![Morphometric separation (principal component analysis) of intact crania representing New Guinea moss-mice. Principal components extracted from a covariance matrix of 10 log-transformed craniodental variables (table 1) separate five highly distinctive phenetic groupings of New Guinea rodents, corresponding to the generic concepts of Pseudohydromys, Neohydromys, Mayermys, Microhydromys (minus “M.” musseri; see Helgen et al. [in press] and account in this paper), and a newly diagnosed genus (Mirzamys). The x-axis (PC1) is largely a shape factor that portrays a decrease in relative size of both M1 and mesopterygoid fossa width from left to right; the y-axis (PC2) is largely a size factor that portrays an overall increase in skull size from top to bottom. In this plot, as in other morphometric plots figured in this paper, ellipses surrounding each cluster are fitted by eye to indicate cohesion, and do not reflect statistical confidence intervals.](https://www.researchgate.net/profile/Kristofer-Helgen/publication/232694538/figure/fig3/AS:267630800011302@1440819564056/Morphometric-separation-principal-component-analysis-of-intact-crania-representing-New_Q320.jpg)
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Chapter 8. Biodiversity and Biogeography of the Moss-mice of New Guinea: A Taxonomic Revision of Pseudohydromys (Muridae: Murinae)
Abstract and Figures
Morphological investigations involving nearly all available museum material representing New Guinea “moss-mice” (rodents traditionally classified in the genera Pseudohydromys, Neohydromys, Mayermys, and Microhydromys) reveal outstanding undiagnosed taxic diversity (a minimum of 16 species, versus the eight species previously described) and allow for redefinition of generic boundaries among these little-studied rodents. Apart from Microhydromys Tate and Archbold, 1941 (comprising two species, as recently revised by Helgen et al., in press), herein we recognize two genera of New Guinea moss-mice: Pseudohydromys Rümmler, 1934 (now incorporating Neohydromys Laurie, 1952, Mayermys Laurie and Hill, 1954, and “Microhydromys” musseri Flannery, 1989) and a newly described genus, Mirzamys. Species of Pseudohydromys are recorded from montane areas throughout New Guinea (elevations spanning 600 to at least 3800 meters), including the mountain ranges of the Central Cordillera, the Huon Peninsula, and the North Coastal ranges. We diagnose and review 12 species of Pseudohydromys, including six species described as new. The new genus Mirzamys is erected to accommodate two newly described species of small terrestrial rodents from middle and upper montane forests and subalpine grassland edges (1900–3450 m) in the mountains of central New Guinea. Together these two new species represent a distinctive hydromyin lineage that resembles the terrestrial New Guinea hydromyin genera Pseudohydromys and Paraleptomys in various traits. Ecological attributes of all recognized moss-mice taxa, both previously and newly described, are reviewed in light of all information currently available about their biology.
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... In this study, we describe a new genus and species of large-bodied murine based on mandibular remains recovered at Liang Bua in mostly Holocene sediments. This taxon shares various morphological similarities with insectivorous but phylogenetically unrelated murine rodents that evolved independently on Sulawesi, the Philippines, and New Guinea, colloquially known either as "shrew rats" or "moss mice" (Rowe et al., 2016;Helgen & Helgen, 2009). Shrew rats and Based on information in Musser (1981), Musser & Boeadi (1980), Kitchener et al. (1991a,b), and Suyanto (1998). ...
... 75 moss mice are terms associated with murines that generally inhabit tropical evergreen forests, adopt a carnivorous diet, and occupy a range of ecological niches usually filled by shrews. These murines sometimes evolve traits reflecting these dietary niches, including a reduced number of molars, simplified occlusal pattern, longer and narrower snout, larger and more complex olfactory turbinals, small molar size relative to the size of mandible and maxilla, and proodont incisors reflecting an insectivorous or vermivorous diet (Musser, 1982;Musser & Heaney, 1992;Musser & Durden, 2014;Helgen & Helgen, 2009;Martinez et al., 2018;Charles et al., 2013). This adaptation repeatedly appears in the highly diversified Murinae with at least six major carnivorous lineages found within (1) the Echiothrix Division on Sulawesi, (2) the genus Crunomys found on the Philippines and Sulawesi, (3) the Chrotomys Division on the Philippines, (4) the Hydromys Division from Australo-Papua, (5) two species within the genus Mus, and (6) the Praomys Division in Africa (Supporting Information Appendix Table 1) (Heaney et al., 2016;Rickart et al., 2019;Musser & Durden, 2002;Esselstyn et al., 2012;Esselstyn et al., 2015;Helgen & Helgen, 2009;Rowe et al., 2016). ...
... These murines sometimes evolve traits reflecting these dietary niches, including a reduced number of molars, simplified occlusal pattern, longer and narrower snout, larger and more complex olfactory turbinals, small molar size relative to the size of mandible and maxilla, and proodont incisors reflecting an insectivorous or vermivorous diet (Musser, 1982;Musser & Heaney, 1992;Musser & Durden, 2014;Helgen & Helgen, 2009;Martinez et al., 2018;Charles et al., 2013). This adaptation repeatedly appears in the highly diversified Murinae with at least six major carnivorous lineages found within (1) the Echiothrix Division on Sulawesi, (2) the genus Crunomys found on the Philippines and Sulawesi, (3) the Chrotomys Division on the Philippines, (4) the Hydromys Division from Australo-Papua, (5) two species within the genus Mus, and (6) the Praomys Division in Africa (Supporting Information Appendix Table 1) (Heaney et al., 2016;Rickart et al., 2019;Musser & Durden, 2002;Esselstyn et al., 2012;Esselstyn et al., 2015;Helgen & Helgen, 2009;Rowe et al., 2016). The identification and description of an endemic shrew rat from Flores not only reveals a greater species and ecomorphological diversity on the island and in Wallacea, but may also contribute towards understanding past and current anthropogenic impacts on murine extinction. ...
... There have been no previous reports of helminths from M. obiensis. The nematodes Heterakis fieldingi Smales, 1996, Protospirura kaindiensis Smales, 2001 and an Odilia sp. have been reported from P. murinus, as has Rictularia mackerrasae Mawson, 1971 from P. murinus reported as Neohydromys fuscus Laurie (see Smales, 2001Smales, , 2006Helgen & Helgen, 2009). Encapsulated ascaridid larvae and a heligmonellid, tentatively identified as Odilia emanuelae (Mawson, 1961), have been reported from Pseudohydromys germani (Helgen) as Mayermys ellermani Laurie & Hill (see Smales, 2006;Helgen & Helgen, 2009). ...
... The nematodes Heterakis fieldingi Smales, 1996, Protospirura kaindiensis Smales, 2001 and an Odilia sp. have been reported from P. murinus, as has Rictularia mackerrasae Mawson, 1971 from P. murinus reported as Neohydromys fuscus Laurie (see Smales, 2001Smales, , 2006Helgen & Helgen, 2009). Encapsulated ascaridid larvae and a heligmonellid, tentatively identified as Odilia emanuelae (Mawson, 1961), have been reported from Pseudohydromys germani (Helgen) as Mayermys ellermani Laurie & Hill (see Smales, 2006;Helgen & Helgen, 2009). ...
... Host terminology follows Helgen & Helgen (2009). Classification of the Heligmonellidae and the terminology of the synlophe ridges and bursal rays follow Beveridge et al. (2014), Durette-Desset & Digiani (2012, 2015 and Digiani & Durette-Desset (2014). ...
Bisastrongylus multiovorumgen. nov., sp. nov. (Nematoda: Heligmonellidae: Nippostrongylinae) is described from a murid, Melomys obiensis, from the Moluccas, Indonesia. The new genus differs from all other Nippostrongylinae genera in having a synlophe of 12 unequal ridges with a type A carene, the dorsal ridge 1 being larger than the ventral ridge 1'. Pentastomid larvae, an acanthocephalan, Plagiorhynchus sp., and the nematodes Rictularia sp., a spirurid and the nippostrongylin Soricstrongylus obreensisgen. nov., sp. nov. were collected from the murid Pseudohydromys murinus from Central Province, Papua New Guinea. The new genus is distinguished by a combination of characters including a synlophe of 11–13 ridges at mid body, left ventral ridges larger and a sub frontal axis of orientation. The assemblage of P. murinus is discussed and a re-evaluation of nippostrongylin Odilia sp., previously reported from P. murinus indicates that the specimens were likely a Parasabanema sp.
... Whereas there are, in Australia, around 60 species of native rodents in the family Muridae that make up around 20% of the current extant mammalian fauna, the number of species of mice and rats in New Guinea is still not known in detail 304 Records of the Australian Museum (2020) Vol. 72 although there appears to be at least 100 species (Tate, 1951;Flannery, 1995aFlannery, , 1995bMusser & Carleton, 2005;Aplin, 2006;Musser et al., 2008;Musser & Lunde, 2009;Helgen, 2005aHelgen, , 2005bHelgen & Helgen, 2009;Helgen et al., 2010;Aplin & Ford, 2013;Rowe et al., 2008Rowe et al., , 2019Roycroft et al., 2020) with only a few species occurring in both Australia and New Guinea. ...
... 303). In more recent decades, New Guinea murines have been subject to considerable clarifying taxonomic review and revision (e.g., Flannery, 1995aFlannery, , 1995bMenzies, 1996;Musser & Carleton, 2005;Helgen, 2005aHelgen, , 2005bMusser et al., 2008;Helgen & Helgen, 2009;Musser & Lunde, 2009;Helgen et al., 2010), and murine tribes and divisions, including those of the New Guinea species, have been investigated extensively using molecular phylogenetic methods (e.g., see Lecompte et al., 2008;Smissen & Rowe, 2018;Rowe et al., 2019;Roycroft et al., 2020). In the current study, we follow the systematic arrangement as detailed by Roycroft et al. (2020) which includes the following five divisions and two tribes in New Guinea and adjacent islands. ...
... Data for number of fetuses of dissected specimens at the Australian National Wildlife Collection, Canberra (ANWC) and the South Australian Museum, given with specimen numbers (e.g., 2F-= 2 fetuses). Dwyer, 1975;Flannery, 1995aFlannery, , 1995bHelgen & Helgen, 2009;Helgen, 2005aHelgen, , 2005bHelgen, 2007;Helgen et al., 2010;Jackson & Woolley, 1993;McPhee, 1988;Menzies, 1996;Musser & Lunde, 2009;Musser et al., 2008;Olson, 1982;Taylor & Horner, 1973;Taylor et al., 1990;and Woolley & Furby, 1996. being 2.5-3.5 µm (Figs 5b, 6b-d; Table 2) although a slightly longer apical hook and ventral processes of 5 µm was present in Uromys caudimaculatus (Fig. 6e) whereas the sperm tail ranged from 90-130 µm (see Table 2). ...
In New Guinea there are around 100 species of native rodents in the family Muridae that are members of two tribes, the Hydromyini and Rattini, and five divisions—the Hydromys, Uromys, Mallomys, Pogonomys, and Rattus divisions. Here we review their basic reproductive biology so far as it can be determined from the material available. We find that females of most species in the Hydromys and Uromys divisions have 4 nipples, most species in the Pogonomys division have 6, whereas in the Mallomys division nipple number across species ranges from 2 to 6, and in the Rattus division from 4 up to 12. The number of fetuses observed in pregnant individuals in species of all of the hydromyine divisions was generally between 1 and 3 but in three species in the Rattus division up to 6, or even occasionally more, occurs. In males, the relative testes mass (RTM) of most species in the Hydromys, Uromys, Mallomys, and Rattus divisions was usually between 1 and 3% of body mass, whereas in the Pogonomys division it varied markedly from only around 0.4% in Hyomys goliath up to 5% in two species of Pogonomys. The spermatozoa of species in the Hydromys and Uromys divisions, like in the Australian species of these divisions, contained a head with an apical hook together and two ventral processes, whereas in the Pogonomys and Mallomys divisions marked interspecific differences occurred with some having a sperm head with an apical hook and ventral processes but in others there were no ventral processes but a long apical hook. Sperm tail length of most species was generally between 90 and 130 µm but Chiruromys and Xenuromys had sperm tail lengths of 150–153 µm. Male accessory sex glands were generally similar across the species except for that of the preputial glands which appeared to be absent in species of Pogonomys and Chiruromys but very large in Hyomys. The findings of large relative testes mass in Pogonomys and long sperm tails in Chiruromys and Xenuromys suggest selection for high levels of intermale sperm competition and hence multimale breeding systems in these species, whereas the variation in preputial gland size suggest interspecific differences in social organization.
... The mtDNA data also suggests an early split within D. goodfellowi, resulting in the large genetic divergence between the morphologically similar D. g. goodfellowi and D. g. beurgersi in eastern Papua New Guinea. This is likely to coincide with major river valleys that dissect the Central Cordillera in south-east New Guinea and appear to form major contemporary barriers to gene flow in Dendrolagus (Eldridge et al., 2018) and other groups of mid-high evaluation taxa including birds (Irestedt et al., 2009;Irestedt et al., 2015), other mammals (Meredith et al., 2010;Westerman et al., 2012;Helgen & Helgen, 2009) and insects (Janda et al., 2016;Toussaint et al., 2014). ...
Tree-kangaroos (genus Dendrolagus) are a morphologically distinctive genus of specialized, arboreal macropodids confined to the wet forests of New Guinea and northeast Australia. A distinct Goodfellow’s group, containing up to four species, has long been recognized. Resolving the relationships of taxa within the group has been hampered by limited samples of most taxa. Here we supplement published genetic data from high quality tissue samples with molecular data generated from museum specimens to improve taxon and geographic coverage. This includes specimens of the previously unsampled D. g. goodfellowi, the holotype and paratype of D. deltae, and additional specimens of D. matschiei, D. spadix and D. g. buergersi. DNA sequence data were generated from three mitochondrial loci. Phylogenetic analysis improved the resolution of relationships within the Goodfellow’s group, with the morphologically similar D. g. goodfellowi and D. g. buergersi being recovered as sister taxa, while D. pulcherrimus was the sister to the closely related, but morphologically and ecologically distinct, D. spadix and D. matschiei. Despite being sister to D. g. buergersi, D. g. goodfellowi was highly divergent. However, the two are morphologically very similar and we recommend retaining the taxonomic status quo (recognizing them as two subspecies of a single species) until improved sampling and a more thorough analysis is possible. The problematic D. deltae was confirmed as a junior synonym of D. matschiei.
... However, the morphological configuration of H. bellwoodi-combining an elongate lower jaw diastema, low angled lower incisor and relatively small molars for the size of the jaw-is reminiscent of some species of the Sulawesian genus Bunomys (e.g., B. chrysocomus) and some New Guinean species of Rattus (e.g., R. verecundus), and seen in more extreme expression in New Guinean species of Leptomys and Paraleptomys (Taylor et al., 1982;Musser, 1991Musser, , 2014Musser et al., 2008), for example. All of these taxa are terrestrial animalivores, feeding on snails, earthworms and insects, and perhaps in some cases, small vertebrates, and their morphological adaptations are plausibly related to the demands of this lifestyle (Musser & Heaney, 1992;Helgen & Helgen, 2009;Musser & Durden, 2014;Musser, 2014;Rowe et al., 2016;Veatch et al., 2023). In contrast, the short lower jaw diastema and larger molars of H. funderus are more reminiscent of taxa with a herbivorous diet, and in particular, of species known to feed primarily on fruits or large seeds that require heavy gnawing activity to penetrate tough skin or nut casing followed by heavy mastication to process fibrous flesh or pith (e.g., Samuels, 2009). ...
... They host rodent communities with closely related species often inhabiting different elevation zones (Roycroft et al., 2022). Elevation-related morphological changes were already reported in several species of Pseudohydromys (Helgen & Helgen, 2009). Also, the elevational gradients in Andes host a rich fauna of sigmodontine rodents (Parada et al., 2015), including genera with different elevation specialists (D'Elía et al., 2008;Steppan, 1998). ...
Several rodent genera radiated in Ethiopian highlands during Pleistocene, giving rise to species specialized to different elevation zones. The zonation is accompanied by vegetation change from savannahs or forests in lower elevations, through the mid-elevation belt of ericaceous shrublands to Afroalpine moorlands on the mountain tops. Such evolution on elevation gradient is expected to promote phenotypic adaptations and several genera are known to include highly distinctive high-elevation specialists. In this study, we systematically examined shape and size differentiation of skull and mandible in three genera, Lophuromys, Arvicanthis and Stenocephalemys, each including from six to eight species inhabiting different elevation zones on both sides of the Great Rift Valley. Employing computed tomography, three-dimensional virtual models, landmark-based geometric morphometrics and a powerful toolkit of partial least squares analysis, we demonstrated the elevation-related differentiation is a prominent feature of skull and mandible shape variation in all three genera. We identified the parts of skull and mandible which change most between the successive zones. These include, notably, orbits and insertion areas of masticatory muscles, which suggests adaptive shifts in visual perception and diet. We also showed that while the same parts are modified in all three genera, they are modified in different ways and to a different degree. The evidence for size differentiation was mixed despite the universal trend of larger size in higher elevations. Our study confirms the high-elevation specialists represent the most distinctive forms, which is especially true for the species from Bale and Arsi Mountains, East of the Great Rift Valley.
... Wam is a noun prefix applied to names of terrestrial mammals; Nok is a noun prefix applied to names of bats. Bonaccorso 1998) provides an overly simplistic caricature of the complexity of New Guinea mammal diversity, as renewed detailed systematic reviews continue to demonstrate (e.g., Helgen and Flannery 2004, Woolley 2005, Helgen 2005b, 2007a, Musser et al. 2008, Musser and Lunde 2009, Helgen and Helgen 2009, Parnaby 2009). Currently unnamed or unrecognized biological species are known in almost every Melanesian mammal genus (Helgen 2007a, 2007b. ...
We undertook a survey of mammal diversity and indigenous knowledge of mammals in Hewa country, in the vicinity of Wanakipa village, Southern Highlands Province, during July 2008. Specimens collected during this survey document the occurrence of 22 mammal species in the immediate area, including one monotreme, 6 marsupials, 7 rodents, and 8 bats. Two species of conservation concern, the Eastern Long-Beaked Echidna Zaglossus bartoni (IUCN Critically Endangered) and Goodfellow's Tree Kangaroo Dendrolagus goodfellowi (IUCN Endangered) were documented by trophy skulls and bones kept by Hewa hunters. Three mammals are apparently reported from Southern Highlands Province for the first time: the bent-wing bat Miniopterus macrocneme, the long-eared bat Nyctophilus microtis, and a rodent species in the Rattus rattus species complex. The presence of this last species, an invasive rodent trapped distant from an immediate village commensal context, provides an interesting indication of recent environmental change in the highlands of Papua New Guinea. Hewa informants convincingly communicated their familiarity with more than 20 additional mammal species, including those restricted to higher elevations than we visited during our survey, including one of special conservation concern, the tree kangaroo Dendrolagus [dorianus] notatus (IUCN Endangered). Hewa folk taxonomy suggests that three tree kangaroo species may occur in the region, one of which may be the Lowland Tree Kangaroo (Dendrolagus spadix) or even a currently undocumented population or unrecognized species. The mammal fauna of Wanakipa appears similar in composition to better surveyed areas of similar elevation situated along the southern margin of the central cordillera in Southern Highlands Province, such as Mt. Sisa and Mt. Bosavi.
INTRODUCTION
The word “Hewa” is a somewhat generic term that refers to a group of shifting cultivators who speak one of the Sepik Hill Stock languages of the Sepik Ramu Phylum (Lewis 2009). They number fewer than 2,000 people and inhabit roughly ca. 65,000 hectares of hilly and sub montane forest in the uppermost catchment of the Strickland River. A wall of limestone cliffs that rise abruptly to over 2,300 m in the south effectively separates the Hewa from the Duna, Paella, and Ipili cultures. As is often the case with people living on the margins of larger societies, the “Hewa” answer to several different names given to them by their more powerful neighbors. They are the Hewa to the Duna, but the Sismen to the Min speakers on the western side of the Strickland River. Those living around the village of Wanakipa now typically refer to themselves as Hewa.
Unlike their highland neighbors, the Hewa do not occupy fertile valleys and are instead scattered throughout the mountains. They prefer to cut their gardens at elevations between 500–1,000 meters and traditionally it was this band of forest that experienced the greatest disturbance from cultivation. The village of Wanakipa is a recent phenomenon. Although Wanakipa has been on the map as a government station since Papua New Guinea's independence in 1975, when Thomas first visited the site in 1988 only four families were living at the station. There was no school or medical aid post. At that time, the Hewa considered the site to be too hot and malarial, preferring to establish their homes at higher elevations. Their attitudes changed in 1990 with the intervention of the Lutheran Mission and the construction of an airstrip. The airstrip enabled the mission to establish and supply a medical aid post. Access to medicine, though variable, has attracted the Hewa to this formerly undesirable site. Today, Wanakipa has become a village, complete with an aid post, school, and weekly market.
However, outside of Wanakipa, the majority of the Hewa continue to live in scattered households. Their rugged environment and low density settlement pattern has discouraged road building and makes it difficult to provide government services. In these more traditional circumstances, Hewa infant mortality and life expectancy are likely to approach pre-contact rates (Gillett 1991:22). Unlike their highland neighbors, the Hewa do not grow coffee or participate significantly in the modern cash economy. They remain subsistence-oriented horticulturists and traditional environmental knowledge (TK) is still an important aspect of their culture.
Here we summarize our combined efforts, working in association with Hewa naturalists, to characterize the mammal fauna living in the vicinity of Wanakipa during the July 2008 Conservation International Rapid Assessment biodiversity Survey to the upper Strickland Basin.
MATERIALS AND METHODS
Data collection and sampling methods
Because the majority of Melanesian mammals can only be reliably identified by comparison with series of museum specimens (Flannery 1995, Bonaccorso 1998, Helgen 2007a), scientific efforts to document mammalian biodiversity in New Guinea require the collection and long-term preservation of voucher specimens. We obtained voucher specimens on the 2008 RAP survey by collecting “trophy” skulls and other bones retained by Hewa hunters, and by a combination of live-trapping (Sherman traps) and lethal trapping (snap traps) for small ground mammals, and mist netting of bats.
Protocols for capture and handling of mammals followed standard guidelines established by the American Society of Mammalogists for animal care and use (Gannon et al. 2007). For each specimen prepared as a museum voucher, standard external measurements were taken with a ruler (total length, tail length, hind foot length with and without the claws, ear length, and in the case of bats, forearm length) and Pesola scales (body mass). The sex and maturity of each specimen were assessed in the field, and microhabitat and other ecological data were noted for each specimen wherever possible. Vouchers were prepared as fluid preparations (fixation in 10% formalin, then transferred to 70% ethanol for long-term storage) or as study skins and/or skeletons. For each freshly collected specimen, liver, kidney, and/or muscle tissue was preserved in 95% ethanol for subsequent genetic analyses. Some small mammals were photographed in life in semi-natural settings. Specimens from the Wanakipa/Tualapa RAP survey are deposited in the Division of Mammals at the National Museum of Natural History (USNM), part of the Smithsonian Institution in Washington, D.C., along with their associated temporal, geographical, ecological, and mensural data, and genetic samples.
Common names generally follow Flannery (1995) or (for bats) Bonaccorso (1998), but in a few cases (among rodents) where taxonomic changes have necessitated a change in vernacular usage, common names follow Musser and Carleton (2005). Conservation classifications provided for each species (Critically Endangered, Endangered, Least Concern, etc.) derive from the current rankings on the IUCN Red List (www.iucnredlist.org), most recently revised as part of the Global Mammal Assessment, an effort by mammalogists worldwide to summarize the current state of knowledge individually relevant to the conservation of every living mammal species (Schipper et al. 2008).
Study sites
The mammal survey focused on three sites within several hours' walking distance of the village of Wanakipa (airstrip at 05°15.425′S, 142°31.297′E, situated at 807 m) in Southern Highlands Province (Table 21.1):
Tualapa (July 11–23, 2008)
A camp and study site was established at Tualapa (05°17.003′S, 142°29.849′E), situated at 1,115 m, in a setting of kunai grassland and forest habitats (see Takeuchi, Chapter 9, this volume).
We established a trap-line in forest and kunai grass for ‘removal trapping’ of small mammals. Fifty medium-sized Sherman live traps and 25 ‘museum special’ snap traps were set each night, for 10 nights, on the forest floor and in vegetation and low trees up to 3 meters above the ground. These traplines resulted in 21 catches (= 3% trap success) representing five rodent species (Melomys lutillus, Melomys rufescens, Rattus exulans, Rattus foersteri, Rattus ‘rattus’; see species accounts below). Two 60 meter pitfall-lines were established and at each line we set 10 pitfall traps, approximately 60 centimeters deep and 5 meters apart, and monitored them for 6 nights. Pitfall traps collected only the small rodent Lorentzimys nouhuysii (see species account below). Mist-nets set by day for catching birds were also monitored at night for bat-catching. Ten nights of netting yielded series of Nyctimene papuanus, Paranyctimene sp., and Syconycteris australis, and a single specimen of Macroglossus minimus (see species accounts below). Two harp traps were set up along enclosed forest trails for a total of 10 trap nights, but no bats were captured in the harp traps.
Mammals were also documented at Tualapa by working with Hewa hunters. Hunters sourced series of the bats Miniopterus macrocneme and Miniopterus magnater from a cave near Tualapa, and speared a specimen of the large rat Uromys caudimaculatus (see species accounts below). We also obtained “trophy” jaws and crania of various game animals from hunters at Tualapa, some of which were likely brought from the broad vicinity of the village of Wanakipa, including Zaglossus bartoni, Echymipera kalubu, Echymipera rufescens, Dendrolagus goodfellowi, Phalanger gymnotis, Phalanger mimicus, Spilocuscus maculatus, Uromys caudimaculatus, and Dobsonia moluccensis.
Table 21.1. Mammal species documented by specimens at Wanakipa stations. ‘Trophy’ refers to specimens collected as trophy material only. Other records reflect animals collected during the survey. Greatest effort was invested in surveying mammals at Tualapa (see text).
Umge (July 15–17)
A “Fly Camp” above Tualapa was set up at Umge (05°18.245′S, 142°30.704′E), situated at 1,438 m in montane forest (see Takeuchi, Chapter 9, this volume). Helgen visited this camp briefly while Opiang and Thomas stayed at Tualapa. Two nights of trapping with 10 Sherman and 10 museum-special traps yielded no captures. One night of mist-netting with two mistnets around a fruiting fig yielded series of Syconycteris australis and Paranyctimene sp.
Putuwe (July 23–26)
At the close of our survey we camped for 3 nights at a site named Putuwe (05°13.867′S, 142°31.933′E), at 570 m, situated at the junction of the Lagaip and Uruwabwa rivers, amongst village homes and gardens. Mist-nets set by day for catching birds were also monitored at night for bat-catching, resulting in series of Nyctimene papuanus and Syconycteris australis, and a single capture of Nyctophilus microtis. Specimens of Rattus exulans were collected in village homes and Melomys lutillus was collected in a sweet potato field. Two specimens of the bandicoot Echymipera kalubu killed by village dogs were salvaged as museum specimens (see species accounts for all, below). Pitfall traplines were also set up (as at Tualapa) for two nights, but yielded no captures.
RESULTS
Annotated species list
Order Monotremata, Family Tachyglossidae (Echidnas)
Zaglossus bartoni (Thomas, 1907) (Eastern Long-Beaked Echidna)
IUCN Red List Status: Critically Endangered: “Listed as Critically Endangered due to a suspected continuing population decline of at least 80% over the last three generations (i.e., the last 45–50 years) based on direct observation in parts of its range, declines in area of occupancy (reports from hunters), and actual levels of exploitation due to hunting.” (Leary et al. 2008a).
This largest of extant monotremes (5–10 kg) occurs in forests and upland grasslands from sea level (albeit rarely so low) to an elevation of at least 4,150 m along the length of New Guinea's central cordillera and in the outlying mountain ranges of the Huon Peninsula (Flannery and Groves 1998). It is now rare across much of its geographic range, probably because of widespread and intensive human hunting with the aid of dogs, and it remains common today only in areas of low human population density (George 1978, Flannery 1995). The subspecies in the Hewa region is Z. b. diamondi, the distribution of which stretches from Papua New Guinea's Eastern Highlands to the Paniai (= Wissel) Lakes of Papua Province, Indonesian New Guinea (Flannery and Groves 1998).
The species is hunted and eaten by Hewa, both with and without dogs. Several trophies were collected from Hewa hunters at Tualapa. Some Hewa informants told us that they distinguished two kinds of echidna (apparently using Taku as a general term for both), which they called Wam Taku Haiba (definitely referring to Zaglossus) and Wam Taku Itu. The latter may well be the Short-beaked Echidna, Tachyglossus aculeatus, which occurs along the southern margin of the central cordillera elsewhere in Southern Highlands Province (e.g. at Mt. Bosavi; Leary and Seri 1997), and may occur in Hewa country. Or it may represent an intriguing, very small echidna, possibly an unnamed species, once captured by Opiang at Crater Mountain on the southern margin of the central cordillera in Eastern Highlands Province (an adult female weighing 1.8 kg). It is also possible that Wam Taku Itu is simply another name for Zaglossus bartoni. Whatever it may be, the traditional recognition by the Hewa and some other informants in Papua New Guinea of two or three morphological types of echidna (Opiang, pers. obs.) advocate for continuing field and taxonomic studies of the echidna species of New Guinea.
As so little is known about Zaglossus reproduction (Flannery 1995, Opiang 2009), we asked Hewa informants what they knew about the subject. They told us that the Long-Beaked Echidna has only one baby at a time, but no informant had seen an echidna so young that it could not walk on its own. The youngest echidnas seen were said to be about 4 human fingers long (about 300 mm?).
To follow up on the results of our survey, Opiang and Thomas are now working with the Hewa to locate active echidna dens. These dens will be marked via satellite locator beacons and monitored by the Hewa. This will hopefully begin to shed light on the identity of the ‘second’ Hewa echidna, as well as on poorly understood aspects of echidna biology in New Guinea, such as reproduction. It will most definitely engage the Hewa as partners in echidna conservation.
Order Peramelemorphia, Family Peramelidae (Bandicoots)
Echymipera kalubu (Lesson, 1828)
IUCN Red List Status: Least Concern.
Echymipera kalubu is a medium-sized bandicoot common at lower to middle elevations throughout most of New Guinea. Familiar to hunters, this animal is commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones. It is apparently referred to by the Hewa name Wam Wauma or Wam Tsua Kiwauma (Tsua being apparently a more general term for bandicoots). Hewa informants describe that it is generally found at lower elevations (1,000 m and lower), that it eats fruit, pandanus, and worms, and that it has litters of four or less. Two specimens that were killed by dogs at Putuwe during our survey were salvaged as museum specimens (USNM 585488 and 585613, adult females).
Echymipera rufescens (Peters and Doria, 1875)
IUCN Red List Status: Least Concern.
Echymipera rufescens is a medium-sized bandicoot that occurs at lower to middle elevations throughout most of New Guinea. Familiar to hunters, this animal is commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones. It is apparently referred to by the Hewa name Wam Lokoume or Wam Kimafima.
Order Diprotodontia, Family Macropodidae (Kangaroos)
Dendrolagus goodfellowi Thomas, 1908 (Goodfellow's Tree- Kangaroo)
IUCN Red List Status: Endangered: “Listed as Endangered based on an ongoing population decline of at least 50% over the past three generations (i.e., 30 years) due to actual levels of exploitation from hunting and a decline in habitat quality. It has already been extirpated from significant portions of its range.” (Leary et al. 2008b).
Dendrolagus goodfellowi is a red-brown and golden arboreal kangaroo distributed throughout the eastern half of New Guinea's extensive central cordillera, from the Star Mountains region in the west to Milne Bay in the east (Flannery 1995, Flannery et al. 1996), generally in forest at low to middle elevations (Flannery 1995). Related taxa occur in the North Coastal ranges (Torricelli and Foja Mountains: D. pulcherrimus), the Huon Peninsula (D. matschiei), and in the lowlands of south-central New Guinea (D. spadix) (Menzies 1991, Flannery 1995, Flannery et al. 1996, Helgen 2007b).
Goodfellow's Tree Kangaroo is hunted and eaten in Hewa country, and we collected several trophy skulls and bones kept by local hunters. It is referred to by the Hewa name Wam Inai Tukelo (Inai being apparently a general term for tree kangaroos), and occurs at lower to middle elevations in Hewa country, including around Tualapa. Interestingly, Hewa hunters concurred that there were 3 kinds of tree kangaroos in Hewa country -Wam Inai Tukelo, which they associated with photographs and trophy skulls referable to D. goodfellowi; Wam Inai Maputa, which they stated only occurred high up in the mountains (i.e., ‘cold ples tru’ in Tok Pisin) and recognized in photographs as Doria's Tree Kangaroo (i.e., Dendrolagus [dorianus] notatus, a high elevation tree kangaroo of the central highlands variably recognized as a subspecies of D. dorianus or a distinct, closely-related species); and finally Wam Inai Tabaghali Loi, the accounts of which we cannot immediately associate with a known species. Hunters stated that Wam Inai Tabaghali Loi has a brown body with a darker midline stripe, and is otherwise broadly similar in appearance to D. goodfellowi. In accordance with a commonly known story, Hewa refer to Wam Inai Maputa as Ya, or first born; Wam Inai Tabaghali Loi as Tabaghali, or second born; and Wam Inai Tukelo as Nom, or last born.
If not simply a variant of D. goodfellowi recognized by folk taxonomy, the most straightforward suggestion for this species' identification would be the Lowland Tree Kangaroo, Dendrolagus spadix, a rare species endemic to the lowlands of south-central New Guinea that may well occur in Hewa country, and is similar in size and sometimes appears very similar in coloration to D. goodfellowi. However, in our discussions Hewa informants instead universally associated the name Wam Inai Tabaghali Loi with both study skin and living animal pictures of Matschie's Tree Kangaroo, D. matschiei, in Flannery's Mammals of New Guinea book (Flannery 1995: 134–135, 139), rather than with a figured photographed study skin of D. spadix (Flannery 1995: 139).
Matschie's Tree Kangaroo is thought to be geographically restricted to the mountains of the Huon Peninsula in northeastern New Guinea and to the nearby island of Umboi (Flannery 1995, Flannery et al. 1996), such that it would seem that this identification could be ruled out immediately on geographic grounds. However, it is in this light that we are drawn to revisit an old riddle from the taxonomic history of tree-kangaroos - the 1936 description of Dendrolagus deltae, a tree kangaroo identical or very similar to D. matschiei, supposedly from Mt. Pratt in Southern Highlands Province - recently discussed by Helgen (2007a):
In 1936, Troughton and Le Souef (1936) named Dendrolagus deltae, a tree-kangaroo similar to D. goodfellowi and D. matschiei (and bearing intimate resemblance to the latter), based on two specimens supposedly collected on Mt. Pratt “in the north-east of the Delta Division” (today in Southern Highlands Province) and received via the Taronga Zoo in Sydney [skins and skulls later deposited in the Australian Museum, Sydney]. “Mount Pratt” is an obscure locality, but apparently refers to an outlying peak to the immediate east of Mt. Bosavi (Lidicker and Ziegler [1968:24]; see map in Monckton [1922]; Laurie and Hill [1954:151] gave the coordinates as “6°31′S, 143°38′E”). Laurie and Hill (1954), Lidicker and Ziegler (1968), and Ziegler (1978:135) credited the type locality of deltae as valid, but Groves (1982) rejected it, pointing out the intimate similarity of D. deltae to D. matschiei and citing earlier doubts of the locality's authenticity as voiced by Kirsch and Calaby (1978). Groves (1982:180) wrote “the reason for the description of Dendrolagus deltae is a complete mystery. The answer is perhaps the uncritical acceptance by the authors of the type locality … the locality of the holotype as reported must have been wrong…” However, in light of Bosavi's endemism, and because the Dendrolagus matschiei species-complex (sensu Flannery 1993; Flannery et al. 1996) shows a clear tendency toward differentiation in areas offlying the central cordillera (witness D. pulcherrimus in the Foja and North Coastal Ranges, D. matschiei in the Huon Peninsula, D. spadix in the southern lowlands, an unidentified taxon in the Arfak Mountains-Aplin in litt.), it now seems rash to reject out-of-hand the original information presented by Troughton and Le Souef (1936), as recent reviewers have done (e.g. Flannery et al. 1996:9; Martin 2005; Groves 2005), at least until stronger negative evidence is available (e.g. as far as I am aware, there have been no modern expeditions to Mt. Pratt; cf. Leary and Seri 1997:86). The more ornate stripe-patterning of the back and tail in goodfellowi/pulcherrimus is more highly derived than the simpler pattern in matschiei and deltae, and it is not inconceivable that these latter forms could represent morphologically-conservative montane taxa within the matschiei complex, isolated to the north and south of the eastern central cordillera, respectively. At least some average differences are apparent between the type series of deltae and correctly provenanced specimens of matschiei (Lidicker and Ziegler 1968; Groves 1982; my examinations); these and other purported differences require closer critical attention. Certainly it is difficult to understand why the type series of deltae (which originated during the time when the Delta region was first being seriously explored), if truly from the Huon Peninsula, was misattributed to an obscure peak in the Kikori River Basin from which no other mammal specimens have ever been collected (and I can find no references, past or current, to a similarlynamed peak on the Huon). To me at least, the riddle of the Delta tree-kangaroo lives on for now.
If Dendrolagus deltae really is a tree kangaroo similar to D. matschiei that occurs along the southern margin of the central cordillera in Southern Highlands Province, this offers an additional possible explanation for the identification of the third Hewa tree kangaroo. Only additional work in collaboration with the Hewa, ideally involving tracking of tree kangaroos and study of tree kangaroo skulls and teeth retained by hunters throughout the area, will help to choose between these hypothetical identities of Wam Inai Tabaghali Loi. Is this Hewa designation another name for either D. goodfellowi or D. [dorianus] notatus, is it perhaps D. spadix, or is it even possibly the tree kangaroo that Troughton and Le Souef (1936) named D. deltae? Or perhaps yet another explanation awaits us. Tree kangaroos are important animals in the culture of both the Hewa and New Guinea conservation biologists; we hope to work together to solve this riddle of the third Hewa tree kangaroo, second born.
Order Diprotodontia, Family Phalangeridae (Cuscuses)
Phalanger gymnotis (Peters and Doria, 1875) (Ground Cuscus)
IUCN Red List Status: Least Concern.
The Ground Cuscus is a medium-sized to large (mass ca. 2–5 kg) terrestrial cuscus that is widespread in lowland and montane forest (sea level to 2,700 m elevation) throughout New Guinea. Familiar to hunters, this animal is commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones. It is referred to by the Hewa name Wam Wai.
Phalanger mimicus (Thomas, 1922) (Southern Common Cuscus)
IUCN Red List Status: Least Concern.
The Southern Common Cuscus is a medium-sized (mass ca. 2–4 kg) arboreal cuscus that occurs throughout much of southern New Guinea at relatively low elevations (sea level to at least 800 m) (Norris and Musser 2001). This animal is commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones. It is apparently referred to by the Hewa name Wam Nabli. Hunters describe that it occurs in both primary and secondary forest at lower elevations, and that it eats the leaves of the plants referenced by the Hewa names Me Neki (scientific name Pasania sp.), Me Paghai (Pandanus sp.), Me Tsaghal (Ficus sp.), and Me Tial (Castanopsis acuminatissima).
Spilocuscus maculatus (Desmarest, 1818) (Common Spotted Cuscus)
IUCN Red List Status: Least Concern.
The spotted cuscus is a relatively large possum (3–6 kg) that occurs throughout lower elevations (sea level to 1,500 m) in most areas of New Guinea. Recent work suggests that the species referred to under this scientific name is actually a complex of species that differ in color, body size, and skull and teeth morphology, some of which may occur sympatrically, such that multiple species will eventually be recognized in place of one (Helgen 2007b). The subspecies in the Hewa region is S. m. goldiei (Ramsay, 1876). Familiar to hunters, spotted cuscuses are commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones at Tualapa and observed several hats made from the fur of this species being worn in Wanakipa. Hewa folk taxonomy distinguishes two kinds of Spilocuscus, known to our informants by the names Wam Kail and Wam Kail Yelekai. The first, Wam Kail, is said to be characterized by females with more and smaller black spots, and males with larger black spots and small white spots. The second, Wam Kail Yelekai, is said to be characterized by females being white with black spots, males black with white spots. At present we understand all of these to be color variants of S. m. goldiei. Hewa informants mention that spotted cuscuses eat the leaves of a number of trees, referenced by the Hewa names Me Paghai (scientific name Pandanus sp.), Me Yat (Ficus drupacea), Me Eli (Diospyros sp.), Me Tsiphlai (Ficus macrocarpa var. latifolia), Me Toghol (Daphniphyllum gracile), and Me Yowal (Casuarina oligodon).
Order Rodentia, Family Muridae (Rats and Mice)
Lorentzimys nouhuysii Jentink, 1911 (Long-Footed Tree-Mouse)
IUCN Red List Status: Least Concern.
The saltatorial mice of the genus Lorentzimys are widespread in New Guinea, but the taxonomy of the genus, in which only a single species is currently recognized (Flannery 1995, Musser and Carleton 2005) is confused and requires comprehensive revision (see Aplin and Kale, Chapter 18, this volume). Four Lorentzimys specimens (USNM 585612, 585614–585616, mass 18 and 19 grams in two adults) were collected in our pitfall traps set in creekside forest at Tualapa. The overall size and reddish chest patch in our series suggest identification as true L. nouhuysii, typically a mouse of lower to middle elevation forest habitats (Helgen, pers. obs.). Hewa informants suggested that Lorentzimys dens at the base of trees and eats insects as well as leaves of Me Patu (Macaranga sp.) and Me Tsaghal (Ficus sp.).
Melomys lutillus (Thomas, 1913) (Grassland Melomys)
IUCN Red List Status: Least Concern.
This small rodent (weighing 31–58 grams in our sample of adults) is common in many open habitats in New Guinea, especially grasslands at lower middle elevations (Menzies 1996). It was commonly trapped at night at Tualapa and Putuwe, always in kunai grassland, sometimes in grassy areas interspersed with regrowth and gardens.
Melomys rufescens (Alston, 1877) (Black-Tailed Melomys)
IUCN Red List Status: Least Concern.
This common and widespread semi-arboreal rat is usually associated with secondary habitats (Flannery 1995, Menzies 1996). Three specimens (USNM 585637–585639, adults weighing 48–60 grams) were trapped on the ground in secondary forest at Tualapa - one along a creek, the others at the base of a tree.
Rattus exulans (Peale, 1848) (Pacific Rat)
IUCN Red List Status: Least Concern.
This small rat, thought to have arrived to New Guinea only in recent millennia (Taylor et al. 1982), was common in disturbed habitats near Wanakipa. It was trapped in the evening and at night in kunai grassland at Tualapa and in village houses at Putuwe (USNM 585618–585623).
Rattus foersteri Rümmler, 1935 (Small Spiny Rat)
IUCN Red List Status: Least Concern.
Helgen (2007b) applied Rümmler's name R. foersteri to eastern New Guinean populations of the rat usually referred to by the name Rattus steini, the taxonomy of which requires a detailed overview. A single specimen (USNM 585625, a young adult female) referred to this species was snap trapped at Tualapa, but the microhabitat was not noted.
Rattus ‘rattus’ (Linnaeus, 1758) (Black Rat complex)
IUCN Red List Status: Least Concern.
We collected a single rat (USNM 585624, adult male) at Tualapa that appears referable to the “Rattus rattus complex” sensu Aplin et al. (2003: 172–173), an invasive species and probably a relatively very recent arrival in New Guinea (Taylor et al. 1982, Flannery 1995). Our specimen has a reddish brown dorsum with long black guard hairs, a yellowish venter with gray hair bases, and a dark and relatively hairy tail. This specimen's relatively large size (190 grams), longer tail than head body length (170 mm tail versus 160 mm HB), and black dorsal surfaces of the feet distinguish it from the native Rattus of similar body size that might be expected to occur in the area, including R. leucopus, R, sordidus, R. steini, and R. novaeguineae. Its long black dorsal guard hairs and large plantar pads preclude its identification as R. argentiventer, another non-native Rattus recorded from New Guinea (Musser 1973, Taylor et al. 1982). This specimen would probably be best identified as Rattus tanezumi under current taxonomy (Musser and Carleton 2005), but ongoing studies of the morphology and genetics of the Rattus rattus complex point to a complex taxonomy involving many taxonomic lineages, the distributions, biological attributes, and names for which are not yet clearly resolved (K. Aplin, in litt.). The Tualapa specimen was trapped in kunai grass alongside Rattus exulans and Melomys lutillus, and though taken in a disturbed habitat, it was trapped several kilometers from the village of Wanakipa and not immediately near any concentrated human habitation. This capture is somewhat unexpected given that the most recent summary of New Guinea records of Rattus rattus, now several decades old now, recorded the species only from satellite island, coastal, and major town and city localities only, with very few exceptions, and no records at all originating from the vicinity of the highlands of Papua New Guinea (Taylor et al. 1982). The presence of this invasive species at Tualapa is one indication of environmental disturbance that is probably new to the highlands in recent decades. Other invasive species have spread in Papua New Guinea in recent decades, including plants, insects, and tree diseases (Leps 2002, Kiapranis and Nimiago 2005), presumably in tandem with expanding and increasing impacts of air travel, industry, international trade, and the expansion of the Highlands Highway, which extends from Lae in Morobe Province in the east to Tari (Southern Highlands Province) and Porgera (Enga Province) in the west and may be a major factor in opening the highlands to invasive species from around the world. Broad survey efforts will be needed to understand both the current distribution of Rattus rattus and potentially other invasive mammal species, and to begin to evaluate their potential ecological impact on native fauna, if any. The spread of Rattus rattus and other invasive murine species has been fingered elsewhere in the decline of native rodents (Smith and Carpenter 2006, Amori et al. 2008, Wyatt et al. 2008, Harris 2009), as on other native fauna, especially birds.
Uromys caudimaculatus (Krefft, 1867) (Mottled-Tailed Giant-Rat)
IUCN Red List Status: Least Concern.
This large rat is widespread in New Guinea, but its current taxonomy is in need of a detailed review (Musser and Carleton 2005, Helgen 2007b), an important priority in New Guinea systematic mammalogy. A single specimen (USNM 585512, a young adult female weighing 220 g) was secured by hunters in forest at Tualapa. This species is commonly hunted and eaten in Hewa country, and we documented several trophy skulls and bones. Hewa informants called it by the Hewa name Wam Ute Tsin, a name also applied by informants to photographs of Xenuromys barbatus, a much rarer large rat of very similar external appearance.
Order Chiroptera, Family Pteropodidae (Fruit-Eating Bats)
Dobsonia moluccensis (Quoy and Gaimard, 1830) (Greater Bare- Backed Bat)
IUCN conservation status: Least concern.
This large (400–600 g), usually cave-roosting bat is one of the most common bats of New Guinea. The subspecies in New Guinea is D. m. magna Thomas, 1905. Familiar to hunters, these bats are commonly hunted and eaten in Hewa country, and we documented a number of trophy skulls and bones at Tualapa. Informants used the Hewa name Nok Teliau for this bat.
Macroglossus minimus (E. Geoffroy, 1810) (Northern Blossom- Bat)
IUCN conservation status: Least concern.
This small blossom bat is widely distributed in the lowlands of New Guinea, but is usually considerably less common than Syconycteris australis (see below). We mistnetted a single specimen at Tualapa (USNM 585523, adult male, forearm 42 mm) in secondary forest. Informants used the Hewa name Nok Semina Mea Mea for this bat (and for Syconycteris australis, see below).
Nyctimene papuanus Andersen, 1910 (Common New Guinea Tube-Nosed Bat)
IUCN conservation status: Least concern.
This was the bat we most commonly mist-netted during our survey; it was netted at Tualapa, Umge, and Putuwe. Our specimens closely matched the description and measurements of Nyctimene papuanus as described in detail by Andersen (1912). This taxon is usually recognized as a subspecies of N. albiventer (Gray, 1863), originally described from Morotai in the North Moluccas, but comparison of these specimens against a large series of near-topotypical albiventer from Halmahera at USNM confirms that our New Guinea specimens are much larger and easily distinguished craniodentally from true albiventer. The taxonomy of Melanesian Nyctimene is highly confused and awaits wholesale taxonomic revision (Nancy Irwin, in litt.). Critical review of museum specimens indicates that several currently unrecognized species, at least one of which is without any scientific name, are incorporated amongst series of specimens usually identified as “N. albiventer” from New Guinea (Aplin and Kale, Chapter 18, this volume, Nancy Irwin, in litt.; Helgen, pers. obs.). Informants used the Hewa name Nok Semina Kotnalia for this bat.
Paranyctimene sp. (Unstriped Tube-Nosed Bat)
IUCN conservation status: Least concern.
We mistnetted a series of specimens of a species of Paranyctimene (USNM 585520–585522, 585680–585684, forearms 53–57 mm) in primary forest at Umge (alongside fruiting figs and a trickling stream), where it was most common, and in secondary forest at Tualapa. These specimens seem to be larger than typical P. raptor, and may represent P. tenax (see Bergmans 2001). The taxonomy of Paranyctimene, an endemic New Guinea pteropodid genus, is highly confused, and awaits wholesale taxonomic revision (Nancy Irwin, in litt.).
Syconycteris australis (Peters, 1867) (Common Blossom-Bat)
IUCN conservation status: Least concern.
This common and widespread nectarivorous bat (forearm 42–48 in our sample of adults) was mistnetted at Tualapa (in secondary forest), Umge (in primary forest amongst fruiting figs and a trickling stream), and at Putuwe (in gardens and regrowth). Informants used the Hewa name Nok Semina Mea Mea for this bat (and for Macroglossus minimus, see above). This was the second most commonly mistnetted bat on our survey after Nyctimene papuanus.
Order Chiroptera, Family Miniopteridae (Bent-Winged Bats)
Miniopterus macrocneme Revilliod, 1914 (Long-Legged Bent- Winged Bat)
IUCN conservation status: Data deficient (based on taxonomic uncertainty; Bonaccorso and Reardon 2010; see below).
A sizeable series representing a relatively small species of Miniopterus (forearm 40–47) with relatively long legs (tibia 16–19 mm) was collected from a cave near Tualapa. Close inspection of skulls and external features confirms that this series corresponds to the species referred to as Miniopterus macrocneme under current taxonomy (Peterson 1981, Hill 1983). This species was cohabitating with the larger bentwing bat Miniopterus magnater, a cave-roosting association previously reported by Bonaccorso (1998:390).
As far as we know, there are no previously published records of M. macrocneme from Southern Highlands Province (at least not as compiled by Flannery [1995] and Bonaccorso [1998]), which is surprising, as this is one of the more common bats of central New Guinea. It is apparently especially common in caves in the central cordillera of Western and West Sepik (Sandaun) Provinces (Flannery and Seri 1990, Flannery 1995). The lack of previously published records from Southern Highlands Province reflects the overall paucity of attention given to insectivorous bats in New Guinea to date, both in terms of field survey efforts and in taxonomic reviews of available museum material (Helgen 2007b, Armstrong and Aplin, Chapter 19, this volume).
Species boundaries and nomenclature of the species of Miniopterus within New Guinea and across the broader Australasian region remain confused and in need of detailed systematic review, ideally incorporating qualitative anatomical, morphometric, and genetic comparisons across a large array of museum samples representing all major islands and regions and all named forms or putative taxa. At minimum, six biological species occur on the New Guinea mainland (Bonaccorso 1998, Simmons 2005), but the appropriate scientific names, actual geographic distributions across New Guinea and broader Australasian islandscapes, and phylogenetic relationships among these species remain poorly understood, and additional species are likely to be recognized after detailed taxonomic review. At the moment, the taxonomic status of M. macrocneme is classified by the IUCN as Data Deficient owing to the taxonomic uncertainty that prevails amongst Melanesian Miniopterus. As Bonaccorso and Reardon (2010) explained, “The urgent priority … is to resolve Miniopterus taxonomy and identification in order to understand distribution, abundance, habitat requirements, ecology, and threats …” (Bonaccorso and Reardon 2010). This is a major priority for systematic mammalogy in the region.
Miniopterus magnater Sanborn, 1931 (Western Bent-Winged Bat)
IUCN conservation status: Least Concern.
A small series of M. magnater (USNM 585772–585776, forearms 49–51 mm, tibiae 19–22 mm) were collected from the same cave as our series of M. macrocneme (see above). Identification of these specimens as M. magnater is based on studies of external features and skulls in comparison with the revisionary overviews produced by Peterson (1981) and Hill (1983).
Order Chiroptera, Family Vespertilionidae (Evening Bats)
Nyctophilus microtis Thomas, 1888 (Papuan Big-Eared [or Long- Eared] Bat)
IUCN conservation status: Least concern.
We mist-netted a single specimen of N. microtis (USNM 585718, adult male, forearm 39 mm) amongst village environs (homes, gardens, and regrowth) at Putuwe. The identification of this specimen as N. microtis was confirmed by Harry Parnaby, global authority on Nyctophilus taxonomy (e.g., Parnaby 1987, 1988, 2002, 2009). As far as we know, there are no previously published records of N. microtis from Southern Highlands Province (cf. Flannery 1995, Bonaccorso 1998), but its presence in the province was certainly to be expected. Nyctophilus microtis is the most commonly recorded Nyctophilus in New Guinea, and has been documented in neighboring provinces of Papua New Guinea, including Western, Gulf, and Chimbu Provinces. It is widely distributed in New Guinea, and occurs from sea level to at least 2,600 m (Flannery 1995, Bonaccorso 1998).
Order Artiodactyla, Family Suidae (Pigs)
Sus scrofa Linnaeus, 1758 (Feral Pig)
Apparently introduced to New Guinea by humans several thousand years ago, pigs occur as both domesticated and wild-living (feral) populations throughout the island of New Guinea (Hide 2004). Wild pigs are important game animals in Hewa country, and domesticated pigs are kept as property at Wanakipa as throughout New Guinea.
Other species present in the region
Overall, all species encountered during our survey were expected to occur in the region and our overall results, though based on a short survey, suggest that the Hewa mammal fauna is similar to faunas documented at low to middle elevations at other sites situated along the southern margin of the central cordillera in Southern Highlands Province, such as Mt. Sisa (Dwyer 1990) and Mt. Bosavi (Leary and Seri 1997).
In addition to the 22 species we documented based on specimen-backed evidence, interviews with knowledgeable Hewa informants revealed their familiarity with at least as many additional mammal species. For many of these we were able to apply Hewa names to taxonomic names, based on Hewa informants' recognition of photographs and specimens, and on convincing discussions of the appearance and habits of animals (Table 21.2). Several of these additional mammals are species that are usually restricted to habitats at higher elevations than we were able to visit during our brief survey, such as the tree kangaroo Dendrolagus [dorianus] notatus and the cuscus Phalanger carmelitae, or are difficult to trap without extensive, focused effort (especially the amphibious murines Hydromys and Parahydromys; Helgen 2005a). However, most of the species described by Hewa informants (Table 21.2) likely occur in the vicinity of Wanakipa and the camps from which our survey was based (Tualapa, Umge, Putuwe); our brief visit was simply insufficient to encounter them in the field or to document most of them with captured specimens or trophy material retained by hunters.
CONSERVATION RECOMMENDATIONS
The results of our survey provide a better understanding of the mammal fauna of a corner of Southern Highlands Province previously little-investigated by mammalogists, and establish the importance of Hewa lands as a region where some of Papua New Guinea's largest and most endangered native mammals (such as Zaglossus bartoni and Dendrolagus goodfellowi) occur. As we have emphasized elsewhere (Helgen 2007c, Helgen and Opiang, Chapter 20, this volume), few firm steps can be taken with regard to conservation, management, and/or protection of mammals in this region without a more intricate understanding of their distribution, abundance, and threats that they face. Additional survey efforts and more focused research on individual focal species are needed before many specific recommendations for conservation and management be made; below, we highlight priorities for further work in Hewa country specifically and for New Guinea mammalogy in general.
Indigenous knowledge. Collaborations between biologists and indigenous communities are a fundamentally important aspect of biodiversity survey efforts and broader efforts aimed at biodiversity conservation in New Guinea. Since 1988, Thomas has worked with the Hewa to conserve their bio-cultural heritage (Thomas 2009a). This partnership has focused on the impact of human disturbance on biodiversity, as this “humans as a source of disturbance” approach fit both the Western and Hewa view of the dynamics of traditional gardening. Using birds - a visible and widely appreciated indicator of biological diversity, they have developed a vehicle for the cross-cultural communication of environmental knowledge that enables both the Hewa and the conservation community to engage issues surrounding sustainable development and conservation at the headwaters of the Strickland River (Thomas 2010a, 2010b). In the process, they have also developed the Papuan Forest Stewards Initiative (Thomas 2009b and Chapter 8, this volume) to secure their bio-cultural heritage for future generations. Since traditional environmental knowledge is no longer a sign of backwardness in Hewa society but rather a source of income, an enthusiastic cadre of Hewa naturalists, willing to share their knowledge, has emerged. Collaborating with Hewa naturalists has already revealed many unanswered questions about the mammals with which they share their lands that are conducive to cooperative study, including animals as important to New Guinea conservation biologists as tree kangaroos and echidnas. This survey is the first in what we anticipate will become a steady stream of partnerships between the Hewa and scientists that will work to conserve the bio-cultural heritage of this region for now and in the future.
Basic biodiversity survey efforts. To better understand the immediate fauna of the Wanakipa area, basic biodiversity surveys, such as the survey work discussed herein, will need to continue, incorporating more sites and additional habitats (particularly in higher elevation habitats). Our brief survey in Hewa country has probably documented a relatively modest fraction (probably less than 25%) of the actual Hewa mammal fauna. As indicated by discussions with Hewa informants, further surveys will undoubtedly record many additional mammal species in the area, some of which, like various kangaroo and wallaby species (Table 21.2), are species of conservation concern.
Focal species of conservation concern. Targeted ecological studies in collaboration with the Hewa are needed to assess in greater detail the distribution, abundance, and threats faced by what we can label “focal species” for conservation in the region, especially the Long-Beaked Echidna (Zaglossus bartoni) and Goodfellow's Tree Kangaroo (Dendrolagus goodfellowi), which are likely to be targets of concerted hunting pressure. Wallabies, tree kangaroos, and echidnas have probably disappeared from most areas with high human population density and high hunting pressure throughout New Guinea (Bulmer and Menzies 1972, George 1978, Flannery 1992, 1995; Flannery and Groves 1998, Martin 2005). The fact that Zaglossus, at least one tree kangaroo species, and apparently several other macropodids (Table 21.2), all good proxy taxa for understanding hunting pressure, still persist in the vicinity of Wanakipa is a good sign that hunting has not yet resulted in mammal extirpations in the area. On the other hand, the only reason we know that these species occur at Wanakipa is because they are represented in multiple trophy jaw collections held by local hunters, demonstrating that concerted hunting pressure is likely acting on these animals.
Taxonomy. Appropriate conservation prioritization for most New Guinea mammals still cannot be evaluated competently without comprehensive taxonomic revisions first laying the groundwork required for understanding taxonomic boundaries, delineating geographic distributions, and identifying characteristics that will allow for straightforward taxonomic identification of specimens. Biologically diverse Melanesian mammal genera especially in need of sweeping taxonomic revisionary overviews include the rodent genera Paramelomys, Uromys, Pogonomys, and Rattus, miniopterid bats (Miniopterus), the pteropodid genera Nyctimene and Paranyctimene, and marsupial genera such as Petaurus, Cercartetus, Distoechurus, and Echymipera, all of which occur in Hewa country. Additional biological surveys will continue to add much to our knowledge of mammal taxonomy and distribution in New Guinea, but sufficient museum specimens now exist to undertake these principal revisionary goals for most of the island of New Guinea. The taxonomy in use within the enormously valuable standard volumes currently available for New Guinea mammals (e.g. Flannery 1995, Bonaccorso 1998) provides an overly simplistic caricature of the complexity of New Guinea mammal diversity, as renewed detailed systematic reviews continue to demonstrate (e.g., Helgen and Flannery 2004, Woolley 2005, Helgen 2005b, 2007a, Musser et al. 2008, Musser and Lunde 2009, Helgen and Helgen 2009, Parnaby 2009). Currently unnamed or unrecognized biological species are known in almost every Melanesian mammal genus (Helgen 2007a, 2007b, Helgen, pers. obs.). Only once these most taxonomically complex of Melanesian mammal genera are reviewed taxonomically across the sum total of specimens available in world museums will a clearer understanding of mammalian historical biogeography, ecological complexity, and geographic and taxon-based conservation prioritization emerge.
Table 21.2. Knowledge of mammals in Hewa country reported by Hewa informants. We report here the Hewa names that we have been able to most satisfactorily associate with taxonomic names of New Guinea mammals, based on Hewa informants' recognition of photographs and specimens, and on convincing discussions of animals' appearance and habits. Information listed as “Informant Knowledge about Local Occurrence and Habits” represents statements from informants about where the species lives, or where it is most likely to be encountered in Hewa country, along with selected notes regarding diet and behavior. Wam is a noun prefix applied to names of terrestrial mammals; Nok is a noun prefix applied to names of bats.
continued
Invasive species. The nature and impacts of the spread of invasive mammal species in New Guinea, such as rats in the Rattus rattus complex, deserves further study, as brought to light in the present survey. What are the current distributions of invasive rodents and other invasive mammal species in New Guinea, how and where are they spreading, and what impacts do they have on human health, agriculture, and on native fauna and flora? These are important questions that, at least as far as we are aware, are not currently being studied in New Guinea, and will require considerable field work to address.
ACKNOWLEDGMENTS
We particularly thank the Hewa, our local assistants and collaborators, for their hospitality and sharing their traditional knowledge for us. For assistance and camaraderie in the field, we especially thank Steve Richards, Max Kuduk, Wayne Takeuchi, Wayne Maddison, Bruce Beehler, Robert Sine, and Leo Legra. For assistance with our survey, and with the processing and preparation of specimens, we thank the staff of the Papua New Guinea Department of Environment and Conservation, staff of the Papua New Guinea Institute for Biological Research, David Schmidt, Linda Gordon, and Craig Ludwig of the Division of Mammals at the Smithsonian Institution, and Harry Parnaby. For assistance with information requests and reviews, we especially thank Ken Aplin, Harry Parnaby, Darrin Lunde, Guy Musser, Pat Woolley, and Tim Flannery. We thank collection management staff at the American Museum of Natural History (New York), the Australian Museum (Sydney), the Australian National Wildlife Collection (Canberra), the Bernice P. Bishop Museum (Honolulu), the Natural History Museum (London), the National Museum and Art Gallery of Papua New Guinea (Port Moresby), the National Museum of Sweden (Stockholm), and the University of Papua New Guinea (Port Moresby) for facilitating our taxonomic research on New Guinea mammals mentioned in this report.
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The predatory attack itself may differ between that of a carnivorous and omnivorous species, given the shift in lifestyles. I reviewed the information on 63 carnivorous rodent species. The approach here is to compare the predatory attack of carnivorous grasshopper mice (Onychomys torridus and Onychomys leucogaster) with that of the deer mouse (Peromyscus maniculatus) to identify specific differences. The predatory attack of a carnivorous rodent differs from that of the omnivorous rodent in its ability to overcome the flight initiation distance of potential prey, biting capabilities, use of its sensory modalities, seizure of prey, early and rapid development of a proficiency in its attack, vigor of its attack, and resistance to inhibition of the attack. The grasshopper mouse intimidates other rodents with which it competes for prey by a combination of its highly aggressive attack and advertisement of its presence. Focusing on the importance of overcoming the flight initiation distance of potential prey, the distinct and separate nature of the predatory attack, and behavior that reduces competition for prey can provide a framework for describing the evolution of a carnivorous lifestyle in other mammals. (PsycInfo Database Record (c) 2021 APA, all rights reserved).
... Intensive fieldwork coupled with detailed genetic and morphological analyses continue to show that vertebrate species diversity, and especially endemism, remains significantly underestimated in many regions of the world (Helgen & Helgen 2009;Hotaling et al. 2016;Grismer et al. 2017). This pattern is most pronounced in developing tropical countries; however, recent work in Australia has also revealed or reinforced hotspots of localised and evolutionarily divergent vertebrate lineages (Hoskin & Couper 2013;Rosauer et al. 2016Rosauer et al. , 2018Bourke et al. 2017). ...
Over the last decade, the combination of biological surveys, genetic diversity assessments and systematic research has revealed a growing number of previously unrecognised vertebrate species endemic to the Australian Monsoonal Tropics. Here we describe a new species of saxicoline velvet gecko in the Oedura marmorata complex from Groote Eylandt, a large island off the eastern edge of the Top End region of the Northern Territory. Oedura nesos sp. nov. differs from all congeners in combination of moderate size, and aspects of tail morphology and colouration. It has not been reported from the nearby mainland regions (eastern Arnhem Land) suggesting it may be an insular endemic, although further survey work is required to confirm this. While Groote Eylandt is recognised as a contemporary ecological refuge for declining mammal species of northern Australia, newly detected endemic species suggest it may also be of significance as an evolutionary refuge for many taxa, especially those associated with sandstone escarpments.
Forty-four species of mammal were identified from Rofaifo territory, Mt. Erimbari, Eastern Highlands Province (altitude range 1900-2300 m). These comprised 16 marsupials, 20 rodents, four bats, pig, dog, domestic cat and cattle. Pig, dog and cattle were present as domesticated animals only; cats were present as domesticated and feral animals. Bats were not surveyed in detail. The water rat Crossomys monktoni was almost certainly present (based on descriptions) and an additional dasurid is a possibility. Skins of one additional species of Phalanger, one of Dactylopsila and one or two of Dendrolagus were obtained as trade items by the people. Some people described a failed attempt to establish the echidna Tachyglossus aculeatus at Mt. Erimbari.
A new large marsupial glider, Petaurus abidi, from the Torricelli Mountains of northwestern coastal Papua New Guinea is described and figured, and morphologically compared with the three species of the genus previously known. The new form differs from both named races of the generally larger P. australis primarily in possession of a well developed dark mid-dorsal head-and-back stripe, an unelongated ear, noticeably smaller claws, a longer narrower palate, and greater maxillary dental measurements. It differs from P. breviceps and P. norfolcensis primarily by being larger in all cranial and dental measurements, as well as in essentially all external dimensions. However, similarities in colouration and pattern, in ear and claw proportions, and in various skull characters, as well as, to some extent, biogeographical considerations, suggest the new species' closest intrageneric relationship is with one of these two smaller species. The colloquial name recommended for this new species is Northern Glider.
The distribution of three species of Rattus, three of Melomys and one of Leptomys among habitats classed according to successional phase (i.e. garden to forest) was studied by trapping. Species were added progressively to the succession and, in broad outline, those that produced more young preceded those that produced less young. Melomys lutilIus and R . leucopus were restricted to areas of grassy regrowth and of low secondary growth (ca 10-12.5 yr) respectively. Other species remained in the assemblage from the successional phase at which they entered. All species had appeared within 20 yrs of the start of succession.
Developmental constraints presumably operate by influencing patterns of variability: when development causes some features to vary more than others and when the level of variability is correlated with evolutionary change, then development can be said to constrain evolution. This idea was tested by examining the relationship between tooth variation and three other factors: developmental processes, tooth function, and evolutionary change. Data came from two lineages of viverravid carnivorans (Viverravidae, Carnivora) from the Paleogene of North America. Variability in cusp position was significantly correlated with position in the developmental cascade, with the amount of intercusp growth (when growth is relatively greater in some cusps than others), and with amount of evolutionary change. This indicates that tooth development exerts a local constraint on phenotypic variability and on the evolutionary response to functional selection, but comparative data suggest that the developmental constraint itself may evolve. Intense directional or stabilizing selection may modify the developmental cascade so that the constraint is either removed or modified to permit new evolutionary patterns. Thus development does not constrain evolution in an absolute sense, but rather introduces modifiable patterns of covariance among crown features. Both development and function seem to play important, intertwined roles in coordinating evolutionary change in mammalian molars.
The order Rodentia is the most abundant and successful group of mammals, and it has been a focal point of attention for compar ative and evolutionary biologists for many years. In addition, rodents are the most commonly used experimental mammals for bio medical research, and they have played a central role in investi gations of the genetic and molecular mechanisms of speciation in mammals. During recent decades, a tremendous amount of new data from various aspects of the biology of living and fossil rodents has been accumulated by specialists from different disciplines, ranging from molecular biology to paleontology. Paradoxically, our understanding of the possible evolutionary relationships among different rodent families, as well as the possible affinities of rodents with other eutherian mammals, has not kept pace with this information "explosion. " This abundance of new biological data has not been incorporated into a broad synthesis of rodent phylo geny, in part because of the difficulty for any single student of rodent evolution to evaluate the phylogenetic significance of new findings from such diverse disciplines as paleontology, embryology, comparative anatomy, molecular biology, and cytogenetics. The origin and subsequent radiation of the order Rodentia were based primarily on the acquisition of a key character complex: specializations of the incisors, cheek teeth, and associated mus culoskeletal features of the jaws and skull for gnawing and chewing.
