Darrin P. Lunde

THE MAMMALS OF PARACOU, FRENCH GUIANA: A NEOTROPICAL LOWLAND RAINFOREST FAUNA PART 2. NONVOLANT SPECIES

Abstract This report describes the results of nonvolant mammal inventory fieldwork at Paracou, a lowland rainforest locality in northern French Guiana, and concludes the faunal analysis introduced by our previous monograph on the bats of Paracou (Simmons and Voss, 1998). Working within a 3–km radius over the course of 202 sampling dates from 1991 to 1994, we recorded a total of 64 nonvolant species by conventional trapping, arboreal platform trapping, pitfall trapping, diurnal and nocturnal hunting, and interviews with local residents. Included in this total species count are 12 marsupials, 9 xenarthrans, 6 primates, 10 carnivores, 5 ungulates, and 22 rodents. Systematic research with nonvolant mammal specimens collected as voucher material resulted in the discovery of new taxa, documented range extensions of previously described species, and helped resolve many longstanding taxonomic problems: (1) Gracilinanus emiliae (Thomas), herein reported for the first time from French Guiana, is redescribed and its known geographic distribution documented; based on examination of type material and original descriptions, G. longicaudus Hershkovitz is considered a junior synonym of G. emiliae, but Marmosa agricolai Moojen is not. (2) A new genus is proposed for Gracilinanus kalinowskii Hershkovitz, a taxon previously known only from eastern Peru, in recognition of its trenchant morphological differences from all other known didelphimorph marsupials. (3) Marmosops parvidens (Tate) and M. pinheiroi (Pine), the latter originally described as a subspecies of the former, are distinct species that occur sympatrically at Paracou; based on examination of type material, other taxa hitherto synonymized with M. parvidens are also judged to be valid species, including M. juninensis (Tate) and M. bishopi (Pine). (4) Monodelphis brevicaudata (Erxleben), M. glirina (Wagner), and M. palliolata (Osgood) are all distinct species diagnosable by unique combinations of morphological traits; based on examined specimens, M. brevicaudata (with type locality emended herein as Kartabo, Guyana) appears to be endemic to the Guiana subregion of Amazonia and to include both bicolored and tricolored phenotypes; a neotype from Cayenne, French Guiana, is designated to fix the application of Viverra touan Shaw as the oldest available name for the tricolored form. (5) Saguinus midas (Linnaeus) and S. niger (E. Geoffroy), currently treated as synonyms or conspecific races, are unambiguously diagnosable species that do not appear to be sister taxa; a neotype is designated to conserve current usage of niger E. Geoffroy for the black-handed tamarin of southeastern Amazonia. (6) Two new small species of Neacomys are described from material collected at Paracou; their diagnostic attributes are documented by detailed comparisons with other like-sized congeners from northern South America. (7) Nectomys melanius Thomas is recognized as a species distinct from N. squamipes (Brants) and N. palmipes J. A. Allen and Chapman; however, N. parvipes Petter is not a valid taxon and is herein synonymized with N. melanius. (8) The diagnostic characters of Neusticomys oyapocki (Petter and Dubost), a species previously known only from the holotype, are reevaluated and illustrated from freshly collected material. (9) Oecomys auyantepui Tate and O. paricola (Thomas), previously treated as synonyms, are valid species distinguished by consistent cranial differences and occupy allopatric ranges north and south of the Amazon, respectively. (10) A critical examination of small Oecomys specimens from Paracou and other Guianan localities supports the conclusions of other investigators that O. rutilus Anthony and O. bicolor (Tomes) are unambiguously diagnosable species. (11) Oligoryzomys fulvescens (Saussure) and O. microtis (J. A. Allen), currently regarded as valid allopatric species occurring north and south of the Amazon, respectively, are difficult to diagnose unambiguously and may be conspecific; new information is provided about the hitherto ambiguous type locality of the latter taxon. (12) Rhipidomys nitela Thomas is reported from French Guiana for the first time and its previously unpublished diagnostic differences from other congeners are tabulated and discussed. (13) A lectotype is designated for Coendou melanurus (Wagner), and the species is redescribed based on all known specimens in North American and European museums; diagnostic differences between this species and C. insidiosus (Olfers) are illustrated for the first time. (14) A red-rumped agouti (Dasyprocta) is designated as the neotype of Mus aguti Linnaeus to preserve current usage of Dasyprocta prymnolopha (Wagler) for the black-rumped agouti. (15) The diagnostic differences between red and green acouchies (Myoprocta) are discussed and a neotype is designated for Cavia acouchy Erxleben to fix the application of that name to the red species; other nominal taxa of Myoprocta are identified as red or green acouchies based on examination of type material and original descriptions. (16) The diagnostic morphological traits of Proechimys cuvieri Petter and P. guyannensis (E. Geoffroy) are reevaluated and discussed based on character variation in topotypical (French Guianan) material. Analyses of our sampling results indicate that distinct sets of nonvolant species are effectively sampled by different inventory methods, and that increased sampling effort with any method generally results in more species. Although the rate of discovery of new species always decreases with increasing sample size, none of our graphs of species accumulation indicate that an asymptotic value was reached with any method. Instead, nonparametric statistical extrapolations suggest that the Paracou nonvolant mammal fauna consists of somewhere between 69 and 74 species; by implication, our nonvolant inventory is about 86–93% complete. Most missing species are probably marsupials and rodents, but one or two expected primate species might have been locally extirpated by hunters prior to our fieldwork. In terms of higher taxonomic composition, the Paracou nonvolant mammal fauna is typical of those found throughout the humid Neotropical lowlands. However, a quantitative analysis of nonvolant faunal similarity at the species level among 12 exemplar rainforest inventories first clusters the Paracou list with others from the Guiana subregion of Amazonia, next with lists from elsewhere in Amazonia, and lastly with Central American lists. Pairwise similarity values likewise show an obvious positive correlation between faunal resemblance and geographic proximity within the Neotropical rainforest biome. At least 24 species (38%) of the Paracou nonvolant fauna are Amazonian endemics, but 18 (28%) are essentially pan-Neotropical in distribution; the remaining 22 species exhibit a variety of distributional patterns that suggest past connections among different sets of currently disjunct rainforested regions. Species richness comparisons among nonvolant faunal inventories are complicated by a variety of familiar problems including inconsistent methodology, presence or absence of certain key habitats, and uneven sampling effort. A conservative interpretation of sampling results from La Selva (Costa Rica), Paracou, and Manu (Peru), however, suggests progressive increases in richness of about 23% from Central America to the Guianas, and of about the same amount from the Guianas to western Amazonia; over the entire gradient (Central America to western Amazonia), the net increase in observed richness is at least 50%. Whereas rodents are consistently the most diverse clade in all well-sampled nonvolant faunas, rankings of other orders by relative richness exhibit considerable site-to-site variation, at least some of which appears to reflect real geographic differences in taxonomic diversity rather than sampling artifacts. Nonvolant rainforest mammals are hard to classify into trophic guilds due to behavioral plasticity and incomplete knowledge of relevant natural history. Preliminary guild comparisons among three exemplar faunas, however, suggest that the Paracou nonvolant community is substantially less diverse in arboreal frugivores and more diverse in terrestrial animalivores than are nonvolant communities at some Central American and western Amazonian sites. Subsistence and recreational hunting has clearly affected local populations of some nonvolant mammals at Paracou; whereas popular game species (e.g., large primates) were seldom sighted, density compensation may explain high local densities of certain other taxa (e.g., Potus flavus and Cuniculus paca). Patterns of differential habitat use between closely related nonvolant species at Paracou were mostly observed within the terrestrial granivore/frugivore guild. Combining these results with those previously reported for the sympatric bat fauna, we recorded a total of 142 mammalian species at Paracou. By statistical extrapolation from our sampling data, the entire local community perhaps contains 155–168 species; because the known French Guianan rainforest mammal fauna contains at least 167 species for which suitable habitat is present in our study area, such estimates are plausible. By implication, our inventory is perhaps 85–92% complete overall. A synthesis of biogeographic information analyzed in this monograph and by Simmons and Voss (1998) suggests that faunal turnover with increasing geographic distance is much higher for nonvolant mammals than for bats, a necessary consequence of observed group differences in endemicity: whereas many nonvolant rainforest mammals have geographic ranges bounded by obvious topographic or habitat discontinuities (e.g., large rivers, xeromorphic vegetation), most rainforest bats are geographically widespread. Not surprisingly, most of the taxa that usefully define a Guianan center of mammalian endemism are nonvolant species. The geographic limits of Guianan endemism appear to be rem

On the contents of Gracilinanus gardner and creighton, 1989, with the description of a previously unrecognized clade of small didelphid marsupials

Abstract Five nominal species of small didelphid marsupials previously referred to Gracilinanus differ conspicuously from the type species (G. microtarsus) and from all of the other valid taxa that we recognize as members of that genus (G. aceramarcae, G. agilis, G. dryas, G. emiliae, G. marica). These anomalous forms can be distinguished morphologically from Gracilinanus (in the strict sense just defined) by lacking maxillary palatal vacuities, a secondary foramen ovale, and a rostral process of the premaxillae; in addition, P3 is taller than P2, and accessory cusps are often present on C1. A new genus, Cryptonanus, is described to contain these forms, all of which are provisionally recognized as valid species: C. agricolai, C. chacoensis, C. guahybae, C. ignitus, and C. unduaviensis. Separate and combined phylogenetic analyses of nonmolecular data and nuclear gene sequences suggest that Cryptonanus and Gracilinanus (sensu stricto) are reciprocally monophyletic and closely related, although they were not consistently recovered as sister taxa in any analysis. Available specimen records document that Cryptonanus is widely distributed in mostly unforested tropical, subtropical, and temperate biomes south of the Amazon River (from ca. 7°S in the Brazilian state of Ceará to ca. 34°S in the Argentinian province of Buenos Aires), but significant range extensions could be expected from pitfall trapping in extralimital savanna landscapes. Scant field data suggest that species of Cryptonanus may often be associated with wet or seasonally inundated grasslands, an unusual habitat for small didelphids.

A Revision of the Didelphid Marsupial Genus Marmosa Part 1. The Species in Tate's ‘Mexicana’ and ‘Mitis’ Sections and Other Closely Related Forms

Abstract We revise the nominal species of mouse opossums currently synonymized with Marmosa mexicana Merriam, 1897, and M. robinsoni Bangs, 1898, which include all of the trans-Andean taxa currently assigned to the nominotypical subgenus of Marmosa. In addition, we redescribe two other species that appear to be closely related to M. mexicana and M. robinsoni based on morphological or molecular citeria: M. rubra Tate, 1931, and M. xerophila Handley and Gordon, 1979. Based on first-hand examination of holotypes and other material (about 1500 specimens in total), we additionally recognize M. isthmica Goldman, 1912, and M. simonsi Thomas, 1899 (both currently synonymized with M. robinsoni), and M. zeledoni Goldman, 1917 (currently synonymized with M. mexicana), as valid species. For each of the seven species recognized as valid herein (M. mexicana, M. zeledoni, M. isthmica, M. robinsoni, M. xerophila, M. simonsi, M. rubra), we describe and illustrate diagnostic external and craniodental characters, tabulate measurement data from adult specimens, list all known examples of sympatry, and map geographic ranges based on specimens examined. The species newly recognized as valid herein, all of which occur in Central America and/or northwestern South America, substantially increase the known diversity of trans-Andean mouse opossums, but it is not currently known whether or not these represent a distinct radiation within the genus Marmosa.

Description of a New Genus and Species of Rodent (Murinae, Muridae, Rodentia) from the Khammouan Limestone National Biodiversity Conservation Area in Lao PDR

Abstract Saxatilomys paulinae, a new genus and species of murid rodent in the Dacnomys Division is described. It is based on two whole specimens and 14 individuals represented by fragments recovered from owl pellets. The samples come from the Khammouan Limestone National Biodiversity Conservation Area in Khammouan Province in central Lao PDR. This tower karst landscape is part of the Quy Dat limestone massif, which extends eastward into north-central Vietnam (Binh Tri Thien Province). The new genus and species is morphologically (and probably phylogenetically) allied to species of Niviventer and Chiromyscus, which are also members of the Dacnomys Division, but its semispinous dark gray upperparts, dark frosted gray underparts, large, extremely bulbous footpads, and a combination of derived and primitive cranial and dental traits exclude it from membership in Niviventer, Chiromyscus, or any other described genus of Indo-malayan murid. The new species is likely petricolous, and is part of a small but unique community of small nonvolant mammals containing the petricolous gymnure, Hylomys megalotis, and hystricognath, Laonastes aenigmamus. All three species have been collected only in forested, rocky habitats of the Khammouan Limestone, but comparable environments in the adjacent Vietnamese portion of the Quy Dat limestone massif may harbor these same species specialized for living in forested, karstic landscapes.

Description of a New Genus and Species of Rodent (Murinae, Muridae, Rodentia) from the Tower Karst Region of Northeastern Vietnam

Abstract Tonkinomys daovantieni, a new genus and species of murid rodent in the Dacnomys Division, is described. It is represented by 14 adults collected from talus habitats in the forested tower karst landscape of the Huu Lien Nature Reserve of northeastern Vietnam. The combination of semispinous, dense, grayish black fur covering upperparts; a dark gray venter; gray ears; a thick, bicolored tail considerably shorter than length of head and body; and large, extremely bulbous footpads is unlike any other species of Indomalayan murid. Body size and build of the new rat, along with some cranial features, are similar to the Thai Leopoldamys neilli, but other cranial traits coupled with molar occlusal patterns resemble morphology in species of the Indomalayan Niviventer, Chiromyscus, and Saxatilomys. The new species is petricolous, includes insects in its diet, and was found only in talus composed of large limestone blocks. Its distribution in the reserve is likely patchy. Whether this limestone rat is restricted to the extensive karst regions of northeastern Vietnam or also occurs in southern China and elsewhere in the northern karst landscapes of Indochina, and Vietnam in particular, will be known only by conducting surveys in limestone regions outside of northeastern Vietnam.

A Review of Peroryctes broadbenti, the Giant Bandicoot of Papua New Guinea

ABSTRACT The giant bandicoot, Peroryctes broadbenti (Ramsay, 1879), is represented in museum collections by 23 specimens collected at 12 localities in the lowlands of the southeastern peninsula (the “Papuan Peninsula”) of Papua New Guinea. Available data on P. broadbenti are reviewed, including its comparative anatomy and morphological variability, taxonomic relationships, geographic and elevational distribution, dietary and reproductive traits, and conservation status. Despite previous confusion between this species and P. raffrayana (Milne-Edwards, 1878), the two species are readily distinguished by a suite of external, cranial, and dental characters. Diagnostic characters are enumerated and illustrated, and comparisons drawn with other New Guinean bandicoots. Generic distinction of Peroryctes Thomas, 1906, in cranial morphology from other New Guinean bandicoots is also reviewed. A striking degree of sexual dimorphism is documented in both body size and dentition for P. broadbenti; these comparisons are set in context by a review of sexual dimorphism among bandicoots in general.

Systematic Reviews of New Guinea Coccymys and “Melomys” Albidens (Muridae, Murinae) with Descriptions of New Taxa

Abstract A new species of the New Guinea endemic murine genus Coccymys is described, based on a small sample from Mt. Dayman and Mt. Simpson in the Maneau Range at the far eastern end of the Owen Stanley Ranges of eastern Papua New Guinea, and two specimens from the western portion of the Owen Stanleys, one from Smiths Gap near Mt. St. Mary, the other from Bulldog Road in the Wau area. Coccymys kirrhos, n. sp., is a vicariant relative of C. shawmayeri, which occurs in the Central Cordillera of Papua New Guinea extending from Mt. St. Mary in the east to the Telefomin region in the west. Coccymys shawmayeri in turn is the eastern montane vicariant of the western New Guinea C. ruemmleri, so far recorded only from the Snow Mountains in western New Guinea (Papua Province of Indonesia) and the eastern end of the Star Mountains over the border in the western section of Papua New Guinea. Coccymys ruemmleri and C. shawmayeri are regionally sympatric in western Papua New Guinea where the former is apparently restricted to high altitudes on the Star Mountains and the latter occurs at lower altitudes in the highlands bounding the Telefomin Valley. The ranges of C. shawmayeri and C. kirrhos, n. sp., overlap at the western section of the Owen Stanley Ranges, and both species have been caught at Bulldog Road, but in different years. This linearly distributed trio of species has been found only in the montane forests and alpine grasslands of the Central Cordillera—there are no records from mountains on Vogelkop Peninsula and the Huon Peninsula, nor from any of the north coast ranges. The new species is described within the context of rediagnosing the genus Coccymys, and documenting morphometric and geographic limits of C. ruemmleri and C. shawmayeri based on most specimens stored in collections of museums. This material consists primarily of museum study skins and accompanying skulls, some fluid-preserved specimens, skeletal fragments from modern samples of owl pellets, and Holocene and Late Pleistocene fossils (for C. ruemmleri only). All species of Coccymys are nocturnal and scansorial; stomach contents from samples of C. shawmayeri indicate the diet consists of seeds, fruit, and arthropods. The taxon albidens is represented by six modern examples collected at 2800 m and 3225 m in 1938 from the northern slopes of the Snow Mountains of western New Guinea, and three Late Pleistocene fossils obtained from the same region. The species was initially described as a Melomys (Tate, 1951); later an alliance with Coccymys was suggested (Flannery, 1990; Menzies, 1990; Musser and Carleton, 1993), but restudy of anatomical traits (derived solely from stuffed skins with accompaning skulls) reveals a degree of morphological divergence not only from species in Coccymys but from any other “Old Endemic” New Guinea murine. A new genus, Brassomys, sampled by only six modern specimens and three Late Pleistocene fossils, is proposed to embrace albidens. Morphological attributes of that species are contrasted primarily with those characterizing Coccymys, and secondarily with the genera Melomys, Paramelomys, Mammelomys, Protochromys, Abeomelomys, and Pogonomelomys. Biological aspects of albidens are unknown; however, particular external, cranial, and dental traits in combination strongly suggest the species is a nocturnal, arboreal/scansorial invertebrate predator.

Chapter 10. Descriptions of New Species of Crocidura (Soricomorpha: Soricidae) from Mainland Southeast Asia, with Synopses of Previously Described Species and Remarks on Biogeography

Abstract The species of Crocidura occurring in southern China, northern Myanmar and Vietnam, are reviewed and defined based on morphology and multivariate analyses. Three new species are described: C. cranbrooki from northern Myanmar, C. annamitensis from the Annamite Mountains of Vietnam, and C. guy from the Viet Bac limestone region in northeastern Vietnam. Enhanced comparative descriptions, distribution, and remarks on taxonomy are provided for each of the other nine species previously reported from the region: C. fuliginosa, C. attenuata, C. sokolovi, C. vorax, C. rapax, C. indochinensis, C. wuchihensis, C. kegoensis, and C. zaitsevi.

Systematic Review of New Guinea Leptomys (Muridae, Murinae) with Descriptions of Two New Species

Abstract Two new species of the endemic New Guinea rodent genus Leptomys Thomas, 1897, are described: L. paulus, indigenous to the montane forests in the Owen Stanley Range in eastern New Guinea, and L. arfakensis, known only from the Arfak Mountains on the Vogelkop Peninsula in western New Guinea. These descriptions are presented within a taxonomic review of Leptomys based on all known specimens in collections of the worlds museums. In addition to the new species, the morphological attributes and geographic distributions of three others are documented. Leptomys elegans occurs in southcentral and southeastern New Guinea both north and south of the Central Cordillera, ranging from the Kikori River Basin in the west to the southern slopes of the Owen Stanley Range, then extending round to the north side of the Cordillera in the Maneau Range, and in the outlying highland block of Mount Victory. Leptomys signatus is recorded only from the Fly and Kikori River drainages south of the Central Cordillera. Leptomys ernstmayri is found in montane forests of the eastern Central Cordillera (Aroa River to Mt. Karimui), reaches to the outlying mountains on the Huon Peninsula and the Adelbert Range, and also occurs in the isolated Foja Mountains of far western New Guinea. Judged by variation in qualitative and morphometric external, cranial, and dental traits associated with available samples, L. arfakensis, n. sp., is likely more closely related to L. elegans and L. signatus, all with large bodies and relatively short tails, than to L. ernstmayri and L. paulus, n. sp., that are characterized, among other features, by smaller body size and relatively longer tails. There is significant phenetic divergence among the geographic samples identified here as L. ernstmayri, and additional inquiry, especially utilizing data from molecular sources, is required to determine whether this intersample variation reflects the presence of separate species, each endemic to the Huon Peninsula, Adelbert Range, Central Cordillera, and possibly the Foja Mountains, or instead represents montane variation within a single morphologically variable species.

A survey of small mammals from Huu Lien Nature Reserve, Lang Son Province, Vietnam

ABSTRACT A survey of the small mammals of Huu Lien Nature Reserve, Lang Son Province, Vietnam (21°40′52″N, 106°20′28″E) resulted in the capture of 187 specimens representing 21 species of bat, insectivore, and rodent: Suncus etruscus, Rhinolophus pearsonii, Rhinolophus pusillus, Rhinolophus stheno microglobosus, Aselliscus stoliczkanus, Hipposideros armiger, Hipposideros larvatus, Harpiocephalus harpia, Pipistrellus paterculus, Scotomanes ornatus, Callosciurus erythraeus, Dremomys rufigenis, Niviventer fulvescens, Niviventer langbianis, Tonkinomys daovantieni, Rattus andamanensis, Rattus losea, Rattus tanezumi, Mus caroli, Rhizomys pruinosus, and Atherurus macrourus. In addition Ratufa bicolor and Tamiops sp. are reported from observations. The recently described genus and species Tonkinomys daovantieni is currently known only from material collected during the present survey and is apparently endemic to karst habitats in northeastern Vietnam.