pithecus travel

Amazing 13-Million-Year-Old Ape Skull Discovered

The remarkable skull is so well preserved, scientists can see the young ape’s unerupted teeth and an impression of its brain.

More than 13 million years ago in what’s now northern Kenya, an infant ape ended up dead in a lush forest, its body blanketed in ashfall from a nearby volcanic eruption.

Millions of years later, scientists uncovered the baby ape’s skull, the best-preserved of its kind ever found, and got an extraordinarily glimpse into the early stages of ape evolution.

“We’ve been looking for ape fossils for years—this is the first time we’re getting a skull that’s complete,” says Isaiah Nengo , the De Anza College anthropologist who led the discovery, supported by a National Geographic Society grant and the Stony Brook University-affiliated Turkana Basin Institute .

Roughly the size of a lemon, the skull belongs to a newly identified species of early ape named Nyanzapithecus alesi . Some of its features resemble those of today’s living Old World monkeys and apes, and the face bears a striking resemblance to today’s infant gibbons.

What’s more, N. alesi offers insight into early apes’ brains, the team reports in their study, published today in Nature . With a volume of about seven tablespoons, N. alesi ’s brain cavity was more than double that of other Old World monkeys from the time.

And the skull’s intact braincase, which has preserved impressions of the brain’s exterior surface, also contains the infant ape’s unerupted adult teeth.

A Lucky (Smoke) Break

After diverging from Old World monkeys’ ancestors between 25 and 28 million years ago, apes diversified near the middle of the Miocene epoch. Many of those lineages, however, died out roughly 7 million years ago amid a bout of natural climate change. Modern great apes and humans are the descendants of one of the surviving Miocene ape lineages.

The details of this evolutionary tale have remained murky, however, in part because early apes lived in rainforests, which rarely offer conditions favorable to fossilization. Until N. alesi , only one other Miocene ape skull had been found with the braincase, or neurocranium, intact.

“For those species where we have any of the cranium at all, often they’ll have jaws, the face, and sometimes the very beginning of the forehead bone,” says Brenda Benefit , a New Mexico State University anthropologist who reviewed the study before publication. “You do not get a complete neurocranium—that’s unheard of.”

Finding N. alesi took determination and a fantastic stroke of luck. The Leakeys, a family of preeminent paleoanthropologists, had previously excavated northern Kenya’s Napudet dig site. When Nengo took over excavations in 2013, few had high hopes that he would find much of note.

But one day in early 2014, one of the expedition’s assistants, John Ekusi, walked away from the crew to smoke a hand-rolled cigarette. The rest of the team grew puzzled as they watched Ekusi from afar: After a few minutes, he began circling something on the ground that had captured his attention.

Ekusi told the rest of the crew that he may have spotted the head of an elephant femur, gesturing to a rounded bony surface peeking out of the rock. Closer inspection revealed a far rarer find: a small ape skull, only gently squashed from its true-to-life proportions. The crew broke out into a dance in excitement.

The skull of N. alesi , partially excavated after careful removal of loose sand and rocks with dental picks and brushes.

With night fast approaching, however, the crew was forced to rebury the skull and wait until the next morning to excavate it. “I tell you, nobody was sleeping that night,” Nengo says.

Getting Into an Ape’s Head

Dating the sediment layer around the fossil told the team that the ape skull is about 13 million years old. But even with the skull’s fantastic preservation, initial inspection of the prepared fossil couldn’t confirm where the skull belonged on the primate family tree.

To pin that down, Nengo and his colleagues needed glimpses of its adult teeth, which hadn’t yet erupted. So the team took their find to the European Synchrotron Radiation Facility in Grenoble, France, where technicians subjected it to high-powered x-rays that allowed them to peer into the skull without damaging it.

The scans gave Nengo’s team 3-D reconstructions of the teeth, and their distinctive shapes firmly placed the skull among the nyanzapithecenes, an extinct sister group to gibbons, great apes, and humans.

“If they had not done [the synchrotron scans], they never would have been able to identify this,” says Benefit. “To me, that’s a miracle of modern technology.”

Now that N. alesi has been unveiled, Nengo is brimming with ideas for what aspect to study next. He and his colleagues are gearing up to examine the brain impressions on the skull’s interior. They are also doubling back to the ape’s exquisitely preserved ear and are working to reconstruct how N. alesi would have looked in life.

Nengo also plans to go back to Napudet, to track down other fossils he saw hints of in the ancient rock.

“That’s the plan,” he says. “There’re a few interesting things to do.”

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• v.41(6); 2020 Nov 18

Mitogenomic phylogeny of the Asian colobine genus Trachypithecus with special focus on Trachypithecus phayrei (Blyth, 1847) and description of a new species

Christian roos.

1 Primate Genetics Laboratory, German Primate Center, Leibniz Institute for Primate Research, Goettingen 37077, Germany

2 Gene Bank of Primates, German Primate Center, Leibniz Institute for Primate Research, Goettingen 37077, Germany

3 Chances for Nature (CfN), Goettingen 37073, Germany

Kristofer M. Helgen

4 Australian Museum Research Institute, Australian Museum, Sydney, New South Wales 2010, Australia

5 Natural History Museum, London SW7 BD, UK

Roberto Portela Miguez

Naw may lay thant.

6 Wildlife Conservation Society (WCS) - Myanmar Program, Yangon 11041, Myanmar

7 Fauna & Flora International (FFI) - Myanmar Programme, Yangon 11201, Myanmar

Aung Ko Lin

Khin mar yi.

8 Popa Mountain Park, Nature and Wildlife Conservation Division, Forest Department, Popa 05242, Myanmar

9 World Wide Fund for Nature (WWF) - Myanmar, Yangon 11191, Myanmar

Zin Mar Hein

Margaret nyein nyein myint, tanvir ahmed.

10 Department of Zoology, Jagannath University, Dhaka 1100, Bangladesh

Dilip Chetry

11 Primate Research and Conservation Division, Aaranyak, Guwahati, Assam 781028, India

E. Grace Veatch

12 Department of Anthropology, Emory University, Atlanta, GA 30322, USA

13 Department of Anthropology, Yale University, New Haven, CT 06511, USA

Neil Duncan

14 Department of Mammalogy, American Museum of Natural History, New York, NY 10024, USA

Pepijn Kamminga

15 Naturalis Biodiversity Center, Leiden 2333 CR, The Netherlands

Marcus A. H. Chua

16 Lee Kong Chian Natural History Museum, National University of Singapore, Singapore 117377, Singapore

17 Department of Environmental Science and Policy, George Mason University, Fairfax, VA 22030, USA

Christian Matauschek

Zhi-jin liu.

18 Key Laboratory of Animal Ecology and Conservation Biology, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China

19 Center for Excellence in Animal Evolution and Genetics, Chinese Academy of Sciences, Kunming, Yunnan 650223, China

Tilo Nadler

20 Cuc Phuong Commune, Nho Quan District, Ninh Binh Province, Vietnam

Peng-Fei Fan

21 School of Life Sciences, Sun Yat-Sen University, Guangzhou, Guangdong 510275, China

Le Khac Quyet

22 Center for Biodiversity Conservation and Endangered Species, Ho-Chi-Minh City, Vietnam

Michael Hofreiter

23 Institute for Biochemistry and Biology, University of Potsdam, Potsdam 14476, Germany

Dietmar Zinner

24 Cognitive Ethology Laboratory, German Primate Center, Leibniz Institute for Primate Research, Goettingen 37077, Germany

25 Leibniz Science Campus Primate Cognition, Goettingen 37077, Germany

26 Department of Primate Cognition, Georg-August-University, Goettingen 37083, Germany

Frank Momberg

27 Fauna & Flora International (FFI) – Asia-Pacific Programme, Yangon 11201, Myanmar

Associated Data

Mitochondrial genome sequences were submitted to GenBank and are available under accession Nos. {"type":"entrez-nucleotide-range","attrs":{"text":"MT806030-MT806070","start_term":"MT806030","end_term":"MT806070","start_term_id":"1945584014","end_term_id":"1945584574"}} MT806030-MT806070 .

Trachypithecus , which currently contains 20 species divided into four groups, is the most speciose and geographically dispersed genus among Asian colobines. Despite several morphological and molecular studies, however, its evolutionary history and phylogeography remain poorly understood. Phayre’s langur ( Trachypithecus phayrei ) is one of the most widespread members of the genus, but details on its actual distribution and intraspecific taxonomy are limited and controversial. Thus, to elucidate the evolutionary history of Trachypithecus and to clarify the intraspecific taxonomy and distribution of T. phayrei , we sequenced 41 mitochondrial genomes from georeferenced fecal samples and museum specimens, including two holotypes. Phylogenetic analyses revealed a robustly supported phylogeny of Trachypithecus , suggesting that the T. pileatus group branched first, followed by the T. francoisi group, and the T. cristatus and T. obscurus groups most recently. The four species groups diverged from each other 4.5–3.1 million years ago (Ma), while speciation events within these groups occurred much more recently (1.6–0.3 Ma). Within T. phayrei , we found three clades that diverged 1.0–0.9 Ma, indicating the existence of three rather than two taxa. Following the phylogenetic species concept and based on genetic, morphological, and ecological differences, we elevate the T. phayrei subspecies to species level, describe a new species from central Myanmar, and refine the distribution of the three taxa. Overall, our study highlights the importance of museum specimens and provides new insights not only into the evolutionary history of T. phayrei but the entire Trachypithecus genus as well.

INTRODUCTION

Trachypithecus is the most speciose and geographically widespread genus among Asian colobines ( Anandam et al., 2013 ; Groves, 2001 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ). Species of the genus are mainly found in Southeast Asia, from Bhutan, Assam (India), and Bangladesh in the west, through Myanmar, Thailand, Cambodia, and Laos to Vietnam and Southern China in the east, but also occur in large parts of the Sundaland region (Malay Peninsula, Sumatra, Borneo, Java, and some smaller islands). At present, 20 species of Trachypithecus are recognized ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 , 2019a ; Rowe & Myers, 2016 , Zinner et al., 2013 ), but until recently, different classifications with generally lower species numbers and varying species assemblies have been proposed ( Brandon-Jones, 1984 , 1995 , 1996 ; Brandon-Jones et al., 2004 ; Groves, 2001 ; Napier, 1985 ; Napier & Napier, 1967 , 1994 ; Oates et al., 1994 ; Roos et al., 2007 ; Weitzel & Groves, 1985 ). With increasing knowledge, particularly from genetic studies, a clearer picture of the evolutionary history of these primates has been obtained, which has also informed taxonomic revisions of the genus ( Geissmann et al., 2004 ; He et al., 2012 ; Karanth, 2008 , 2010 ; Karanth et al., 2008 ; Liedigk et al., 2009 ; Liu et al., 2013 , 2020 ; Nadler et al., 2005 ; Osterholz et al., 2008 ; Perelman et al., 2011 ; Roos & Zinner, 2021 ; Roos et al., 2007 , 2008 , 2019a ; Thant et al., 2013 ; Wang et al., 2012 , 2015 ; Wangchuk et al., 2008 ; Zhang & Ryder, 1998 ).

Based on differences and similarities in genetics, phenotype, ecology, and behavior, members of the genus are classified into four species groups ( Anandam et al., 2013 ; Osterholz et al., 2008 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 , Zinner et al., 2013 ). The T. pileatus group contains three species ( T. pileatus , T. geei , and T. shortridgei ), the T. francoisi group contains seven species ( T. francoisi , T. delacouri , T. ebenus , T. hatinhensis , T. laotum , T. leucocephalus , and T. poliocephalus ), the T. cristatus group contains six species ( T. cristatus , T. auratus , T. germaini , T. margarita , T. mauritius , and T. selangorensis ), and the T. obscurus group contains four species ( T. obscurus , T. barbei , T. crepusculus , and T. phayrei ) ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ). According to genetic data, the T. pileatus group diverged first, followed by the T. francoisi group, with the T. cristatus and T. obscurus groups most recently ( Roos & Zinner, 2021 ; Roos et al., 2019a ). Generally, mitochondrial and nuclear sequence data have provided consistent gene trees ( Roos et al., 2019a ), indicating limited gene flow, at least among the few species investigated so far. For the Indochinese gray langur ( T. crepusculus ), however, studies indicate that it is likely of hybrid origin ( Liedigk et al., 2009 ; Roos et al., 2019a ). Although various phylogenetic studies on Trachypithecus are available, they are generally limited to only a few species or individual species groups, or are based on short sequences of mitochondrial or nuclear DNA ( Geissmann et al., 2004 ; He et al., 2012 ; Karanth, 2008 , 2010 ; Karanth et al., 2008 ; Liedigk et al., 2009 ; Liu et al., 2013 , 2020 ; Nadler et al., 2005 ; Osterholz et al., 2008 ; Perelman et al., 2011 ; Roos et al., 2007 , 2008 , 2019a ; Thant et al., 2013 ; Wang et al., 2012 , 2015 ; Wangchuk et al., 2008 ; Zhang & Ryder, 1998 ). Thus, a well-supported and complete species-level phylogeny for the genus is still missing.

Phayre’s langur ( T. phayrei ) is a member of the T. obscurus group ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ). The species is one of the most widely distributed of the genus, but also one of the least studied in terms of ecology, behavior, genetics, and systematics. The species contains two subspecies, T. phayrei phayrei ( Blyth, 1847 ) and T. p. shanicus ( Wroughton, 1917 ) ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ). Until recently (e.g., Groves, 2001 ), T. phayrei included a third subspecies, T. p. crepusculus ( Elliot, 1909 ), but based on its putative hybrid status ( Liedigk et al., 2009 ; Roos et al., 2019a ), it has since been elevated to species level ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ). Nuclear sequence data suggest a closer relationship between T. crepusculus and T. barbei than T. phayrei ( Roos et al., 2019a ), hence supporting the separation of T. crepusculus from T. phayrei . The geographical distribution of the remaining subspecies of T. phayrei is poorly defined and based on only a few georeferenced museum specimens. Interestingly, according to the currently proposed distribution of T. phayrei ( Bleisch et al., 2020 ; Figure 1 ), both subspecies seem to have crossed several large rivers ( T. p. phayrei : west and east of the Ayeyarwaddy (=Irrawaddy) River; T. p. shanicus : west and east of the Chindwin, Ayeyarwaddy, and Thanlwin (=Salween) rivers). However, distribution across such large rivers is questionable as the ranges of other arboreal primates in the region are restricted by such barriers (e.g., T. leucocephalus and T. francoisi : Burton et al., 1995 ; Jiang et al., 1991 ; T. germaini and T. margarita : Nadler et al., 2005 ; Roos et al., 2008 ; T. geei and T. pileatus : Chetry et al., 2010a ; Ram et al., 2016 ; Wangchuck et al., 2008 ; Pygathrix spp.: Nadler et al., 2003 ; Hylobatidae: Chetry et al., 2010b ; Fan et al., 2017 ; Thinh et al., 2010a , 2010b ). While DNA sequence data could potentially clarify whether these distribution ranges are real, few molecular genetic studies on the intraspecific relationships of T. phayrei have been reported. Based on mitochondrial DNA, He et al. (2012) showed a clear distinction between both subspecies, while Thant et al. (2013) revealed that a population from central Myanmar (location 6 in Figure 1 ) could neither be assigned to T. p. phayrei nor to T. p. shanicus , suggesting a potential third lineage of T. phayrei .

Distribution of Trachypithecus phayrei according to IUCN Red List ( Bleisch et al., 2020 )

Numbers indicate sample locations for genetic analysis: 1: Letsegan, 2: Kin, 3: Dudaw-Taung, 4: Ramree Island, 5: near Mount Arakan, 6: Mount Popa, 7: 30 miles northwest of Toungoo, 8: Bago Yoma, 9: South Zamayi Reserve, 10: Myogyi Monastery, 11: Panlaung-Pyadalin Cave Wildlife Sanctuary, 12: Mount Yathae Pyan, 13: Yado, 14: Ho Mu Shu Pass, 15: Gaoligong Mountains National Park, 16: Cadu Ciaung, 17: Ngapyinin, 18: Lamaing, 19: Nattaung, 20: Gokteik, and 21: Se’en (for additional information see Supplementary Table S1). Underlined sites refer to type localities of examined holotypes (16: Presbytis melamera , 21: Pithecus shanicus ).

In the current study, we aimed to establish a complete species-level phylogeny and time-calibrated tree for the genus Trachypithecus . We further investigated the taxonomic diversity and geographical distribution of the species T. phayrei . We generated 41 mitochondrial genomes (mitogenomes) via polymerase chain reaction (PCR) followed by Sanger or high-throughput shotgun sequencing using fecal samples from captive and wild animals and tissue samples from historical museum specimens.

MATERIALS AND METHODS

Ethics statement.

We obtained tissue samples from museum specimens collected between 1886 and 1955 (Supplementary Table S1). Fecal material from captive and wild animals was collected during routine cage cleaning and field surveys, respectively, without disturbing, threatening, or harming the animals. Field surveys in Myanmar were permitted by the Forest Department, Myanmar. Body and craniodental measurements were taken solely from museum specimens. All research complied with protocols approved by the Animal Welfare Body of the German Primate Center (Germany) and adhered to the legal requirements of the habitat countries in which research was conducted. We conducted the study in compliance with the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) and the principles of the American Society of Primatologists for the ethical treatment of nonhuman primates.

Sample collection

Fecal samples from wild animals ( n =6) were collected during fieldwork in Myanmar and fecal samples from captive, but wild-born animals were provided by the Endangered Primate Rescue Center, Vietnam ( n =4), Singapore Zoo, Singapore ( n =2), Dhaka Zoo, Dhaka, Bangladesh ( n =1), and Mandalay Zoo, Mandalay, Myanmar ( n =1). Fresh fecal samples were stored in 80% ethanol until further processing. Dried tissue samples (ca. 5×5 mm) from museum specimens were obtained from the Natural History Museum (NHMUK), London, UK ( n =18), American Museum of Natural History (AMNH), New York, USA ( n =5), Naturalis Biodiversity Center (RMNH), Leiden, The Netherlands ( n =1), and the Zoological Reference Collection (ZRC) of the Lee Kong Chian Natural History Museum, Singapore ( n =1). Specimens from the NHMUK included holotypes of Pithecus shanicus Elliot, 1909 (NHMUK.ZD.1914.7.8.5; T. p. shanicus ) and Presbytis melamera Wroughton, 1917 (NHMUK.ZD.1888.12.1.64; synonym of T. p. phayrei ), and a paratype of Presbytis geei Khajuria, 1956 (NHMUK.ZD.1956.379; T. geei ). Furthermore, from the National Center for Biotechnology Information (NCBI) Sequence Read Archive (SRA), we downloaded Illumina raw sequencing reads of two langur specimens, for which nuclear genomic data ( Liu et al., 2020 ), but no published mitogenomes, were available. Details on specimens examined, including their origin, geographic coordinates, sample type, and sequencing data, are provided in Supplementary Table S1.

Mitogenome sequencing and assembly

DNA from fecal samples was extracted in a laboratory dedicated to handle fecal material with various precautions to avoid cross-sample contamination (e.g., separate and UV light decontaminated working areas, protective clothing, negative controls during DNA extraction and PCR amplifications). DNA extraction was performed with a First-DNA All Tissue kit (Gen-Ial, Germany) following Liedigk et al. (2015) . DNA concentration was determined with a NanoDrop ND-1000 spectrophotometer (Thermo Fisher Scientific, USA). Complete mitogenomes were amplified via 20 overlapping PCR products, each 1.0–1.2 kb in length. Details on PCR set-up and cycling conditions are outlined in Roos et al. (2011) and Liedigk et al. (2012 , 2015) . The PCR products were visualized on 1% agarose gels stained with ethidium bromide, then purified with a Qiagen PCR Purification kit (Qiagen, Germany), followed by Sanger sequencing on an ABI 3130xl sequencer (Applied Biosystems, USA) using a BigDye Terminator Cycle Sequencing kit (Applied Biosystems, USA) and both amplification primers. Sequence electropherograms were checked by eye with 4Peaks 1.8 (www.nucleobytes.com) and mitogenomes were manually assembled in SeaView 4.5.4 ( Gouy et al., 2010 ). Annotation was conducted with Geneious 11.1.3 (https://www.geneious.com/).

DNA from museum samples was extracted using a column-based method that specifically recovers small DNA fragments ( Dabney et al., 2013 ; Rohland et al., 2004 ). To reduce the risk of environmental (including human) and cross-sample contamination, DNA extraction and library preparation were performed in an ancient DNA laboratory, in which all standards for such laboratories were implemented (e.g., UV light decontamination before and after use, positive air pressure, separate sterile working areas, protective clothing, and negative controls during DNA extraction and sequencing library preparation). After extraction, the DNA concentration was measured with a Qubit 4.0 fluorometer (ThermoFisher Scientific, USA), and DNA quality and degradation status were checked on a Bioanalyzer 2100 (Agilent Technologies, USA). Genomic DNA (50 ng) was then subjected to shotgun library preparation with a NEBNext Ultra II DNA Library Prep kit (New England Biolabs, USA) following the standard protocols of the supplier. However, due to the degraded status of the DNA, DNA fragmentation prior to library preparation was omitted. After end repair, adapter ligation, and ligation cleanup (without size selection), libraries were indexed with multiplex oligos and then cleaned with the purification beads supplied in the kit. Libraries were also prepared from the pooled negative controls. Library concentration and size distribution were measured with a Qubit fluorometer and bioanalyzer, respectively, and molarity was quantified via quantitative PCR using the NEBNext Library Quant kit (New England Biolabs, USA). Sequencing was conducted on an Illumina HiSeq 4000 (50 bp or 100 bp single-end reads) at the NGS Integrative Genomics (NIG) unit of the University Medical Center Göttingen, Germany, or on an Illumina NextSeq (75 bp paired-end reads) at the University of Potsdam, Germany. Raw sequencing reads were demultiplexed with Illumina software. Subsequent bioinformatic analyses were performed with the Geneious package. First, we trimmed and quality-filtered the reads with BBDuk 37.64 in the BBTools package (https://jgi.doe.gov/data-and-tools/bbtools/) and removed duplicate reads with Dedupe 37.64 (BBTools package); both filtering steps were conducted with standard settings. For assembly, reads were mapped onto the mitogenome of a closely related Trachypithecus spp. (Supplementary Table S1) using the Geneious assembler with standard settings. All newly produced mitogenomes were manually checked and then annotated with Geneious.

To generate mitogenomes from published Illumina sequencing reads deposited in the NCBI SRA, we downloaded the data and randomly selected 20 million reads. Read processing, filtering, mitogenome assembly, and annotation were performed as described for museum samples.

Phylogenetic analyses

For phylogenetic reconstructions, our dataset was expanded with additional mitogenome sequences from GenBank (Supplementary Table S1). The final dataset was comprised of 72 sequences, including 53 Trachypithecus sequences and sequences from various other colobines ( Semnopithecus , Presbytis, Rhinopithecus , Pygathrix , Nasalis , Simias , Colobus , Piliocolobus , and Procolobus ) and non-colobines ( Macaca , Papio , Theropithecus , Chlorocebus , Hylobates , Pongo , Gorilla , Pan , and Homo ). Sequences were aligned with Muscle 3.8.31 ( Edgar, 2010 ) in AliView 1.18 ( Larsson, 2014 ) and manually checked. The generated alignment had a length of 16 969 bp, including 7 197 parsimony-informative and 1 499 parsimony-uninformative variable sites.

Phylogenetic trees were reconstructed with the maximum-likelihood (ML) algorithm in IQ-TREE 1.5.2 ( Nguyen et al., 2015 ) and Bayesian inference (BI) in MrBayes 3.2.6 ( Ronquist et al., 2012 ). For all reconstructions, the optimal substitution model (GTR+I+G), as determined with ModelFinder ( Chernomor et al., 2016 ; Kalyaanamoorthy et al., 2017 ) in IQ-TREE under Bayesian Information Criterion (BIC), was applied. The BI tree was reconstructed via two independent Markov Chain Monte Carlo (MCMC) runs, each for one million generations with tree and parameter sampling every 100 generations and a burn-in of 25%. To check for convergence of all parameters and adequacy of burn-in, we investigated the uncorrected potential scale reduction factor (PSRF) ( Gelman & Rubin, 1992 ), as calculated by MrBayes. The BI posterior probabilities (PP) and consensus phylogram with mean branch lengths from the posterior density of the trees were also calculated in MrBayes. Node support for the ML tree was obtained from 10 000 ultrafast bootstrap (BS) replications ( Minh et al., 2013 ). All phylogenetic trees were visualized and edited in FigTree 1.4.2 (http://tree.bio.ed.ac.uk/software/figtree/).

Divergence times were calculated with the BEAST 2.4.8 package ( Bouckaert et al., 2014 ). We applied a relaxed log-normal clock model of lineage variation ( Drummond et al., 2006 ) and used a Yule tree prior and the selected best-fit model of sequence evolution (GTR+I+G). To calibrate the molecular clock, we constrained 10 nodes with hard minimum and soft maximum bounds using gamma-distributed priors. These 10 nodes refer to the divergence of (1) Hominoidea vs. Cercopithecoidea, (2) Hominidae vs. Hylobatidae, (3) Pongo vs. Gorilla + Pan + Homo , (4) Gorilla vs. Pan + Homo , (5) Pan vs. Homo , (6) Cercopithecinae vs. Colobinae, (7) African vs. Asian Colobinae, (8) Chlorocebus vs. Papionini, (9) Macaca vs. African Papionini, and (10) Papio vs. Theropithecus . A detailed discussion on the selected node constraints is available in Roos et al. (2019b) and details on prior settings are listed in Supplementary Table S2. We ran BEAST analyses for 100 million generations with tree and parameter sampling every 5 000 generations. The adequacy of 10% burn-in and convergence of all parameters were assessed with Tracer 1.6 (http://tree.bio.ed.ac.uk/software/tracer/). We combined the sampling distributions of two independent runs with LogCombiner 2.4.8 and summarized trees with a burn-in of 10% in TreeAnnotator 2.4.8 (both programs are part of the BEAST package).

Morphometric analyses

External measurements (head-body length, tail length, hind foot length, and ear length) were taken from original museum specimen labels (15 adult males, 11 adult females, 14 young, subadults or adults of unknown sex), reflecting measurements taken on fresh specimens in the field (Supplementary Table S3). Eighteen cranial and dental measurements were taken on the skulls of 22 museum specimens (12 adult males, five adult females, five subadults) with hand-held calipers to the nearest 0.1 mm (Supplementary Table S3). A Kruskal-Wallis analysis of variance (ANOVA) by Ranks was conducted to determine significant differences between taxa (alpha=0.05), followed by post-hoc pair-wise population comparisons of traits (Mann-Whitney U test, with Bonferroni correction for multiple testing in Statistica TM 13.5.0.17). Principal component analyses (PCAs) were computed using a combination of dental (molar lengths and widths), and cranial (skull length, condylobasal length, zygomatic width, orbit width, C-M3 length, upper canine width, upper palate breadth, anterior palatal foramina length and width, palatilar length, and braincase breadth and height) measurements in RStudio 1.2.5033 ( RStudio Team, 2020 ). All measurement values were standardized by subtracting the mean and dividing by the standard deviation before multivariate analysis. Principal components were extracted from a covariance matrix.

Mitogenomic data

Of the 41 newly sequenced Trachypithecus mitogenomes, 14 were produced from fecal material via conventional PCR followed by Sanger sequencing, 25 were generated from museum samples via high-throughput shotgun sequencing, and two were assembled from published high-throughput shotgun sequencing reads ( Liu et al., 2020 ). For museum samples, we obtained 7.0–53.9 million raw sequence reads per sample; after quality filtering and duplicate removal, we retained 2 299–30 433 reads mapped to the Trachypithecus spp. reference mitogenomes, resulting in 100% coverage and an average sequencing depth of 7–96 (for detailed information see Supplementary Table S1). For the two mitogenomes generated from published sequences, we obtained 100% coverage and an average sequencing depth of 97 and 342, respectively. All 41 mitogenomes contained 22 tRNA genes, 2 rRNA genes, 13 protein-coding genes, and a control region in the order typically found in mammals. All protein-coding genes were correctly transcribed without any premature stop codons and tRNAs exhibited typical secondary structure, indicating that our mitogenomes are likely free from nuclear mitochondrial DNA sequences (numts).

The ML and BI phylogenetic trees revealed identical branching patterns with strong node support (BS 100%, PP 1.0; Figure 2 , Supplementary Figure S1). Only the relationships among the three clades found in T. phayrei were not well resolved (BS 67%, PP 0.97). Likewise, the phylogenetic position of Semnopithecus among Asian colobines and basal position of Rhinopithecus among odd-nosed monkeys were supported by BS values of 88% and 98%, respectively, with PP for both nodes of 1.0 (Supplementary Figure S1).

Mitogenomic tree showing phylogenetic relationships and divergence times among mitochondrial lineages of Trachypithecus (A) and detailed view on T. phayrei (B)

Node bars indicate 95% highest posterior densities (HPDs). Node supports of <100% ML BS and <1.0 BI PP are given at respective nodes. In A, species group assignment is given on the right; * : T. crepusculus , a member of the T. obscurus group according to phenotype and nuclear sequence data. In B, sample labels contain individual ID and sample location number (as in Figures 1 , ​ ,5, 5 , Supplementary Table S1). Clade assignment is given on the right. Complete ultrametric tree including all non- Trachypithecus taxa and details on estimated divergence times are provided in Supplementary Figure S1 and Table S4, respectively.

In Trachypithecus , the T. pileatus group branched first, ca. 4.49 million years ago (Ma) (95% highest posterior densities (HPDs): 3.91–5.10) ( Figure 2 , Supplementary Figure S1 and Table S4). The remaining taxa diverged 3.87 (3.35–4.38) Ma into a clade containing the T. obscurus and T. cristatus groups, and a clade subsuming the T. francoisi group and T. crepusculus . The T. obscurus and T. cristatus groups split 3.24 (2.78–3.70) Ma, and the T. francoisi group separated from T. crepusculus 3.06 (2.60–3.51) Ma. Speciation events within the four species groups occurred over a prolonged period, from 1.62 (1.31–1.94) Ma to 0.29 (0.22–0.36) Ma. In the T. pileatus group, T. shortridgei diverged from T. pileatus and T. geei 1.62 (1.31–1.94) Ma, and the latter two separated 0.57 (0.43–0.71) Ma. In the T. francoisi group, the southern taxa ( T. laotum , T. hatinhensis , and T. ebenus ) split from the central ( T. delacouri ) and northern taxa ( T. francoisi , T. leucocephalus , and T. poliocephalus ) 1.36 (1.15–1.56) Ma, and the central and northern taxa diverged 1.03 (0.86–1.20) Ma. Among the southern taxa, T. laotum separated from T. hatinhensis and T. ebenus 0.59 (0.47–0.72) Ma, while the latter two diverged 0.33 (0.24–0.42) Ma. Among the northern taxa, T. poliocephalus split from T. francoisi and T. leucocephalus 0.52 (0.42–0.63) Ma, followed by separation of T. francoisi and T. leucocephalus 0.29 (0.22–0.36) Ma. In the T. cristatus group, the mainland taxa ( T. germaini and T. margarita ) diverged from the central Sundaland ( T. cristatus and T. selangorensis ) and Javan taxa ( T. auratus and T. mauritius ) 1.25 (1.07–1.45) Ma and the latter two clades split 0.95 (0.79–1.11) Ma. Speciation events in these three clades occurred 0.87 (0.70–1.06) Ma (mainland clade), 0.29 (0.22–0.36) Ma (central Sundaland clade), and 0.72 (0.59–0.87) Ma (Javan clade). In the T. obscurus group, T. obscurus diverged first 1.60 (1.37–1.84) Ma and T. barbei separated from T. phayrei 1.40 (1.19–1.61) Ma.

For T. phayrei , we obtained three major clades, which separated within a short period, 0.93–0.99 (0.79–1.13) Ma ( Figure 2B ). The samples from Bangladesh and Myanmar, west of the Ayeyarwaddy and Chindwin rivers (locations 1–5; Figures 1 , ​ ,5), 5 ), grouped in the West clade. Those from the central dry zone of Myanmar and neighboring Kayah-Karen Mountains, east of the Ayeyarwaddy River and west of the Thanlwin River (locations 6–13), formed the Central clade, and those from the Shan Plateau and neighboring China (locations 14–21) clustered in the East clade. In the Central clade, we found two subclades, with one containing samples from the central dry zone (locations 6–9; Central clade A) and the other containing samples from the western foothills of the Kayah-Karen Mountains (locations 10–12; Central clade B). One historical sample from Yado (location 13) clustered with Central clade A and not, as expected, with the geographically closer Central clade B. At location 10, the Myogyi Monastery, we found haplotypes of the Central B and East clades. The holotypes of Pithecus shanicus (location 21) and Presbytis melamera (location 16) both nested within the East clade.

Geographical distribution of mitochondrial clades found in Trachypithecus phayrei

Sample locations are numbered as in Figures 1 , ​ ,2 2 (see also Supplementary Table S1) and colored according to their mitochondrial clade assignment. Limits of the Central clade to the northeast and East clade to the southwest, depicted in light green, are not yet firmly resolved. Samples from locations 6–9 form Central clade A, while those from locations 10–12 cluster in Central clade B. Note, at location 10, haplotypes of the Central and East clades were found. Museum specimen from location 13 cluster unexpectedly with Central clade A (see Results).

Morphometric data

Based on the mitogenomic division of T. phayrei into three rather than two clades, we investigated whether morphological features supported such a division. We found that males, but not females, from the West clade had significantly shorter tails than those from the other two clades ( post-hoc test: P <0.025; Figure 3 , Supplementary Tables S5–S6). For molar measurements, ungrouped morphometric comparisons using PCA for all available specimens with full molar complements (both adults and subadults, comparable in this case because these teeth do not change in size as the cranium matures) demonstrated that all three clades occupied distinct molar-dimension morphospace ( Figure 4 , Supplementary Figure S2 and Table S7), even with sex and age variation within each group (Supplementary Figure S3). PCAs based on combined craniodental measurements also separated each clade (Supplementary Figure S3 and Table S8), with molar dimensions important in facilitating morphometric separation, even when both sexes and subadult skulls were included. Cranial measurements alone could not separate the three clades, thus demonstrating the importance of dental size and proportion in clade distinction, despite their overall cranial similarity. However, direct comparisons of skulls revealed useful, if subtle, skull characters in distinguishing the three groups (see Systematic biology, below).

Head-body length (A), tail length (B), and tail/head-body length ratio (C) of adult male and female Trachypithecus phayrei representing West, Central, and East clades (median, quartiles, min-max)

Numbers in brackets: sample sizes; post-hoc pair-wise population comparisons of traits; Mann-Whitney U -test, with Bonferroni correction for multiple testing: * : P <0.025, ( * ): P <0.05; see Supplementary Table S6.

Morphometric comparisons (principal component analysis performed on 12 molar measurements) among Trachypithecus phayrei individuals representing West (red), Central (blue), and East (yellow) clades

Shown is projection of specimen scores of first and second principal components, with variance explained by each component (graphical depictions of third principal component appear in Supplementary Figure S2 and underlying statistics are provided in Supplementary Table S7.)

In the current study, we inferred a robust mitochondrial species-level phylogeny of the genus Trachypithecus . In contrast to earlier studies, which examined only fragments of the mitogenome and/or a few species ( Geissmann et al., 2004 ; He et al., 2012 ; Karanth, 2008 , 2010 ; Karanth et al., 2008 ; Liedigk et al., 2009 ; Liu et al., 2013 ; Nadler et al., 2005 ; Osterholz et al., 2008 ; Roos et al., 2007 , 2008 ; Thant et al., 2013 ; Wang et al., 2012 , 2015 ; Wangchuk et al., 2008 ; Zhang & Ryder, 1998 ), we included full-length mitogenomes of all 20 currently recognized species ( Anandam et al., 2013 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ), including two name-bearing types. Based on this dataset, we resolved the branching patterns among and within species groups, except for the radiation within T. phayrei , with strong nodal support.

However, as mitochondrial DNA is only maternally inherited, the evolutionary history of the genus remains incomplete ( Avise, 2000 ). Unfortunately, nuclear sequence data for species of Trachypithecus are still scarce, but the few available generally result in a tree topology identical to that obtained from mitogenomes ( Liu et al., 2020 ; Perelman et al., 2011 ; Roos et al., 2019a ). The only exception known so far is the phylogenetic position of T. crepusculus , which constitutes a distant relative of the T. francoisi group in mitogenome phylogenies ( Figure 2 ), while nuclear sequence data support its membership in the T. obscurus group ( Liedigk et al., 2009 ) and specifically as sister taxon to T. barbei ( Roos et al., 2019a ). This tree discordance is most likely the result of ancient hybridization ( Liedigk et al., 2009 ; Roos et al., 2019a ).

According to estimated divergence times, the four species groups (and the mitochondrial lineage of T. crepusculus ) separated in the Pliocene, while speciation events within all species groups occurred on similar time scales in the Early Pleistocene, suggesting that Trachypithecus speciation in Southeast Asia has been largely influenced by extrinsic factors such as changes in forest cover and/or sea level ( Hallet & Molnar, 2001 ; Heaney, 1986 ; Miller et al., 2005 ).

Distribution and taxonomy of “ T. phayrei ”

For T. phayrei , we obtained three geographically segregated mitochondrial clades, which does not reflect the current classification of T. phayrei into two subspecies across its distribution ( Anandam et al., 2013 ; Bleisch et al., 2020 ; Roos, 2021 ; Roos et al., 2014 ; Rowe & Myers, 2016 ; Zinner et al., 2013 ) (compare Figures 1 , ​ ,5). 5 ). According to our data, the geographical distribution of the three clades appears to be delineated by large rivers and/or specific habitat types. The West clade is distributed in Bangladesh and Myanmar, west of the Chindwin and Ayeyarwaddy rivers ( Figure 5 ). Although we had no genetic samples from India, specimens from this country would most likely fall into the West clade as well (on both geographical and morphological grounds). The region is dominated by tropical rainforests as well as tropical dry deciduous forests ( Murray et al., 2020 ). The Central clade is restricted to the central dry zone of Myanmar and the western foothills of the Kayah-Karen Mountains between the Ayeyarwaddy and Thanlwin rivers. The region consists of open woodland in the northern part and tropical dry deciduous forest in the southern part ( Murray et al., 2020 ). A specimen from Yado (location 13) in the Kayah-Karen Mountains clustered with the Central clade, but unexpectedly with Central clade A and not with the geographically closer Central clade B. We can exclude contamination in the laboratory as this specimen was not processed with any sample from the Central clade. There are also no indications of incorrect field records, so our findings remain obscure. Hence, the northeastern boundary of the Central clade remains poorly defined, though it might extend into the Kayah-Karen Mountains. The East clade is found on the Shan Plateau and in neighboring China, between the Ayeyarwaddy and Thanlwin rivers, with the southwestern limit probably extending into the Kayah-Karen Mountains. The region is dominated by typical Shan State tropical mixed forest ( Murray et al., 2020 ). According to its currently assumed distribution ( Bleisch et al., 2020 ; Figure 1 ), T. phayrei is also found east of the Thanlwin River, but we found no evidence for this, as specimens from east of the Thanlwin River from China and Thailand did not cluster with T. phayrei but instead fell into the T. crepusculus mitochondrial clade (C.R., unpublished data). Likewise, there is no known evidence for the presence of T. phayrei between the Chindwin and Ayeyarwaddy rivers, an area that may instead be occupied by T. shortridgei . At location 10, Myogyi Monastery, we found haplotypes of the Central clade B and East clade. The semi-habituated langurs at the monastery are fed by monks and visitors ( Quyet et al., 2019 ), and exhibit phenotypical features of individuals of both the Central and East clades. We suspect that pet monkeys from further to the northeast were released at the site and interbred with the resident langurs, or less likely, that both populations overlap here naturally.

Taxonomically, the West clade corresponds to the nominate form T. p. phayrei ( Presbytis phayrei Blyth, 1847 ) with the type locality “Arakan” (=Rakhine State, Myanmar), while the East clade corresponds to populations usually considered to represent subspecies T. p. shanicus ( Pithecus shanicus Wroughton, 1917 ) with the type locality “Hsipaw, Northern Shan States”. For T. p. phayrei , Presbytis barbei Blyth, 1863 from the “interior of Tippera hills” (=Tripura State, India), Semnopithecus holotephreus Anderson, 1878 from an unknown locality, and Presbytis melamera Elliot, 1909 from “Cadu Ciaung, Bhamo, North Burma” are generally regarded or listed as synonyms (e.g., Groves, 2001 ; Napier, 1985 ; Pocock, 1939 ). However, the type locality of melamera (location 16) is east of the Ayeyarwaddy River and geographically close to that of shanicus (location 21). We sequenced the mitogenomes of these two holotypes and found that both clustered in the East clade, suggesting that melamera is not a synonym of T. p. phayrei , but instead is a senior synonym of T. p. shanicus . Wroughton (1918 , 1921) also concluded that his shanicus (named in 1917) is morphologically identical with melamera . Pocock (1928) recognized similarities in the coloration of both holotypes and their close geographical distance but kept them separate because of the absence of a parting on the forehead in the melamera holotype, although this is probably because the individual was a subadult. The taxonomic name for the East clade is thus T. p. melamera ( Elliot, 1909 ), with shanicus Wroughton, 1917 as a junior synonym. For the Central clade, however, no taxonomic name is yet available.

The three mitochondrial clades of T. phayrei diverged almost 1 Ma, a similar time scale as other speciation events within Trachypithecus , and are geographically confined by large rivers and/or different habitat types (ecological adaptation). Furthermore, the members of the three clades are diagnosably different in external morphology (see morphometric data). Following the phylogenetic species concept ( Cracraft, 1983 ), we elevate the two recognized subspecies to species status, i.e., T. phayrei and T. melamera , and describe and name the taxon constituting our “Central clade” as a new species.

Systematic biology

Order Primates Linnaeus, 1758

Family Cercopithecidae Gray, 1821

Subfamily Colobinae Jerdon, 1867

Genus Trachypithecus Reichenbach, 1862

Trachypithecus phayrei ( Blyth, 1847 )

English name: Phayre’s langur.

Synonyms: Presbytis barbei Blyth, 1863 ; Semnopithecus holotephreus Anderson, 1878 .

Distribution: East Bangladesh, Northeast India (Assam, Mizoram, and Tripura), and West Myanmar, west of the Chindwin and Ayeyarwaddy rivers ( Figure 5 ).

Conservation status: Currently listed as Endangered ( Bleisch et al., 2008a ), but reassessment required.

Trachypithecus melamera ( Elliot, 1909 )

English name: Shan State langur.

Synonyms: Pithecus shanicus Wroughton, 1917 .

Distribution: East Myanmar (Shan States) and Southwest China (West Yunnan), between the Ayeyarwaddy and Thanlwin rivers, with the southwestern limit probably extending into the Kayah-Karen Mountains ( Figure 5 ).

Conservation status: Currently listed as Endangered ( Bleisch et al., 2008b ), but reassessment required.

Trachypithecus popa sp. nov.

Popa langur

Holotype: NHMUK ZD.1914.7.19.3 (adult male, stuffed skin and skull, left zygomatic arch slightly damaged; Figures S4–S6), collected by Guy C. Shortridge on 11 September 1913. Head-body length (HBL): 600 mm, tail length (TL): 800 mm, hindfoot length (HFL): 174 mm, ear length (EL): 33 mm, body mass (BM): 7.9 kg. Mitogenome GenBank accession No.: {"type":"entrez-nucleotide","attrs":{"text":"MT806047","term_id":"1945584252"}} MT806047 .

Type locality: Mount Popa, Myingyan District, Myanmar (N20°55’, E95°15’, 4 961 feet=1 512 m a.s.l.) (location 6 in Figures 1 , ​ ,5 5 ).

Paratypes: NHMUK ZD.1914.7.19.4 (adult male, stuffed skin and skull) collected at the type locality by Guy C. Shortridge on 27 September 1913. HBL: 580 mm, TL: 795 mm, HFL: 161 mm, EL: 32 mm, BM: 8.2 kg. NHMUK ZD.1914.7.19.5 (adult female, stuffed skin) collected at the type locality by Guy C. Shortridge on 3 September 1913. HBL: 540 mm, TL: 780 mm, HFL: 152 mm, EL: 30 mm, BM: 7.0 kg. NHMUK ZD.1917.4.24.1 (adult male, stuffed skin and skull) collected at South Zamayi Reserve, 60 miles north of Pegu by J.M.D. Mackenzie on 10 March 1916. HBL: 498 mm, TL: 795 mm, HFL: 168 mm, EL: 33.5 mm, BM: 7.7 kg. NHMUK ZD.1937.9.10.4 (subadult male, stuffed skin and skull) collected 30 miles northwest of Toungoo by J.M.D. Mackenzie on 8 January 1928. HBL: 508 mm, TL: 785 mm, HFL: 165 mm, EL: 31 mm. NHMUK ZD.1937.9.10.5 (subadult male, stuffed skin and skull) collected 30 miles northwest of Toungoo by J.M.D. Mackenzie on 8 January 1928. HBL: 509 mm, TL: 795 mm, HFL: 165 mm, EL: 31 mm. AMNH M-54770 (juvenile male, skull) collected at Camp Pinmezali, Pegu Yoma by John C. Faunthorpe on 27 April 1924. RMNH MAM.59807 (adult male, stuffed skin with skull in situ ) collected at Yado, Mount Cariani, Tounghoo (=Taungoo) District, Myanmar (800–1 000 m) by Leonardo Fea in December 1887 (field number: 40). HBL: 555 mm, TL: 750 mm.

Etymology: The English name for Trachypithecus popa is Popa langur. Mount Popa is a major landmark of the Myingyan District in Myanmar, and the place where the designated holotype was originally collected. The specific name “popa” is used as a noun in apposition.

Description: The species is dark brown or gray-brown on the dorsum, with a sharply contrasting gray or whitish venter. Hands and feet are black. From above the elbow, the arms on the dorsal side gradually darken to black hands. The pale underside extends onto the chin and down to the inner side of the arms and thighs. The tail is paler than the back, notably at the base and underside. The face is black with a wide fleshy-white muzzle and broad white rings fully encircling the eyes. The hairs on the head are raised to a crest or are at least long and irregularly structured, but with no parting or whorl behind the brows present. This crest of hair and the forward-facing whiskers give the head a rhomb-like shape ( Figure 6 , Supplementary Figures S4–S6). Body measurements (median and range) are: males ( n =5) HBL: 562 (498–600) mm, TL: 795 (775–858) mm, HFL: 168 (144–178) mm, EL: 32 (30.0–33.5) mm, BM: 7.9 (7.7–8.2) kg; females ( n =3) HBL: 585 (540–589) mm, TL: 780 (720–784) mm, HFL: 156 (152–160) mm, EL: 30 (20–32) mm, BM ( n =1): 7.0 kg (Supplementary Table S3).

Photos of Trachypithecus phayrei (A, B), Trachypithecus popa sp. nov. (C, D) and Trachypithecus melamera (formerly T. p. shanicus ) (E, F)

A: Adult female T. phayrei at Yangon Zoo, Myanmar (photo by Tilo Nadler); B: Adult male T. phayrei from Lawachara National Park, Bangladesh (photo by Tanvir Ahmed); C, D: Subadult male T. popa from Mount Popa, Myanmar (photo by  Lay Win); E: Adult female T. melamera at Mandalay Zoo, Myanmar (photo by Tilo Nadler), F: Adult female T. melamera with offspring from Gaoligong Mountains National Park, China (photo by Chi Ma).

Diagnosis: Overall, Trachypithecus popa sp. nov. is externally more similar to T. phayrei than to T. melamera . Body coloration in all three species is variable, but generally more fawn in T. melamera and more brownish to gray in Trachypithecus popa sp. nov. and T. phayrei . In Trachypithecus popa sp. nov. and T. phayrei , but not in T. melamera , the pale venter sharply contrasts with the back. The hands and feet are black in all three species. In Trachypithecus popa sp. nov. , the arms (dorsal side) gradually darken to the hands from above the elbow, while in T. phayrei , they gradually darken from below the elbow. In T. melamera , the lower arms are not darker than the upper arms. In Trachypithecus popa sp. nov. and T. phayrei , the hairs on the head are raised to a crest or are at least long and irregularly structured, while T. melamera has a whorl or a parting behind the brows. Whiskers are laterally directed in T. phayrei , but forward directed in Trachypithecus popa sp. nov. and T. melamera . The direction of the whiskers in combination with the hairs on the head gives the head of T. phayrei a triangular shape, versus a rhomb-like shape for Trachypithecus popa sp. nov. and a round shape for T. melamera . All three species have a fleshy-white muzzle, which is wider in Trachypithecus popa sp. nov. and T. melamera . In T. melamera , the white around the eyes is restricted to the inner side, while in T. phayrei , the white normally encircles the eyes fully, although it is sometimes restricted to the inner side. In Trachypithecus popa sp. nov. , the eyes are always fully encircled with broad white eye-rings. Males of T. phayrei have significantly shorter tails than males of the other two species ( Figure 3 , Supplementary Tables S5–S6).

Cranially, Trachypithecus popa sp. nov. has a slightly longer skull, especially relative to its width, than in T. phayrei and T. melamera ; this is achieved by a slight anterior elongation of the facial region of the skull relative to these taxa, rendering Trachypithecus popa sp. nov. slightly more prognathic in lateral and dorsal views and creating a more rectangular shape of the bony palate in ventral view (vs. a more square palate in T. phayrei and T. melamera ). The teeth are, on average, larger in Trachypithecus popa sp. nov. than in T. phayrei and T. melamera , and molar measurements are the clearest means for separating the skulls of this new taxon from its closest relatives (Supplementary Tables S3, S7; Figure 4 , Supplementary Figures S2–S3); in particular, the third molar (M3/m3) appears larger overall in Trachypithecus popa sp. nov. when skulls are directly compared. PCAs using molar measurements and combined craniodental measurements separated T. phayrei , T. melamera , and Trachypithecus popa sp. nov. , but cranial measurements alone did not separate them ( Figure 4 , Supplementary Figures S2, S3).

Distribution: Between the Ayeyarwaddy and Thanlwin rivers in the central dry zone of Myanmar and into the western foothills of the Kayah-Karen Mountains ( Figure 5 ). The northeastern limit is undefined (see Discussion), but the species may occur throughout the Kayah-Karen Mountains. This species is endemic to Myanmar.

Conservation status: As evident from historical records (museum specimens and travel notes), the species was once widespread in the central dry zone of Myanmar. Only two of these populations are known to have survived (location 6: Mount Popa, location 8: Bago Yoma), while all others are considered possibly extirpated. However, during recent fieldwork, three new populations (locations 10–12) were discovered. At location 10, Myogyi Monastery, the langur population is estimated at 75–100 individuals ( Quyet et al., 2019 ), but these langurs are probably hybrids between Trachypithecus popa sp. nov. and T. melamera . The populations at location 11, Panlaung-Pyadalin Cave Wildlife Sanctuary, and location 12, Mount Yathae Pyan, consist of 46–96 individuals ( Quyet et al., 2019 ) and 20–30 individuals (A.K.L. and A.L. pers. observation), respectively. The population at Bago Yoma (location 8) contains about 22 individuals (A.K.L. pers. observation) and at Mount Popa (location 6), field surveys conducted in 2019 revealed a population size of 111 individuals (Thaung Win pers. communication). Mount Popa was declared a national park (Popa Mountain Park) in 1989 and has an area of 128.54 km 2 , including 26.97 km 2 classified as suitable to highly suitable for langurs ( Thant, 2013 ; Thant et al., 2013 ).

Throughout its range, Trachypithecus popa sp. nov. is threatened by hunting, habitat loss, degradation, and fragmentation caused by agricultural encroachment, illegal/unsustainable timber extraction, and disturbances caused by collection of non-timber products and free cattle grazing ( Quyet et al., 2019 ; Thant et al., 2013 ). Considering a total population size of 199–259 individuals (excluding the possible hybrid population at Myogyi Monastery) in the four disjunct populations and the dramatic habitat loss over the last century, we propose to classify Trachypithecus popa sp. nov. as Critically Endangered (CR) as it meets the IUCN Red List criteria B1a and B1b (i-v) ( IUCN, 2001 ). Furthermore, Trachypithecus popa sp. nov. needs to be added to the national and international lists of threatened species (IUCN, CITES). Improved protected area management, in particular improved law enforcement, in Popa Mountain Park and Panlaung-Pyadalin Cave Wildlife Sanctuary is essential to stabilize the two largest known populations. Mount Yathae Pyan is an isolated karst hill. This population could be protected through the designation of a community-protected area (CPA). The population status of the species in Bago Yoma is poorly understood and additional surveys are urgently required. The forests in Bago Yoma are severely degraded and fragmented, but could still provide the largest, contiguous habitat if deforestation and forest degradation are reversed through improved forest protection and restoration.

Comments: Except for species of the T. pileatus group, the natal coat of Trachypithecus spp. is generally yellowish, orange, or light brown ( Anandam et al., 2013 ; Rowe & Myers, 2016 ). Trachypithecus popa sp. nov. may be an exception as photos show an infant with creamy white fur coloration (Supplementary Figure S7).

CONCLUSIONS

We present a robust mitogenomic species-level phylogeny of the genus Trachypithecus , thus providing new insights into the evolutionary history of the genus and forming a basis for future work. Based on our investigations of T. phayrei , we illuminated the intraspecific taxonomy of the species, resulting in the elevation of two known subspecies to species level, renaming of one subspecies, description of a new species, and largely refined distributional ranges for all three species. Including the proposed taxonomic changes, the genus Trachypithecus now contains 22 species, with Myanmar home to a total of 20 non-human primate species ( Trachypithecus popa sp. nov. , T. phayrei , T. melamera , T. barbei , T. obscurus , T. crepusculus , T. shortridgei , T. pileatus , Presbytis femoralis , Rhinopithecus strykeri , Macaca mulatta , M. fascicularis , M. arctoides , M. assamensis , M. leonina , Hoolock hoolock , H. leuconedys , H. tianxing , Hylobates lar , and Nycticebus bengalensis ; Fan et al., 2017 ; Mittermeier et al., 2013 ; Roos et al., 2014 ; Rowe & Myers, 2016 ), of which Trachypithecus popa sp. nov. and probably H. leuconedys are endemic to the country. Trachypithecus germaini , commonly listed for Myanmar (e.g., Anandam et al., 2013 ; Groves, 2001 ; Roos et al., 2014 ; Rowe & Myers, 2016 ), is actually not present in the country. Its putative occurrence in Myanmar is based on the incorrect assignment of Pithecus pyrrhus atrior as a synonym of T. germaini instead of T. barbei ( Geissmann et al., 2004 ; C.R., unpublished data).

Overall, our study reaffirms that museum collections are a valuable source for genetic and taxonomic investigations of primates, particularly as modern high-throughput sequencing technologies allow the analysis of highly damaged DNA, which is typically extracted from such material. Future studies on Trachypithecus should also include nuclear sequence data and multiple individuals per species and should focus on the three polytypic species of the genus, i.e., T. pileatus , T. cristatus , and T. obscurus .

DATA AVAILABILITY

Scientific field survey permission information.

Permission for fieldwork in Myanmar was granted by the Forest Department, Myanmar.

 SUPPLEMENTARY DATA

Supplementary data to this article can be found online.

COMPETING INTERESTS

The authors declare that they have no competing interests.

AUTHORS’ CONTRIBUTIONS

C.R. and F.M. conceived and designed the study. N.L., A.K.L., A.L., C.M., D.M., and L.K.Q. collected samples in the field. R.P.M., N.D., P.K., and M.A.H.C. provided valuable samples from their museum collections. N.M.L.T., N.L., A.K.L., A.L., K.M.Y., P.S., Z.M.H., M.N.N.M., T.A., D.C., L.K.Q., T.N., P.F., and F.M. provided field data. C.R., M.U., L.Y., M.L., Z.L., and M.H. generated the data. C.R., K.M.H., R.P.M., E.G.V., and D.Z. analyzed the data. C.R., K.M.H., and D.Z. wrote the paper. All authors discussed the data and read and approved the final version of the manuscript.

ACKNOWLEDGEMENTS

We are grateful to the Myanmar Forest Department for permitting fieldwork. L.K.Q. and A.K.L. wish to thank the Management Board of the Panlaung-Pyadalin Cave Wildlife Sanctuary, and Kyaw Naing Oo and Win Hlaing for support during fieldwork. We thank the late Alan Mootnick and the staff of Singapore Zoo, Dhaka Zoo, and Mandalay Zoo for proving fecal samples as well as Christiane Schwarz and Michaela Preick for their excellent work in the laboratory. Many thanks also to Thaung Win and Chi Ma for langur photos, Alain Dubois for taxonomic advice, and two anonymous reviewers for their helpful comments on an earlier version of the manuscript.

Funding Statement

This study was supported by the Margot Marsh Biodiversity Foundation, Primate Action Fund, Helmsley Charitable Trust, and Critical Ecosystem Partnership Fund

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The largest island in the Bay of Naples and the site of the first and most northerly Greek base in the west. See colonization, greek. The acropolis was in continuous use between the mid‐8th and the 1st cents. bc. An emporion rather than an apoikia, Pithecusae was settled by Chalcidians and Eretrians (see chalcis; eretria). Throughout the second half of the 8th cent. it served as a large and vital ‘pre‐colonial’ staging‐post—with a stable population numbered in thousands—at the western end of the route from the Aegean and the Levant. The suburban industrial complex has yielded abundant evidence for early metallurgical production; competent local versions of Euboean Late Geometric pottery (see pottery, greek) were produced en masse, and expatriate Protocorinthian potters were also active.

From:   Pithēcūsae   in  Oxford Dictionary of the Classical World »

Subjects: Classical studies

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Ardipithecus kadabba

• Author(s) Fran Dorey
• Updated 17/12/19
• Read time 2 minutes
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Background of discovery

5.8 to 5.2 million years ago

Important fossil discoveries

Eleven specimens, from five localities in Ethiopia, were discovered between 1997 and 2000. They represent at least five individuals and include teeth, jaws, hand, toe, arm and collar bones. The type specimen is a right lower jaw fragment, ALA-VP-2/10. Most of the fossils date to 5.6-5.8 million years old, however one of the toe bones is dated at 5.2 million years old. There is some concern over the classification of the toe bone to this species, as it was found 15 kilometres away from the other fossils and is younger in age. They were classified as a subspecies Ardipthecus ramidus kadabba.

In 2002, six teeth were found at Asa Koma in the Middle Awash. They date to between 5.6 and 5.8 million years old. Distinct features of these teeth led the finders to place all the fossils into a new species Ardipithecus kadabba rather than a subspecies of Ardipithecus ramidus.

What the name means

The name is derived from the local Afar language. ‘Ardi’ means ‘ground’ or ‘floor’, and is combined with the Latinised Greek word ‘pithecus’, meaning ‘ape’. The species name kadabba means ‘oldest ancestor’ in the Afar language.

Distribution

Eastern Africa in the Middle Awash, Ethiopia

Relationships with other species

The scientists that discovered the remains claim this species is a direct human ancestor and the earliest species yet discovered on the human branch of the family tree. Those that discovered Orrorin tugenensis dispute this claim as they believe their find is a better candidate for direct human ancestry. Some scientists assign these remains to the subspecies Ardipithecus ramidus kadabba, because it shares many similarities to Ardipithecus ramidus , but has more primitive, or ape-like, teeth features.

Key physical features

• similar in size to modern chimpanzees

Body size and shape

• similar to modern chimpanzees
• the structure of the toe bones suggests that this species may have been bipedal. However, some scientists debate whether this fossil should be included with this species as it was found about 15 kilometres away from the other fossils and is dated several hundred thousand years younger.

Jaws and teeth

• some primitive dental features such as thick tooth enamel and relatively large canines compared to humans.
• some features of the teeth show a movement away from the primitive ape-like condition, such as molars that are larger than those of chimpanzees, a tendency towards incisiform lower canines and hominin-like upper pre-molars.

There is no evidence for any specific cultural attributes, but they may have used simple tools similar to those used by modern chimpanzees, including:

• twigs, sticks and other plant materials that were easily shaped or modified. These may have been used for a variety of simple tasks including obtaining food.
• unmodified stones, that is stones that were not shaped or altered before being used. These tools may have been used to process hard foods such as nuts.

Environment and diet

Fossil evidence from the site indicates the area was a mosaic of woodland and grasslands with lakes, swamps and springs. The discovery that this species lived in a forest environment challenged the theory about what kind of environment fostered the evolution of bipedalism. Did bipedalism evolve to take advantage of new open grassland environments, as was once believed, or did it first evolve in the trees?

The large back molars and narrower incisors (compared to chimpanzees) suggest that the diet included more fibrous foods than just fruit and leaves.

The Australian Museum respects and acknowledges the Gadigal people as the First Peoples and Traditional Custodians of the land and waterways on which the Museum stands.

Image credit: gadigal yilimung (shield) made by Uncle Charles  Chicka  Madden

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• Encyclopedia of Life

Pithecia pithecia Guianan saki

Geographic Range

White-faced sakis ( Pithecia pithecia ) are located in Brazil, and remote parts of Venezuela. Their range also encompasses most of French Guiana, Guyana and Suriname. They live along the Cuyuni river basin, east of the Caroni River, and south of the Orinoco River. ( Veiga and Marsh, 2008 )

• Biogeographic Regions

White-faced sakis are arboreal and live in both upland and lowland rainforests. Although they can inhabit very wet and very dry forests, they prefer areas with an abundance of fruit trees and watering holes. This species is most common at canopy heights of 15 to 25 m. They will also spend time foraging on the ground and at low levels in the understory foliage (3 to 15 m). Overnight sleeping areas typically are larger trees in the canopy with a wealth of foliage for cover. ( Anzelc, 2009 ; Cunningham and Janson, 2007 )

• Habitat Regions
• terrestrial
• Terrestrial Biomes

Physical Description

White-faced sakis exhibit sexual dimorphism, with larger males, and sexual dichromatism. Males have a black coat with white fur that surrounds their face. Female sakis have a shorter, brownish grey coat with two vertical lines from their eyes to their nose. Females may also have orange brown colored fur that emerges around the chest area and continues down to their abdomen. At birth males and adult females are very similar in appearance. A gradual color change over 3.5 to 4 years occurs, in which male sakis become all black with bright white faces. Sakis have long bushy tails, which are used for balance while jumping from tree to tree. The tails are not used for grasping objects or branches. Average adult mass is 1.8 kg; however, a slight sexual dimorphism separates males (2.38 kg) from females (1.76 kg). ( Anzelc, 2009 ; Fleagle and Meldrum, 1988 ; Norconk, 2006 )

• Other Physical Features
• endothermic
• bilateral symmetry
• Sexual Dimorphism
• male larger
• sexes colored or patterned differently
• male more colorful
• Average mass 1.871 kg 4.12 lb

Reproduction

Sakis are known to be monogamous in captivity (zoo environments) although Waters (1995) indicates there have been exceptions in the wild. Anzelc (2009) suggests that monogamy in the wild is less common than expected, and is less common when groups are larger than 2 to 3 individuals. Groups of 4 to 6 are not uncommon, and can include more than one adult breeding male or female. This suggests polygamous or polyandrous mating system, depending on the breakdown of adults in the group. ( Anzelc, 2009 ; Waters, 1995 )

• Mating System
• polyandrous

Males and females live in small groups. Despite practicing monogamy in zoos, a study of wild sakis in Venezuela found that some sakis were not monogamous. In wild groups, males will make calls to the females during mating season instead of as an alarm call. Males reach sexual maturity in approximately 32 months. Females are about the same age, but can take several months more. It isn't until the females' ovarian cycle is regular that they are determined sexually mature. Gestation periods for sakis average 146 days, and females bear 1 offspring at a time. Saki siblings from the previous year or 2 may help care for a newborn saki. ( Norconk, 2006 )

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• Breeding interval Sakis breed once per year.
• Breeding season Sakis breed in the spring.
• Average number of offspring 1
• Average gestation period 146 days
• Average age at sexual or reproductive maturity (male) 32 months

Females sakis are the predominant caretakers. Infants stay attached to their mother's thigh from birth to 1 month. From age one to four months, the young shift to a dorsal position where they can achieve greater mobility. The mothers carry their infants for the first 3 months. After the infant is around the age of 5 months, the mother will stop carrying it. They feed, protect, and nurture young until they are ready to be on their own. However, infants observe one birthing event prior to leaving their family group. ( Norconk, 2006 ; Waters, 1995 )

• Parental Investment
• female parental care
• extended period of juvenile learning

Lifespan/Longevity

In the wild, white-faced sakis have been known to live about 15 years. One wild-caught saki in captivity lived to the age of 36, spending over 28 of those years in captivity. ( de Magalhaes and Costa, 2009 )

• Range lifespan Status: captivity 36 (high) years
• Average lifespan Status: wild 15 years

Sakis are social but live in small groups of just 2 to 4 individuals. The groups travel together daily, and can easily move 1 to 2 km per day. Most movement occurs in the early morning and early hours of the afternoon. They spend about 9 hours on the move. These activity bouts are relatively shorter than related monkeys, who may be active 10 to 12 hours per day. Sakis are adept leapers aiding in predator avoidance. Male and female sakis exhibit grooming and mating behaviors, most common between mothers and infants. Male and female sakis teach each other how to raise the young. In captivity sakis have been known to carry group member's infant. ( Anzelc, 2009 ; Walker, 2005 ; Waters, 1995 )

Females in captivity start to reproduce much earlier in age than they would in the wild, which leads to earlier mortality. After reaching at least 37 months, the survival rate of the individual greatly increases. The longer the female waits to reproduce, the longer she will live. One example of late birthing was observed in an 18-year-old saki who gave birth in a zoo. This saki lived in the wild in its native habitat until she was captured late in life altering her behavior from those reared in captivity. ( Waters, 1995 )

• Key Behaviors
• territorial
• Range territory size 1 to 15.2 km^2

Groups of sakis in Suriname have been known to use a relatively small home range of 10 hectares. Relocated groups utilize much larger home ranges, and reports of 68 to 152 ha were typical. These sakis will mark and defend their territory by a series of activities. Anzelc (2009) summarizes them as "scent gland (sternal/gular/anogenital) rubbing, urine-washing, and territorial calls ....and agonistic interactions, using grunts, trills, branch and body shakes, piloerection, and fast pursuits to threaten and displace extra-group members." ( Anzelc, 2009 )

Communication and Perception

Sakis live in small groups ranging from 2 to 4 individuals; however larger groups of 6 or more have been reported and may include more than one adult breeding male or female. To establish territory they have loud vocal calls usually performed in duets of monogamous males and females. These duets strengthens their courtship bond. They also socialize by grooming on one another. White-faced sakis will scent-mark an area. Males rub their chests on trees. They usually choose trees with edible fruit to excite females and to try to stimulate courtship behavior during breeding season. ( Anzelc, 2009 ; Lehman, et al., 2001 ; Setz and Gaspar, 1997 )

• Communication Channels
• Other Communication Modes
• scent marks
• Perception Channels

Food Habits

Sakis eat the seeds of fruiting bodies. They spend 95 to 99% of total consumption time eating and breaking open the seeds. Year-round, they prefer to eat seeds 38 to 88% of the time. Leaves are also an important source of food. They eat the young leaves of plants during the dry season when fruits are not plentiful. Given this diet, their intake of fats are extremely high, but their intake of proteins are low. On occasion, they have been known to consume insects and flowers when fruit is scarce. ( Anzelc, 2009 ; Norconk and Conklin-Brittain, 2004 )

• Primary Diet
• Animal Foods
• Plant Foods
• seeds, grains, and nuts

When a terrestrial predator, such as red-tailed boas , are near sakis will first make an alarm call. Then they will group together and mob the predator in hopes of driving it away. Other terrestrial predators include a weasel called tayras , jaguars , green anacondas , and ocelots . Their biggest threats are avian predators. Because of their size, sakis are easy prey to the harpy eagle , which are known to attack large primates. A study reported more alarm calls when there is an avian threat, such as an eagle or vulture. When a bird of prey is spotted sakis make the alarm call, which is echoed by group member, and then they stay completely motionless. After time has elapsed, sakis might slip out undetected, heading for lowest parts of the canopy. They try to remain as concealed as possible in the canopy. ( Norconk and Gleason, 2002 )

• Anti-predator Adaptations
• tayra ( Eira barbara )
• jaguars ( Panthera onca )
• green anacondas ( Eunectes murinus )
• ocelots ( Leopardus pardalis )
• red-tailed boa ( Boa constrictor )
• vultures and harpy eagles ( Harpia harpyja )

Ecosystem Roles

Saki have parasites typical to that of new world monkeys and non-human primates. For example a common parasites are roundworms ( Pterygodermatites nycticebi ). Heartworms ( Dirofilaria immitis ) are present in this species, too. They can also get diseases such as diabetes and the Mayaro virus (which is found in mammals that live in trees). ( Gamble, et al., 1998 ; Thoisy, et al., 2003 )

• Ecosystem Impact
• disperses seeds
• roundworms ( Pterygodermatites nycticebi )
• heartworms ( Dirofilaria immitis )

Economic Importance for Humans: Positive

White-faced sakis are charaismatic organisms that attract high interest in zoos, however they are recently being exploited for their charisma. There is a market for these monkeys as pets, which is detrimental to the sakis. They are hunted as a source of food by locals. This hurts the population of sakis, because they don't reproduce quickly enough to replace the individuals killed and captured. ( "White-Faced Saki Monkey", 2012 )

• Positive Impacts
• body parts are source of valuable material

Economic Importance for Humans: Negative

Sakis may carry diseases which can be transferred to humans including the hepatitis virus and the naturally occurring herpes virsus (HSV-1). However, they are not a major disease transmitter. ( Schrenzel, et al., 2003 )

• Negative Impacts
• carries human disease
• causes or carries domestic animal disease

Conservation Status

This species is not currently listed by IUCN and is of little concern for conservation managers. However, due to habitat destruction and the pet trade, this status could change. It is listed in Appendix II of CITES, indicating that the species could become threatened if trade or import and/or export is not regulated. ( Veiga and Marsh, 2008 )

• IUCN Red List Least Concern
• US Federal List No special status
• CITES Appendix II
• State of Michigan List No special status

Contributors

Nicole Grubich (author), Radford University, Karen Powers (editor), Radford University, Kiersten Newtoff (editor), Radford University, Melissa Whistleman (editor), Radford University.

living in the southern part of the New World. In other words, Central and South America.

uses sound to communicate

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

Referring to an animal that lives in trees; tree-climbing.

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

either directly causes, or indirectly transmits, a disease to a domestic animal

uses smells or other chemicals to communicate

to jointly display, usually with sounds in a highly coordinated fashion, at the same time as one other individual of the same species, often a mate

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

parental care is carried out by females

an animal that mainly eats leaves.

A substance that provides both nutrients and energy to a living thing.

an animal that mainly eats fruit

an animal that mainly eats seeds

An animal that eats mainly plants or parts of plants.

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

Having one mate at a time.

having the capacity to move from one place to another.

the area in which the animal is naturally found, the region in which it is endemic.

the business of buying and selling animals for people to keep in their homes as pets.

chemicals released into air or water that are detected by and responded to by other animals of the same species

Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).

having more than one female as a mate at one time

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

breeding is confined to a particular season

remains in the same area

reproduction that includes combining the genetic contribution of two individuals, a male and a female

associates with others of its species; forms social groups.

uses touch to communicate

Living on the ground.

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

movements of a hard surface that are produced by animals as signals to others

uses sight to communicate

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

2012. "White-Faced Saki Monkey" (On-line). Oregon Zoo. Accessed April 09, 2012 at http://www.oregonzoo.org/discover/animals/white-faced-saki-monkey .

Anzelc, A. 2009. The Foraging and Travel Patterns of White-Faced Sakis in Brownsberg Nature Park, Suriname: Preliminary Evidence for Goal-Directed Foraging Behavior . Kent, Ohio: Kent State University, Master's thesis, 194 oo.. Accessed April 24, 2012 at http://www.personal.kent.edu/~mnorconk/pdfs/Anzelc-thesis-7-20-09b.pdf .

Cunningham, E., C. Janson. 2007. Integrating information about location and value of resources by white-faced saki monkeys (Pithecia pithecia). Animal Cognition , 10/3: 293-304.

Fleagle, J., D. Meldrum. 1988. Locomotor behavior and skeletal morphology of two sympatric Pitheciine monkeys, Pithecia pithecia and Chiropotes satanas. American Journal of Primatology , 16/3: 227-249.

Gamble, K., J. Fried, G. Rubin. 1998. Presumptive dirofilariasis in a pale-headed saki monkey (Pithecia pithecia). Journal of Zoo and Wildlife Medicine , 29/1: 50-54.

Lehman, S., W. Prince, M. Mayor. 2001. Variations in group size in white-faced sakis (Pithecia pithecia): Evidence for monogamy or seasonal congregations. Neotropical Primates , 9/3: 96-100.

Lloyd, M., J. Susa, J. Pelto, A. Savage. 1995. Gestational diabetes mellitus in a white-faced saki (Pithecia pithecia). Journal of Zoo and Wildlife Medicine , 26/1: 76-81.

Norconk, M. 2006. Long-term study of group dynamics and female reproduction in Venezuelan Pithecia pithecia. International Journal of Primatology , 27/3: 653-674.

Norconk, M., A. Rosenberger, P. Garber. 1996. Adaptive Radiations of Neotropical Primates . New York: Plenum Press.

Norconk, M., N. Conklin-Brittain. 2004. Variation on frugivory: The diet of Venezuelan white-faced Sakis. Internation Journal of Primatology , 25/1: 1-26.

Norconk, M., T. Gleason. 2002. Predation risk and antipredator adaptations in white-faced sakis, Pithecia pithecia. Pp. 169-183 in L Miller, ed. Eat or be eaten: predator sensitive foraging among primates . Cambridge, United Kingdom: Press Syndicate of the University of Cambridge.

Richard-Hansen, C., C. Fournier-Chambrillon. 2001. Abundance, use of space, and activity patterns of white-faced sakis (Pithecia pithecia) in French Guiana. American Journal of Primatology , 55/4: 203-221.

Schrenzel, M., K. Osborn, A. Shima, R. Klieforth, G. Maalouf. 2003. Naturally occuring fatal herpes simplex virus 1 infection in a family of white-faced saki monkeys (Pithecia pithecia pithecia). Journal of Medical Primatology , 32/1: 7-14.

Setz, E., J. Enzweiler, V. Solferini, M. Amendola, R. Berton. 1999. Geophagy in the golden-faced saki monkey (Pithecia pithecia chrysocephala) in the Central Amazon. Journal of Zoology , 247/1: 91-103.

Setz, E., D. Gaspar. 1997. Scent-marking behaviour in free-ranging golden-faced saki monkeys, Pithecia pithecia chrysocephala: Sex differences and context. Journal of Zoology , 241/3: 603-611.

Sussman, R., J. Phillips-Conroy. 1995. A survey of the distribution and density of the primates of Guyana. International Journal of Primatology , 16/5: 761-791.

Thoisy, B., J. Gardon, R. Salas, J. Morvan, M. Kazanji. 2003. Mayaro virus in wild mammals, French Guiana. Emerging Infectious Diseases , 9/10: 1326-1329.

Thoisy, B., I. Vogel, J. Reynes, J. Pouliquen, B. Carme, M. Kazanji, J. Vie. 2001. Health evaluation of translocated free-ranging primates in French Guiana. American Journal of Primatology , 54/1: 1-16.

Veiga, L., L. Marsh. 2008. "IUCN Red List of Threatened Species" (On-line). Pithecia pithecia. Accessed February 23, 2012 at www.iucnredlist.org .

Walker, S. 2005. Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela. American Journal of Primatology , 66/4: 369-387.

Warren, K., M. Norconk. 1993. : Physical and chemical properties of fruit and seeds eaten by Pithecia and Chiropotes; in Surinam and Venezuela. International Journal of Primatology , 14/2: 207-227.

Waters, S. 1995. A review of social parameters which influence breeding Pithecia pithecia in white-faced saki in captivity. International Zoo Yearbook , 34/1: 147-153.

de Magalhaes, J., J. Costa. 2009. A database of vertebrate longevity records and their relation to other life-history traits. Journal of Evolutionary Biology , 22/8: 1770-1774.

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New Research

Scientists Now Know Where the Largest Ape to Ever Exist Sits in Primate Family Tree

Proteins from a 1.9 million-year-old molar show that the 10-foot-tall ‘Gigantopithecus’ is a distant relative to modern orangutans

Jason Daley

Correspondent

Western scientists first learned about extinct giant ape species Gigantopithecus blacki—the largest primate to ever exist—in 1935 when an anthropologist came across some of its massive molars in Chinese drug stores selling them as dragon teeth. Since then, researchers have identified thousands of teeth and a few partial jawbones from the creature. With these pieces in hand, they’ve tried to fit the bigfoot-like ape into the primate family tree. Without any usable DNA, however, the task has been difficult.

Now, using proteins in dental enamel, researchers report they've finally found how the Gigantopithecus fits into the great ape puzzle, according to a new study published in the journal Nature .

According to a press release , DNA has been key in helping scientists map out the messy relationships between primates and hominids that lived within the past 50,000 years. But in fossils older than that, DNA is very difficult to extract and scientists have only done it successfully in a few rare cases, including in one 400,000-year-old hominin specimen.

Gigantopithecus remains are estimated to be between 300,000 to 2 million years old, placing its reign at some point during the Pleistocene epoch.

No Gigantopithecus DNA has ever been recovered. That’s why an international team of researchers used techniques from an emerging field called proteomics to get molecular information from the Gigantopithecus molar in the new study.

In traditional DNA sequencing , pieces of the DNA molecule are put through a process that copies its sequence of nucleotides and puts them back together into a full genome. The quality and completeness of the genome, however, depends on how well-preserved the original sample of DNA is. Most DNA degrades much more quickly, especially in hot, humid climates.

But in proteomics, researchers more or less reverse-engineer DNA by looking at the proteins preserved in teeth, which last much longer. Because each protein is made up of amino acids, and because each amino acid is encoded by a three-letter DNA sequence, researchers can produce snippets of ancient DNA by analyzing the proteins. Last September, the technique was used to properly place a 1.7-million-year-old species of wooly rhinoceros in its family tree, proving that the method could be used to understand ancient animals.

Researchers applied the protein-mining technique to a 1.9 million-year-old molar from Gigantopithecus found in a Chuifeng cave in China. Gretchen Vogel at Science reports the team dissolved tiny amounts of enamel from the tooth and then analyzed it using mass spectrometry. They were able to identify 500 peptides, or short chains of amino acids, from six different proteins.

Bruce Bower at Science News reports that five of those proteins still occur in extant ape and monkey species. The team compared the accumulated differences in the proteins to those animals, finding that the massive Gigantopithecus is a distant relative of modern orangutans. The two lineages likely diverged from a common ancestor over 10 million years ago.

“Until now, all that was known about this species was based on the morphology of the many teeth and the few mandibles found, typical of a herbivore," study author Enrico Cappellini, an evolutionary geneticist at the University of Copenhagen, says in the press release. “Now, the analysis of ancient proteins, or palaeoproteomics, has allowed us to reconstruct the evolutionary history of this distant relative.”

The success of this technique has big implications for the future of paleoanthropology. Because many of the fossilized remains of ancient hominins come from tropical and subtropical areas, like East Africa, southern Africa and Indonesia, there’s little chance that viable DNA has survived. But the protein trick changes everything.

“Until now, it has only been possible to retrieve genetic information from up to 10,000-year-old fossils in warm, humid areas,” Welker tells Katie Hunt at CNN . “This is interesting, because ancient remains of the supposed ancestors of our species, Homo sapiens, are also mainly found in subtropical areas, particularly for the early part of human evolution. This means that we can potentially retrieve similar information on the evolutionary line leading to humans.”

The team also says that they may be able to look at more than just molars. It could be possible to analyze protein sequences in the bones of apes and hominins that lost their viable DNA long ago.

While the study tells researchers a little bit about Gigantopithecus’s origins, Capellini tells Hunt that it doesn’t shed much light on what the massive ape looked like or how it behaved.

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Jason Daley is a Madison, Wisconsin-based writer specializing in natural history, science, travel, and the environment. His work has appeared in Discover , Popular Science , Outside , Men’s Journal , and other magazines.

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Khabarovsk, Russia

You are here, about khabarovsk.

Khabarovsk is one of the most significant and beautiful cities of Russia's Far East. It stands on the right bank of the Amur River along the scenic Trans-Siberian railway and almost touches the Chinese border.

The city of Khabarovsk played a crucial role in East - Russian history and is famous for its historic sights, monuments of architecture of different eras, religious buildings, lovely parks, gardens, and artificial lakes which surprise its visitors with impressive fountain shows.

Khabarovsk History

Founded in 1858, the city is now loved by Chinese travelers and those who are going on iconic train journeys along the world's longest railway from Siberia. After days of relentless taiga, people reach this vibrant city with multiple attractions, plenty of historical sights from the tsarist-era, and a number of places to try traditional Russian cuisine. Khabarovsk is indeed a charming city that deserved to be on your travel itinerary. Especially, if you are the legendary Trans-Siberian is on your travel radar.

We suggest beginning your Khabarovsk tour from the famous monument erected in honor of Nikolay Muravyov - Amursky, one of the best-known explorer of East Siberia, a general, and the founder of the city.

Continue your Khabarovsk trip with a riverside walk along the picturesque Nevelsky Embankment and pass through the third tallest church (35 feet) in all Russia - Spaso-Transfiguration Cathedral standing on top of a hill. Take in the spectacular location and view of this Cathedral and its classic golden domes, dominating the city skyline and being visible from a large distance.

Your Trans-Siberian itinerary would become even better if you include a visit to the famous Khabarovsk Bridge as well. The railway bridge goes over the Amur River and is considered to be the longest bridge on the Trans-Siberian route.

Best Things to Do in Khabarovsk

• Stop by the Nikolay Muravyov - Amursky monument
• Visit the gorgeous Spaso-Transfiguration Cathedral
• Take a picture by the renown Khabarovsk Bridge over the Amur River

Top Attractions in Khabarovsk

The Kamchatka peninsula is perhaps one of the most beautiful locations in the world. With about 300 volcanoes, 29 of which are still active, the mountains dazzle visitors.

All Attractions in Khabarovsk

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Explore Khabarovsk

Plan Your Trip to Khabarovsk: Best of Khabarovsk Tourism

Essential khabarovsk.

Khabarovsk Is Great For

Shopaholics

Eat & drink

• Intourist Hotel
• AZIMUT City Hotel Khabarovsk
• Boutique-Hotel Khabarovsk City
• Restaurant Dom by Rakusa
• Amur Restaurant
• Sultan Bazar
• Restaurant Sopka
• Pizzeria VDROVA
• Teach Me How to Drink
• The Bridge Across the River Amur
• Khabarovsk Regional Museum Named After N.I. Grodekov
• Lenin Square

Understand [ edit ]

Overlooking the confluence of the Amur and Ussuri Rivers, Khabarovsk is the second largest city in the Russian Far East , approaching 600,000 residents and growing. It is also the capital of both Khabarovsk Krai and the Far Eastern Federal District . Unlike Vladivostok, the city has never been closed to foreigners, and retains a distinct international feel, rare for the Russian provincial centers – a feeling propped up by an increasing Asian presence with arrivals from Asian countries now numbering over a million each year. In turn, Asians come here to experience a piece of Europe close to home, with the fortunate effect that the city is spending huge swaths of money renovating the city, in which old classical buildings were spared much of the destructive effects of the 1917-23 civil war, to provide its visitors with just that feeling. From a European's perspective, Soviet city planning has unmistakably taken its toll, but it is still far more attractive than your average Siberian city.

Climate [ edit ]

The climate is temperate and monsoonal, with a cold, dry winter and a hot and humid summer. The average temperature for a full year is just 2°C, but covers over wide span of monthly averages ranging from a bone chilling −20°C in January to a quite warm +21°C average in July. The city sees an average of 686 mm precipitation in a year, but unfortunately the lions' share falls in the warm summer months. The number of sunny days per year is 70, which is higher than Moscow's 54. Climate-wise, end of May - early June or end of August - early September are the best time for a visit.

Get in [ edit ]

By plane [ edit ], by train [ edit ].

• 48.49659 135.07283 2 Khabarovsk railway station ( Habarovsk 1 ), Leningradskaya, 58 , ☏ +7 4212 38-39-40 . Khabarovsk station, listed as Habarovsk 1 in most train schedules, is a major stop on the Trans-Siberian Railway . There are several trains each day bound for Vladivostok (800 km) and Moscow (about 8500 km) along the main Trans-Siberian line. Other options include trains #386 or #035 to Blagoveshchensk , #325 for Tynda, #667э for Komsomolsk , #943э Vanino , all on the Baikal-Amur Mainline . Vanino is an interesting option as it allows ferry connections to Sakhalin and further on to Wakkanai in Japan – more details in the Russia to Japan via Sakhalin itinerary. The international trains are Khabarovsk- Harbin , ongoing twice a week and Khabarovsk- Pyongyang on special days.  

By boat [ edit ]

• 48.46904 135.0581 3 River Port , Shevchenko 1 .  

If you want to go to places upstream on the Amur river, the Meteor speedboats will often be your transport of choice, but only during the summer when the river is navigable. However, in 2008, the water level in the river was at a historic low, so that the Meteor traffic had to be stopped. If Meteor traffic functions normally, you can go some 1,000 km downstream to the Ul'chi municipal district (rayon), a region mostly inhabited by indigenous Ul'chi people.

• Fuyuan – In spring and summer there are daily hydrofoil services to Fuyuan in northeastern China , departing from the ferry terminal facing the Amur river.
• Komsomolsk – If you are heading for the BAM line up north, an interesting option is to take a hydrofoil cruising up the Amur river to Komsomolsk (6 hours), and catch a train from there.

Get around [ edit ]

The best thing to start with is to walk around the center of the city. Have a nice walk from Lenin Square to the Amur River via the main street, Muravieva-Amurski. You will find all sorts of shops and places to eat.

By tram [ edit ]

The city has a network of four tram lines (there is no line 3 or 4). The most useful section for visitors is the stretch of the network running from the main railway station along Amursky Boulevard, before making a left turn down Volochaevskaya St. (near the market), and crossing Muravyov-Amursky Street one block west of Lenina Square, it then continues south intersecting Lenina Street roughly at its halfway point, before a stop at the botanical gardens (Lines 1, 2 & 6). The remainder of the network mainly extends into the sleepy suburbs. Line 5 serves the North, Line 1 and 2 the South along Krasnorechenskaya St.

By bus [ edit ]

The electric trolleybuses also has a few useful sections for visitors, Line 1 runs between the Airport and Komsomolskaya Square (River promenade, Museum cluster) along Karla Marksa and Mureava Amursky streets.

The regular bus number 1, is a useful circle line. It starts at the Railway station, turns down Seryshev street (a block north of Amursky Boulevard) until it reaches the river park at Lenin Stadium. Turns down Komsomolskaya Street (and square) and runs south until Lenina Street. It then runs the entire length of Lenina street before north at the City History Museum and returns to the train station.

Major destinations [ edit ]

• Airport 18, T1, T2, T4
• Botanical Gardens' 9, 25, 29, 33, 54
• City History Museum 1, 54, 56, 57, T5
• Komsomolskaya Square 1, 9, 14, 19, 29, 34, 38, 55, 56, T1, T3
• Lenina Square 14, 19, 21, 29, 34, 38, 55, 56, T1, T3
• Railway Station 1, 6, 7, 11, 13, 20, 22, 24, 26, 34, 54, 57, T2, T5
• Slavy (Glory) Square 1, 9, 29, 33, 34, 56

See [ edit ]

The Far Eastern Museums [ edit ]

There is a fantastic cluster of top notch museums along Shevchenko Street, just behind the tall blue-domed Church of Theotokos on Komsomolskaya Square towards the river and stadium. Not only are the museums some of the best in the far east, they also make their home in some impressive century-old buildings dating back to before the revolution. After a visit, the nice river promenade is just a short walk away, so you can wash all that new found knowledge away with some pivos in good company.

Tugged away just across the next street behind the military museum, you also find the Archeology Museum on Turgeneva street.

Learn [ edit ]

The Pacific National University [dead link] , formally a Polytechnic Institute, is now a full fledged university, with over 21.000 students enrolled. Has a single Masters programme in Computer Sciences in cooperation with a German university, which is taught in English.

The Far Eastern State University of Humanities [dead link] offers a summer course in Russian language in July as well as courses during the academic year.

The Far-Eastern State Medical University [dead link] is a major medical institution in Eastern Siberia.

The Far-Eastern State University of Railways being one of the largest universities includes the course of Russian-Americam Programme.

The Far-Eastern State Scientific Library is an old and beautiful Art-Nouveau building in the city's center and has American, German and Japanese centers.

Japanese Center in Khabarovsk offers course of Japanese language as well as participation in business seminars

Connect [ edit ]

Phones [ edit ].

Khabarovsk has the usual set of Russian mobile operators:

GSM 900/1800:

• Beeline ( by Vympelcom ), ☏ +7 4212 64-90-64 .  
• Megafon , toll-free: +7 800 333 0500 .  
• MTS ( Mobile TeleSystems ), toll-free: +7 800 333 0890 .  
• Skylink , Dzerzhiskogo, 4 ( Near Amur hotel. ), ☏ +7 4212 74-44-44 . The all-Russian CDMA operator, having less subscribers, than GSM operators, but popular for faster and cheaper mobile Internet service.  
• Megafon , ☏ +7 800 333 0500 . new standart of mobile internet.  

Check roaming prices, especially for mobile Internet, before using any non-Russian SIM card. Some mobile connection standards are not supported in Russia , e.g. those for Japan and the United States .

If you're staying in Russia for a week or more, it's definitely worth it to buy a local SIM card, but be aware, that a passport is needed for that. The easiest way to refill a local mobile account is to use an ATM. Most ATMs have bilingual interfaces, allowing numerous kinds of payments, including those for mobile services by local operators. You can also do it through terminals spread all over the city - like Qiwi or mobile shops.

Post [ edit ]

The General post office at 28 Muravyov-Amurskiy St. If you plan on calling anyone, Khabarovsk is UTC +10 (or 7 hours ahead of Moscow).

The post-office at the railway station is located on 13 Leningradsky per. about 200 m from the station building.

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