<Emphasis Type="Italic">Tarsius wallacei</Emphasis>: A New Tarsier Species from Central Sulawesi Occupies a Discontinuous Range

On the basis of distinguishing characteristics of various genetic markers, pelage color, tail tuft, and vocalizations, we describe a new species of the genus Tarsius Storr 1780. The new taxon Tarsius wallacei sp. nov. occupies a disjunct range in Central Sulawesi, Indonesia. The two isolated populations differ significantly in body size, but are alike in color, tail tuft dimensions, vocalizations, and genetic composition. Morphologically, the new species is similar to other Sulawesi lowland tarsiers. In the field, it can be distinguished from its congeners via a characteristic duet song and its yellow-brown pelage coloration and a copper-colored throat. Genetic analyses prove Y-chromosomal and mitochondrial DNA sequences and also microsatellite allele frequencies to be absolutely diagnostic.     

Molecular Phylogenetics and Chronometrics of Tarsiidae Based on 12S mtDNA Haplotypes: Evidence for Miocene Origins of Crown Tarsiers and Numerous Species within the Sulawesian Clade

We report new mitochondrial DNA sequence data from tarsiers sampled from several populations, including the extreme northeast and southwest of the range of the Tarsius tarsier species complex, the most extensive sampling ever reported for this taxon. Our results provide the opportunity to produce the first ever molecular chronometric analysis of Tarsiidae. These results date the age of crown tarsiers, minimally, to the middle Miocene, and each of the 3 tarsier species groups, Tarsius bancanus, T. syrichta, and the T. tarsier complex, to the early or middle Miocene. Thus, each of these 3 species has evolved in isolation for a period of time that is consistent with that which would be expected for multiple speciation events. Our analysis of the Tarsius tarsier complex reveals 5 subclades, each of which is interpreted to represent a haplogroup at, or above, the species level, a result that is consistent with current hypotheses about numerous cryptic species within this species complex. The implications for conservation within the Sulawesi biogeographic region are that Sulawesi is subdivided into numerous subregions of endemism and that, by extrapolating the example of cryptic tarsier species to other taxa, biodiversity may be underestimated by an order of magnitude. The practical realties of conservation in Sulawesi are such that it is most reasonable to assume that anthropogenic extinctions are occurring, and that some species will go extinct before they have even been identified.     

Geographical Variation in Duet Songs of Sulawesi Tarsiers: Evidence for New Cryptic Species in South and Southeast Sulawesi

Geographic populations of Sulawesi tarsiers differ conspicuously in the acoustic structure of the loud duet calls produced by mated pairs. There is evidence that the differences in call features relate to species differentiation. Duet calls can thus be used as a tool to identify cryptic species and for their monitoring in the field. We recorded duets from various geographically isolated populations of Tarsius tarsier in south and southeast Sulawesi to survey the species identity of the tarsiers in these regions. We compared their calls with the duet calls of the Manado tarsier (= Tarsius spectrum), Tarsius dianae, and the Togian Island tarsier. In addition, we recorded and analyzed the duet call of the tarsier from Peleng Island, Tarsius pelengensis. Qualitative characteristics of duet songs of both sexes, analysis of frequency and temporal parameters, and multivariate statistics show 4 distinct vocal forms that predict the existence of 4 new species. These are found on Selayar Island, on the south mainland (Bantimurung), on the southeast mainland (Puwato), and on the Islands of Kabaena and Buton. Our analysis of the call of the Peleng tarsier revealed striking similarities with the call of Tarsius dianae. Further morphological analyses are needed to support the proposed new species in south and southeast Sulawesi and to clarify the taxonomic affinity between the central Sulawesi tarsier species Tarsius pelengensis and T. dianae.     

The Genera and Species of Tarsiidae

We revise the taxonomy of the primate family Tarsiidae. We classify extant tarsiers in 3 genera—Tarsius, Cephalopachus, and Carlito—each of which originated in the Miocene, or earlier, and each of which is allopatrically distributed within a distinct biogeographic region: Sulawesi, Sundaland, and Greater Mindanao, respectively. Within the genus Tarsius, formerly regarded as a single species, Tarsius spectrum, we recognize 8 allopatric and parapatric species, 9 if the inclusion of pumilus is warranted, and note that more are likely to be described in the near future. We restrict Tarsius tarsier, the senior taxon of the genus, to the island of Selayar, off the tip of the southwestern peninsula of Sulawesi. In doing so, it is required that we resurrect Tarsius fuscus Fischer 1804, for the population of tarsiers from the southwestern peninsula near the city of Makassar. We note that neither Cephalopachus nor Carlito has been the subject of anywhere near as much field research as has Tarsius; thus we question if the currently accepted α taxonomy for these genera is based on knowledge or ignorance.     

The Social System of Lariang Tarsiers <Emphasis Type="Italic">(Tarsius lariang)</Emphasis> as Revealed by Genetic Analyses

Previous sociobiological studies of tarsiers were invariably based on field observations. Sulawesi tarsiers are known for monogamous or facultative polygynous social mating systems, but, to date, no data exist to describe the genetic mating system. We here bring together behavioral studies and molecular tools for the first time to elucidate mating behavior and kinship within a tarsier population. We investigated the social system of the recently described Tarsius lariang, which researchers have never studied before. Between September and November 2005, we conducted field observations and sampling in central Sulawesi, Indonesia, where this species is endemic. Ten of 11 social groups of the focal population were composed of 1 adult male, 1 adult female, and putative offspring. To enlighten genetic relationships, we used 12 microsatellite loci and mitochondrial DNA sequences of 26 captured and sampled Lariang tarsiers for parentage and relatedness analyses. A significant number of young were the offspring of the sampled group adults, suggesting a predominantly monogamous social and genetic mating system. There is evidence for extrapair young in groups in which adult pairs exhibit close relationships, leading to the assumption that extrapair mating is solicited to avoid inbreeding. Ten of 11 social groups lived in monogamous social systems, indicating monogamy to be the rule in Tarsius lariang.     

Duet Vocalizations among Different Populations of Sulawesi Tarsiers

A most conspicuous vocal behavior of Sulawesi tarsiers is the loud calls that males and females coordinate into duets. Differences in the acoustic structure of this display relate to specific differentiation in Tarsius spectrum and T. dianae. More recent studies on dueting behavior indicated the existence of a new species of tarsiers on the Togian Islands in Tomini Bay. I analyzed the duet calls of the Togian tarsier to assess the differences in acoustic structure of duet calls between this putative new species and T. spectrum or T. dianae. Discriminant function analysis revealed that Togian tarsiers, T. spectrum and T. dianae, are clearly separated by acoustic characteristics in songs. The degree of separation of the Togian tarsiers from the mainland species support them being a distinct species.     

A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric-morphometric method

Tarsius is an extant genus of primates endemic to the islands of Southeast Asia that is characterized by enormously enlarged orbits reflecting its nocturnal activity pattern. Tarsiers play a pivotal role in reconstructing primate phylogeny, because they appear to comprise, along with Anthropoidea, one of only two extant haplorhine clades. Their fossils are extremely rare. Here, we describe a new species of Tarsius from the Middle Miocene of Thailand. We reconstructed aspects of its orbital morphology using a geometric-morphometric method. The result shows that the new species of Tarsius had a very large orbit (falling within the range of variation of modern Tarsius) with a high degree of frontation and a low degree of convergence. Its relatively divergent lower premolar roots suggest a longer mesial tooth row and therefore a longer muzzle than in extant species. The new species documents a previous unknown Miocene group of Tarsius, indicating greater taxonomic diversity and morphological complexity during tarsier evolution. The current restriction of tarsiers to offshore islands in Southeast Asia appears to be a relatively recent phenomenon.     

Preliminary Data on the Behavior,Ecology, and Morphology of Pygmy Tarsiers (<Emphasis Type="Italic">Tarsius pumilus</Emphasis>)

We here present the first behavioral, morphological, and ecological data on a living group of highland pygmy tarsiers, Tarsius pumilus, and compare them with the lowland tarsier species of Sulawesi. Pygmy tarsiers were previously known only from 3 museum specimens and had not been seen alive since the first specimens were collected in 1917 and 1930. As part of a 2-mo exploratory study, we recently located a group consisting of ≥4 pygmy tarsiers in the mossy cloud forest of Lore Lindu National Park, Central Sulawesi, Indonesia. We captured 1 adult female and 2 adult male group members. The tarsiers weighed a mean of 50 g, less than half the weight of adult lowland tarsiers. Our behavioral observations indicate that these tarsiers live in small groups that return to the same sleeping tree each morning. Though our data are preliminary, pygmy tarsier body proportions appear to differ from the lowland Sulawesian tarsier species, with relatively long hind limbs compared to total body length. We also observed differences in grouping and communication behaviors. Unlike lowland tarsiers, pygmy tarsiers possess multiple adult males per group and rarely vocalize or scentmark. These differences may correlate with variables unique to their high altitude habitat, especially a reduction in food and tree resources, given that tree density declined sharply along an altitudinal gradient. Our preliminary study identifies the importance of altitudinal ecological gradients to tarsier behavioral ecology in Central Sulawesi, providing questions and predictions for future research directions.     

Positional behavior and social organization of the philippine tarsier (Tarsius syrichta)

This study provides preliminary data on the positional behavior and habitat use of the Philippine tarsier,Tarsius syrichta, which was studied in its natural habitat on the island of Leyte. The locomotor behavior ofT. syrichta is quite comparable to the other species of tarsiers but frequency of use of different postures and support types does seem to distinguish this species from the others.Tarsius bancanus appears to be the most specialized, using vertical clinging postures the most frequently and sitting the least;Tarsius dianae uses vertical clinging the least frequently and sitting the most, andT. syrichta is intermediate. The Philippine tarsier has a small home range similar to that ofT. spectrum. On the other hand, this species likely does not have the small family groups typical of Sulawesi tarsiers but instead uses a noyau system likeT. bancanus. This data supports some apsects ofNiemitz’s (1977) model of anatomical and behavioral specialization among tarsier species.     

A Novel Reproductive Mode in Frogs: A New Species of Fanged Frog with Internal Fertilization and Birth of Tadpoles

We describe a new species of fanged frog (Limnonectes larvaepartus) that is unique among anurans in having both internal fertilization and birth of tadpoles. The new species is endemic to Sulawesi Island, Indonesia. This is the fourth valid species of Limnonectes described from Sulawesi despite that the radiation includes at least 15 species and possibly many more. Fewer than a dozen of the 6455 species of frogs in the world are known to have internal fertilization, and of these, all but the new species either deposit fertilized eggs or give birth to froglets.     

Origin of the Sulawesi macaques (Cercopithecidae: Macaco) as suggested by mitochondrial DNA phylogeny

One of the sharpest biogeographical transitions in the world occurs between the Indonesian islands of Borneo and Sulawesi; this transition is demarcated by Wallace's line. Macaque monkeys represent an interesting anomaly to faunal distributions in this region as they occur on both sides of Wallace's line, with Macacafascicularis, M. nemestrina and other species to the west and seven Sulawesi species to the east. We have investigated macaque evolution and dispersal in the Sunda region and Sulawesi using phylogenetic analysis of mitochondrial DNA sequences. Female philopatry of macaques, which causes sharp geographic clustering of maternally inherited mitochondrial DNA haplotypes, makes mitochondrial phytogenies particularly useful for investigating ancient patterns of dispersal. Results of this study suggest the following: (1) M. fascicularis is not a sister taxon to any species of Sulawesi macaque; (2) haplotypes of some M. nemestrina have a sister relationship to northern and central Sulawesi macaques, while haplotypes of other M. nemestrina have a sister relationship to soudiern Sulawesi macaques; (3) Sulawesi was probably colonized by macaques twice, once to the base of the northern peninsula now occupied by M. hecki and once to the southwestern peninsula now occupied by M, mama; and (4) within north/central and southern Sulawesi, patterns of dispersal are largely consistent with contemporary and past geography of the island, with the exception of a geographically discontinuous relationship between M. nigra and a portion of M. tonkeana from a region in northwest central Sulawesi.     

A New Species of Muscicapa Flycatcher from Sulawesi,Indonesia

The Indonesian island of Sulawesi, a globally important hotspot of avian endemism, has been relatively poorly studied ornithologically, to the extent that several new bird species from the region have been described to science only recently, and others have been observed and photographed, but never before collected or named to science. One of these is a new species of Muscicapa flycatcher that has been observed on several occasions since 1997. We collected two specimens in Central Sulawesi in 2012, and based on a combination of morphological, vocal and genetic characters, we describe the new species herein, more than 15 years after the first observations. The new species is superficially similar to the highly migratory, boreal-breeding Gray-streaked Flycatcher Muscicapa griseisticta, which winters in Sulawesi; however, the new species differs strongly from M. griseisticta in several morphological characters, song, and mtDNA. Based on mtDNA, the new species is only distantly related to M. griseisticta, instead being a member of the M. dauurica clade. The new species is evidently widely distributed in lowland and submontane forest throughout Sulawesi. This wide distribution coupled with the species'' apparent tolerance of disturbed habitats suggests it is not currently threatened with extinction.     

Taxonomic revision of shrews of the genus Crociduras from the Sunda Shelf and Sulawesi with description of two new species (Mammalia: Soricidae)

The skulls of 387 shrews of the genus Crocidura sampled in peninsular Malaysia, Sumatra, Borneo, Java and Sulawesi were submitted to principal component and stepwise discriminant analyses. These analyses helped to delineate morphological taxa in this species-rich genus of mammals. Most morphologic groups could be attributed to described species, except one taxon from Sumatra and one from Sulawesi, which are described and named as new. Most of the 21 species recognized in this paper are endemic to one major island. Although Sulawesi has never been connected to the mainland, it supports at least six species, followed by Sumatra (5–6 species), Java and the Malay Peninsula (4 spp) and Borneo (3 spp). C. monticola is apparently the only widespread species whose distribution range covers the entire Malay Archipelago except the Philippines and Sulawesi. In contrast, the continental C. fuliginosa enters only marginally into the Sunda Shelf: its southernmost record is on the Malay Peninsula. This interpretation is completely different from the classical view that C. fuliginosa is a cosmopolitan species occupying the whole of Southeast Asia. Identification keys, tables of measurements and discriminant functions provided in this work may aid in identification of the various species and subspecies of Crocidura living in the Malaya Archipelago.     

Infanticide by a male spectral tarsier (<Emphasis Type="Italic">Tarsius spectrum</Emphasis>)

As part of a long-term field study on the behavioral ecology and conservation of a population of spectral tarsiers (Tarsius spectrum) in Tangkoko Nature Reserve, Sulawesi, Indonesia, I observed the first known instance of infanticide in this species. A young infant was killed by a neighboring male. Neither the epiphenomenal aggression hypothesis, social pathology hypothesis, nutritive benefits of cannibalism hypothesis, nor sexual selection hypothesis are supported by this observation. The only hypothesis that could not be rejected outright, on the basis of this single observation, was the competition for limited resources hypothesis.     

Small,odd and old: The mysterious Tarsius pumilus is the most basal Sulawesi tarsier

In this study, we present the first genetic evidence of the phylogenetic position of Tarsius pumilus, the mountain tarsier of Sulawesi, Indonesia. This mysterious primate is the only Eastern tarsier species that occurs exclusively in cloud forests above 1800 m.a.s.l. It exhibits striking morphological peculiarities—most prominently its extremely reduced body size, which led to the common name of ‘pygmy tarsier’. However, our results indicate that T. pumilus is not an aberrant form of a lowland tarsier, but in fact, the most basal of all Sulawesi tarsiers. Applying a Bayesian multi-locus coalescent approach, we dated the divergence between the T. pumilus lineage and the ancestor of all other extant Sulawesi tarsiers to 9.88 Mya. This is as deep as the split between the two other tarsier genera Carlito (Philippine tarsiers) and Cephalopachus (Western tarsiers), and predates further tarsier diversification on Sulawesi by around 7 Myr. The date coincides with the deepening of the marine environment between eastern and western Sulawesi, which likely led to allopatric speciation between T. pumilus or its predecessor in the west and the ancestor of all other Sulawesi tarsiers in the east. As the split preceded the emergence of permanent mountains in western Sulawesi, it is unlikely that the shift to montane habitat has driven the formation of the T. pumilus lineage.     

Herpetofaunal Richness and Community Structure of Offshore Islands of Sulawesi,Indonesia1

We examined the distribution and broad habitat associations of the herpetofauna on three offshore islands of southeast Sulawesi, Indonesia. A total of 74 amphibian and reptile taxa were recorded, comprising 13 frogs, 29 lizards, 29 snakes, 1 freshwater turtle, and 1 crocodile. Of the total taxa, 38 percent were endemic to Sulawesi, 13 were new undescribed taxa. Range extensions were also recorded for one taxon previously not known from Sulawesi. Herpetofauna of these islands is largely derived from that of mainland Sulawesi, and as for Sulawesi generally, is depauperate compared with herpetofaunal assemblages in Borneo, Java, and Thailand. Taxon richness was much higher in minimally disturbed forest and forest habitats with only moderate disturbance levels than in highly modified or disturbed habitats, such as secondary forests, plantations, and villages. Disturbed habitats were characterized by widespread, habitat generalists and human commensals. Forests were characterized by endemic and habitat‐specialist taxa. Little discrimination of taxon composition or endemism was found between minimally and moderately disturbed forest habitats. These results reaffirm the need for more general biological survey and research in this region. Taxa most likely to be displaced by human impacts tend to be endemic taxa, for which there exists little or no ecological information. The similarity in the herpetofaunal community structure between habitats with minimal and moderate disturbance levels has important implications for our understanding of ecological resilience in tropical herpetofaunal communities.     

Five additions to the list of Sepsidae Diptera for Vietnam: Perochaeta cuirassa sp. n., Perochaeta lobo sp. n., Sepsis spura sp. n., Sepsis sepsi Ozerov, 2003 and Sepsis monostigma Thompson, 1869

A recent collecting trip to Vietnam yielded three new species and two new records of Sepsidae (Diptera) for the country. Here we describe two new species in the species-poor genus Perochaeta (Perochaeta cuirassasp. n. andPerochaeta lobosp. n.) and one to the largest sepsid genus Sepsis (Sepsis spurasp. n.) which is also found in Sumatra and Sulawesi. Two additional Sepsis species are new records for Vietnam (Sepsis sepsi Ozerov, 2003; Sepsis monostigma Thompson, 1869). We conclude with a discussion of the distribution of Perochaeta and the three Sepsis species.     

THE BIOGEOGRAPHY OF SULAWESI REVISITED: IS THERE EVIDENCE FOR A VICARIANT ORIGIN OF TAXA ON WALLACE'S “ANOMALOUS ISLAND”?

Sulawesi, the largest island in the Indonesian biodiversity hotspot region Wallacea, hosts a diverse endemic fauna whose origin has been debated for more than 150 years. We use a comparative approach based on dated phylogenies and geological constraints to test the role of vicariance versus dispersal in the origin of Sulawesi taxa. Most divergence time estimates for the split of Sulawesi lineages from their sister groups postdate relevant tectonic vicariant events, suggesting that the island was predominantly colonized by dispersal. Vicariance cannot be refuted for 20% of the analyzed taxa, though. Although vicariance across Wallace's Line was only supported for one arthropod taxon, divergence time estimates were consistent with a "tectonic dispersal" vicariance hypothesis from the East in three (invertebrate and vertebrate) taxa. Speciation on Sulawesi did not occur before the Miocene, which is consistent with geological evidence for more extensive land on the island from that time. The Pliocene onset of periodic sea-level changes may have played a role in increasing the potential for dispersal to Sulawesi. A more extensive taxon sampling in Wallacea will be crucial for refining our understanding of the region's biogeography and for testing hypotheses on the origin of taxa on its most important island.     

Ancient lakes as hotspots of diversity: a morphological review of an endemic species flock of <Emphasis Type="Italic">Tylomelania</Emphasis> (Gastropoda: Cerithioidea: Pachychilidae) in the Malili lake system on Sulawesi,Indonesia

The viviparous freshwater gastropod Tylomelania (Caenogastropoda: Cerithioidea: Pachychilidae) endemic to the Indonesian island Sulawesi has radiated extensively in two ancient lake systems. We here present the first systematic species-level review of taxa in the five lakes of the Malili lake system, which contains the most diverse and best studied freshwater fauna on Sulawesi. Our results indicate a significantly higher diversity of Tylomelania in these lakes than previously perceived based on morphological evidence for delimiting the taxa. We describe nine new species, thus increasing the number of taxa known from the Malili lakes to 25. Tylomelania species are inhabiting all available substrates in the lakes, and the diversity of habitats is reflected in an unparalleled range of radula types in this closely related group. Several species show a high intraspecific variability in some characters, and their closer investigation will probably lead to the discovery of more cryptic species. As it is, this species flock on Sulawesi is among the largest freshwater mollusc radiations known. Since the Malili lake system also contains other large endemic species flocks of e.g. crustaceans and fishes, it is a major hotspot of freshwater biodiversity in Asia to become a conservation priority. Handling editor: K. Martens     

Phylogeny and biogeography of water skinks of the genus Tropidophorus (Reptilia: Scincidae): a molecular approach

Phylogenetic relationships of the Oriental semiaquatic lygosomine skinks of the genus Tropidophorus were inferred from 1219 base positions of mitochondrial 12S and 16S rRNA genes. Results of the phylogenetic analyses incorporating data for representatives of other lygosomine genera indicated that the basal phylogenetic split within Tropidophorus separated a clade of continental Indochinese species exclusive of T. cocincinensis and T. microlepis from one comprising T. cocincinensis , T. microlepis and species from Borneo, Sulawesi and the Philippines. Of the latter group, the two continental species form the sister taxon to a clade comprising the island species. Diversification among species in Indochina and among Borneo, the Philippines and Sulawesi was likely concentrated in the Miocene, with no apparent dispersal among these regions during the Pleistocene. The body depression recognized in several Indochinese species is likely to have occurred twice in parallel as an adaptation to saxicolous habitats.