and right jugal, with left M1 and right M1–M2, and associated mandibles; NHM (Mammalogy) 71.1558/ M15706, 71.1558/ M15707, 71.1558/unnumbered, NHM (Palaeontology) M15735, M15736, M15737, M15738, M15739, M15740, M42029, mandibles. Blackie’s Cave: UF 172782, partial skull. Fig Tree Cave: UF 172759, 172760, edentulous mandibles. Hutia Cave: UF 172751, nearly complete edentulous skull lacking jugals, with associated edentulous mandibles, innominates, and tibia; UF 172752, 172753, 172758, partial skulls; UF 172755, 172763, edentulous mandibles. Patton’s Fissure: UF 18763, 21406, partial skulls; UF 21405, palate; UF 18647, 18650, 21385, mandibles with p4–m3; UF 18557, 18617, mandibles with p4–m2; UF 21395, mandible with p4; UF 21385, 22850, mandibles with m1–m3; UF 18542, 18588, 18597, 18618, 18619, 18670, 18671, 21351, 21355, 21395, 22853, edentulous mandibles. Peter Cave: UF 22855, partial skull; UF 61291, nearly complete skull with right M1–M2 and left M1; UF 22854, edentulous mandible. Pollard Bay Cave: UF 22886, palate; UF 18557, mandible with i, p4–m2. Shearwater Cave 2: UF 172792, 2 cervical vertebrae. Spot Bay Cave: UF 172764, skull; UF 172767, mandible with p4– m3; UF 172766, mandible with m2. Grand Cayman: Agouti Cave: UF 172870, partial skull; UF 172871, mandible with p4–m3; UF 172872, mandible with p4–m2; UF 172875, mandible with i, m2–m3; UF 172873, 172874, 172876– 172882, edentulous mandibles. Barn Owl Cave: UF 22875, 22877, 22878, palates; UF 22872, mandible with p4–m3; UF 22871, mandible with p4–m2; UF 22876, mandible with p4–m1; UF 22873, 22874, 22879, edentulous mandibles. Big Ear Cave: UF 162850, mandible with i1, m1. Chisholm Cow Well: UF 172821, premaxilla with I1; 172820, mandible with p4; UF 172836, edentulous mandible. Crab Cave: UF 22865, maxilla with P4; UF 22858, mandible with p4– m3; UF 22857, 22860, 22862, 22863, edentulous mandibles. Crocodile Canal: UF 61147, i1; UF 61149, 61150, 61154, femora. Dolphin Cave: UF 172920, nearly complete skull with left and right M1; UF 172921, edentulous partial skull; UF 172911, 172912, 172848, 172862, 172927– 172929, 411275–411277, edentulous mandibles. Furtherland Farms Cow Well: UF 172798, 172799, humeri; UF 172801, 172802, 172814, innominates; UF 172800, 172815, tibia. Miller’s Cave: UF 172089, partial skull; UF 172890, maxilla with P4–M1; UF 172896, mandible with i1, p4, m3; UF 172897, mandible with p4–m2; UF 172898, mandible with p4–m1; UF 172891, mandible with m1–m2; UF 172892–172895, 172899–172902, edentulous mandibles. Old Man Cave: UF 22880, nearly complete skull lacking left premaxilla, portion of left maxilla, both jugals, and all teeth; UF 22881, mandible with p4, m2. Queen Elizabeth II Botanic Park: UF 172947, mandible with p4–m3. Tadarida Cave: UF 61293, partial skull; UF 172837, mandible with m2; UF 172840, mandible with i1. Little Cayman: Agave Cave: UF 172794, humerus; UF 172793, femur. Franz’s Shelter: UF 172795, edentulous maxilla; UF 172796, 172797, edentulous mandibles. Weary Hill Cave: UF Environmental Archaeology Collection (uncat.), edentulous mandible. [This list of Referred Material does not include all Capromys fossils from the Cayman Islands, only the most complete cranial and mandibular specimens. Postcranial specimens are listed if they represent the only records of Capromys from a particular site. There is also a large sample of isolated teeth and postcranial elements not listed here.]</p> <p>DIAGNOSIS: Differs from all described Cuban subspecies of Capromys pilorides in: smaller overall size; shorter maxillary and mandibular toothrows, resulting from smaller, lowercrowned cheekteeth with reduced amount of cement on anterior and posterior edges; maxillary toothrows strongly convergent anteriorly and nearly parallel from the anterior margin of P4 to the border between M1 and M2; narrow, anteriorly directed dorsal maxillary process; smaller orbit (about half the height of infraorbital foramen), owing to strong inflation of frontals posteriorly between orbits and parietal suture (frontals greater in height than parietals), and deeper zygomatic arch ventral to orbit; more constricted internal narial opening, which is rounded rather than triangular in ventral outline; shorter and less inflated auditory bullae; and shorter mandibular symphysis with reduced posterior margin.</p> <p>DESCRIPTION: One of the most characteristic features of the Cayman Capromys is the com- paratively short maxillary toothrows relative to the total length of the skull. The mandibular toothrows are also correspondingly short. The mean alveolar lengths of the upper toothrows and lower toothrows are very similar (tables 3, 4). Although the majority of partial skulls, isolated maxillae, and mandibles are edentulous, there are specimens that preserve partial or complete upper and lower dentitions (figs. 10–11, 16–17). In these specimens, the teeth are somewhat compressed anteroposteriorly, and they have a comparatively thin layer of cement on the anterior and posterior margins of the individual teeth. We were only able to take dental measurements of the upper cheekteeth (P4 and M1–M3) on a few specimens, including the holotype, but were able to measure a larger sample of lower cheekteeth (tables 3, 4).</p> <p>The upper toothrows are strongly convergent anteriorly, with an average palatal width of only 3.3 mm opposite P4 (table 3). The convergence of the upper toothrows continues posteriorly to approximately the level of the border between M1 and M2, with the toothrows essentially parallel between P4 and M1. Posterior to M1, the toothrows begin to diverge laterally, reaching their greatest width at the posterior palatal margin. The convergence of the upper toothrows is also evident dorsal to the toothrows, as the internal narial opening is constricted both vertically and transversely, and is rounded rather than triangular in ventral outline. The root capsule for the incisor is barely visible on the external surface of the premaxilla and maxilla. The origin for the incisor root capsule is located immediately anterior to P4 and just dorsal to the maxillary root of the zygomatic. The root capsules of all four cheekteeth are visible as external swellings on the maxilla.</p> <p>The ascending or dorsal process of the maxilla is relatively thin and oriented noticeably anteriorly. The dorsoventral height of the infraorbital foramen is almost twice that of the orbit. The small size of the orbit compared to other Capromys is a result of both the deep zygomatic arch ventral to the orbit and the inflated frontals dorsal to the orbit. The zygomatic arch, composed primarily of the jugal and also the dorsal maxillary process anteriorly, is vertically deep. The jugal fossa is rather large and oriented at approximately 45° to the ventral margin of the zygomatic arch. A well-developed jugal spine projects posteroventrally from the jugal. The frontals are noticeably inflated, from the anterior edge of the orbits posteriorly to the frontoparietal suture (fig. 12). This inflation is also evident on the portion of the frontals that comprises the internal wall of the orbits, and is especially prominent ventral to the postorbital processes. Only the anterior third of the frontals is not inflated, between the anterior edge of the orbits and the nasofrontal suture. In lateral view, the frontal inflation is emphasized by noticeable depressions both anteriorly dorsal to the anterior edge of the orbits and posteriorly along the frontoparietal suture. Because of this inflation, the width of the frontals posterior to the postorbital processes is greater than their width anterior to these pro- cesses (table 3). The temporal crests are well defined and nearly meet posteriorly along the midline of the skull, although they do not form a sagittal crest. The auditory bullae are comparatively short anteroposteriorly, giving them a somewhat circular to elliptical outline. The bullae are not inflated.</p> <p>The posterior margin of the m3 alveolus in the mandible is oriented at a slight angle to the long axis of the toothrow. The mandibular symphysis is quite broad and well developed, but is truncated posteriorly with a reduced posterior margin. A thin ridge of bone extends posteriorly along the ventral margin of the mandible from the symphysis to approximately the level of m1 or m2. In side view, this thin elongate ridge gives the mandible the appearance of being deeper vertically. The mandible is relatively deep below the cheekteeth, indicating that the cheekteeth are quite hypsodont. The condyloid process is wide at its base, and the area for insertion of the masseter on the labial portion of the ramus ventral to the condyloid process is expanded in area, especially vertically.</p> <p>The upper incisors are relatively broad and highly arched. As with most other capromyids, the cheekteeth are rootless and ever growing, but are relatively small compared to other Capromys. The p4 is longer and narrower than the other cheekteeth, and has a rounded anterior margin. The nature of the enamel band along the anterior lingual margin is quite variable: some specimens have no trace of a third reentrant in the anterior position, while other specimens have a notice- able infolding of the enamel band along the anterior margin of p4, and the average condition is a slight, medially directed infolding of the enamel band along the anterior lingual margin of p4. If this anterior infolding of the enamel band is not considered a true reentrant, then there are only two lingual reentrants on p4. The anterior lingual reentrant is the best developed of the two lingual reentrants. This anterior reentrant extends medially well beyond the median long axis of the tooth, and is anterior in position with respect to the single labial reentrant. The posterior reentrant is less well developed than the anterior reentrant, and does not extend medially past the median long axis of p4. The m3 is oriented at a slight angle to the other cheekteeth in the mandible, such that in an edentulous mandible the posterior margin of the m3 alveolus is also oriented at a slight angle to the long axis of the toothrow as opposed to the other three cheekteeth, which are perpendicular to this axis. The occlusal surfaces of the upper and lower dentition are flat and horizontal.</p> <p>MORPHOMETRICS: The new subspecies displays some overlap in the observed range of most cranial measurements seen in a sample of 14 modern skulls of Cuban Capromys pilorides pilorides (tables 3, 4). The Cuban sample has a total skull length ranging from 85–106 mm (mean = 95 mm), whereas six complete or semicomplete skulls of C. pilorides lewisi range from 88–96 mm in total length (mean = 92 mm). However, there is no overlap in the alveolar length of the upper toothrows between these two samples (C. pilorides pilorides: 21–24 mm, mean = 22 mm; C. pilorides lewisi: 18–20 mm, mean = 19 mm).</p> <p>The largest available measurement series for the Cayman Capromys sample is mandibular alveolar toothrow length (Grand Cayman, n = 29; Cayman Brac, n = 13; Little Cayman, n = 2). Too few specimens are available from Little Cayman to permit statistical morphometric comparison with the other islands, but comparison between the Grand Cayman and Cayman Brac measurement series shows no statistically detectable size difference between samples from these two islands (Welch twosample t-test not assuming equal variance: Grand Cayman, mean = 19.81 mm; Cayman Brac, mean = 19.33 mm; p = 0.240). The entire available Cayman Capromys sample (mean = 19.70 mm) can therefore be pooled for analysis with Cuban Capromys samples.</p> <p>TABLE 3 Comparative measurements (in mm) of the skull and upper dentition of Capromys pilorides lewisi from the Cayman Islands (combined data for Grand Cayman and Cayman Brac), and Capromys pilorides pilorides</p> <p>from mainland Cuba. Abbreviations: M, mean; R, range; and N, sample size.</p> <p>The pooled Cayman Capromys sample has a significantly smaller mandibular alveolar toothrow length compared to a sample of C. pilorides pilorides from mainland Cuba measured from museum collections (n = 39; mean = 21.56 mm; p &lt;0.001; appendix 2), to measurement data for the type series of C. pilorides doceleguas reported in Varona (1980) (n = 7; mean = 23.23 mm; p &lt;0.001), and to measurement data for mandibular toothrow length (likely to be very close to alveolar toothrow length) for the type series of C. pilorides gundlachianus reported in Varona (1983) (n = 6; mean = 21.80 mm; p = 0.001). These represent the only unpooled measurement series available for direct statistical comparison with our Cayman Capromys sample. Borroto Páez et al. (1992) provided pooled measurement series and summary statistics for mandibular alveolar toothrow length for the other two described subspecies of C. pilorides, C. pilorides relictus (n = 18; mean = 22.30 mm) and C. pilorides ciprianoi (n = 19; mean = 22.63 mm), and reported that neither subspecies showed a statistically significant difference in this measurement from their sample of C. pilorides pilorides. We therefore interpret our Cayman Capromys sample as almost certainly also having a significantly smaller mandibular alveolar toothrow length than these two subspecies. The smaller toothrow lengths in Cayman Capromys appear to be a result of the more compressed molariform teeth (premolars and molars) with reduced cement on the anterior and posterior margins of the individual teeth.</p> <p>Among the larger species of extinct Cuban capromyids in the general size range of C. pilorides, Silva Taboada et al. (2007) recognized two species in the genus Macrocapromys, M. acevedo and M. latus, both of which have since been placed in Capromys (Borroto-Páez et al., 2012). Neither of these large species of Capromys is known from a complete skull, but measurements of the upper toothrow of C. acevedo and C. latus (range = 21–24 mm) are larger than the upper toothrow length of C. p. lewisi (range = 18–20 mm) (Silva Taboada et al., 2007).</p> <p>MORPHOLOGICAL COMPARISON WITH CUBAN CAPROMYS: In addition to size, Capromys pilorides lewisi differs from Cuban subspecies of C. pilorides in several cranial and dental characters. Cuban C. pilorides are characterized by having the infraorbital foramen considerably greater in height than the orbit compared to species of Mesocapromys and Mysateles, in which the infra- orbital foramen and orbit are similar in height. This character is even more extreme in the Cayman Capromys, in which the orbit is even smaller than in Cuban C. pilorides owing to the deeper zygomatic arch ventral to the orbit and the dorsal inflation of the frontals (figs. 12, 13).</p> <p>TABLE 4</p> <p>Comparative measurements (in mm) of the mandible and lower dentition of Capromys pilorides lewisi from the</p> <p>Cayman Islands (combined data for Grand Cayman and Cayman Brac), and Capromys pilorides pilorides from mainland Cuba. Abbreviations: M, mean; R, range; and N, sample size.</p> <p>One of the most diagnostic features of Capromys pilorides lewisi, and one in which this subspecies differs from Cuban C. pilorides, is the strong anterior convergence of the upper toothrows. The maxillary toothrows in C. pilorides lewisi are highly convergent anteriorly, and are essentially parallel between the P4s and M1s (figs. 10A, B, 11A, B). In contrast, the upper toothrows of Cuban C. pilorides are not as convergent anteriorly, diverging gradually from anterior to posterior (fig. 10C). Another characteristic feature of C. pilorides lewisi related to the convergence of the upper toothrows is the strongly constricted internal narial opening, which is smaller in both the vertical and transverse dimensions than in Cuban C. pilorides.</p> <p>The frontals are noticeably inflated in Capromys pilorides lewisi from the anterior edge of the orbits posteriorly to the frontoparietal suture, and are particularly inflated dorsal to the orbits (fig. 12). In lateral view, this frontal inflation is delineated anteriorly by a depression dorsal to the anterior margin of the orbits and posteriorly by a depression along the frontoparietal suture. In contrast, the frontals of Cuban C. pilorides typically show no dorsal inflation and are essentially flat, with a smooth transition between the frontals and parietals and no depression along the frontoparietal suture. Because of this inflation, the width of the frontals is somewhat greater in C. pilorides lewisi than in Cuban C. pilorides, even though Cuban samples are larger in most other cranial measurements (tables 3, 4). However, some variation is observed in dorsal inflation of the frontals between Cuban Capromys individuals (fig. 13). The width of the frontals posterior to the postorbital processes is greater in C. pilorides lewisi than their width anterior to these processes, the opposite of the condition in Cuban C. pilorides in which the frontals are broader anterior to the postorbital processes. The auditory bullae also differ between Cayman and Cuban Capromys; the auditory bullae in C. pilorides lewisi are rather small, anteroposteriorly short, with a rounded shape, and show no evidence of inflation, compared to Cuban C. pilorides in which the bullae are larger, longer, somewhat compressed laterally, and moderately inflated.</p> <p>GENETICS: Extraction and amplification of UF 18588 from Patton’s Fissure on Cayman Brac successfully yielded the entire mitogenome (15,908 base pairs). Maximum likelihood and Bayesian analyses of hutia mitogenome data generated congruent phylogenetic trees, showing UF 18588 as the sister taxon to the Capromys pilorides mitogenome sample available on Gen- Bank (fig. 18). The monophyly of the Cayman + Cuban Capromys clade is supported by extremely strong Bayesian approximated posterior probability values (1) and bootstrap values for maximum-likelihood analysis (1).</p> <p>TABLE 5</p> <p>Average pairwise estimates of mitogenome sequence divergence (main value) and cytochrome b sequence</p> <p>divergence (value in parentheses) between a specimen of Capromys from Cayman Brac (UF 18588)</p> <p>and GenBank data for extant hutia species.</p> <p>Sequence divergence between UF 18588 and C. pilorides is low (table 5). Estimated divergence from the mainland Cuban C. pilorides sequence used in our analyses is only 0.5% across the entire mitogenome, and only 0.6% across the entire 1140 base pair cytochrome b (cyt b) gene (table 5).</p> <p>REMARKS: The larger of the two hutia taxa present in the Quaternary record of the Cayman Islands is clearly referable to the genus Capromys on the basis of the considerably greater dorsoventral diameter of the infraorbital foramen relative to that of the orbit, the relatively thin and anteriorly oriented dorsal process of the maxilla, the presence of a small medial enamel infolding (anteroflexid) along the anterior lingual margin of p4, and its overall large size (Kratochvil et al., 1978; Díaz-Franco, 2001; Silva Taboada et al., 2007; Borroto-Páez and Mancina, 2011). Our genetic analysis also confirms genus-level assignment of the extinct Cayman population to Capromys.</p> <p>Capromys is the most abundant and widespread fossil mammal in Quaternary deposits on the Cayman Islands, and the specimens of Capromys reported from Little Cayman are the only fossil vertebrates recorded from that island (Morgan, 1994a). It was the first fossil vertebrate reported from the Cayman Islands (Moyne, 1938; Westermann, 1953), and the former occurrence of Capromys on the Cayman Islands has been widely recognized by previous authors who have reviewed fossil material from the islands without providing a formal description of the taxon (Varona, 1974; Morgan, 1977, 1994a; Steadman and Morgan, 1985; Morgan and Woods, 1986; Harvey et al., 2016). The only exception to this is Patton (1966), who mentioned only Geocapromys as occurring on Cayman Brac, but investigation of material that he collected reveals that this sample also contains Capromys.</p> <p>Except for its presence in the Quaternary record of the Cayman Islands, Capromys is restricted to Cuba and its satellite islands (Isla de la Juventud and offshore archipelagos), where only one extant described species, C. pilorides, is currently assigned to the genus. Capromys garridoi, a second named species that was described by Varona (1970) from a single individual collected from Cayo Majá, Archipiélago de los Canarreos, has been reinterpreted as a misidentified specimen of C. pilorides (Silva Taboada et al., 2007; Borroto-Páez and Mancina, 2011). Based on information provided by C.B. Lewis, Westermann (1953) and Varona (1974) interpreted Capromys fossils from Cayman Brac as conspecific with C. pilorides; other authors have conversely left the Cayman Capromys in open</p> <p>0.04</p> <p>nomenclature, or have considered it to represent a distinct but undescribed species. Cayman Capromys specimens can be differentiated from Cuban Capromys according to both qualitative morphological characters and statistical morphometric size differences. Several of these differences may represent evolutionary adaptations to the distinct ecological conditions of the Cayman Islands. The reduction in body size may be explicable on biogeographic grounds, consistent with vertebrate body size decreasing with land area on other oceanic islands (Burness et al., 2001). Compared to Cuban Capromys, the Cayman Capromys has cheekteeth that are vertically shorter (even though both taxa have rootless teeth) and with a reduced occlusal surface area, and has a reduced mandibular symphysis possibly indicative of altered biomechanical bite forces, all of which might be associated with differences in dietary ecology caused by differing vegetation structure and diversity between the Cayman Islands and Cuba (Brunt, 1994; Proctor, 1994). Comparable differences in crown height and area of cheekteeth are also shown between other Quaternary hutia taxa (Plagiodontia, Hyperplagiodontia) that likely differed in their trophic ecology (Hansford et al., 2012).</p> <p>Considerable morphological variation has been documented between different mainland and insular Cuban Capromys populations, but the relationship between this morphological variation and genetic differentiation between populations is complicated and very poorly understood, leading to confusion over the taxonomic status and relationships between allopatric Capromys populations. Five Cuban subspecies of C. pilorides have been proposed, all of which have been diagnosed morphologically on the basis of craniodental and soft tissue characters: C. pilorides pilorides (Cuban mainland), C. pilorides relictus (northern Isla de la Juventud), C. pilorides ciprianoi (southern Isla de la Juventud), C. pilorides doceleguas (Archipiélago de las Doce Leguas), and C. pilorides gundlachianus (Archipiélago de Sabana) (Varona, 1980, 1983; Borroto Páez et al., 1992; Silva Taboada et al., 2007; Borroto-Páez and Mancina, 2011). Based on analysis of the first 415 base pairs of cyt b for four of these five subspecies, however, the two putative subspecies from Isla de la Juventud (C. pilorides ciprianoi and C. pilorides relictus) show a low level of divergence (0.4%) similar to that observed within other subspecies of C. pilorides (0.0–0.5%) (Woods et al., 2001; Borroto-Páez et al., 2005); C. pilorides ciprianoi has therefore been interpreted as a junior synonym of C. pilorides relictus by some authorities (e.g., Woods et al., 2001), but is retained as a valid taxon by others (e.g., Silva Taboada et al., 2007).</p> <p>Cyt b divergence data have also been used to propose the existence of an undescribed subspecies from Cayo Campo, Archipiélago de los Canarreos (Woods et al., 2001), and a further three offshore Cuban populations have been proposed to also represent distinct but unnamed subspecies (Borroto-Páez et al., 2012). A Capromys specimen studied by Borroto-Páez et al. (2005) from Cayo Ballenato del Medio, Archipiélago de Sabana-Camagüey, which was reportedly morphologically similar to individuals of C. pilorides, showed a markedly higher level of sequence divergence within the first 415 base pairs of cyt b (5.5%–6.4%) compared with levels of divergence seen between samples from the five named C. pilorides subspecies (0.4%–1.9%). This specimen has been variously interpreted as representing a previously unrecognized cryptic Capromys species (Borroto-Páez et al., 2005), or as representing the existing subspecies C. pilorides gundlachianus (Kilpatrick et al., 2012), potentially elevated to species level as C. gundlachianus (Woods and Kilpatrick, 2005); however, it remains undescribed because the skull of the only available specimen is damaged. Most recently, analysis of three mitochondrial genes (cyt b, COI, 12s rRNA) by Upham and Borroto- Páez (2017) demonstrated a primary cyt b divergence of 5.2% between populations all previously considered to represent C. pilorides pilorides from eastern and western mainland Cuba, which is probably consistent with species-level divergence. These authors also demonstrated a further divergence of 2.0% within the western clade between populations from mainland Cuba and from Isla de la Juventud and nearby Cayo Cantiles. However, these patterns of genetic divergence have not yet been related to named taxonomic units within Capromys.</p> <p>Although the morphological differentiation shown by the Cayman Capromys population led Morgan (1994a) to consider that it had likely diverged from the Cuban Capromys source population early in the Pleistocene, sequence divergence demonstrated in our ancient DNA analysis between UF 18588 and a C. pilorides sample from central mainland Cuba is surprisingly low in comparison to divergences between other hutia taxa (table 5). Insight into the taxonomic status of the extinct Cayman Capromys population is provided by consideration of sequence divergence within the cyt b region, for which previously published data are available across a wider sample of extant hutia taxa. Whereas estimated sequence divergence across the entire cyt b gene is only 0.6% between UF 18588 and the mainland Cuban C. pilorides sequence used in our analysis, cyt b sequence divergence between the two recognized extant Geocapromys species (G. brownii and G. ingrahami), which constitute the other congeneric hutia taxon pair included in our analysis, is almost an order of magnitude greater at 5.5%. Conversely, levels of estimated divergence for the first 415 base pairs of cyt b reported by Borroto-Páez et al. (2005) between the named allopatric subspecies of C. pilorides vary between 0.4%–1.9%, and are therefore much more comparable to the estimated divergence between UF 18588 and mainland Cuban C. pilorides in our analyses. Indeed, the values reported by Borroto-Páez et al. (2005) may represent underestimates of sequence divergence across the entire cyt b region, as the first half of cyt b evolves at a slower rate than the second half (Irwin et al., 1991; Spotorno et al., 2004). Levels of divergence across the entire cyt b region between the three subspecies of Plagiodontia aedium (1.0%– 2.9%) are also greater than seen between UF 18588 and mainland Cuban C. pilorides (Brace et al., 2012).</p> <p>Available sequence divergence data therefore do not support recognition of the Cayman Capromys population as a distinct species, as although it is genetically distinct from mainland Cuban C. pilorides, it shows much lower divergence from this population compared with any interspecific divergence values between welldefined species seen across the Capromyinae, with genetic variation instead consistent with evolutionarily recent subspecies-level differentiation. As this extinct population is demonstrably morphologically distinct from all described Cuban subspecies of C. pilorides, we recognize it as a new subspecies, C. pilorides lewisi. Given the confusion over the taxonomic status of different allopatric Capromys populations, however, it is currently impossible to reconstruct the precise evolutionary affinities of C. pilorides lewisi either to other named subspecies of C. pilorides or to the various putative unnamed but apparently distinct taxa within the genus that have been identified by different authors, and this must await further sampling of the genus across its Quaternary distribution.</p> </div>	https://treatment.plazi.org/id/03AA87B0FFCAFF8FFD3410CAFEAFFBAE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Morgan, Gary S.;Macphee, Ross D. E.;Woods, Roseina;Turvey, Samuel T.	Morgan, Gary S., Macphee, Ross D. E., Woods, Roseina, Turvey, Samuel T. (2019): Late Quaternary Fossil Mammals From The Cayman Islands, West Indies. Bulletin of the American Museum of Natural History 2019 (428): 1-81, DOI: 10.1206/0003-0090.428.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2019/issue-428/0003-0090.428.1.1/Late-Quaternary-Fossil-Mammals-from-the-Cayman-Islands-West-Indies/10.1206/0003-0090.428.1.1.full
03AA87B0FFFDFF93FF6A16F9FEF3FE05.text	03AA87B0FFFDFF93FF6A16F9FEF3FE05.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Geocapromys caymanensis Morgan & Macphee & Woods & Turvey 2019	<div><p>Geocapromys caymanensis, new species</p> <p>Figures 19–24</p> <p>HOLOTYPE: UF 21388, nearly complete skull with right M3, lacking the left premaxilla, both nasals, and portions of the left and right zygomatic arches (figs. 19C, 20C).</p> <p>TYPE LOCALITY: Patton’s Fissure, near Spot Bay, northern coast of Cayman Brac, Cayman Islands. The holotype was recovered from layer 7 (120–140 cm below the surface), which has been radiocarbon dated at 13,230 ± 135 yr BP.</p> <p>ETYMOLOGY: The name caymanensis refers to the Cayman Islands.</p> <p>AGE: Late Pleistocene–Holocene (see Radiocarbon Dating).</p> <p>DISTRIBUTION: Known only from Cayman Brac and Grand Cayman.</p> <p>REFERRED SPECIMENS: Cayman Brac: Fig Tree Cave: UF 172761, edentulous mandible. Hutia Cave: UF 172756, nearly complete edentulous skull, lacking nasals, left jugal, and left auditory bulla. Patton’s Fissure: UF 18835, partial skull with left M3, lacking nasals, anterior portion of frontals, left zygomatic arch, and braincase; UF 18836, partial skull, lacking rostrum, teeth, and zygomatic arches; UF 18768, 21358, partial skulls with associated mandibles; UF 21365, partial skull lacking teeth, nasals, zygomatic arches, and braincase; UF 21389, braincase; UF 21398, rostrum with left M2–M3 and right M1–M3; UF 61098, partial skull with left and right M3 and anterior portion of frontals, lacking nasals, zygomatic arches, and braincase; UF 61084, 61086–61089, rostra with edentulous palates; UF 61083, 61090, rostrums; UF 17025, 18553, 18768, 18833, 18836, 21390, 21392, 61099, 61100, partial skulls; UF 61093, palate with right M1–M3; UF 61094, maxilla with M1– M3; UF 61095, palates with left and right M2– M3; UF 21403, associated edentulous maxilla and mandible; UF 61058, 61059, 61078, 61079, mandibles with p4–m3; UF 18821, mandible with p4–m3; UF 61052, mandible with p4–m1; UF 61053, 61060–61062, mandibles with m1– m3; UF 61054, 61063, 61064, mandibles with m1–m2; UF 61049, 61050, 61055, 61065, 61066, mandibles with m2–m3; UF 21403, 61048, 61051, 61057, 61080, 61082, edentulous mandibles; UF 18651, 18652, 18670–18678, mandibles. Peter Cave: UF 22856, partial skull lacking teeth, nasals, zygomatic arches, and braincase; UF 23707, partial skull. Pollard Bay Cave: UF 17025, 18553, partial skulls; UF 18542, mandible; UF 17048, mandible with i1. Spot Bay Cave: UF 61272, partial skull with left M1–M2 and right P4–M3; UF 61273, partial skull with right M1; UF 21674, palate with left M1–M3 and right M1; UF 172780, partial skull with left otic region; UF 172768, edentulous palate; UF 61256, mandible with i1, p4–m3; UF 61257, 172769, mandibles with p4–m2; UF 172775, mandible with m1–m3; UF 172770, mandible with i1, m1–m2; UF 172776, mandible with i1, m2–m3; UF 61260, 61261, 61269–61271, 172772–172774, 172777– 172779, edentulous mandibles. Grand Cayman: Agouti Cave: UF 172883, mandible with i1, m1; UF 172884–172886, edentulous mandibles. Barn Owl Cave: UF 23742, mandible with p4–m2; UF 23741, mandible with m2–m3; UF 23738, 23739, 23745, edentulous mandibles. Bodden Cave: UF 23749, partial maxilla with P4; UF 23747, 23748, edentulous palates. Chisholm Cow Well: UF 172831, 172832, humeri; UF 172833, 172834, ulnae; UF172835, tibia. Crab Cave: UF 23729, partial maxilla with P4; UF 23722, mandible with p4–m1; UF 23712–23721, 23723–23727, edentulous mandibles. Dolphin Cave: UF 172941, palate with left P4–M2 and right M1– M2; UF 172858, palate with left M1; UF 172913, 172930, 172931, edentulous palates; UF 172859, 172914, 172916, 172932, mandibles with p4–m3; UF 172850, 172852, mandibles with p4–m2; UF 172851, 172936, 172937, mandibles with m1– m3; UF 172933, mandible with m1–m2; UF 172917, mandible with m1; UF 172853–172857, 172915, 172918, 172919, 172922, 172923, 172925, 172935, 172938, 172942, 172943, 411278, 411279, edentulous mandibles. Furtherland Farms Cow Well: UF 172544, nearly complete edentulous skull, lacking nasals, left premaxilla, and jugals; UF 172808, partial skull; UF 172805, mandible with i1, p4–m3; UF 172806, mandible with i1, p4–m2; UF 172807, mandible with i1, p4–m1; UF 172545, associated mandibles with left i1, p4 and right i1; UF 172546, edentulous mandible. Miller’s Cave: UF 172903, edentulous mandible; UF 172904, 17290, humeri. Old Man Cave: UF 23743, partial edentulous skull with maxillae and frontals (juv.); UF 23745, 23746, 172903, edentulous mandibles. Queen Elizabeth II Botanic Park: UF 172949, palate with left and right P4. Tadarida Cave: UF 172839, edentulous mandible.</p> <p>DIAGNOSIS: Geocapromys caymanensis is a small species of Geocapromys, slightly larger than G. ingrahami, similar in size to G. thoracatus, and smaller than G. brownii and G. columbianus. The most distinctive feature of G. caymanensis is the strong anterior convergence of the upper toothrows, with left and right P4 separated by less than 2 mm anteriorly and medially. This anterior convergence gives the upper toothrows of G. caymanensis a gentle but noticeable curvature from anterior to posterior. The dorsal surface of the frontals is comparatively narrow and essentially parallel sided from the nasofrontal suture posteriorly to the frontoparietal suture. The anterior portion of the frontals shows an incipient inflation from the orbit anterior to the nasofrontal suture, particularly on the lateral surface. The postorbital processes are weak and bluntly triangular. The dorsal process of the maxilla is vertical and broad anteroposteriorly. The lateral jugal fossa on the zygomatic arch is comparatively small and narrow. The posteroventral edge of the jugal lacks a spine. The zygomatic arch in not rotated ventrally. In lateral view, the ventral surface of the zygomatic arch is level with the alveolar margin of the toothrow. The auditory bullae are rounded and short anteroposteriorly, but show moderate inflation.</p> <p>DESCRIPTION: The description of Geocapromys caymanensis is based primarily on one nearly complete skull (the holotype, UF 21388) and three partial skulls (UF 18835, 18836, 21365) from Patton’s Fissure on Cayman Brac, and a nearly complete but edentulous skull (UF 172544) from the Furtherland Farms Cow Well on Grand Cayman (figs. 19, 20). Three of the skulls preserve M3 but are otherwise edentulous. Geocapromys has unrooted, ever-growing, highcrowned (hypseledont) teeth that are not firmly rooted in the skull, which results in most of the teeth falling out of the skulls and jaws upon death. Large samples of isolated teeth are often found in fossil sites that contain Geocapromys. The descriptions and illustrations of the teeth are based on other specimens listed above.</p> <p>The most distinctive feature of Geocapromys caymanensis is the extreme anterior convergence of the upper toothrows (figs. 19, 21). The left and right P4 nearly meet at the anterior margin of the toothrow. The width of the palate remains very narrow for the entire length of the P4s, with left and right toothrows separated by less than 2 mm at the alveolar margin between P4 and M1. Based on measurements of palatal width (table 6), the palate is actually slightly narrower at the alveolar margin between P4 and M1 (1.7 mm, mean of five individuals) than anterior to P4 (1.9 mm, mean of five individuals). Posterior to the middle of M1, the toothrows begin to diverge laterally, with the distance between toothrows reaching a width of about 5 mm at the posterior margin of the palate. This is similar to the posterior width of the palate in the two other small species of Geocapromys, G. ingrahami and G. thoracatus. Because of their strong anterior convergence at the level of P4 and rather sharp divergence posterior to M1, the upper toothrows have a gentle but noticeable curvature from anterior to posterior, particularly obvious on the lateral alveolar margin.</p> <p>Another diagnostic character of Geocapromys caymanensis is the narrow, parallel-sided frontals on the dorsal surface of the skull (fig. 19). Compared to other species of Geocapromys, the frontals of G. caymanensis are especially narrow anteriorly, from the level of the least interorbital breadth anteriorly to the nasofrontal suture, and are only slightly broader posteriorly between the postorbital processes and the frontoparietal suture. Measurements that reflect the narrow frontals are the breadth of the frontals both anterior to the supraorbital processes (interorbital breadth) and posterior to the supraorbital processes (postorbital breadth). These frontal measurements are narrower than the same measurements in all other species of Geocapromys, even though G. caymanensis is not the smallest species in the genus (tables 6, 7). Although narrow transversely, the anterior frontals in the Cayman Geocapromys are moderately inflated, from dorsal to the anterior edge of the orbit anteriorly to the nasofrontal suture. This anterior frontal inflation is primarily obvious laterally along the internal margin of the orbit and dorsal to the superior process of the maxilla, but not along the midline where the frontals are essentially flat. The widest portion of the frontals occurs at the level of the postorbital processes, which are small and bluntly triangular. Compared to other Geocapromys, the postorbital processes are weak in G. caymanensis. The temporal crests are low, rather weak, U-shaped, and do not meet to form a sagittal crest; they are separated by 3–4 mm on the posterior surface of the parietals, just anterior to their connection with the nuchal crest.</p> <p>The dorsal process of the maxilla is preserved in only one skull of G. caymanensis (holotype; fig. 19C), in which this process is vertical and relatively broad anteroposteriorly. The lateral jugal fossa (a character present on the posteroventral margin of the zygomatic arch ventral to the orbit in capromyines) is preserved in only one of the Geocapromys skulls from Cayman Brac (UF 18835), in which this fossa is narrow and its ventral margin lacks a jugal spine. In lateral view, the zygomatic arch in this skull of G. caymanensis is not noticeably downturned or rotated ventrally. The ventral edge of the zygomatic arch is only slightly inclined posteroventrally and is on essentially the same level as the alveolar margin of the toothrow. In ventral aspect, the pterygoid region is constricted anteroposteriorly. This compressed pterygoid region is indicative of a rather foreshortened braincase that is also noticeably downturned ventrally. The auditory bullae are comparatively short, rounded, and somewhat inflated. In posterior view, the auditory bullae extend ventrally to the occipital condyles.</p> <p>MORPHOMETRICS: As for Capromys, the largest available measurement series for the Cayman Geocapromys sample is the mandibular alveolar toothrow length (Grand Cayman, n = 41; Cayman Brac, n = 30). Although Geocapromys skulls from Grand Cayman and Cayman Brac are otherwise morphologically very similar, statistically detectable size differences are present between samples from these two islands (Welch two-sample t-test not assuming equal variance: Grand Cayman, mean = 15.25 mm; Cayman Brac, mean = 16.52 mm; p &lt;0.001). Sample series from each island were therefore compared separately with morphometric data for other extant and extinct Geocapromys populations measured from museum collections: G. brownii (n = 17, mean = 19.11 mm), G. columbianus (n = 40, mean = 18.76 mm), G. ingrahami (sample comprising both recent specimens from East Plana Cay, and Quaternary fossil specimens from Abaco, Crooked Island, and Exuma: n = 57, mean = 16.12 mm), and G. thoracatus (n = 12, mean = 14.72 mm) (appendix 2). Our Grand Cayman Geocapromys sample has a significantly smaller mandibular alveolar toothrow length compared to G. brownii (p &lt;0.001), G. columbianus (p &lt;0.001) and G. ingrahami (p &lt;0.001), and a significantly greater mandibular</p> <p>alveolar toothrow length compared to G. thoracatus (p = 0.016). Our Cayman Brac Geocapromys sample has a significantly smaller mandibular alveolar toothrow length compared to G. brownii (p &lt;0.001) and G. columbianus (p &lt;0.001), a significantly greater mandibular alveolar toothrow length compared to G. thoracatus (p &lt;0.001), and does not differ statistically in size from G. ingrahami (p = 0.089).</p> <p>MORPHOLOGICAL COMPARISON WITH OTHER SPECIES OF GEOCAPROMYS: Geocapromys is the</p> <p>most widespread genus in the Capromyinae, known as either a living animal or from historical museum specimens or fossils from the Cayman Islands, Bahamas, Cuba, Jamaica, and Little Swan Island. Geocapromys caymanensis is a rather small species of Geocapromys, comparable in size to the smallest of the three living species, G. ingrahami from the Bahamas (tables 6, 7). Among the three extant species of Geocapromys, G. caymanensis is most similar to the Bahamian hutia G. ingrahami and least similar to the Jamaican hutia G. brownii.</p> <p>Geocapromys ingrahami survives today as a wild population on a single island, East Plana Cay in the southern Bahamas, with translocated populations also established on Little Wax Cay and Warderick Wells Cay (Clough, 1972; Turvey et al., 2017). Two extinct subspecies have been described based on fossils from the Bahamas: G. ingrahami irrectus from Crooked Island, with referred samples from Eleuthera and Long Island, and G. ingrahami abaconis from Abaco (Lawrence, 1934). Both subspecies were distinguished from the living G. ingrahami primarily on the basis of their larger size (e.g., longer alveolar toothrows). Numerous additional extinct populations of G. ingrahami have been reported since, mostly from islands on the Great Bahama Bank, although these samples were not referred to either of the two extinct subspecies (Morgan, 1989a). Geocapromys caymanensis overlaps with G. ingrahami in many of the most important cranial measurements indicative of size, including total length and condylobasal length of the skull, and breadth of the skull at the auditory meatus. The general size similarity in Geocapromys individuals from these two geographically separate island groups may be explained by the relatively small land area of islands in both regions compared to the much larger islands of Cuba and Jamaica; this may have resulted in common constraints on maximum body size attainable in both regions (cf. Burness et al., 2001). The Bahamian and Cayman species of Geocapromys differ in the degree of anterior convergence of upper toothrows, which are highly convergent in G. caymanensis, and convergent but less extreme in G. ingrahami. Measurements of the anterior palatal width show that the toothrows are more strongly convergent anteriorly in these two species than in G. brownii and G. thoracatus (table 6). The frontals of G. ingrahami are slightly broader than in G. caymanensis, which has narrower frontals compared with all other species in the genus. G. ingrahami shows no evidence of the anterior frontal inflation that is characteristic of G. caymanensis, and also has more prominent, sharply triangular postorbital processes compared to the small, blunt processes of G. caymanensis. The dorsal process of the maxilla is vertical in both G. caymanensis and G. ingrahami but is noticeably broader in the Cayman species. These two species share several characters of the zygomatic arch that may also indicate a close phylogenetic relationship, including the narrow jugal fossa, lack of a jugal spine, and the minimal ventral rotation of the entire zygomatic arch with the ventral edge of the arch level with the alveolar margin of the toothrow. Both G. ingrahami and G. caymanensis also have a foreshortened braincase that is downturned ventrally. The short braincase is reflected ventrally in the constricted pterygoid regions of these two species. G. ingrahami has comparatively larger auditory bullae that are more inflated than in the Cayman species.</p> <p>TABLE 6</p> <p>Comparative measurements (in mm) of the skull and upper dentition of Geocapromys caymanensis,</p> <p>and other living and extinct species of Geocapromys from Jamaica, Bahamas, and Little Swan Island. Abbreviations: M, mean; R, range; and N, sample size.</p> <p>TABLE 7</p> <p>Comparative measurements (in mm) of the mandible and lower dentition of Geocapromys caymanensis, and other living and extinct species of Geocapromys from Jamaica, Bahamas, and Little Swan Island. Abbreviations: M, mean; R, range; and N, sample size.</p> <p>Geocapromys caymanensis differs in size and most diagnostic cranial characters from the largest living species in the genus, G. brownii from Jamaica. Measurements in tables 6 and 7 reveal no overlap in cranial measurements between the smaller G. caymanensis and the larger G. brownii. The strong anterior convergence of the upper toothrows in G. caymanensis is not observed in G. brownii. The characters of the dorsal surface of the frontals also differ considerably between these two species. In G. brownii, the frontals are very broad and strongly inflated anterior to the postorbital processes, especially laterally, whereas the frontals are quite narrow and constricted posterior to the postorbital processes. Overall, the frontals in G. brownii are much broader anterior to the postorbital processes than posterior to these processes (table 6). In G. caymanensis, the frontals are comparatively narrow and almost parallel sided, exhibit only a minor degree of anterior inflation, and are slightly broader posterior to the postorbital processes. G. brownii has rather weak V-shaped temporal crests that meet to form a prominent sagittal crest on the parietals about midway between the frontal and occipital sutures. The temporal crests are also weak in G. caymanensis, but tend to be more U-shaped, converge (but do not meet) much farther posteriorly on the parietals, and do not form a sagittal crest. G. caymanensis has a foreshortened and ventrally downturned braincase compared to G. brownii, in which the braincase is more elongated. The comparative length of the braincase is especially obvious in the pterygoid region, in which the distance between the posterior edge of the toothrow and the anterior edge of the auditory bulla is much shorter in G. caymanensis than in G. brownii. The dorsal process of the maxilla is vertical in G. caymanensis, whereas this process is more posteriorly oriented in G. brownii. The posterior orientation of the dorsal process of the maxilla in G. brownii is related to the overall ventral rotation of the zygomatic arch, with the ventral edge of the arch located markedly ventral to the alveolar margin and occlusal surface of the upper teeth. The zygomatic arch is not rotated ventrally in G. cay- manensis; the ventral edge of the arch is instead located approximately level with the alveolar margin of the toothrow. G. brownii has a large, broad jugal fossa with a prominent jugal spine located on the posteroventral margin of the jugal; a jugal spine is absent in G. caymanensis, and the jugal fossa is narrow. The auditory bullae are larger and more elongated in G. brownii, versus shorter and rounded in G. caymanensis.</p> <p>The Little Swan Island hutia, G. thoracatus, is somewhat larger than G. caymanensis in most cranial measurements (table 6), and shares many morphological features with the larger Jamaican G. brownii, to which G. thoracatus appears to be closely related (Morgan, 1985). This species is now extinct but survived on Little Swan Island until the 1950s, and is represented by skins, skulls, and skeletons in several museum collections (Morgan, 1989b; Tonge, 2014). Unlike G. caymanensis, G. thoracatus does not exhibit a strong anterior convergence of the upper toothrows. This species has a small spinous process that projects 1–2 mm beyond the posterior palatal margin along the midline; this process is not observed in any other species of the genus, including G. caymanensis. One of the most char- acteristic features of G. thoracatus compared to its congeners is its comparatively small teeth. The frontals are broad in G. thoracatus, especially posterior to the postorbital processes, and there is no evidence of anterior inflation of the frontals, compared with G. caymanensis, which has narrow, parallel-sided frontals with a minor degree of inflation anteriorly. Features of the dorsal process of the maxilla and zygomatic arch of G. thoracatus are similar to those of G. brownii, although less pronounced. G. thoracatus has a broad, posteriorly oriented dorsal process of the maxilla, a broad jugal fossa with a well-developed jugal spine, and a ventrally rotated zygomatic arch, all of which are quite distinct from G. caymanensis; the Cayman species instead has a vertical dorsal process of the maxilla, a narrow jugal fossa lacking a jugal spine, and an absence of ventral rotation of the zygomatic arch. G. thoracatus has longer and narrower auditory bullae than G. caymanensis, which has shorter, rounded, and more inflated bullae.</p> <p>Four extinct species of Geocapromys have been named from Cuban fossil deposits, but only G. columbianus is now considered valid (Silva Taboada et al., 2007). G. columbianus is not well described or illustrated, with little cranial material having been figured in the literature (Silva Taboada et al., 2007; Díaz-Franco and Jiménez Vázquez, 2008). In his original description of G. columbianus, Chapman (1892) characterized this species as having strongly convergent upper toothrows, with a measurement of “0.04 in” (= 1.0 mm) between the anterior margins of the P4s. He also noted that the left and right than the Cayman and Cuban species of Geocapalveoli essentially meet between the upper pre- romys. Although not specifically mentioned in molars. Allen (1917) obtained additional fossil the description, the illustration of the holotype specimens of G. columbianus from Cuba and of G. columbianus shows a rather strong curvaconfirmed several of the morphological features ture of the upper toothrows, especially along the of this species reported by Chapman (1892), in lateral margin (Chapman, 1892: fig. 3), a feature particular the strong anterior convergence of also characteristic of G. caymanensis but not the upper toothrows. The Cayman species also quite as pronounced. Chapman (1892) did not has strongly convergent upper toothrows, with provide a measurement of the alveolar length of the anterior margins of the P4s separated by 1.9 the toothrow in the holotype of G. columbianus mm (mean of five individuals; table 6), but this because the palate is broken off posterior to M2. convergence is apparently not quite as extreme as Silva Taboada et al. (2007) presented selected in the Cuban species. The toothrows in G. cay- measurements of Geocapromys columbianus, manensis are also highly convergent between P4 including several cranial measurements of a sin- and M1, separated by only 1.7 mm (mean of five gle skull. The condylobasal length of this skull individuals; table 6); however, the left and right (65.9 mm) is about 12% larger than the only toothrows do not nearly meet between the P4s as skull of G. caymanensis on which this measurein G. columbianus (Chapman, 1892: 314–315, fig. ment could be taken, the holotype (58.4 mm). 3). The strong anterior convergence of the upper The breadth at the auditory meatus in the single toothrows may indicate a close phylogenetic skull of G. columbianus is 28.1 mm, versus a relationship between G. caymanensis and G. mean of 26.4 mm (range of 25.2–27.1 mm) for columbianus. The Bahamian hutia G. ingrahami three specimens of G. caymanensis, about 10% also demonstrates an anterior convergence of smaller than the Cuban species. Silva Taboada et the upper toothrows, although to a lesser degree al. (2007) listed a mean of 16.5 mm (range of 14.4–18.2 mm; 48 specimens) for the alveolar length of the upper toothrow of G. columbianus. Although the mean for the alveolar length of the upper toothrow in five specimens of G. caymanensis is slightly larger at 16.9 mm, the upper limit for the observed range is 17.6 mm, somewhat less than in G. columbianus. Based on the wide range of values, we suspect Silva Taboada et al. (2007) included some juveniles in their sample, whereas we excluded juveniles, thus increasing the mean length of the alveolar toothrow compared with the Cuban sample. Overall, the available measurements suggest that G. columbianus is about 10% larger than the Cayman species. Other cranial characters of G. columbianus that differ from G. caymanensis are: the posterior orientation of the dorsal process of the maxilla, a slight ventral rotation of the zygomatic arch, the presence of a small jugal spine, and the near convergence of the temporal crests on the parietals to form a short low sagittal crest. In contrast, G. caymanensis has a vertical dorsal process of the maxilla, no ventral rotation of the zygomatic arch, no jugal spine, and the temporal crests do not meet.</p> <p>REMARKS: The species of Geocapromys form a monophyletic group that is distinguished from Capromys by a number of external and cranial characters (Morgan, 1985; 1989b). External characters that distinguish Geocapromys from Capromys include: shorter tail, reduced first digit on the front foot, and shorter finer fur. Dental and cranial features that characterize Geocapromys are: more procumbent incisors, origin of the upper incisor root capsule high on maxilla above P4, broad vertically or posteriorly oriented dorsal process of the maxilla, 30° inclination of all cheekteeth, and presence of an anteroflexid on p4. Among the three living or recently extinct species of Geocapromys, G. ingrahami is distinguished from G. brownii and G. thoracatus by the following characters that are considered derived for G. ingrahami: more convergent upper toothrows, constricted pterygoid region, shortened and more inflated braincase, and inflated auditory bullae. The extinct taxa of Geocapromys from Cuba and the Cayman Islands possess most of the derived characters present in G. ingrahami, although the expression of these characters varies significantly between species. The shared possession of these derived characters suggests that G. caymanensis and G. columbianus are probably more closely related to G. ingrahami than they are to G. brownii or G. thoracatus, and these species were previously associated as the ingrahami species group within Geocapromys (Morgan, 1985).</p> <p>From a biogeographic standpoint, the extinct G. columbianus from Cuba represents a plausible source population for G. caymanensis. Only G. columbianus and G. caymanensis exhibit extreme anterior convergence of the upper toothrows, which is almost certainly a derived feature indicating a close relationship between these species. Considering Cuba’s central location between the Bahamas to the north and the Cayman Islands to the south, as well as the long history and evolutionary diversity of capromyine rodents in Cuba, it seems most likely that G. columbianus or a precursor probably gave rise to both G. caymanensis and G. ingrahami sometime prior to the Late Pleistocene.</p> <p>Geocapromys occurs in almost every cave deposit excavated in both Cayman Brac and Grand Cayman (see list of Referred Specimens). No fossils of Geocapromys are known from Little Cayman, although only limited fossil exploration has been conducted on that island. As in his initial assessment of the status of the then-undescribed Nesophontes populations from Grand Cayman and Cayman Brac, Morgan (1994a) proposed that Geocapromys populations from these two islands varied in size, and should therefore be interpreted as representing two distinct species. We support Morgan’s (1994a) observation by revealing statistically detectable size differences between Geocapromys samples from Grand Cayman and Cayman Brac, but, as with the allopatric Cayman Nesophontes populations, these samples are otherwise morphologically indistinguishable and lack any island-specific unique morphological synapomorphies. We are therefore reluctant to taxonomically differentiate the two allopatric Cayman Geocapromys populations in the absence of further information about their phylogenetic relationships, and we encourage future investigation of the evolutionary history of these populations using ancient DNA techniques.</p> </div>	https://treatment.plazi.org/id/03AA87B0FFFDFF93FF6A16F9FEF3FE05	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Morgan, Gary S.;Macphee, Ross D. E.;Woods, Roseina;Turvey, Samuel T.	Morgan, Gary S., Macphee, Ross D. E., Woods, Roseina, Turvey, Samuel T. (2019): Late Quaternary Fossil Mammals From The Cayman Islands, West Indies. Bulletin of the American Museum of Natural History 2019 (428): 1-81, DOI: 10.1206/0003-0090.428.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2019/issue-428/0003-0090.428.1.1/Late-Quaternary-Fossil-Mammals-from-the-Cayman-Islands-West-Indies/10.1206/0003-0090.428.1.1.full
