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Home , Cricetidae, Handleyomys alfaroi, Hispid cotton rat

Occasional Papers Museum of Texas Tech University Number 308 31 May 2012

Sm a l l Ma m m a l Su r v e y o f Ch i q u i b u l Fo r e s t Re s e r v e , Ma y a Mo u n t a i n s , Be l i z e , 2001

An d r e w En g i l i s , Jr., Ro n a l d E. Co l e , a n d Ti m Ca r o

Ab s t r a c t

The Chiquibul Forest Reserve is located in western in the Maya Mountains and protects one of the largest patches of rainforest remaining in Central America. We conducted inventories of small in the forest reserve from 4 July through 8 August 2001. Our five trapping sites were centered within a few kilometers of the Las Cuevas Field Station. In total, we ran 3,686 trap-nights capturing 154 specimens (4.2% trap success) representing 15 of small mammals. We ran mist nets for ten nights capturing 39 phyllostomid bats representing six species. Heteromys desmarestianus was the most abundant trapped; it was captured at a rate four times more frequently than all other species. Ototylomys and were next in abundance. Almost all species of and bats were in a high state of reproductive activity. Our efforts confirmed an additional eleven species to the Chiquibul Forest Reserve, five non-volant: Cryptotis mayensis, fulvescens, Handleyomys rostratus, Sigmodon toltecus, Nyctomys sumichrasti, and six volant: Carollia sowelli, Sturnira lilium, Artibeus jamaicensis, Dermanura toltecus, D. watsoni, Centurio senex. The biogeographic affinities of small mammals of the Maya Mountains lay more nested within the Central American faunal group – less so than the Yucatan faunal region. The discovery of Cryptotis mayensis in the Maya Mountains provided a significant range and ecological extension. We present preliminary character traits to separate Sigmodon toletcus from S. hispidus.

Key words: Belize, Chiroptera, Cryptotis, Maya Mountains, Museum of Wildlife and Fish Biology, node trapping, Rodentia, Sigmodon, small mammals

In t r o d u c t i o n

The Chiquibul Forest Reserve is located in the forests associated with the granitic soils north of the Maya Mountains of western Belize and protects one of Maya Mountains (Murie 1935) and lowlands of Belize the largest patches of rainforest remaining in Central and the Yucatan Peninsula (Herskovitz 1951; Disney America (Harcourt 1996). Limited inventories of small 1968; Bersot 2003). Few researchers have undertaken mammals in this region (at least those that included inventories in rainforest habitat associated with lime- voucher specimens) have focused primarily on the pine stone soils further to the south (but see Rabinowitz and 2 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y

Nottingham 1989; Caro et al. 2001a; Kelly and Caro ment diversity used in an assessment of overall species 2003). Recent efforts to describe the small mammal richness in this region (Caro et al. 2004). Recent, and fauna were focused on factors affecting trapping suc- at times sweeping, taxonomic revisions impacting cess (Kelly and Caro 2003), landscape-scale patterns the fauna of this region compel us to provide a list of of mammalian diversity (Caro et al. 2001b), and the species currently known from the region. Subsequent effectiveness of using flagship species in establishing to our efforts, the construction and operation of a hy- reserves (Caro et al. 2004). The only previous trap- droelectric dam in the region has resulted in an influx ping reports from this region failed to secure vouchers of people into the northern part of the Chiquibul For- to confirm the identification of small mammals in the est Reserve (Conservation Strategy Fund 2000). The Chiquibul (Caro et al. 2001b; Kelly and Caro 2003). future of this fauna is uncertain. The specimens reported herein were collected to docu-

St u d y Ar e a a n d Me t h o d s

The Chiquibul Forest Reserve, Cayo District, river or beside a pond; 100 traps within the forest in- western Belize, is surrounded by the fully protected terior; and 100 traps along forest edge, usually a trail; Chiquibul National Park (Fig. 1). The vegetation thus totaling 300 total traps/site. Snap traps and some is a mosaic of subtropical evergreen and deciduous small live traps were set to enhance capture of arboreal seasonal forest (Brewer and Webb 2002; Brewer et al. mammals, being tied to trees and vines 1- 3 m above 2003), although stands of pines (Pinus) occur in the ground; others were placed under or on fallen logs, northern sector (Hartshorn et al. 1984). Some blocks near burrows, or on small trails. Traps were set of the Chiquibul Forest Reserve have been, and are still for three days and three nights at each site and were being, selectively logged for commercially important baited with a variety of baits including pureed banana, species such as mahogany (Swietenia macrophylla) and peanut butter, rolled oats, banana slices, and in large cedar (Cedrela odorata) on a >40-year rotational basis Tomahawks only, sardines. Traps were baited every (Bird 1998). Also, a large part of the Chiquibul Forest morning and left open for 24 hours. They were checked Reserve suffered losses of trees from Hurricane Hattie twice a day. Samples of each species in these transects in 1961 (Wolffsohn 1967). Rainfall averages about were kept as vouchers. 1500 mm/year, with the rainy season starting in June and continuing through January (Caro et al. 2001). Our In addition, two pitfall arrays with drift fences period of trapping was conducted at the beginning of were set by herpetologists sampling frogs in the forest the rainy season from 4 July through 8 August 2001. at the Aguada trapping locality (Fig. 1). We were able to piggy-back upon their efforts from 18 July to 1 August, We sampled five collecting sites roughly centered checking the pitfalls twice per day, collecting small within a few kilometers of the Las Cuevas Field Station mammals. Each pitfall array comprised five 1-gallon (Fig. 1; Table 1). Each site was sampled using tapping buckets at 5 m intervals and connected with a plastic nodes along a transect. Transects were placed where drift fence approximately 20 m in length. habitat conditions appeared favorable for small mam- mals, and comprised five nodes (ca. 50 m intervals) To sample bats, we ran two 2 x 10 m mist nets selected on the basis of animal sign, diversity of forest for ten nights, set in active flyways determined by floor structure, and liana growth. At each node, we set presence of water (streams or forest ponds) or gaps in twenty traps, including six Museum Special or Victor the forest (natural or manmade). Each net was opened snap traps, eight Sherman live-traps (23 x 8 x 8 cm), prior to sunset and left open through a portion of the three similar custom-made wire mesh traps (30 x 8 x night and checked at one-hour intervals until closed. 8 cm), two medium-sized Tomahawk traps (40 x 13 x When checked, bats were either released or collected 13 cm or 40 x 17 x 17 cm), and one large Tomahawk as voucher specimens. Nets were then closed for the trap (65 x 22.5 x 22.5 cm). In each of our five sites, we day. We also recorded and identified all species of set three transects with this combination and number mammals encountered by visual and or auditory means of traps. At each site we set 100 traps along a stream, while trapping and working in the field. En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 3

Orange Caribbean Walk

Belize City Belize

Belmopan

San Ignacio

17 S

Stann Creek Cayo

Sea Toledo 89 W N

Figure 1. Location of Las Cuevas Field Station and general study area (indicated by star) in Maya Mountains, Belize. Italic names refer to political districts. Lower map shows 1-km grids of study area. Collecting sites are as follows: (1) Las Cuevas Field Station, (2) Aguada Creek, (3) Monkey Tail Branch, (4) San Pastor, (5) Millionario. 4 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y

Table 1. Collecting localities in the Chiquibul Forest Reserve (refer to Fig. 1) and collecting effort per site. Trap, net, and pitfall nights reflect the number of each unit times the number of nights open (e.g., two nets open for three consecutive nights = six net nights).

Site Collecting/Trapping Locality (as written on specimen tags) Trap nights Net nights Pitfall nights

Belize: Cayo District, Las Cuevas Research Station, 16.732869°N 88.985952°W, 1 314 6 550m.

Belize: Cayo District, 3.3km E, 1.1km N of Las Cuevas Research Station, 2 843 6 14 Aguada Creek, 16.742952°N 88.955533°W, 600m.

Belize: Cayo District, Monkey Tail Branch, 5.4 km E, 0.5 km N of Las Cuevas 3 830 6 Research Station, 16.740202°N 88.934717°W, 465m.

Belize: Cayo District, San Pastor, 1.0 km W, 0.4 km N of Las Cuevas Research 4 846 6 Station, 16.726200°N 88.990233°W, 590 m.

Belize: Cayo District, Millionario, 2.1 km N, 2.1 km W of Las Cuevas Research, 5 853 6 Resumadero Road, 16.756950°N 9 89.012833°W, 600m.

Relative abundance of non-volant mammals was or alcohol preserved specimens. The bulk of material expressed as the number of individuals/100 trap nights. was deposited at the Museum of Wildlife and Fish Bi- We did not record capture effort for netted bats. In ology, University of California, Davis (WFB), with a the field, mammal identifications were aided by Reid subset left at the Las Cuevas Field Station, Belize, as a (1997) and Emmons and Feer (1990). Determinations reference collection. We opportunistically combed and were primarily undertaken using known reference picked ectoparasites from collected mammals. These materials and comparisons with specimens at the US were deposited in the Bohart Museum of Entomol- National Museum and American Museum of Natural ogy, University of California, Davis, awaiting further History. in general follows Wilson and determinations by experts. The museum operates Reeder (2005); deviations from this reference are noted off of an animal care and use protocol established at where applicable. the University of California, Davis (protocol number 15331). We obtained collecting and export permits All collected mammals were preserved as study from the Belize Government prior to trapping and skins plus skulls, complete skeletons, partial skeletons, obtaining specimens.

Re s u l t s

In total, we ran 3,686 trap-nights capturing 154 Rodents were also in a high state of reproductive activ- (4.2% trap success) representing 15 species ity across all species. We recorded 33 species of mam- of small mammals (Table 2). Mist nests captured 39 mals through trapping and observations (Table 3). individuals representing six species of bats (all Phyl- lostomidae; no Vespertillionidae were netted). Several We experienced little or no problems with ants female bats contained embryos indicating reproductive raiding our traps, and rain impacted only one trap night. activity and potential timing of births for the beginning Large forest cockroaches (Blaberidae) and beetles of the rainy season. Heteromys desmarestianus was the (Scarabaeidae) sometimes tripped small live traps. most abundant mammal trapped; it was captured at a We include measurements (external and cranial) for rate four times more frequently than all other species. all specimens collected (Appendix). Ototylomys phyllotis was next in abundance (Table 2). En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 5

Table 2. Results of small mammal trapping, Maya Mountains, Belize.

Collecting Site Total % abundance/ Field Station Monkey Tail San Pastor Millionario Aguada Captures 100 traps Didelphis virginiana 0 0 2 1 1 4 0.11% Didelphis marsupialis 0 0 5 2 2 9 0.24% Philander opossum 0 1 0 1 0 2 0.05% Marmosa mexicana 0 4 2 0 3 9 0.24% Cryptotis mayensis 0 0 0 0 5 5 0.14% Heteromys desmarestianus 10 12 0 11 20 53 1.44% 1 0 0 0 0 1 0.03% Handleyomys alfaroi 3 0 1 4 6 14 0.38% Handleyomys rostratus 1 0 1 0 2 4 0.11% Sigmodon toltecus 4 0 4 6 0 14 0.38% Nyctomys sumicrasti 0 1 0 0 1 2 0.05% Ototylomys phyllotis 18 5 5 0 5 33 0.90% nudicaudatus 1 1 0 0 1 3 0.08% Leopardus pardalis 0 0 1 0 0 1 0.03% TOTALS 38 24 21 25 46 154 4.18%

Table 3. List of mammals observed and trapped in the Chiquibul Forest Reserve, 4 July – 8 August 2001.

Didelphis marsupialis - Common Opossum Handleyomys rostratus - Long-nosed Rice Rat Didelphis virginiana - Virginia Opossum Oligoryzomys fulvescens - Fulvous Colilargo Philander opossum - Gray Four-eyed Opossum Tylomys nudicaudatus - Peters' Climbing Rat Marmosa mexicana - Mexican -opossum Ototylomys phyllotis - Big-eared Climbing Rat Cryptotis mayensis - Yucatan Small-eared Nyctomys sumichrasti - Sumichrast's Vesper Rat Carollia sowelli - Silky Short-tailed Bat Sigmodon toltecus - Hispid Sturnira lilium - Little Yellow-shouldered Bat Dasyprocta punctata - Agouti Artibeus jamaicensis - Jamaican Fruit-eating Bat Agouti paca - Paca Dermanura watsoni - Thomas's Fruit-eating Bat Nasua nasua - Coati Dermanura tolteca - Toltec Fruit-eating Bat Felis onca - Jaguar Centurio senex - Wrinkle-faced Bat Felis concolor - Mountain Lion Alouatta pigra - Black Howler Monkey Felis pardalis - Ocelot Ateles geoffroyi - Geoffroy's Spider Monkey Tapirus bairdii - Baird's Tapir Dasypus novemcinctus - Nine-banded Armadillo Dicotyles tajacu - Collared Peccary Sciurus deppei - Deppe's Squirrel Tayassu pecari - White-lipped Peccary Heteromys desmarestianus - Desmarest's Spiny Pocket Mouse Mazama temama - Central American Red Brocket Handleyomys alfaroi - Alfaro's Rice Rat 6 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y

Sp e c i e s Acc o u n t s

ORDER DIDELPHIMORPHIA M. mitis, and M. robinsoni provided very helpful Family Didelphidae identification criteria for subadult animals. We re- Didelphis virginiana Kerr 1792 examined the skulls and pelage traits of the subadults Virginia Opossum and concur with Voss that seven subadult specimens are M. mexicana. Specimens examined.—Nine individuals were captured in Tomahawk live traps which were baited Two adult males (WFB 4233 and 4305) were with a mix of fruit, vegetable matter, and peanut butter. both old animals. WFB 4233 had a very large skull Seven of the nine captures were in secondary forest with well-developed supraorbital ridges, atypical for habitat and the other two in forest edge habitat. Just M. mexicana (Rossi et al. 2010). Furthermore this one of the nine (WFB 4321) was kept as a voucher specimen had reduced palatine fenestrae, a large upper specimen. It was a subadult male with left testis = 4 toothrow (8.9 mm), a very large CB length (40.0 x 3 mm. mm), and the black mask did not extend to touch the ears. These latter character traits coupled with the large Didelphis marsupialis Linnaeus 1758 supraorbital ridge are inconsistent with measurements Common Opossum and traits reported for mexicana; in fact this specimen’s measurements exceeded those reported for M. mexicana Specimens examined.—Four individuals were (Rossi et al. 2010). In our opinion WFB 4233 exhibits captured using Tomahawk live traps; two were retained more traits consistent with those reported for M. isth- as specimens, WFB 4292 and WFB 4313. Both were mica rather than M. mexicana (Rossi et al. 2010). In a subadults, left testis = 11 x 8 mm and 6 x 8 mm, respec- recent communication with R. Voss, he stated that our tively. The two released animals also were subadults. animals were unusually old, and both being males, can One individual was captured in forest edge habitat and exhibit larger inflected supraorbital ridges and were all the other three were captured in secondary forest. clearly M. mexicana. Specimen WFB 4233 probably represents an aberrant M. mexicana, but our diagnosis Marmosa mexicana Merriam 1897 will remain referred, Marmosa cf. mexicana, until it can Mexican Mouse Opossum be further resolved. The second specimen of adult male (WFB 4305) had less clearly developed supraorbital Specimens examined.—Nine individuals were ridges and metrics fell more within those reported for collected (WFB 4233, 4269, 4289, 4301, 4302, 4305, adult male M. mexicana. 4317, 4319, and 4323). Of the nine specimens ex- amined only two were adults as determined by tooth Remarks.—All of the subadult M. mexicana eruption patterns and size. specimens were captured in trees rather than on the ground. Six were caught in snap traps tied to branches Diagnosis.—Our collections of Marmosa exhibit and lianas and the seventh was caught in a box trap, two distinct age classes, an arboreal cohort of subadults also tied to a tree limb. WFB 4233 and 4305 both and a terrestrial cohort of adult males. Our first incli- were adult males with well developed, active testes. nation was to consider these as separate species based One of our two captures was made with a ground set on body size and craniodental characteristics (e.g., M. box trap and the second was captured in a pitfall. Both mexicana and M. robinsoni). Publication of a revision animals were heavily parasitized with botfly larvae of Central American Marmosa (Rossi et al. 2010) led (family Oestroidea). WFB 4305 had recently survived to uncertainty as to our identification and animals were an apparent attack with part of its facial skin missing sent to the American Museum of Natural History for and healed. The two old males present interesting determination. All specimens where determined as behavioral questions relating to terrestrial dispersal in M. mexicana by R. Voss (pers. comm.). Rossi et al.’s this arboreal species. (2010) thorough re-evaluation of Marmosa mexicana, En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 7

Philander opossum (Linnaeus 1758) calis, the other known shrew in the area, by its grayer Gray Four-eyed Opossum pelage and longer tail, longer and less circumscribed , and larger size. Our animals fall well Specimens examined.—Two specimens were within all other cranial metrics defined forC. mayensis collected (WFB 4306 and 4337); both were males, (Woodman and Timm 1993) (Appendix). The upper a subadult and adult respectively. WFB 4306 was pelage coloration imparts a reflective nature (well captured in a box trap set in forest edge habitat. WFB described by Woodman and Timm 1993). We could 4337 was captured by the semi-feral house cat living not see the tricolored coloration of the dorsal hairs on the grounds of the research station. described for this species (Choate 1970; Woodman and Timm 1993). The specimens were too small in all ORDER SORICOMORPHA measurements reported for C. magna and were larger Family Soricidae than C. hondurensis. Cryptotis mayensis (Merriam 1901) Yucatan Small-eared Shrew Remarks.—This species has a very limited distri- bution and is poorly represented in collections. Most Specimens examined.—Five specimens were col- records are from the Yucatan Peninsula (Woodman and lected and examined (WFB 4223, 4234, 4243, 4271, Timm 1993). C. mayensis is reported to be the only 4274). Cranial and body measurements fall within member of the C. nigrescens group to be restricted to those reported by Woodman and Timm (1993) for this lowlands where most records fall below 100m in dry species. All were taken from pitfall traps in a wet seep scrub, deciduous forest, and seasonally dry evergreen along Aguada Creek in deciduous semi-evergreen forest forest elevation (Woodman and Timm 1993). Murie at 600 m elevation. The soil was very shallow on top (1935) reported capturing Cryptotis in marshlands of partially exposed limestone rock. The overstory was and a palmetto draw along a pine ridge in the Cayo completely closed shading the forest floor; leaf litter District (he reported the species as C. micrura which covered the forest floor. The understory was sparsely has been synonomized with C. mayensis and C. tropi- vegetated. The one identified female collected was per- calis (Woodman and Timm 1993; Wilson and Reeder forate, indicating recent reproductive activity, however 2005). Choate (1970) examined Murie’s specimens none of the males captured showed signs of being in and assigned them to C. parva tropicalis. We examined reproductive activity (no lateral glands present). Murie’s specimens (from the University of Michigan, Museum of Zoology) and they are correctly assigned Diagnosis.—Specimens were sent to the USNM to C. tropicalis (confirmed in recent communication and determined by N. Woodman. Our animals rep- with N. Woodman). While it remains unclear as to resented a relatively large shrew, dorsal coloration the correct elevation and precise location of Murie’s a dark gray-brown with faint pale spots on the hind specimens, we mapped the general locality of his col- quarters, vent gray. All of the measurements, external lection based on information gleaned from the original and cranial, indicate it to be too large for C. parva (the tag (Fig. 2). In Belize, pine ridges occur from near sea CBL on our C. mayensis skulls are 20.1, 19.7, and 19.6 level to 500 m elevation. He did associate his mm, and tail length ranges from 29 to 31 mm, n = 5 with the pine ridge top which would imply a higher [Appendix]). The cranium is massive when compared elevation than recorded from our specimens of C. with C. tropicalis and the unicuspid teeth are huge, U3 mayensis (Murie 1935). Prior to our collection, there displaces U4 lingually, and U4 is not visible from lateral is only one confirmed whole specimen ofC. mayensis view of the cranium. Our specimens are also among from Belize, a single female collected by Disney on 16 the largest C. mayensis measured to date (N. Wood- February 1966. The animal was collected in lowlands man pers comm.). However, there are few modern at Baking Pot, Cayo District, and was determined by R. museum-quality skins and skulls of this species (many L. Peterson (1968) (Fig. 2). Our collection extends the more are from owl pellets and from archeological and known distribution further south (approximately 100 paleontological sites), so there are few complete skulls km), to higher elevation (600 m) and to include mesic (only 15 total including our specimens) (N. Woodman semi-evergreen rainforest. It is likely more common pers comm.). Cryptotis mayensis differs from C. tropi- and widespread in forests of the Maya Mountains. Orange Walk

8 Occ a s i o n a l Pa p e r s , MBelizeu s e u m o f Te x a s Te c h Un i v e r s i t y

Belmopan San Ignacio

17 S

89 W

Figure 2. Location of whole specimens of Cryptotis shrews collected in/near the Maya Mountains, Belize. Shaded triangle = Baking Pot, C. mayensis (Disney 1966); shaded hexagon = Pine Ridge, C. tropicalis (Murie 1935); shaded star = our 2001 collection near Las Cuevas Field Station, C. mayensis. N

ORDER CHIROPTERA in length. All of our captures were via mist nets set up Family Phyllostomidae in forest edge and rainforest pools generally below 1.5 Carollia sowelli Baker, Solari and Hoffman 2002 m above the forest floor. Silky Short-tailed Bat Artibeus jamaicensis Leach 1821 Specimens examined.—Seven individuals were Jamaican Fruit-eating Bat captured in mist nets (Appendix). Of these, six were adult males and one was an adult female. The lone Specimens examined.—Twenty-five individuals female contained one embryo. Two of the males had were captured, of which 13 were examined and kept measureable testes (6 x 6 mm and 4 x 3 mm). External as voucher specimens (Appendix). Of these, six were characteristics to distinguish C. sowelli from C. brev- adult females and seven were adult males. Three of icauda are poorly developed, but larger overall cranial the six females captured contained one embryo (each) measurements of our animals match those reported for and one showed a placental scar. Of the seven males C. sowelli (Baker et al. 2002; Appendix); thus we refer kept as voucher specimens, four showed mature, active our animals to that species. All of these bats were net- testes (the largest = 9 x 8 mm, the smallest = 3 x 2 mm). ted either in forest edge or at the edge of a rainforest Fifteen of our 25 captures were made in one night using pool. All were netted within one meter of the ground, a mist net set over or near water in primary forest. including two animals captured in the bottom panel of the mist net. Dermanura tolteca (Saussure 1860) Toltec Fruit-eating Bat Sturnira lilium (E. Geoffroy 1810) Little Yellow-shouldered Bat Specimens examined.—One adult male (WFB 4277) with a left testis measuring 6 x 3 mm was col- Specimens examined.—Four adult males and two lected and kept as a voucher specimen. This individual adult females were collected (Appendix). One of the was captured in a mist net set over a pond in primary females was carrying one embryo. Only one of the rainforest. males was examined and its left testis measured 5 mm En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 9

Diagnosis.—This specimen was identified toD. the adults were determined to be reproductively active. tolteca due to its deeply incised interfemoral mem- The males all had scrotal testes and the females were brane, coupled with a forearm of 40 mm and maxillary lactating. Two of these females contained embryos. tooth row of 6.69 mm (Hall 1981). Cranial and tooth Several exhibited sharply defined molt lines. row measurements of this specimen also align well with those reported from and Guatemala Diagnosis.—Two species of Heteromys are re- (Davis 1969). We include this species in the ported from Belize, H. desmarestianus and H. gaumeri Dermanura following genetic relationships separat- (Kirkpatrick and Cartwright 2006). The latter is a ing small Artibeus bats from larger species (Redondo smaller, lowland species restricted to northern Belize et al. 2008). and associated with Yucatan semi-evergreen forest (Schmidt et al. 1989; Bersot 2003). Our specimens Dermanura watsoni Thomas 1901 were assigned to H. desmarestianus based on two key Thomas’s Fruit-eating Bat external characters, lacking hair on the posterior portion of the sole of the hind foot, and possessing a sparsely Specimens examined.—One adult female (WFB haired tail lacking a penicillate tail tip (Schmidt et al. 4244) and one adult male (WFB 4275) were captured 1989). Our measurements, external and cranial, fall and kept as voucher specimens. WFB 4244 contained within the range reported by Rogers and Schmidly one embryo. WFB 4244 was captured in forest edge (1982). near the Las Cuevas Research Station and WFB 4275 was netted at the edge of a primary rainforest pool. Remarks.—This species was the most abundant of the forest floor. It was not captured in open Diagnosis.—This is a small bat with a broad, habitats, but preferred forest interior. Typical localities continuous uropatagia encompassing both legs and included tree falls or debris piles shaded by an over- the terminal edge hairless (unlike D. tolteca). Both story. We captured higher numbers in closed forest specimens lacked a third molar on the maxillary tooth tracts (Aguada site) than recovering logged sites or edge row. Facial stripes were evident on fresh animals; these (Millionario and Monkey Tail) (Fig. 1; Table 1). We were somewhat obscured after the specimens dried. We had no captures at the more recently logged San Pastor include this species in the genus Dermanura following site. These findings might suggest that this species may genetic relationships separating small Artibeus bats be susceptible to logging activities. One specimen was from larger species (Redondo et al. 2008). obtained in the kitchen of the research station. Another partial specimen was found in regurgitation from a Centurio senex Gray 1842 jaguar (Panthera onca) (fresh tracks were associated Wrinkle-faced Bat with the regurgitated rodent).

Specimen examined.—One adult female was Family collected (WFB 4273). No embryos or placental scars Oligoryzomys fulvescens (Saussure 1860) noted. This specimen was captured one meter above Fulvous Colilargo the ground in a mist net set at the edge of a rainforest pool. The characteristic facial pattern and folds quickly Specimens examined.—One adult male was disappeared after the specimen dried. captured in a pitfall trap along Aguada Creek and vouchered (WFB 4230). The animal had scrotal testes ORDER RODENTIA indicating it was reproductively active. Family Heteromyidae Heteromys desmarestianus Gray 1868 Handleyomys alfaroi (J.A. Allen 1891) Desmarest’s Spiny Pocket Mouse Alfaro’s Rice Rat

Specimens examined.—Fifty-three individuals Specimens examined.—Nine males and five were captured of which 18 (7 males and 11 females) females were captured and kept as voucher specimens were kept as voucher specimens (Appendix). All of (Appendix). Three were taken from pitfall traps along 10 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y

Aguada Creek. The remaining 11 were caught in Sher- Sigmodon toltecus (Saussure 1860) man or box traps on the forest floor. All of the adult Toltec Cotton Rat males had scrotal testes. One of the adult females had five embryos and the remaining four adult females were Specimens examined.—Eleven individuals were perforate and contained placental scars. This species captured and retained as vouchers (Appendix). Of was most commonly encountered in damp sites with these, nine were females, five of which either contained intact overstory. Some were associated with dense embryos or placental scars. The two males were both tangles on the forest floor. Formerly placed in the genus scrotal. (Wilson and Reeder 2005), we follow the tax- onomy proposed for Oryzomyine rodents by Weksler Diagnosis.—We examined the literature for ex- et al. (2006) who provisionally assigns this species to ternal characteristics and craniodental measurements Handleyomys, pending further revision. or differences to help confirm the identification of our specimens of Sigmodon; they do not exist. To date only Handleyomys rostratus Merriam 1901 molecular studies have been provided to ascertain the Long-nosed Rice Rat differences between S. toltecus and S. hispidus (Carroll et al. 2005; D. Carroll pers. comm.). We compared Specimens examined.—Five individuals were our small series of Sigmodon from Belize with a series captured and kept as voucher specimens (Appendix). in our collection from Texas, USA, and below pres- Our captures consisted of three adult males and two ent a brief assessment of character differences. The adult females. populations we examined represent two extremes in the geographic distribution of both and there remains Diagnosis.—These animals were easily sepa- a possibility that craniodental and external measure- rable from other Handleyomys and Oryzomys species ments could be clinal; more investigation is needed. by a combination of external characteristics including The Rio Grande River provides the geographic barrier prominent tufts of digital bristles projecting beyond between the two species (Peppers et al. 2002); all of nails of hind toes and orange colored ear hair (both our S. hispidus examined were from north of the river. characters absent in O. cousei). H. rostratus dorsal We found five character traits that helped to distinguish pelage is a brown infused with rich rufous coloration; between toltecus and hispidus: tooth row measure- this contrasts sharply with the white-gray ventral pel- ments, nasal bone extension past incisors, cranium age. O. couesi is paler dorsally, lacks rufous tones and shape, overall dorsal pelage color, and length of rump its ventral coloration is washed with rich buffy tones, spines (Table 4; Fig. 3). blending onto the flanks (this based on comparisons with specimens in the MWFB). Adult H. rostratus is Overall, S. toltecus averaged smaller in nearly all overall smaller than O. cousei. These five specimens measurements but appears to have a somewhat longer were separated from H. alfaroi by their larger size, tail proportionally to the head and body (Table 4). The more rusty dorsal pelage (sepia brown on H. alfaroi) dorsal coloration of toltecus is richer with deep rufous and through cranial features. overtones, absent from S. hispidus. Also the pelage is softer in toltecus, lacking long guard hairs on the rump Remarks.—The three collected males had scrotal for which S. hispidus is named (Fig. 3). The skull of testes and the two females had placental scars, indicat- toltecus, from age class to age class, is shorter, with a ing recent breeding. These individuals were captured higher brain case. The rear of the skull is angled in in a variety of ways and habitats, but all in forest edge. toltecus and more squared in hispidus (Fig. 3). The Two were captured in pitfall traps, one was captured in nasal bones do not project past the incisors as they do the kitchen of the Las Cuevas Research Station, and the with S. hispidus. We noted these traits as consistent remaining two were captured in Sherman traps in semi- differences but these should be vetted with genetic disturbed habitat. For taxonomy we followed Weksler evidence. S. toltecus has been confirmed from Guate- et al. (2006) who provisionally assigns this species to mala approximately 20 km west of the Maya Mountains Handleyomys, pending further revision. (Peppers et al. 2002) En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 11

Table 4. Selected external measurements (in mm; mean in parentheses) and pelage features of Sigmodon collected in Maya Mountains, Belize (WFB specimens) and in Texas, USA (WFB specimens). Refer to Appendix for abbreviations of cranial measurements. Sigmodon toltecus Sigmodon hispidus (Maya Mountains, Belize) (Texas, USA) n =7 n = 14 Tail Length 93 – 114 (141) 93 – 116 (147) Tail % Head and Body 71.3 – 88.2 (76.3) 54 – 81 (71) Hind Foot 26 – 32 (29.6) 27 – 31 (30.3) Ear (notch) 17 – 19 (18.1) 15 – 19 (17.2) GSL 30.5 – 36 (32.4) 31.5 – 38.4 (33.7) CBL 28.3 – 34.6 (30.6) 29.2 – 35.9 (31.6) ZYW 16.6 – 20.8 (18.3) 17.5 – 20.6 (18.6) LIW 4.6 – 5.4 (4.9) 4.4 – 5.3 (4.9) NL 11.3 –12.6 (12.1) 12 – 14.8 (13) M1 – M3 5.4 – 5.8 (5.6) 5.6 – 6.4 (6.1) Distal Nasal Projection even with distal tip premaxilla extends past distal tip of premaxilla Brain case slightly inflated flattened Dorsal pelage sepia with rufous wash, black undercoat light drab sepia, sepia undercoat Belly with rufous wash pale gray and buff Eye ring rufous buff Rump spines 9 – 14 (12) 15 – 25 (20)

Remarks.—Formerly included in the species Sig- of our captures were in traps set in low trees in semi- modon hispidus (Wilson and Reeder 2005), we follow evergreen forest with intact overstory. the phylogenetic relationships proposed by Peppers and Bradley (2000) and Peppers et al. (2002) and con- Ototylomys phyllotis (Merriam 1901) firmed by Henson and Bradley (2009). We captured Big-eared Climbing Rat all of our animals in edge and disturbed habitats, grass being a common habitat element for all captures. We Specimens examined.—Thirty-three animals were did not capture this species in closed forest or second captured of which twenty-three individuals were kept growth habitats. as voucher specimens (Appendix). Of those collected, 13 were males and 10 were females. The captures were Nyctomys sumichrasti (Saussure 1860) equally represented by adults and subadults. All adults Sumichrast’s Vesper Rat were in reproductive condition as determined by scrotal testes (males) and with enlarged nipples and evidence of Specimens examined.—Two individuals (WFB lactation and/or placental scars (females). Ototylomys 4270 and 4272) were captured and kept as voucher were captured equally on the forest floor and in traps set specimens. WFB 4270 was an adult female with pla- on trees and lianas above the ground (up to 2 m above cental scars. WFB 4272 was a subadult male. Both the forest floor). They were most abundant in second 12 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y S. S. hispidus (WFB 6642), lower is lower 6642), (WFB hispidus S. , , not squared anteriorly Nasal Bone projects past incisors past Bone projects Nasal angled of skull Rear (WFB 6642). Lateral skull view: upper is upper view: skull Lateral 6642). (WFB hispidus S. (WFB 4336). (WFB 4336), right is right 4336), (WFB from Texas (WFB 6642) and Belize (WFB 4336) showing pelage and cranial differences. Dorsal view Dorsal differences. cranial and pelage showing 4336) (WFB Belize and 6642) (WFB Texas from Sigmodon S. toltecus S. (WFB 4336), also note smaller tooth row in this image. Study skin lateral view and close-up of rump hairs: upper is toltecus (WFB 6642), lower is S. toltecus Figure 3. Specimens of Specimens 3. Figure is left skulls: of En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 13 growth forest, with some breaks in canopy cover. This ORDER CARNIVORA species was second to Heteromys in abundance. Family Felidae Leopardus pardalis (Linnaeus 1758) Tylomys nudicaudatus (Peters 1868) Ocelot Peter’s Climbing Rat Remarks.—One adult male was captured in a Specimens examined.—Three individuals were large Tomahawk trap and released without measure- captured, of which two (WFB 4237 and 4290) were ments. The ocelot completely filled the trap. Despite kept as voucher specimens. Both were subadult being cramped into such a small place, it was quite males with abdominal testes. The third animal was nimble inside the trap and was formidable to remove. represented by a tail only (which is characteristically Without the availability of tranquilizing drugs, we re- white tipped). The tail was sheared off by a museum leased the cat without taking measurements or assessing special. All three were captured by snap traps set ap- its reproductive condition. proximately two meters above the ground on lianas paralleling tree trunks.

Di sc u ss i o n

At our study site we found relative abundance confirm identification of small mammals. Our efforts dominated by two species, Heteromys desmarestianus add an additional eleven species to the Chiquibul Forest and Ototylomys phyllotis; they accounted for 56% of Reserve checklist, five non-volant: Cryptotis mayensis, total captures. This species pair, or congeners from Oligoryzomys fulvescens, Handleyomys rostratus, Sig- lower altitudes, account for the highest capture rates modon toltecus, Nyctomys sumicrasti, and six volant: across numerous studies in Central America (Disney Carollia sowelli, Sturnira lilium, Artibeus jamaicensis, 1968; Rabinowitz and Nottingham 1989; Caro et al. Dermanura toltecus, D. watsoni, Centurio senex. 2001a, b; Bersot 2003; Klinger 2006). Short-term in- ventories of small mammals in Central America have We believe that two species from previous efforts characteristically shown that in tropical forest habitats, have been erroneously reported from the Chiquabul. total number of captures is often dominated by two spe- Caro et al. (2001b) document two species of Heteromys cies (Fleming 1975; Rabinowitz and Nottingham 1989; from the Chiquibul Forest Reserve, H. desmarestianus Bersot 2003). Biogeographic affinities as documented and H. gaumeri. Our trapping efforts did not yield by the dominance of two typical Central American any specimens of the latter. In Belize, H. gaumeri is rainforest species, coupled with other small mammals relatively common at Shipstern Natural Reserve (200 (Handleyomys, Tylomys, Nyctomys) and observations km to the north in Corozal District) (Bersot 2003). It of Dasyprocta punctata and Mazama temama, indicates is likely that in Belize, H. desmarastinus replaces H. that the mammalian affinities of the Maya Mountains gaumeri in the south and at higher elevations. It is lay nested within Central American faunal group. The unlikely that these species occur sympatrically in the only Yucatan “endemic” found in our study site, Cryp- Chiquibul Forest Reserve; confirmation through vouch- totis mayensis, may not be endemic to the Yucatan, as ers should be obtained in order to confirm the presence our record infers a more widespread geographic and of H. gaumeri in Chiquibul. ecological distribution than previously thought. We recorded two species of Handleyomys from Caro et al. (2001a) report on previous grid- the Chiquibul (H. alfaroi and H. rostratus). The former trapping efforts from the same study site. Using is a rodent of primary forest, the latter of forest edge and research station records, visual reports and trapping clearings. Caro et al. (2001a) also reported two species results, they documented 42 species of mammals from with similar ecological distribution, H. alfaroi from pri- the region. They did not take voucher specimens to mary forests and from forest edge. It 14 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y is likely that the latter represents a misidentification and diversity sampling is the primary goal. Our trap suc- since no vouchers were secured, we cannot confirm the cess is very similar to other transect-based surveys in presences of O. couesi reported for the Chiquibul. The Belize where, in the adjacent Cockscomb Basin Nature status and distribution of O. couesi in Belize is poorly Reserve, Rabinowitz and Nottingham (1989) recorded understood. It was not captured during surveys in the trapping success of 5.23%, and in the Shipstern Nature Shipstern Nature Reserve (Bersot 2003). H. rostratus Reserve Bersot (2003) documented a 3.5% overall trap was the second most common rodent found in the success. nearby Bladen Reserve (Klinger 2006) although it was not recorded by Caro et al. (2001a); we documented Our minor sampling of bats has confirmed several the species at Las Cuevas Field Station. species from the Chiquibul Forest Reserve. Nonethe- less, we believe our sampling of bats underestimates Previous surveys in the area report a very low the diversity present in this reserve, and vespertilionids trap success using grids as the primary trapping method remain poorly documented; more work is needed. (Caro et al. 2001a; Kelly and Caro 2003). We employed node trapping on transects and achieved better results. Accurate biological inventories are central to Using grids, Caro et al. (2001a) captured eight species establishing species baselines and are always important of non-volant mammals with an average of 0.7 ani- in areas where little is known about the resident fauna mals/100 trap nights (total 3,659 trap-nights). Using and flora. Recently, inventories have been given added nodes, we captured 15 species of non-volant mammals impetus because so much of the world is now affected with an average of 4.2 animals/100 trap nights (total by human activities and inventories from relatively 3,686 trap-nights). By coincidence our trapping ef- pristine areas provide a yardstick against which eco- forts were nearly identical. Although we used a more logical change can be measured. This is especially diverse array of traps (ranging from large tomahawks pertinent in subtropical Central American rainforest, a to small museum specials), we believe that the increase habitat that has been heavily degraded and altered, and in trap success is based upon placing transects and in Chiquibul itself where a new hydroelectric dam has nodes in optimal forest sites where clear animal sign been built following an acrimonious debate between was noted. We recommend this method when rapid environmentalists and development groups.

Ac knowledgment s

We thank the Belize Forestry Department for some helpful comments regarding Sigmodon. Douglas permission to work in the Chiquibul Forest Reserve A. Kelt and Irene E. Engilis (University of California, and the Las Cuevas Research Station for logistical sup- Davis) provided helpful comments on the manuscript. port. The following researchers and museums provided Andrew R. Grant (MWFB Student Intern) and Thomas access to specimens and provided technical support: McNamara assisted with small mammal trapping and Linda K. Gordon, USNM (Division of Mammals); Neal specimen preparation. We thank the late Chapal Bol, Woodman, USNM USGS Curator (provided important Barnabas Caro, Emily Fitzherbert, Toby Gardner, critique and determinations on Cryptotis mayensis); Marcella Kelly, and Chris Minty for help in the field Stephen Henshaw and Phil Myers, University of and at the field station. This work was supported by Michigan MNH, provided loans of specimens used funds from the University of California, Davis Center in determinations. Robert Voss (AMNH) provided for Biosystematics Studies and the Selma Herr Fund determinations of Marmosa. Darin Carroll provided for Ornithology. En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 15

Li t e r a t u r e Ci t e d

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Addresses of authors:

An d r e w En g i l i s , Jr. Ti m Ca r o

Museum of Wildlife and Fish Biology Dept. of Wildlife, Fish, & Conservation Biology Dept. of Wildlife, Fish, & Conservation Biology University of California, Davis University of California, Davis One Shields Avenue One Shields Avenue Davis, California USA 95616 Davis, California USA 95616 [email protected] [email protected]

Ro n a l d E. Co l e

Museum of Wildlife and Fish Biology Dept. of Wildlife, Fish, & Conservation Biology University of California, Davis One Shields Avenue Davis, California USA 95616 [email protected] Editor for this manuscript was Robert J. Baker. En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 17 25.5 19.5 13.0 13.0 15.5 19.0 17.5 129.0 140.0 Wt (g) . erupt erupt erupt erupt erupt 8.5 4 4 4 4 4 10.5 MTR 10.37 550 685 730 M M M M M Wt (g) Wt (g) Cryptotis ). . . 3 5.1 5.6 5.81 5.63 5.63 5.55 5.59 14.61 13.91 14.81 M1-M3 M1-M3 M1-M3 (or M 4 . . 9.1 3.3 9.95 3.75 9.98 3.62 3.88 3.84 3.81 3.57 M1-M2 M1-M2 M1-M2 . . NL NL NL 11.5 37.31 23.94 12.11 12.15 38.55 12.26 10.47 18.94 18.28 . 5.2 5.49 5.21 5.01 4.94 6.93 5.46 6.37 LIW LIW LIW 11.27 11.28 10.88 . . 37.4 42.35 ZYW 15.74 43.17 15.54 14.25 15.04 22.15 16.65 23.38 ZYW ZYW . e n d i x CBL CBL CBL 79.65 79.87 27.96 26.25 81.96 27.64 25.12 26.54 38.68 28.72 40.01 pp A 51 46 15 16 47 18 16 17 22 18 26 15 Ear Ear Ear 46 50 18 17 53 16 13 15 22 15 23 16 HF HF Foot . 341 299 311 211 Tail 143 121 144 123 130 134 204 Tail Tail 92 88 93 90 77 85 313 329 335 100 174 178 H&B , Mexican Mouse Opossum H&B H&B Age Age Age SubA SubA SubA SubA SubA SubA SubA SubA SubA SubA Adult Adult F M M M M M M M M M M M Sex Sex Sex Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Material Material Material not erupted. 4 not erupted on either specimen. 4292 4313 4317 4323 4321 4269 4289 4301 4302 4305 4319 4233 4 WFB# WFB# WFB# M Measurements (in mm) for all specimens collected in the Abbreviations Chiquibul for Forest material: Reserve, S+S=skin Cayo District, Belize. Number; MN=Museum measurements: for Abbreviations only. skin=skin only; skel=skeleton only; skull=skull preserved; Alc=alcohol skull; and H&B=Head and Body; HF=Hind Foot; CBL=Condylobasal; ZYW=Zygomatic width; LIW=Least Interorbital Width; M2=Molar 1-Molar 2; M1-M3=Molar 3; MTR=molariform tooth row length, Premolar through Molar NL=Nasal Length; M1- Opossum Virginia 1792 , Didelphis virginiana Kerr Didelphis marsupialis Linnaeus 1758 , Common Opossum M Marmosa mexicana Merriam 1897 Specimen 4289 in collection at Las Cuevas Field Station. 18 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y 255 226 Wt (g) 6 7 7.5 5.5 7.5 Wt (g) . . 19 15 17 14 MTR 14.5 18.5 13.55 Wt (g) PL 8.7 8.8 8.7 . . . 6.8 6.7 7.41 7.29 10.6 10.33 C1-M3 . . M1-M3 5.3 5.4 5.1 MTR . . . 5.8 5.8 5.6 5.46 LIW . . 7.05 7.03 IO 4.6 4.57 4.59 M1-M2 . . . 20.8 CBL 21.33 21.18 20.36. . . ZP 2.4 NL 2.35 2.25 27.72 29.06 . . . 22.7 22.6 GSL 22.37 22.38 . . 7.9 9.3 CB 9.51 7.98 8.98 LIW 39 39 41 41 40 40.4 40.6 . . 29 Forearm CBL 19.58 20.16 28.62 19.69 ZYW 7 5 4 7 8 6 7 . . GSL CBL 58.25 20.89 20.79 57.12 20.76 Tragus 0 0 0 0 0 18 31 17 15 18 31 17 18 18 Ear Ear Ear 11 11 11 11 12 36 12 13 12 35 12 12 14 12 HF HF HF - - - - 7 29 31 29 12 29 31 14 241 256 Tail Tail Tail 64 71 68 71 60 57 58 76 81 63 62 75 218 221 , Yucatan Small-eared Shrew Yucatan , H&B H&B H&B Age Age Age SubA Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult . .. F F M M M M M M M M M M Sex Sex Sex (Merriam 1901) Baker, Solari and Hoffman 2002 , Silky Short-tailed Bat Baker, Alc Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S Skel Skel Skull Material Material Material 4271 4274 4249 4254 4329 4337 4223 4234 4243 4247 4276 4309 4310 4306 WFB# WFB# WFB# , Gray Four-eyed Opossum Philander opossum (Linnaeus 1758) , Gray Four-eyed Cryptotis mayensis ZP=zygomatic breadth; CB=cranial length; skull GS=greatest measurements: Additional Station. Field Cuevas Las at collection in 4274 Specimen plate; IO=interorbital breadth; PL=palatal length. Carollia sowelli Specimens 4254 and 4310 in collection at Las Cuevas Field Station. Additional measurements: C1–M3=tooth row length molar. from canine to last En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 19 36 41 35 44 17 14 12 13 14 49 34 38 41 43 41 48 47 35 13.5 13.5 Wt (g) Wt (g) Wt (g) ...... 10 9.8 10.5 9.93 6.47 6.38 6.69 10.2 10.39 10.29 10.03 10.29 C1-M3 C1-M3 C1-M3 ...... 7.4 8.29 7.37 7.06 5.65 6.81 5.26 7.44 7.48 4.85 7.65 6.98 LIW LIW LIW ...... 17 17.11 16.72 12.89 17.48 12.63 16.43 16.44 12.23 16.88 16.48 17.18 ZYW ZYW ZYW ...... CBL CBL CBL 27.61 27.32 27.21 20.67 27.09 20.22 26.18 26.57 19.32 26.82 26.77 26.55 ...... 27.9 28.2 GSL GSL GSL 28.56 28.67 28.67 21.33 28.56 20.83 27.26 20.28 28.04 28.24 62 58 61 62 39 41 64 38 60 61 64 40 61 65 63 63 61 38.9 37.5 37.1 Forearm Forearm Forearm . . 6 7 6 7 5 4 7 7 7 6 3 7 7 7 6 7 7 7 6 7 Tragus Tragus Tragus 19 21 19 21 16 15 17 18 23 15 13 20 21 21 16 22 21 21 21 22 Ear Ear Ear 11 11 11 11 14 16 15 17 12 17 14 17 18 16 12 16 16 16 16 15 HF HF HF 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Tail Tail Tail 78 84 81 87 56 60 63 55 93 66 54 85 85 86 58 86 76 88 84 87 Toltec Fruit-eating Bat Toltec , H&B H&B H&B Age Age Age Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult F F F F F F F F M M M M M M M M M M M M Leach 1821 , Jamaican Fruit-eating Bat Sex Sex Sex (Saussure 1860) (Saussure Alc Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Skel Skel Skin Skin Skin Material Material Material 4311 4328 4320 4312 4245 4246 4297 4287 4253 4284 4285 4286 4250 4278 4279 4280 4281 4282 4283 4277 WFB# WFB# WFB# , Little Yellow-shouldered Bat Yellow-shouldered 1810) , Little Sturnira lilium (E. Geoffroy Additional measurement: C1-M3=tooth row length from canine to last molar Artibeus jamaicensis Specimens 4282 and 4312 in collection at Las Cuevas Additional Field measurement: Station. C1-M3=tooth row length from canine to last mo - lar. Dermanura tolteca Additional measurement: C1-M3=tooth row length from canine to last molar 20 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y 65 91 58 53 90 62 92 99 74 59 44 98 94 75 60 56 100 49.5 12 Wt (g) 13 Wt (g) 14.5 Wt (g) ...... 6.11 5.25 5.31 5.43 5.34 5.71 5.94 5.84 5.85 5.24 5.82 5.65 5.18 5.14 6.45 M1-M4 C1-M3 C1-M3 ...... 4.25 4.91 4.02 5.02 4.36 4.39 4.57 4.64 4.63 4.79 4.43 4.23 4.92 4.33 4.22 LIW LIW M1-M3 ...... 12.03 14.25 ZYW ZYW 3.34 3.94 3.25 3.48 3.36 3.75 3.59 3.88 3.53 3.49 3.47 3.31 3.33 M1-M2 . 19.4 CBL CBL 16.26 . . . . . NL 15.2 13.7 11.87 17.25 13.71 14.02 12.69 15.42 15.98 15.93 15.41 16.52 16.01 . GSL GSL 20.51 17.82 . . . . . 9.16 9.39 8.52 8.92 9.17 9.57 9.99 8.06 9.96 9.56 9.39 8.76 LIW 10.43 41 39 34.5 . . . . . 18 NT Forearm Forearm 17.9 15.2 17.8 16.59 16.74 16.29 16.04 16.66 17.83 17.15 16.52 ZYW . 6 7 5 . . . . . CBL 30.59 33.87 30.41 28.67 31.01 29.39 31.74 33.55 29.19 32.85 33.66 32.92 30.14 Tragus Tragus 18 19 17 15 17 18 17 15 17 18 19 16 15 19 20 17 17 13 17 17 18 Ear Ear Ear 9 11 36 36 36 10 38 36 35 34 33 35 34 35 31 27 36 36 35 36 34 HF HF HF 0 0 0 171 183 156 132 149 129 180 147 137 173 176 152 131 129 Tail Tail Tail [87] [86] [94] [119] 53 41 56 55 118 116 113 115 118 128 145 128 126 127 136 131 143 155 142 [136] [154] H&B H&B H&B Age Age Age SubA SubA SubA SubA Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult , Desmarest’s Spiny Pocket Mouse Gray 1868 , Desmarest’s F F F F F F F F F F F F F M M M M M M M M Sex Sex Sex Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Skel Skel Skin Material Material Material 4340 4339 4232 4238 4255 4256 4257 4258 4263 4264 4266 4299 4303 4325 4244 4275 4273 4202 4212 4213 4251 WFB# WFB# WFB# , Thomas’s Fruit-eating Bat Thomas’s Thomas 1901 , Dermanura watsoni Additional measurement: C1-M3=tooth row length from canine to last molar Bat Wrinkle-faced Centurio senex Gray 1842 , Heteromys desmarestianus Specimens 4238, 4251, 4299 in collection at Las Cuevas Field Station. En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 21 23 17 26 33 33 21 26 38 41 11.5 27.5 14.5 19.5 19.5 53 55 48 49 Wt (g) 42.5 Wt (g) 9.5 Wt (g) . . 3.8 erupt. 4.6 3.82 3.62 3.86 3.55 3.87 3.62 3.81 3.86 3.74 3.71 3.52 3 3.93 3.77 4.65 M1-M3 M1-M3 M 2.67 M1-M3 . . 3.1 3.08 3.01 3.23 2.85 3.15 3.01 3.02 3.13 3.01 2.93 3.05 3.51 3.02 2.95 3.59 2.79 M1-M2 M1-M2 2.12 M1-M2 . 13 9.9 NL NL 11.4 8.58 9.03 9.65 8.88 9.74 9.33 11.24 11.82 10.71 10.86 13.29 13.07 12.99 12.32 10.71 7.6 NL . 5 5.1 5.2 5.11 4.99 4.85 4.68 5.33 5.29 5.24 4.82 5.42 4.46 5.28 5.02 5.16 4.88 5.03 LIW LIW 3.72 LIW . 11.54 11.45 13.27 12.22 12.35 13.42 14.05 14.45 12.29 16.17 15.13 12.06 16.27 14.55 13.89 12.13 15.08 13.82 ZYW ZYW NT ZYW . . . CBL CBL 20.52 24.35 22.86 21.43 24.91 26.13 26.22 22.77 27.93 27.54 22.27 28.81 27.69 25.38 28.57 24.56 CBL 18.64 14 15 14 15 14 17 16 16 15 20 18 15 15 18 18 17 18 16 14 Ear Ear 12 Ear 23 24 23 25 25 26 25 26 24 30 27 25 31 27 26 24 23 29 24 HF HF 20 HF 75 90 92 89 94 95 90 86 116 119 116 119 118 107 121 142 131 124 131 Tail Tail 91 Tail Fulvous Colilargo , Long-nosed Oryzomys 67 93 84 83 83 93 87 80 84 83 112 105 108 122 128 125 104 125 105 60 H&B H&B H&B Age Age Age SubA SubA Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult F F F F F F F M M M M M M M M M M M M M , Alfaro’s Oryzomys Alfaro’s Allen 1891) , (J.A. Sex Sex Sex Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Material Material Material 4294 4300 4293 4259 4252 4241 4240 4239 4333 4231 4236 4330 4334 4335 4230 4219 4210 4214 4322 4332 WFB# WFB# WFB# (Saussure 1860) , Oligoryzomys fulvescens (Saussure Handleyomys alfaroi Specimen 4334 in collection at Las Cuevas Field Station. Handleyomys rostratus Merriam 1901 22 Occ a s i o n a l Pa p e r s , Mu s e u m o f Te x a s Te c h Un i v e r s i t y 97 81 68 36 79 59 79 111 142 18.5 18.5 Wt (g) 55 53 47 47 60 96 56 45 40 57 31 110 67 21 5.5 5.8 5.7 5.5 5.4 5.6 5.5 erupt. erupt. erupt. 5.79 3 3 3 Wt (g) Wt (g) M1-M3 M M M . . erupt. 4 4 6.66 6.54 6.61 6.76 6.53 6.77 6.34 6.25 6.63 erupt. 3 3.7 3.9 4.1 3.8 3.7 3.6 3.7 3.7 4.49 3 4.16 M1-M3 M1-M3 M M1-M2 M . . 12 12 7.8 9.9 8.6 NL 4.7 11.4 11.3 11.9 12.6 12.4 12.8 4.77 4.84 4.49 4.87 4.61 4.94 4.68 4.55 4.75 3.87 3.49 M1-M2 M1-M2 5 4.9 4.8 4.5 4.7 5.4 4.8 4.6 4.9 4.8 . . 5.03 LIW NL NL 11.61 11.52 10.82 12.86 13.19 14.64 12.78 12.03 12.83 15.26 7.73 10.61 . . 13.9 19.1 17.7 17.1 14.8 13.7 20.8 18.4 16.6 18.4 18.9 ZYW 5.2 5.8 6.2 5.4 5.66 5.55 5.64 5.23 5.98 6.06 LIW 5.72 6.29 LIW . . 23.1 29.8 28.7 26.3 23.3 34.6 30.2 28.3 30.4 32.5 CBL 32.53 17.9 18.2 16.26 16.93 20.34 17.36 16.87 16.46 17.74 19.97 ZYW 14.31 19.58 ZYW . . . 36 . . 31.3 30.5 32.5 30.9 32.3 33.1 GSL CBL 29.45 32.91 31.07 34.41 36.83 32.29 31.47 30.24 32.15 37.75 CBL 25.03 31.65 14 19 17 18 16 18 13 19 18 17 19 Ear 19 22 21 20 22 23 25 23 20 20 28 27 Ear 16 17 Ear 23 31 30 29 27 31 23 31 30 27 29 HF 25 27 26 25 26 28 28 27 26 26 25 27 HF 22 29 HF 64 80 65 93 111 114 112 122 105 101 Tail [25] 99 111 111 110 115 118 101 122 141 120 108 147 Tail 101 Tail [113] 99 151 150 126 126 124 151 171 134 130 152 H&B 73 116 116 110 105 126 120 124 144 152 124 124 165 132 , Big-eared Climbing Rat H&B H&B Age SubA SubA SubA Adult Adult Adult Adult Adult Adult Adult Adult Age Age SubA SubA SubA SubA SubA SubA Adult Adult Adult Adult Adult Adult Adult Adult , Sumichrast’s Vesper Rat Vesper 1860) , Sumichrast’s (Saussure F F F F F F F F F F F F F F F M M M M M M M M M M Sex Sex Sex Merriam 1901 , Toltec Cotton Rat Toltec 1860) , (Saussure S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Skel Material Material Material 4211 4220 4217 4216 4215 4208 4209 4207 4206 4203 4205 4201 4224 4225 4226 4270 4272 4227 4314 4315 4316 4324 4326 4331 4336 WFB# WFB# WFB# Sigmodon toltecus Nyctomys sumichrasti Ototylomys phyllotis En g i l i s e t a l .–Ma m m a l s o f Ma y a Mo u n t a i n s , Be l i z e 23 65 65 35 38 83 50 51 51 55 41 89.5 42 59 Wt (g) Wt (g) . . . erupt. 6.3 6.3 7.32 6.65 6.61 6.39 6.91 3 erupt. erupt. M1-M3 3 3 M M1-M3 M M . . . 4.95 4.77 4.69 4.73 4.33 4.65 4.64 4.43 M1-M2 5.99 5.94 M1-M2 . . . NL 11.97 11.33 13.23 14.25 10.61 10.85 13.54 13.19 NL 11.24 11.36 . . . 5.9 5.3 6.11 6.86 5.57 5.65 5.71 5.44 LIW 7.34 7.86 LIW . . . 17.83 17.23 19.25 16.57 16.83 18.99 17.36 17.97 ZYW 17.39 18.05 ZYW . . . 34.7 CBL 30.82 36.89 30.68 29.99 36.18 32.45 32.28 CBL 32.46 33.18 23 19 20 25 19 19 24 21 27 21 16 Ear 20 21 Ear 28 23 25 26 26 24 26 25 27 24 27 HF 30 30 HF 87 94 112 110 110 131 155 103 164 138 Tail 111 [122] 127 Tail 111 110 114 112 113 143 120 155 122 133 130 134 155 H&B H&B Age Age SubA SubA SubA SubA SubA Adult Adult Adult Adult Adult Adult Adult Adult F F F F F (cont.) M M M M M M M M Sex Sex Alc Alc S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S S+S Material Material 4338 4327 4318 4229 4304 4221 4222 4228 4288 4291 4237 4290 4242 WFB# WFB# Ototylomys phyllotis Specimens 4216, 4242, 4288, 4338 in collection at Las Cuevas Field Station. Climbing Rat nudicaudatus (Peters 1868) , Peter’s Tylomys Pu b l i c a t i o n s o f t h e Mu s e u m o f Te x a s Te c h Un i v e r s i t y

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