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The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1–8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000–30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located. © 2022, The Author(s). 8028-00005B; IP DKRVO 2019-2023, MK000094862; 220457/Z/20/Z, ERC-2013-StG-337574-UNDEAD, ERC-2019-StG-853272-PALAEOFARM; 075-15-2021-1069; European Molecular Biology Organization, EMBO: 217223/Z/19/Z; Vallee Foundation; Velux Fonden; Wellcome Trust, WT; Francis Crick Institute, FCI: FC001595; Horizon 2020 Framework Programme, H2020: 796877; Medical Research Council, MRC; Natural Environment Research Council, NERC: 210119/Z/18/Z, NE/K003259/1, NE/K005243/1, NE/S00078X/1, NE/S007067/1; Cancer Research UK, CRUK; European Research Council, ERC: 852558; Grantová Agentura České Republiky, GA ČR: 15-06446S; Svenska Forskningsrådet Formas: 2018-01640; Knut och Alice Wallenbergs Stiftelse; Vetenskapsrådet, VR: 681396, BELSPO B2/191/P2/ICHIE; Russian Science Foundation, RSF: 16-18-10265-RNF, 20-17-00033, 21-18-00457-RNF, 310763; Science for Life Laboratory, SciLifeLab; Narodowa Agencja Wymiany Akademickiej, NAWA: PPN/PPO/2018/1/00037 This work was supported by grants to P. Skoglund from the European Research Council (grant no. 852558), the Erik Philip Sörensen Foundation and the Science for Life Laboratory, Swedish Biodiversity Program, made available by support from the Knut and Alice Wallenberg Foundation. A.B., L.S., P. Swali and P. Skoglund were supported by Francis Crick Institute core funding (FC001595) from Cancer Research UK, the UK Medical Research Council and the Wellcome Trust. P. Skoglund was also supported by the Vallee Foundation, the European Molecular Biology Organisation and the Wellcome Trust (217223/Z/19/Z). Computations were supported by SNIC-UPPMAX. We also acknowledge support from Science for Life Laboratory, the Knut and Alice Wallenberg Foundation, the National Genomics Infrastructure funded by the Swedish Research Council and the Uppsala Multidisciplinary Center for Advanced Computational Science for assistance with massively parallel sequencing and access to the UPPMAX computational infrastructure. We thank the Yukon gold mining community and First Nations, including the Tr’ondëk Hwëch’in, for continued support of our palaeontology research in the Yukon Territories, Canada. We thank the Danish National High-Throughput Sequencing Centre and BGI-Europe for assistance in sequencing data generation and the Danish National Supercomputer for Life Sciences–Computerome ( https://computerome.dtu.dk ) for computational resources. We thank National Museum Wales for continued sampling support. M. Germonpré acknowledges support from the Brain.be 2.0 ICHIE project (BELSPO B2/191/P2/ICHIE). M.T.P.G. was supported by the European Research Council (grant no. 681396). M.-H.S.S. was supported by the Velux Foundations through the Qimmeq Project, the Aage og Johanne Louis-Hansens Fond and the Independent Research Fund Denmark (8028-00005B). L.D. acknowledges support from FORMAS (2018-01640). D.W.G.S. received funding for this project from the European Union’s Horizon 2020 research and innovation programme under Marie Skłodowska-Curie grant agreement no. 796877. M.P. was supported by the Polish National Agency for Academic Exchange–NAWA (grant no. PPN/PPO/2018/1/00037). V.J.S. was supported by the University of Zurich’s University Research Priority Program ‘Evolution in Action: From Genomes to Ecosystems’. This research was done with the participation of ZIN RAS (grant no. 075-15-2021-1069). We are grateful to the museum of the Institute of Plant and Animal Ecology UB RAS (Ekaterinburg, Russia) for provision of samples. R.P.J. and C.O’D. were supported by the Standing Committee for Archaeology of the Royal Irish Academy through the Archaeological Excavation Research Grant Scheme. E.Y.P., P.N. and V.V.P. are supported by the Russian Science Foundation (grant no. 16-18-10265-RNF and 21-18-00457-RNF). Y.V.K. was supported by the Russian Science Foundation (grant no. 20-17-00033). M.H. was supported by the European Research Council (consolidator grant GeneFlow no. 310763). M.L.-G. was supported by the Czech Science Foundation GAČR (grant no. 15-06446S) and institutional financing of the Moravian Museum from the Czech Ministry of Culture (IP DKRVO 2019-2023, MK000094862). L.S. is supported by the Sir Henry Wellcome fellowship (220457/Z/20/Z). We thank Staatliches Museum für Naturkunde Stuttgart for sample access. L.F. and G.L. were supported by European Research Council grants (ERC-2013-StG-337574-UNDEAD and ERC-2019-StG-853272-PALAEOFARM) and Natural Environmental Research Council grants (NE/K005243/1, NE/K003259/1, NE/S007067/1 and NE/S00078X/1). L.F. was also supported by the Wellcome Trust (210119/Z/18/Z). This research was funded in whole, or in part, by the Wellcome Trust (FC001595). For the purpose of open access, the author has applied a CC-BY public copyright licence to any author accepted manuscript version arising from this submission.

Despite a considerable amount of research, the systematics of the ranid genus Huia have remained unresolved, mostly owing to insufficient sampling and morphological similarities. As currently circumscribed, Huia consists of five species, but multiple studies have consistently demonstrated that it is not a monophyletic genus. However, no study has approached the problem with adequate data and provided a systematically sound solution, leaving the genus to languish in a classification that is phylogenetically incoherent. We generated the most comprehensive sampling of Huia to date, based on extensive fieldwork in Java and Sumatra. Using an integrative taxonomy framework, we analysed four mitochondrial and two nuclear markers and, in conjunction with tadpole morphology, investigated the phylogenetics of Huia and its congeners. Corroborating previous studies, Huia is recovered as a paraphyletic group. Huia cavitympanum emerges as the sister taxon to Meristogenys. The remaining members of Huia form a monophyletic group, sister to the H. cavitympanum + Meristogenys clade. Our extensive geographical sampling in Sumatra and Java reveals multiple highly divergent lineages that potentially represent undescribed species. Using our expanded molecular and morphological dataset, we resolve the paraphyly of Huia by restricting the genus to its type species and propose a new genus to accommodate the strongly supported clade of Sumatran and Javan populations previously belonging to Huia.

Rivers are known to act as biogeographic barriers in several strictly terrestrial taxa, while possibly serving as conduits of dispersal for freshwater-tolerant or -dependent species. However, the influence of river systems on genetic diversity depends on taxa-specific life history traits as well as other geographic factors. In amphibians, several studies have demonstrated that river systems have only minor influence on their divergence. Here, we assess the role of the paleodrainage systems of the Sunda region (with a focus on the island of Sumatra) in shaping the evolutionary history of two genera of frogs (Sumaterana and Wijayarana) whose tadpoles are highly dependent on cascading stream habitats. Our phylogenetic results show no clear association between the genetic diversification patterns of both anurans genera and the existence of paleodrainage systems. Time-calibrated phylogenies and biogeographical models suggest that these frogs colonized Sumatra and diversified on the island before the occurrence of the Pleistocene drainage systems. Both genera demonstrate phylogenetic structuring along a north-south geographic axis, the temporal dynamics of which coincide with the geological chronology of proto Sumatran and -Javan volcanic islands. Our results also highlight the chronic underestimation of Sumatran biodiversity and call for more intense sampling efforts on the island.

Tadpoles of the Vampire tree frog Rhacophorus vampyrus differ substantially from other rhacophorid tadpoles, by having profound modifications in external morphology. The morphological peculiarities of this species likely correlate with their arboreal microhabitat and strict oophagous diet. In this work, we examine buccal and musculoskeletal anatomy and compare them to other rhacophorid and egg-eating larvae. The shape and arrangement of cartilages of the lower jaw are unique among tadpoles, and the lack of a palatoquadrate suspensorium is only known in the distantly related macrophagous tadpoles of the dicroglossid Occidozyga baluensis. The cranial musculature is massive, and the morphology of several mandibular, hyoid, and abdominal muscles could be related to the ingestion and transit of large eggs. In the buccal cavity, conspicuous aspects are the absence of ridges and papillae, and the development of a unique glandular zone in the buccal floor. Finally, observations of the skeletal support of keratinized mouthparts allow us to present a topography-based hypothesis of homology of the conspicuous fangs of these tadpoles.

The current account presents the results of a 14-day amphibian survey at Maliau Basin Conservation Area (MBCA). With a total of approximately 170 man-hrs, 44 species were detected at four study sites during the field period; four more species were later discovered outside the two-week campaign. The results are compared to the results of previous surveys. Apart from adults, we present the first photographic documentation of the larval stages ofChiromantisinexpectatusand BorneanPhrynoidisjuxtaspera, along with a brief tadpole description; the better-known tadpoles of four more species were recorded. The results of our expedition suggest that nine more species are present at MBCA than reported by previous studies. We present an updated list of known species in the MBCA, comprising 61 species. The species accumulation curve over the 14 days period of the core survey did not show signs of asymptotic saturation. We conclude that the definitive species number for MBCA amphibians has the potential to increase with more thorough surveys in the future.

While the island of Borneo is considered a global biodiversity hotspot, the species richness in many groups remains unknown and appears underestimated. During herpetological surveys carried out in the interior of Sarawak, East Malaysia, several individuals of a small species of the genus Ansonia Stoliczka 1870 were collected on the Usun Apau plateau and in the Gunung Hose mountain range (Ansonia sp. Usun Apau). An integrative taxonomic approach comprising phylogenetic (2.4 kb mitochondrial rDNA fragment, Bayesian Inference and Maximum Likelihood, >5.1 % to its closest relative) and morphometric analyses (25 measurements, multivariate ratio analysis and linear discriminant analysis), as well as morphological comparisons support the status of this operational taxonomic unit as a separate taxon at species level. The obtained phylogenetic hypothesis corroborates the two major clades within Ansonia found in previous studies. Within Clade One Ansonia sp. Usun Apau and the enigmatic Ansonia torrentis are part of a monophyletic group of the Bornean species Ansonia hanitschi, Ansonia minuta, Ansonia platysoma, Ansonia spinulifer, Ansonia vidua, and two additional undescribed taxa. This subclade must be considered as the result of an on-island radiation in the complex evolution of Ansonia. The new species is formally described including the identification of diagnostic morphometric traits. Ansonia sp. Usun Apau is endemic to two isolated mountain ridges in central Sarawak and must be considered as a new element of the unique diversity of the Bornean amphibian fauna that is potentially threatened by habitat loss at least in parts of its range.

The tadpoles of many anuran amphibians inhabit lotic habitats and evolved oral devices to adhere to the substratum. Although published anatomical descriptions of rheophilous tadpoles exist, little is known about the modifications in gastromyzophorous tadpoles that possess abdominal suckers and live in torrential sections of streams. We describe the gastromyzophorous tadpoles of Huia cavitympanum and Meristogenys jerboa from torrential streams of Borneo, with special attention to the anatomy of their abdominal suckers and their relations to cranial structures and musculature. One cranium of H. cavitympanum and its associated muscles were computer-reconstructed in three dimensions from serial histological sections. The abdominal sucker and oral sucker comprise a set of muscles and ligaments that attach to internal skeletal structures. Some muscles could be identified to attach directly to soft tissue of the abdominal sucker and most likely contribute to suction. Comparing tadpoles of H. cavitympanum to the closely related gastromyzophorous M. jerboa reveals differences in external and internal features, such as cornu trabeculae fusion and jaw details. Because of phylogenetic uncertainties, it is unclear whether or not this structural complex evolved once or several times convergently in ranids.

Tadpoles of Occidozyga species have been reported to be carnivorous, feeding on insects and other tadpoles. We present photographic evidence for the previously undocumented larval feeding behavior in O. baluensis. Furthermore, we present a detailed anatomical description of the skull, cranial musculature, and gross gut morphology based on three-dimensional reconstructions from serial sections and μCT imagery. The cranial anatomy of larval O. baluensis is highly derived in many characters, with respect to taxa outside the genus Occidozyga, most notably the palatoquadrate and hyobranchial apparatus, that play a major role in tadpole feeding. A large larval stomach was present in the specimens examined, indicative of a macrophagous carnivorous mode of feeding. Because of the relatively small oral orifice, relatively large-sized food items found in the larval stomach, and the tunnel-like arrangement of structures that form the buccal cavity, we hypothesize that suction feeding utilizing strong negative pressure is employed by this species. Furthermore, we propose that force, rather than speed, is the main characteristic of their feeding. The unique features of the study species substantially expand the known morphospace for tadpoles, particularly among the Acosmanura (Pelobatoidea, Pelodytoidea, and Neobatrachia). Except for Microhylidae, acosmanurans previously described possess limited innovative larval morphologies. Larval carnivory has evolved convergently several times in distant anuran clades and shows structural, behavioral, and functional differences in the known examples.

Lea E. Waser, Manuel Schweizer, Alexander Haas, Indraneil Das, André Jankowski, Pui Yong Min, Stefan T. Hertwig (2016): From a lost world: an integrative phylogenetic analysis of Ansonia Stoliczka, 1870 (Lissamphibia: Anura: Bufonidae), with the description of a new species. Orgamisms, Diversity & Evolution 16: 1-17, DOI: 10.1007/s13127-016-0294-2

Ansonia teneritas, sp. nov. Holotype: NMBE 1057364, from a small stream on the upper reaches of the Sungei Julan on the northern side of the Usun Apau Plateau just above the falls of the Sungei Julan Valley, Usun Apau National Park, Sarawak, East Malaysia (Borneo). GPS N 03.03292°, E 114.66647°, 1002 m a.s.l, coll. A. Haas, Pui Yong Min, S.T. Hertwig & A. Jankowski, 28. August 2010. Adult male. Paratypes: NMBE 1057318 – 1057327, ZRC 1.12525- 12526, from the same locality as holotype, coll. A. Haas, Pui Yong Min, S.T. Hertwig & A. Jankowski, 27 and 28 August 2010; ZMH A11568, from Gunung Hose, Sarawak, East Malaysia (Borneo), GPS N 02.23526°, E 113.68693°, 1104 m asl., coll. A. Haas & Y.M. Pui, 25. August 2011. Diagnosis The new species is assigned to the genus Ansonia on the basis of a robustly supported phylogenetic hypothesis (Fig. 3) and the presence of the following morphological characters: body slender, subarticular tubercles indistinct, limbs long and slen- der, webbing membranous, parotid glands absent, and tympanum visible (Inger 1960, 1966). Ansonia teneritas sp. nov. can be distinguished from all congeners by the following combination of characters: (1) SVL in males Description of holotype Habitus very slender (Fig. 5); body slightly wider than head; head depressed (Fig. 6); head shorter than it is wide; rostrum truncate in dorsal view; tip of snout rounded, longer than canthus, protruding in lateral view, clearly projecting beyond tip of mandible; rostral length shorter than eye diameter; nostril lateral, closer to snout tip than to eye; indistinct vertical ridge between tip of snout and centre of maxilla; canthus rounded; lores vertical and slightly concave; eye diameter greater than eye-nostril distance; interorbital distance slightly larger than the upper eyelid; tympanum round and distinct; eye diameter 3.48 times greater than tympanic diameter; distance from eye to tympanum 0.43 mm Fingers slender; tips slightly expanded forming small, spatulate discs; first finger significantly shorter than second; nuptial pads with blackish brown spines on first finger; subarticular tubercles weak and indistinct, exclusively below the proximal articulation of the first phalange, absent below distal phalanges; palm smooth with large, distinct rounded palmar tubercles; tips of all toes rounded; fifth toe slightly longer than third; membranous web reaching disc of first toe, disc on both sides of second toe, third toe with one and two-thirds phalanges free, fourth with three phalanges free, fifth toe with two phalanges free of web. Subarticular tubercles indistinct; two metatarsal tubercles, both oval and slightly raised; tarsal fold absent. Skin of head, dorsum, flanks, and dorsal surfaces of limbs with scattered, isolated, flat, rounded warts of heterogeneous size, warts on temporal and dorsolateral regions not significantly larger (Figs. 5 and 7); warts on head, dorsum, flanks, and limbs often terminating in fine brown keratinous spines or caps; spines below the symphysis of mandibles absent, but one row of very small, spineless tubercles along the ventral face of the lower jaw; skin flaps on posterior thigh near vent absent. Colour in preservative grayish brown with a contrasting dark gray pattern of bands and spots on head and dorsum beginning at the eyes and running to the forehead and shoulders, forming an x with a light spot in its center; irregular light spot below the eye on upper lip; limbs gray brown with contrasting lighter cross bands; venter pale yellowish white with dark mottling on throat, chest, and belly; small, whitish tubercles present on the pectoral region and the venter (Fig. 7). Coloration in life olive-green with a contrasting dark brown pattern of irregular spots and bands, often forming an x-shape between the eyes, on forehead, and on shoulders; larger warts on dorsum yellow to orange, whitish warts ventrolaterally on flanks and from jaw joint to shoulder, an irregular whitish spot below the eye on the upper lip, limbs dark brown with yellow crossbars (Fig. 5); venter pale yellowish white with brown mottling, especially on throat and chest, and interspersed with whitish tubercles. Iris bright red-orange with an irregular network of black reticulations (Fig. 5). Variation The extent of the dark pattern on throat, chest, and venter varies from a few markings on the anterior part of the throat to intense mottling on throat, chest, and belly (Fig. 7). Some specimens possess up to three rows of very small, spineless tubercles along the ventral face of the lower jaws, while in other specimens, such tubercles are lacking. There is no noticeable variation in the described characters between the specimens from Usun Apau National Park and the single female specimen from Gunung Hose, Sarawak. Sexual dimorphism Males are smaller (SVL 1 9.1 6 – 2 1.5 3 mm) than females (21.37 – 24.4 mm); males possess vocal sacs and nuptial pads with numerous blackish brown spines. Females have less extensive webbing: the first toe is fully webbed, the second toe is about one phalange free of web, the third toe about two, the fourth about three to three and a half, and the fifth about two to two and a half free of web. Comparison The following characters distinguish Ansonia teneritas sp. nov. from its other Bornean congeners: Ansonia albomaculata Inger 1960: tympanum obscured by skin; head not depressed; sharp tarsal ridge present; skin of body and limbs uniformly covered with small, round warts; an oblique light band from the posterior corner of the eye to axilla; no contrasting pattern on head, dorsum, or limbs (Inger 1966; Inger and Stuebing 2005). Ansonia echinata: black spines under mandible, large ventrolateral tubercles tipped with small black spines; spinose tubercles on top of snout with black spines, no contrasting pattern on head, dorsum, or limbs (Inger and Stuebing 200 9). Ansonia fuliginea: larger body size (males 32 – 36 mm, females 38 – 44 mm); snout vertical in profile; head not depressed; no contrasting pattern on head, dorsum, or limbs, blackish on sides and underside (Inger 1960, 1966; Inger and Stuebing 2005). Ansonia guibei Inger 1966: snout vertical in profile; head not depressed; first finger long, reaching base of second when adpressed; fourth and second finger equal in length; membranous web reaching tips of third and fifth toes; skin dorsally and dorsolaterally with numerous round warts with clusters of dark spinules; oblique flap of skin present on each side of the vent; no contrasting pattern on head, dorsum, or limbs (Inger 196 6; Inger and Stuebing 20 0 5). Ansonia hanitschi: larger body size (males 20 – 28 mm, females 28 – 35 mm); males with a few spinose tubercles below symphysis of the mandibles; colour pattern on head, dorsum, and limbs obscure and less contrasting (Dring 1983; Inger 1966; Inger and Stuebing 2005); Ansonia latidisca: larger body size (males 35 mm, females 5 5 mm); head not depressed; snout almost vertical in profile; tips of the three outer fingers dilated into truncate discs, that of third as wide as tympanum; forelimb very long and slender; two rows of interorbital, conical tubercles; different colour pattern on head and back consisting of a bright green background with a distinct pattern of blackish brown spots and scattered orange or red warts (Inger 1966; Matsui et al. 2012 Ansonia leptopus: larger body size (males 35 mm, females 55 mm); head not depressed; first finger long, reaching disc of second; dorsolateral warts larger, males with rows of brown spines under mandible; no contrasting pattern on head, dorsum, or limbs (Inger 1966; Inger and Dring 1988; Inger and Stuebing 2005). Ansonia longidigita: larger body size (males 50 mm, females 65 mm); head not depressed; first finger long, reaching tip of disc of second; snout almost vertical in profile; a pair of longitudinal rows of tubercles in interorbital space; males with three to six rows of brown or black spines under mandible; no contrasting pattern on head or dorsum (Inger 1 966; Inger and Dring 1988; Inger and Stuebing 2005; Matsui et al. 2010). Ansonia minuta: head not depressed; distinct tarsal ridge; nuptial pads absent; third and fifth toes webbed to disc in males; more warts on dorsal face of the head; warts on dorsum and flanks not spinose, dark pattern on dorsum less contrasting (Inger 19 60, 1966; Inger and Stuebing 2005). Ansonia platysoma: warts on dorsum and flanks often without or with indistinct keratinous spines or caps; nuptial pads consisting of spines with a paler brown coloration; third toe usually webbed to disc in males; no white pattern on upper lip; no white warts on shoulders or flanks; colour pattern on head, dorsum, and limbs obscure and less contrasting (Inger 1960, 1966). Ansonia spinulifer: larger body size (males 30 – 4 0, females 40 – 45); head not depressed; first finger reaching tip of second when adpressed; dorsal and dorsolateral warts large, elevated and juxtaposed, tipped with one or three strong melanistic spines; males with spinose tubercles un- der mandibular symphysis and lower jaw; adults black on top with a light spot between the shoulders but without a contrasting pattern on head, dorsum, or limbs (Inger and Dring 1988; Inger and Stuebing 2 0 0 5). Ansonia torrentis Dring 1983: larger body size (males 30 – 33 mm); head relatively wide in comparison to body; warts on dorsum without keratinous spines; males with spinose tubercles under mandibular symphysis and lower jaw (Dring 1983). Ansonia vidua: larger body size (females 30 – 34 mm); head not depressed; two low, curved longitudinal ridges present on the interorbital region; skin of back, flanks and upper surfaces of the limbs covered with numerous small, regular, flat, rounded warts resulting in a velvet-like appearance; coloration of head, body and limbs uniformly black-brown with no markings (Hertwig et al. 2014). Etymology The species name teneritas, Latin for ‘ tenderness ’, refers to the gracile body shape and small body size of this species in comparison to congeners. We suggest the English name ‘ Gracile Slender Toad ’. Ecological notes The type locality of Ansonia teneritas sp. nov. is a small tributary of the upper reaches of the Sungei Julan, just above the spectacular falls of the Sungei Julan Valley on the northern side of the Usun Apau plateau (Fig. 8). The Usun Apau plateau is part of the Central Bornean plateau and forms the watershed between the Rejang and Baram river basins in central Sarawak (Hutchison 2005; Rousseau 1977). The Sungei Julan is a tributary of the Baram drainage system. It consist of several tablelands and smaller plateaus with an elevation of approximately 760 – 1 0 0 0 m with sharply cut edges, precipitous cliffs, and deep marginal embayments (Hutchison 2005). The Usun Apau highlands were formed by extensive volcanic activity during the Upper Miocene to Quaternary, and the few mountains that stand over 300 m above the tablelands are relicts of former volcanoes (Hutchison 2005). As a result of their volcanic formation history the high tablelands of the central area are built up of dacite tuff and agglomerate, while the southern mountains are covered by basalt lava (Hutchison 200 5; Campbell 1956). The area covers about 700 sq. km of forest and has been protected as the Usun Apau National Park since 200 5. The small stream at the type locality is presumably permanent and has a moderate gradient. The stream bed consists of alternating small shallow rocky pools (20 to 50 cm depth) and fast-running sections on bedrock (0.5 to 1.5 m in width, depending on the amount of precipitation) (Fig. 8). Below the type locality the stream flows to the Sungei Julan just below the upper tier of the Eastern Julan falls in the steep Sungei Julan Valley. The vegetation at the type locality is primary lower montane forest. On the surrounding hills the forest type changes to kerangas (Bornean heath) forest. All specimens of Ansonia teneritas sp. nov. were collected at night from rocks or low vegetation along the stream. Males call at night from saplings on the stream banks or on bedrock. Adult frogs of the following species were detected in the immediate vicinity of the stream and the surrounding forest: Ansonia longidigita Inger, 1 9 6 0; Hylarana picturata (Boulenger 1 9 2 0); Leptobrachella mjobergi Smith, 1 9 2 5; Limnonectes cf. kuhlii (Tschudi, 1838); Limnonectes ibanorum (Inger 1 9 6 4); Meristogenys sp., Metaphrynella sundana (Peters, 18 67); Microhyla petrigena Inger and Frogner 1979; Occidozyga laevis (Günther 1858); Philautus hosii (Boulenger 1895); Philautus macroscelis (Boulenger 1896); Philautus cf. mjobergi Smith, 1 9 2 5; Philautus tectus Dring, 1 9 8 7; and Staurois parvus Inger and Haile, 1959. The second locality from where Ansonia teneritas is known is the Gunung Hose, a relatively isolated, remote, and poorly explored mountain range in central Sarawak. The Gunung Hose range is composed of volcanic rock forming frequently steep cliff faces (Lee, 2002). The single specimen was found on low vegetation by a deeply entrenched small stream in a selectively logged secondary forest near a logging road. At the same locality adults of Ansonia longidigita Inger, 1960 and Staurois guttatus (Günther, 1858) were found; additionally, Rhacophorus borneensis Matsui, Shimada, and Sudin, 2013 was heard calling. The larval stages and call of the new Ansonia remain undocumented.

Schweizer, M., Guntert, M. & Hertwig, S. T. (2012). Out of the Bassian province: historical biogeography of the Australasian platycercine parrots (Aves, Psittaciformes). —Zoologica Scripta, 42, 13–27. Aridification from mid-Miocene onwards led to a fragmentation of mesic biomes in Australia and an expansion of arid habitats. This influenced the diversification of terrestrial organisms, and the general direction of their radiations is supposed to have been from mesic into drier habitats. We tested this hypothesis in the platycercine parrots that occur in different habitats in Australia and also colonized Pacific islands. We inferred their temporal and spatial diversification patterns using a Bayesian relaxed molecular clock approach based on three nuclear and two mitochondrial genes and model-based biogeographic reconstructions. The Bassian biota was found to be the centre of origin of platycercine parrots and diversification within two of their three clades coincided with the beginning of aridification of Australia. The associated habitat changes may have catalysed their radiation through adaptation to arid environments and vicariance because of the fragmentation of non-arid habitats. The small oceanic islands of Melanesia contributed as stepping stones for the colonization of New Zealand from Australia.

The parrot genus Prioniturus occurs in the oceanic Philippines, Palawan and Wallacea, a geologically dynamic region with a complex history of land and sea. The described taxa of Prioniturus have been variously placed in different assemblages, and different numbers of species have been recognized. However, a phylogenetic framework is so far lacking. This would be the prerequisite to reconstructing dispersal and colonization patterns of Prioniturus across and within Wallacea and the Philippines. Following our robustly supported phylogenetic hypothesis based on two mitochondrial genes, we propose to treat Prioniturus mindorensis comb. nov. as well as Prioniturus montanus and Prioniturus waterstradti as separate species. In Prioniturus discurus discurus and Prioniturus discurus whiteheadi, further studies using additional data and specimens are necessary to clarify their taxonomic status. This result is congruent with other studies demonstrating that alpha diversity of the Philippine avifauna is strongly underestimated. According to our biogeographic reconstruction, Prioniturus has diversified by a complex combination of colonization of islands and subsequent divergence in allopatry among and within island groups. Dispersal between Sulawesi/Wallacea and the Philippines occurred twice and documents a rare case of faunal exchange between these two regions. Zusammenfassung Die Papageien der Gattung Prioniturus sind auf den Philippinen und den Inseln der Wallacea verbreitet. Diese geologisch dynamische Region hat eine komplexe Geschichte der Verteilung von Land und Meer. Einigen der bisher beschriebenen Taxa innerhalb von Prioniturus wurde jeweils ein unterschiedlicher taxonomischer Rang zuerkannt. Obwohl eine phylogenetische Hypothese der Verwandtschaftsverhaltnisse dieser Gruppe bisher nicht vorliegt, wurden zudem unterschiedliche Artengruppen vorgeschlagen. Anhand unserer Analysen von Sequenzdaten zweier mitochondrialer Gene konnten wir die Ausbreitungsgeschichte der verschiedenen Prioniturus-Taxa zwischen den Inseln der Philippinen und der Wallacea sowie der Radiationen innerhalb dieser beiden biogeografischen Zonen rekonstruieren. Nach den Ergebnissen unserer gut aufgelosten phylogenetischen Hypothese sollten P. mindorensis comb. nov. sowie P. montanus und P. waterstradti in den Artrang erhoben werden. Bei P. d. discurus und P. discurus whiteheadi sind allerdings weitere Untersuchungen an umfangreicherem Material notwendig, um den Status beider Formen zu klaren. Dieses Ergebnis stimmt mit anderen Studien uberein, welche zeigen konnten, dass die Alpha-Diversitat der Vogel der Philippinen stark unterschatzt wird. Gemass unserer biogeographischen Rekonstruktion diversifizierte Prioniturus durch eine Folge von mehreren Kolonisierungsereignissen und anschliessender allopatrischer Artbildung zwischen und innerhalb von Inselgruppen. Die nachgewiesene Besiedlung von Sulawesi/Wallacea ausgehend von den Philippinen bzw. umgekehrt dokumentiert einen sonst nur selten nachgewiesenen Faunenaustausch zwischen diesen beiden Regionen.

Hertwig, S.T., Das, I., Schweizer, M., Brown, R., & Haas, A. (2012). Phylogenetic relationships of the Rhacophorus everetti-group and implications for the evolution of reproductive modes in Philautus (Amphibia: Anura: Rhacophoridae). —Zoologica Scripta, 41, 29–46. This study presents the first phylogenetic analysis of the enigmatic Rhacophorus everetti species group and the first description of its unique tadpole. A total sample of 95 12S and 16S mitochondrial rDNA sequences were compiled including new sequence data from 28 rhacophorid species. Based on 1332 and 1407 bp, respectively, and on the gap coding method applied, a new hypothesis about the phylogeny of rhacophorid tree frogs from Sundaland was obtained. While Rhacophorus was uncovered as a polyphyletic assemblage, the monophyly of the Bush Frogs of the genus Philautus, including the Rhacophorus everettigroup, is robustly supported. We, therefore, transfer the everetti-group to the genus Philautus. As a second step, we recognise Philautus macroscelis (comb. nov.) from Borneo and P. everetti (comb. nov.) from Palawan as distinct allopatric species. Molecular and morphological evidence clearly indicates that each is a distinct lineage with a unique ancestry and discrete evolutionary fate. Moreover, close phylogenetic relationships of several Philautus species from Borneo to taxa from outside Borneo were recovered; P. everetti and P. macroscelis being the only one example. These findings indicate a complex biogeographical history of Sundaland Bush Frogs, which can only be explained by repeated dispersal and vicariance events between the Asian mainland and the Sunda islands. Finally, a single tadpole discovered on Gunung Kinabalu was matched genetically to P. macroscelis. Features of its peculiar external morphology suggest that this larva is endotrophic and possibly nidicolous. A comparable reproductive biology was formerly unknown in rhacophorid tree frogs. The presence of a free-swimming tadpole in Philautus challenges the notion that terrestrial direct development represents an apomorphic character unambiguously shared by all members of this genus. The implications for the evolution of reproductive modes in Bush Frogs are discussed in a phylogenetic context.

Aim Parrots are thought to have originated on Gondwana during the Cretaceous. The initial split within crown group parrots separated the New Zealand taxa from the remaining extant species and was considered to coincide with the separation of New Zealand from Gondwana 82-85 Ma, assuming that the diversification of parrots was mainly shaped by vicariance. However, the distribution patterns of several extant parrot groups cannot be explained without invoking transoceanic dispersal, challenging this assumption. Here, we present a temporal and spatial framework for the diversification of parrots using external avian fossils as calibration points in order to evaluate the relative importance of the influences of past climate change, plate tectonics and ecological opportunity. Location Australasian, African, Indo-Malayan and Neotropical regions. Methods Phylogenetic relationships were investigated using partial sequences of the nuclear genes c-mos, RAG-1 and Zenk of 75 parrot and 21 other avian taxa. Divergence dates and confidence intervals were estimated using a Bayesian relaxed molecular clock approach. Biogeographic patterns were evaluated taking temporal connectivity between areas into account. We tested whether diversification remained constant over time and if some parrot groups were more species-rich than expected given their age. Results Crown group diversification of parrots started only about 58 Ma, in the Palaeogene, significantly later than previously thought. The Australasian lories and possibly also the Neotropical Arini were found to be unexpectedly species-rich. Diversification rates probably increased around the Eocene/Oligocene boundary and in the middle Miocene, during two periods of major global climatic aberrations characterized by global cooling. Main conclusions The diversification of parrots was shaped by climatic and geological events as well as by key innovations. Initial vicariance events caused by continental break-up were followed by transoceanic dispersal and local radiations. Habitat shifts caused by climate change and mountain orogenesis may have acted as a catalyst to the diversification by providing new ecological opportunities and challenges as well as by causing isolation as a result of habitat fragmentation. The lories constitute the only highly nectarivorous parrot clade, and their diet shift, associated with morphological innovation, may have acted as an evolutionary key innovation, allowing them to explore underutilized niches and promoting their diversification.

Statistical analyses of 60 cranial and six postcranial characters of 105 domestic cats and 77 wildcats from Slovakia showed that the two forms can be separated on the basis of just a few morphometric traits. To test the distinguishing power of these characters we first pre-classified the specimens of our sample as wildcat or domestic cat on the basis of the cranial volume, which is a non-overlapping diagnostic character in most populations investigated so far. Pearson’s r correlation matrix of all 60 cranial traits showed a high degree of correlation. A linear discriminant analysis, excluding cranial volume, showed that maximal length of M1, distance between foramen rotundum and foramen ovale and diameter of porus acusticus externus was necessary to separate the two forms. When postcranial traits were included, tibia length, femur length and distance between foramen rotundum and foramen ovale sufficed to assign the individuals to the correct group, with just one misclassification. Distinguishing between all four possible groups (domestic/wild versus male/female) was only partially possible, with a minimum 5% misclassification rate always with regard to sex. The results of the principal component analysis underlined that domestic cats and wildcats can be identified unequivocally on the basis of a set of morphometric measurements even in the absence of the best discriminating traits. In comparison with a reference population from Thuringia, Germany that the authors had studied previously, the morphological differences between domestic cats and wildcats are much more pronounced in Slovakia. Moreover, the ranking of the diagnostic value of characters differs between populations. As possible explanations for this finding, we discuss the theory of an east-west gradient in body size caused by a more maritime climate in the west and a more continental one in the east, and that of a lower degree of introgression and hybridization between the two forms in the Slovakian population.

The present study reports the discovery of the hitherto unknown breeding grounds of the large-billed reed warbler Acrocephalus orinus in the Badakhshan region of Tajikistan. A total of eight adult individuals were caught at three riverine woodland sites and breeding was confirmed at one locality. The identity of the birds was confirmed on the basis of partial sequences of the cytochrome b gene. Even among individuals sampled at the same locality, genetic diversity was surprisingly high. This was interpreted as the result of secondary contact between previously isolated or fragmented populations. The breeding range of the large-billed reed warbler is likely small and the species’ habitat is under intense pressure from the collection of firewood and fodder. Consequently, the large-billed reed warbler remains of considerable conservation concern.

We describe the external morphology of tadpoles of Ansonia longidigita Inger, 1960, A. minuta Inger, 1960, and A. platysoma Inger, 1960 from East Malaysia, Borneo. Specimens were collected from small to medium-sized streams with swift current in Sarawak State. Tadpole identities were confirmed by matching tadpole genetic sequences (16S mtRNA) to sequences from adult toads from the same collection sites. Among the three species, A. minuta appears to prefer the strongest current and was collected from fast-flowing waters in association with boulders. Ansonia longidigita larvae inhabit more moderate currents than sympatric A. platysoma. Species microhabitat choices are reflected in their body shape, with A. minuta and A. platysoma being more streamlined than A. longidigita. Colour photos of live specimens and a key to the known Bornean Ansonia tadpoles are provided to facilitate field identification.

The medaka, Oryzias latipes, is a model organism in experimental biology. Its ontogeny and osteology have been studied frequently, but no detailed description of its cranial anatomy exists. Based on manual dissections, histological slide sections, and 3D-reconstructions, we describe the anatomy of the jaw apparatus of O. latipes, focusing on musculature, connective tissue elements, and the nervous system. The soft tissue anatomy of the head of O. latipes is characterized by several derived characters that seem to be related to the modification of the upper jaw and a reduced mobility of its bones, which is an evolutionary novelty of the Beloniformes. These apomorphies could also be influenced by the small size of O. latipes. The maxilla is medial to the premaxilla. The presence of extensive connective tissue elements severely limits the mobility of both bones against each other and against the neurocranium. The external section of m. adductor mandibulae is partly fused with the internal section, originates from the quadrate, and inserts to the lower jaw. The intermediate head of the internal section is narrow and forms a bend along the ventral margin of the orbit. The intramandibular section is a separate narrow muscle between retroarcticular and dentary. These characters need to be verified in other representatives of Beloniformes to test their contribution to the disputed phylogenetic relationships within this taxon and to improve the understanding of the evolution of beak-like jaws in beloniform fishes.

While the western populations of the wildcat (Felis silvestris silvestris) in Germany come into contact with wildcats in France and Switzerland, the eastern distribution area is geographically completely isolated and consists of scattered subpopulations. To investigate population structure, evolutionary relationships and degree of hybridization with domestic cats we analysed the mitochondrial control region of 86 cats in combination with 11 microsatellite loci of 149 cats. According to our microsatellite data, German wildcats are divided into two separate populations corresponding to the western and eastern distribution areas. We found no indication of a further subdivision of the eastern population. German wildcat populations are genetically distinct from domestic cats in the main, but we identified 18.4% of the whole wildcat sample as being of hybrid origin, corresponding to 4.2% of the eastern and 42.9% of the western wildcat population, and 2.7% of the domestic cat sample. The mitochondrial haplotypes form a network of three connected clusters and reveal a high level of genetic diversity, especially within the eastern population. Our findings are explained at best in terms of continuous introgression between domestic cats and wildcat populations and differing degrees of recent hybridization in the various populations. Future conservation efforts should focus on preserving the existing gene flow between the isolated distribution areas, but also on preventing the spread of hybrids and limiting the habitat alterations that lead to increased contact with domestic cats. In conclusion we discuss possible evolutionary reasons for the still traceable genetic integrity of the wildcat despite its long history of interbreeding.

Germany’s large population of wildcats (Felis silvestris silvestris) can be clearly distinguished from domestic cats on the basis of morphological characters. However, an examination of 71 specimens from Thuringia also illustrates the risks involved in using only a few such characters. The most reliable tool for identification in the field are three pelage characters (distinctness of tail bands, stripes on the nape and stripes on the shoulder). Only two morphological characters (intestine length and cranial volume) are unambiguous and demonstrate no overlap in distribution between domestic cats and wildcats. A linear discriminant analysis with forward selection of variables showed that only five skull variables are necessary to distinguish all four groups (subspecies × sex). Additionally, the high degree of correlation between most of the 49 variables examined (as indicated by Pearson’s r correlation matrix) speaks against the utility of measuring such high numbers of characters in the future. Principal component analysis (PCA) enabled the subspecies to be separated clearly. The first PCA axis was highly correlated with variables characterizing overall body size, thus separating male and female into wildcats and domestic cats. Even when the chief differentiating characters are missing, the PCA still resulted in a good separation of subspecies. None of the genetically determined hybrids could have been deciphered unambiguously using the morphological characters still intact after a road death. Hybridization seems to occur whenever wildcats change their ecological function and become field cats. The impulse to hybridize seems to come much more from the wildcat side than the side of feral cats, and deforestation represents the major threat to the wildcat.

Genes selected for a phylogenetic study need to contain conserved information that reflects the phylogenetic history at the specific taxonomic level of interest. Mitochondrial ribosomal genes have been used for a wide range of phylogenetic questions in general and in anuran systematics in particular. We checked the plausibility of phylogenetic reconstructions in anurans that were built from commonly used 12S and 16S rRNA gene sequences. For up to 27 species arranged in taxon sets of graded inclusiveness, we inferred phylogenetic hypotheses based on different a priori decisions, i.e. choice of alignment method and alignment parameters, including/excluding variable sites, choice of reconstruction algorithm and models of evolution. Alignment methods and parameters, as well as taxon sampling all had notable effects on the results leading to a large number of conflicting topologies. Very few nodes were supported in all of the analyses. Data sets in which fast evolving and ambiguously aligned sites had been excluded performed worse than the complete data sets. There was moderate support for the monophyly of the Discoglossidae, Pelobatoidea, Pelobatidae and Pipidae. The clade Neobatrachia was robustly supported and the intrageneric relationships within Bombina and Discoglossus were well resolved indicating the usefulness of the genes for relatively recent phylogenetic events. Although 12S and 16S rRNA genes seem to carry some phylogenetic signal of deep (Mesozoic) splitting events the signal was not strong enough to resolve consistently the inter-relationships of major clades within the Anura under varied methods and parameter settings. Zusammenfassung Zur Anwendung in einer phylogenetische Analyse mussen die ausgewahlten Gene konservierte und detektierbare Information zum untersuchten phylogenetischen Niveau enthalten. Ribosomale Gene des Mitochondriums wurden fur ein breites Spektrum phylogenetischer Fragestellungen bei verschiedenen Gruppen und insbesondere bei Froschlurchen eingesetzt. Wir untersuchten die Frage, ob Rekonstruktionen der Anuren-Phylogenie, basierend auf 12S und 16S rRNA Gensequenzen, plausibel sind. An einer Auswahl von 27 Arten, arrangiert in Taxa-Gruppen abgestufter Hierarchie, rekonstruierten wir phylogenetische Hypothesen unter verschiedenen, a priori festgelegten Bedingungen. Dazu gehorten die Auswahl verschiedener Alinierungsmethoden und—parameter, der Umgang mit variabel alinierten Positionen, die Auswahl der Algorithmen zur Baumkonstruktion sowie die Auswahl alternativer Modelle der Sequenzevolution. Die Methoden und Parameter der Alinierung und der Rekonstruktion, sowie die Auswahl der Taxa, hatten bedeutenden Einfluss auf die Resultate. Daraus resultierte eine grose Anzahl alternativer Topologien, in denen nur sehr wenige Knoten in allen Analysen Unterstutzung fanden. Ausschluss variabel alinierter Positionen ergaben Topologien mit niedrigem Grad der Auflosung. Die Sequenzen enthielten ein gewisses Signal fur die Monophylie von Discoglossidae, Pelobatoidea, Pelobatidae und Pipidae. Der Knoten Neobatrachia wurde deutlich unterstutzt. Die robuste Auflosung intragenerischer Phylogenien von Bombina und Discoglossus weisen auf eine besondere Eignung der Gene fur die Untersuchung junger Aufspaltungsereignisse hin. Obwohl 12S und 16S rRNA-Gene eine heterogene Unterstutzung fur wenige fruhe (mesozoische) phylogenetische Ereignisse zeigten, war das Signal nicht geeignet, um die Beziehungen der Taxa hoherer Ordnung der Anura unter variierten Parametern und Analysemethoden konsistent aufzulosen.

This study presents a cladistic analysis of the Cyprinodontiformes based on a comparative anatomical examination of 95 species. Seventy-eight characters of musculature, connective tissue elements and nervous system of the jaw apparatus are described, in many cases for the first time. In order to investigate the contribution of such soft tissue traits, the new data have been analysed separately and in combination with an extensive data matrix of morphological, primarily osteological characters produced by Costa using the nona software. The inclusion of soft tissue characters led to the description of numerous additional apomorphies and to an improved resolution of the consensus topologies in the combined analysis. Though the consensus of the separate tree searches was limited in resolution due to a relatively high degree of homoplasy and probably to rapidly evolving adaptive transformations, phylogenetic hypotheses mostly congruent to recent phylogenetic studies were yielded. These findings are evidence for the usability of cranial soft tissue characters as a reliable additional source of information for cladistic studies of fish phylogeny. With regard to the highly contentious deep splitting events within the Cyprinodontiformes and the monophyly of the Aplocheilidae, the consideration of the new data supported a new hypothesis that agrees with the palaeogeographical framework of the break-up of Gondwana. The genus Aplocheilus from India represents the sister group to all remaining taxa of the Cyprinodontiformes, while the second branch separates the genus Pachypanchax from Madagascar. A comparable biogeographical pattern has been found congruently in different groups of teleostean freshwater fishes. The Valenciidae from Europe are closely related to the Nearctic taxon Funduloidea. The placement of the genus Fluviphylax remained unresolved because it is not only characterized by the possession of several derived characters but also by numerous plesiomorphic ones.

A new species of stream toad of the genus Ansonia is described from Gunung Murud, Pulong Tau National Park, of northern Sarawak, Malaysia, Borneo. Ansonia vidua, sp. nov., is morphologically distinguished from its Bornean congeners by the following combination of characters: medium size (SVL of adult females 33.5-34.4 mm); body uniformly black-brown in life; absence of a visible pattern on dorsum or limbs; presence of two low interorbital ridges; shagreened skin on dorsum, sides and upper surfaces of the limbs with numerous homogeneously small, rounded warts; first finger shorter than second; reduced webbing between the toes and an absence of a sharp tarsal ridge. Uncorrected genetic distances between related taxa ofgt; 4.3% in 16S rRNA gene support its status as a hitherto undescribed species.

A new brightly-coloured (olive and red) species of microhylid frog of the genus Calluella Stoliczka 1872 is described from the upper elevations of Gunung Penrissen and the Matang Range, Sarawak, East Malaysia (Borneo). Calluella capsa, new species, is diagnosable in showing the following combination of characters: SVL up to 36.0 mm; dorsum weakly granular; a faint dermal fold across forehead; toe tips obtuse; webbing on toes basal; lateral fringes on toes present; outer metatarsal tubercle present; and dorsum greyish-olive, with red spots; half of venter bright red, the rest with large white and dark areas. The new species is the eighth species of Calluella to be described, and the fourth known from Borneo. A preliminary phylogeny of Calluella and its relatives is presented, and the new taxon compared with congeners from Malaysia and other parts of south-east Asia.

Aim The Neotropical parrots (Arini) are an unusually diverse group which colonized South America in the Oligocene. The newly invaded Neotropics may have functioned as an underused adaptive zone and provided novel ecological opportunities that facilitated diversification. Alternatively, diversification may have been driven by ecological changes caused by Andean uplift and/or climate change from the Miocene onwards. Our aim was to find out whether Arini diversified in a classical adaptive radiation after their colonization of South America, or whether their diversification occurred later and was influenced by more recent environmental change. Location Neotropics. Methods We generated a time-calibrated phylogeny of more than 80% of all Arini species in order to analyse lineage diversification. This chronogram was also used as the basis for the reconstruction of morphological evolution within Arini using a multivariate ratio analysis of three size measurements. Results We found a concentration of size evolution and partitioning of size niches in the early history of Arini consistent with the process of adaptive radia- tion, but there were no signs of an early burst of speciation or a decrease in speci- ation rates through time. Although we detected no overall temporal shifts in diversification rates, we discovered two young, unexpectedly species-rich clades. Main conclusions Arini show signs of an early adaptive radiation, but we found no evidence of the slowdown in speciation rate generally considered a feature of island or lake radiations. Historical processes and environmental change from the Miocene onwards may have kept diversification rates roughly constant ever since the colonization of the Neotropics. Thus, Arini may not yet have reached equilibrium diversity. The lack of diversity-dependent speciation might be a general feature of adaptive radiations on a continental scale, and diversification processes on continents might therefore not be as ecologically limited as in isolated lakes or on oceanic islands.

The tree-frog family Rhacophoridae is a major group contributing to the high pecies richness and reproductive diversity among vertebrates of Sundaland. Nonetheless, rhacophorid evolution, specially on Borneo, has not been studied within a phylogenetic context. In this study, we examine the phylogenetic relationships of 38 (out of 41) Bornean species of Rhacophoridae, in combination with data from previous phylogenetic studies. In the final super matrix of 91 species, we analyse sequence data from two mitochondrial and three nuclear genes. The resulting trees show the genus Rhacophorus as a paraphyletic assemblage. As a consequence, we transfer Rhacophorus appendiculatus and R. kajau to two other genera and propose the new phylogeny-based combinations- Kurixalus appendiculatus and Feihyla kajau, respectively. Furthermore, we use our phylogenetic hypotheses to reconstruct the evolution of reproductive modes in rhacophorid tree frogs. Direct development to the exclusion of a free larval stage evolved twice independently, once in an ancestor of the Pseudophilautus + Raorchestes clade in India and Sri Lanka, and once within Philautus in Southeast Asia. The deposition of egg clutches covered by a layer of jelly in Feihyla is also present in F. kajau and thus confirms our generic reassignment. The remarkably high diversity of rhacophorid tree frogs on Borneo is the outcome of a complex pattern of repeated vicariance and dispersal events caused by past changes in the climatic and geological history of the Sunda shelf. We identified geographic clades of closely related endemic species within Rhacophorus and Philautus, which result from local island radiations on Borneo.

As a result of strong artificial selection, the domesticated dog has arguably become one of the most morphologically diverse vertebrate species, which is mirrored in the classification of around 400 different breeds. To test the influence of breeding history on the genetic structure and variability of today's dog breeds, we investigated 12 dog breeds using a set of 19 microsatellite markers from a total of 597 individuals with about 50 individuals analysed per breed. High genetic diversity was noted over all breeds, with the ancient Asian breeds (Akita, Chow Chow, Shar Pei) exhibiting the highest variability, as was indicated chiefly by an extraordinarily high number of rare and private alleles. Using a Bayesian clustering method, we detected significant genetic stratification within the closely related Schnauzer breeds. The individuals of these three recently differentiated breeds (Miniature, Standard and Giant Schnauzer) could not be assigned to a single cluster each. This hidden genetic structure was probably caused by assortative mating owing to breeders' preferences regarding coat colour types and the underlying practice of breeding in separate lineages. Such processes of strong artificial disruptive selection for different morphological traits in isolated and relatively small lineages can result in the rapid creation of new dog types and potentially new breeds and represent a unique opportunity to study the evolution of genetic and morphological differences in recently diverged populations.

Vicariance is thought to have played a major role in the evolution of modern parrots. However, as the relationships especially of the African taxa remained mostly unresolved, it has been difficult to draw firm conclusions about the roles of dispersal and vicariance. Our analyses using the broadest taxon sampling of old world parrots ever based on 3219 bp of three nuclear genes revealed well-resolved and congruent phylogenetic hypotheses. Agapornis of Africa and Madagascar was found to be the sister group to Loriculus of Australasia and Indo-Malayasia and together they clustered with the Australasian Loriinae, Cyclopsittacini and Melopsittacus. Poicephalus and Psittacus from mainland Africa formed the sister group Of the Neotropical Arini and Coracopsis from Madagascar and adjacent islands may be the closest relative of Psittrichas from New Guinea. These biogeographic relationships are best explained by independent colonization of the African continent via trans-oceanic dispersal from Australasia and Antarctica in the Paleogene following what may have been vicariance events in the late Cretaceous and/or early Paleogene. Our data support a taxon pulse model for the diversification of parrots whereby trans-oceanic dispersal played a more important role than previously thought and was the prerequisite for range expansion into new continents. (C) 2009 Elsevier Inc. All rights reserved

The bizarre larvae of Leptobrachella mjobergi are fossorial and live in the gravel beds of small streams. These tadpoles are vermiform in body shape. Here we present details on their skeleton and musculature, particularly of the head. The entire cranium and its associated musculature are reconstructed in three dimensions from serial histological sections. The hyobranchial apparatus is highly reduced. The head of the L. mjobergi larva is more mobile than in other anuran species. This mobility can largely be ascribed to the exclusion of the notochord from the cranial base and an articulation of the foramen magnum floor with the atlas of the tadpole. The articulation is unique among anuran species, but design parallels can be drawn to salamanders and the articulation between atlas and axis in mammals. In L. mjobergi, the atlas forms an anterior dens that articulates with the basal plate in an accessory, third occipital articular face. The muscle arrangements deviate from the patterns found in other tadpoles: For instance, epaxial and ventral trunk muscles reach far forward onto the skull. The post-cranial skeleton of L. mjobergi is considerably longer than that of other anurans: it comprises a total of 35 vertebrae, including more than 20 post-sacral perichordal centra. Despite a number of features in cranial and axial morphology of L. mjobergi, which appear to be adaptations to its fossorial mode of life, the species clearly shares other features with its megophryid and pelobatid relatives.

The megophryid frogs Leptobrachella brevicrus, Leptolalax dringi and Megophrys dringi are species exclusively known from highly localised areas in isolated mountain ranges on Borneo. The tadpoles and adults in this study were collected at the shared type locality for the three species in Gunung Mulu National Park, Sarawak, Malaysia (Borneo). The species identities of larvae were determined via comparison to syntopic adults using DNA barcoding techniques based on partial 16S rRNA mitochondrial gene sequences. The genetic data supported the status of the three taxa as valid species. Descriptions of colouration in life and after preservation, external morphological features, morphometric measurements and ecological notes in comparison to congeneric species are supplied. The tadpoles of L. brevicrus and L. dringi show similar adaptations to a fossorial lifestyle. These include an elongated, vermiform body, a relatively long tail and small eyes. Both were found in the gravel beds of a small mountain stream. In contrast, the larvae of M. dringi are adapted to occupying and feeding at the surface of pools within the stream.

A new brightly-coloured (olive and red) species of microhylid frog of the genus Calluella Stoliczka 1872 is described from the upper elevations of Gunung Penrissen and the Matang Range, Sarawak, East Malaysia (Borneo). Calluella capsa, new species, is diagnosable in showing the following combination of characters: SVL up to 36.0 mm; dorsum weakly granular; a faint dermal fold across forehead; toe tips obtuse; webbing on toes basal; lateral fringes on toes present; outer metatarsal tubercle present; and dorsum greyish-olive, with red spots; half of venter bright red, the rest with large white and dark areas. The new species is the eighth species of Calluella to be described, and the fourth known from Borneo. A preliminary phylogeny of Calluella and its relatives is presented, and the new taxon compared with congeners from Malaysia and other parts of south-east Asia.

This communication reports the discovery of the hitherto unknown larval forms of Rhacophorus rufipes and R. penanorum, andre-describes the tadpole of R. dulitensis. Tadpoles of all three species were discovered at Gunung Mulu National Park, Sarawak(Borneo), Malaysia. The identity of the larvae was determined by DNA barcoding techniques using partial 16S rRNA mito-chondrial gene sequences. Larval DNA sequences matched those of syntopic adults of respective species. Detailed descriptionsof external morphology and colouration in life are provided along with ecological notes. The tadpole of R. rufipes and R.dulitensis can be classified as generalized, benthic-nectonic type, whereas tadpoles of R. penanorum show adaptations typical for a lotic, rheophilous lifestyle.