Primate Conservation 2019 (33): 65-73

A New Tarsier Species from the Togean Islands of Central Sulawesi,

Indonesia, with References to Wallacea and Conservation on
Sulawesi

Myron Shekelle1, Colin P. Groves2†, Ibnu Maryanto3, Russell A. Mittermeier4, Agus Salim5 and Mark S. Springer6

Department of Anthropology, Western Washington University, Bellingham, WA, USA

1

2
School of Archaeology and Anthropology, Australian National University
Canberra, Australia
3
Museum Zoologicum Bogoriense, LIPI, Cibinong, Indonesia
4
Global Wildlife Conservation, Austin, TX, USA
5
PT. Hatfield Indonesia, Jakarta, Indonesia
6
Department of Evolution, Ecology, and Organismal Biology, University of California, Riverside, CA, USA

Abstract: We describe and name a new species of tarsier from the Togean Islands, in Tomini Bay, bounded by the northern and
eastern peninsulas of Sulawesi. In doing so, we highlight how 25 years of sustained research on the alpha taxonomy of Sulawe-
sian tarsiers, Tarsius, have helped to identify key conservation priorities in the Sulawesian region of the Wallacea Biodiversity
Hotspot.

Key words: Biodiversity, bioacoustics, cryptic species, duet call, Togean form, taxonomy, hotspots

Introduction grown from one to twelve (including the species described

here). Based on variation in their calls, Shekelle and Lek-
The impending global environmental catastrophe of bio- sono (2004) predicted that 16 or more cryptic species would
diversity loss has been termed the “sixth mass extinction” eventually be described from Sulawesi and its surrounding
(for example, Leakey and Lewin 1996), and led Myers et al. islands. Nietsch and Niemitz (1993) reported tarsiers from
(2000) to propose that global conservation priorities should the Togean Island chain, and believed that they might be tax-
focus especially on the “hotspots” method for identifying onomically distinct. The suggestion to taxonomically sepa-
global conservation priorities; with only so much time and rate Togean Island tarsiers was supported by further work on
resources, conservation efforts should be spent “where excep- the role of vocalizations in species differentiation among the
tional concentrations of endemic species are undergoing Sulawesi tarsiers by Nietsch and Kopp (1998) and Nietsch
exceptional loss of habitat” (p.853; see also Mittermeier and (1999). Shekelle et al. (1997) conducted playback tests in the
Rylands 2017). Wallacea is one of the top 25 global hotspots wild that concurred with the findings of Nietsch, i.e. Togean
for biodiversity conservation, and Sulawesi is by far the larg- Island tarsiers appeared to be taxonomically separable on the
est landmass within it, and by some measures the highest basis of vocalizations. Shekelle (2003) reported that genetic
regional priority for conservation (Wilson et al. 2006). data were consistent with the hypothesis that tarsier acous-
In the time since Niemitz (1984) revised tarsier taxonomy tic forms, including the Togean form, were distinct species,
and MacKinnon and MacKinnon (1980) published the first a finding also supported by Shekelle (2008) and Shekelle et
report of wild tarsiers from Sulawesi—wherein they noted al. (2010). The hypothesis that tarsier acoustic groups are
the presence of a duet call with geographically-structured distinct species was robustly supported in a fine-grained phy-
variation and the likelihood of much unrecognized taxonomic logeographic study of T. dentatus and T. lariang by Merker et
variation—the number of recognized tarsiers on Sulawesi has al. (2009), who tested sequence data from mtDNA, y-chromo-
some, and microsatellites against bioacoustics and morphol-
†Colin P. Groves 1942-2017 ogy along a transect that passed through the species boundary.

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Table 1. Measurements of type specimens compared with two recently

described species T. spectrumgurskyae and T. supriatrnai.

Figure 1. Skulls of three type specimens (left to right), Tarsius supriatnai, T.

spectrumgurskyae and T. niemitzi. Photos by Myron Shekelle.

We present a new molecular phylogeny for Tarsiidae based on and relatively dark pelage with dark gray facial fur, particu-
12S mtDNA sequences that offers further support that Togean larly in adults (Fig. 3). Atypical for tarsier species endemic to
Island tarsiers are taxonomically separable from other known small islands, T. niemitzi does not have a reduced tail tuft (see
tarsier taxa. Shekelle et al. 2008b). VOCALIZATIONS: Spectrograms of
the Togean form were published by Nietsch and Kopp (1998)
Tarsius niemitzi sp. nov. (Fig. 4). Its duet is structurally simple, possibly the simplest
of all known tarsier duets (Nietsch and Niemitz 1993). One
Holotype: MZB 32654 (Fig. 1, Table 1), adult male. female call is followed by two or three male calls. The female
Museum Zoologicum, Bogoriense, Cibinong, West Java, note is a downward modulated whistle, with a maximum fre-
Indonesia. Collector: Simson Katiandagho, 1 October 2009. quency of 12−13 kHz, and a minimum frequency around 6
Type locality: Desa Benteng, Kecamatan Togean, Kabu- kHz. Each female note has a duration of about 0.5 seconds
paten Tojo Una-Una, Central Sulawesi, Indonesia. Geo- and notes are repeated at the rate of about 1 call per 1.5 sec-
graphical coordinates for Desa Benteng, Togean Island are onds. The male note resembles a temporally compressed ver-
0°23'36.7"S, 122°01'37.8"E. sion of the female call: maximum frequency is around 10−11
Hypodigm: Only the type specimen available. kHz, and the minimum frequency is 5 kHz or lower. The
Description: MORPHOLOGY: Surveys of wild popu- duration of the male note is only about 0.15 seconds. Male
lations have indicated that body weight and tail length fall calls occur in a phrase of 2 or 3 notes that gradually ascend
within the range of a number of other Tarsius species, includ- in maximum frequency. Togean tarsiers are unique among
ing T. tarsier, T. fuscus, T. dentatus, T. spectrumgurskyae, and known tarsier acoustic forms in that they respond in play-
T. supriatnai. Body weights and tail lengths may well over- back experiments to all other tarsier duet calls by duetting
lap with all species of Tarsius, except T. pumilus and possi- themselves (Fig. 5). GENETICS: Shekelle et al. (2008a)
bly some offshore taxa, such as T. sangirensis. Body weight: reported genetic evidence showing Togean Island tarsiers to
female 104−110 g (n = 2); male 125−138 g (n = 3). Tail be a monophyletic clade with an apomorphic 2 base pair dele-
length: female = 245−261 mm (n = 2); male 246−258 mm (n tion in the 12S gene that is diagnostic of the clade. Shekelle
= 3) (data from Shekelle 2003). Tarsius niemitzi is similar to T. et al. (2010) used a subset of their previous alignment, which
dentatus in having darkly pigmented skin, particularly the tail, eliminated the region with the 2 bp deletion from the analysis.

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 New species of tarsier on the Togean Islands

Figure 2. Top right: Southeast Asia and Australia, with red box indicating
contents of Top Left. Top left: Sulawesi, with red box indicating contents of
bottom. Bottom: The Togean Islands in the U-shaped bay of Tomini, bordered
in the north and west by the northern peninsula, and the south by the eastern
peninsula of Sulawesi, Indonesia. Expected species distribution circled in red, Figure 3. Niemitz’s tarsier Tarsius niemitzi sp. n. from Malenge Island,
approximate location of type locality marked with red X. Sulawesi. Photo by Myron Shekelle, illustration by Stephen D. Nash.

Figure 4. Spectrograms for six species of Eastern tarsiers (adapted from Shekelle 2008). The Sejoli form might be conspecific with T. supriatnai.

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Shekelle et al.

Methods

DNA sequences for the 12S rRNA gene in Tarsius, Car-

lito, and Cephalopachus were downloaded from GenBank
and imported into Geneious Prime 2019.1.1 (Kearse et al.
2012). Partial sequences <750 bp in length were discarded.
The remaining 29 sequences were aligned in Geneious with
MAFFT (Katoh et al. 2002; Katoh and Standley 2013). The
resulting alignment encompassed 968 bp for 11 tarsiid spe-
cies. We used PhyML 3.3.20180214 (Guindon et al. 2010) to
construct a maximum likelihood phylogram and to perform a
bootstrap analysis with 500 pseudoreplicates. We employed
a GTR + Γ model of sequence evolution with SPR branch
swapping. We also analyzed the 12S alignment with MrBayes
3.2.6 (Huelsenbeck and Ronquist 2001) and a GTR + Γ model
of sequence evolution. The Bayesian analysis was run for
1,000,000 generations after a burnin of 100,000 generations
with four heated chains. A molecular dating analysis was per-
formed with the mcmctree program (Yang and Rannala, 2006)
in PAML (Yang 2007). Given the absence of appropriate tar-
siid fossils for calibrating the tree, we employed secondary
Figure 5. Results of playback tests (adapted from Shekelle 2008). calibrations for three different nodes. First, crown Tarsiidae
was calibrated with a minimum age of 20.4 Ma and a maxi-
Nevertheless, Togean Island tarsiers were still recovered as mum age of 25.2 Ma based on five recent timetree studies that
a monophyletic clade. We performed a new alignment after are available on timetree.org (Fabre et al. 2009; Shekelle et
downloading the sequences from GenBank, and again find a al. 2010; Springer et al. 2012; dos Reis et al. 2012; Pozzi et
robustly supported monophyletic clade (Fig. 6). al. 2014). We also employed minimum and maximum age
Diagnosis: The duet call, and the species provenance and constraints for Carlito to Cephalopachus (min = 4.8 Ma, max
genetics are diagnostic. = 18.6 Ma) and crown Tarsius (min = 3.4 Ma, max = 11.1
Distribution: Togean Islands, except for Una Una (Fig. Ma) based on Shekelle et al. (2010). The mcmctree analy-
2). sis was run for 100,000 generations after 10,000 generations
Etymology: Niemitz’s tarsier. This species is named of burnin. We performed the analysis with autocorrelated
in honor of Dr. Carsten Niemitz, universally regarded as the rates and soft-bounded constraints that allowed for 2.5% tail
father of tarsier field biology. Although his field work was probabilities.
preceded by that of Fogden (1974), it was Niemitz who first
dedicated a substantial part of his career to the systematic Results
study of wild tarsiers. Niemitz and his student, Alexandra
Nietsch, surveyed tarsiers in Central Sulawesi. Subsequently, Figure 6A shows the timetree that resulted from the
reportedly acting on a tip from an Indonesian government mcmctree analysis. The maximum-likelihood phylogram
official, Rolex Lameanda, Nietsch traveled to the Togean (log-likelihood = -2835.37431) from the PhyML analysis of
Islands to survey tarsiers there and, with Niemitz, was the first 12S rRNA mtDNA is shown in Figure 6B. The phylogram and
to report their presence and the likelihood of their taxonomic timetree are both rooted between Tarsius and Carlito + Cepha-
distinctiveness (Nietsch and Niemitz 1993). We have found lopachus following Groves and Shekelle (2010) and Springer
no evidence that Tarsius togeanus (as per Surjadi and Supri- et al. (2012). Bootstrap support percentages (PhyML) and
atna 1998, p.281; Ito et al. 2008, p.234) and T. togianensis (as posterior probabilities (MrBayes) are also shown on the time-
per Gursky 2007, p.5) are names arising from formal descrip- tree for clades that are supported above the 50% bootstrap and
tions, and they are as such nomina nuda. 0.50 posterior probability levels. Relationships among differ-
Local Name: Bunsing, tangkasi, podi ent species of Tarsius are not well resolved, but the Togean
Field work: Nietsch and Niemitz (1993), Shekelle et al. Island tarsiers form a well-supported monophyletic clade
(1997), Nietsch and Kopp (1998), Nietsch (1999), Shekelle (bootstrap support = 92%, posterior probability = 0.98). This
(2003, 2008). support level is comparable to that for T. dentatus (92.8%)
Material examined: There is no material that we know and T. wallacei (90.0%).
of other than the type specimen. Shekelle (2003) observed
and photographed several individuals that were captured and
released.

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 New species of tarsier on the Togean Islands

Table 2. Specimens in the phylogenetic analysis.

Figure 6. A (above). Time-calibrated phylogeny based on mcmctree (Yang and

Rannala 2006) with bootstrap support percentages (PhyML) above branches,
and Bayesian posterior probabilities below branches for clades with support
values >50% (bootstrap percentages) and 0.50 (posterior probabilities). The
timescale is in millions of years.. See Table 2 for specimen ID and GenBank
accession numbers. B (below). Maximum likelihood phylogram based on
PhyML 3.3.20180214 (Guindon et al. 2010) analysis of 12S rRNA sequences genetic markers, will find strong support for the taxonomic
with a GTR + Γ model of sequence evolution.
separability of T. niemitzi, as Merker and others did for T. lari-
ang. We strongly encourage further work on tarsier taxonomy
Discussion using holistic approaches with complementary data sets.
Extant tarsiers are classified in three genera, following
We describe a new species of tarsier from the Togean Groves and Shekelle (2010), that are allopatrically distrib-
Islands of Central Sulawesi, Indonesia. It was long suspected uted in distinct biogeographic regions: Tarsius is found on
on the basis of bioacoustic (Nietsch and Niemitz 1993) and Sulawesi and surrounding islands; Cephalopachus is found
biogeographic (Shekelle and Leksono 2004) evidence, and on a restricted subset of Sundaland, principally southern
later mtDNA sequence (Shekelle et al. 2008). Merker et al. Sumatra and the island of Borneo; Carlito is found on islands
(2014) have argued for a cautious interpretation of mtDNA of the southern Philippines that were a single Ice Age land-
trees, particularly in tarsiers. Indeed, Merker et al. (2009) mass, sometimes called Greater Mindanao. Tarsius contains
made a far stronger case for the taxonomic separability for twelve species (in order of seniority): T. tarsier (= spectrum),
the species that Merker and Groves described in 2006, on T. fuscus, T. sangirensis, T. dentatus (= dianae), T. pumilus,
the basis of morphological, anatomical, and acoustic analy- T. pelengensis, T. lariang, T. tumpara, T. wallacei, T. spec-
ses, than we make here. But as with their experience, we are trumgurskyae, T. supriatnai, and now T. niemitzi. Burton and
cautiously optimistic that subsequent analyses, using more Nietsch (2010) reported three more unnamed taxa from the

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Southeastern peninsula of Sulawesi. Shekelle and Leksono Specifically, the T. tarsier species complex (i.e. all of the spe-
(2004) predicted Sulawesi would ultimately be shown to be cies of Tarsius except for T. pumilus), listed as Lower Risk /
home to at least 16 distinct tarsier taxa. The speciose alpha Near Threatened (LR/nt) in 2000, would likely be listed as
taxonomy of Tarsius stands in contrast with that of Cepha- Vulnerable (VU), were it classified as a single species (Fig. 7,
lopachus and Carlito. We question if this contrast is based Table 3). With reclassification, taxa in this species complex
upon knowledge or ignorance of the alpha taxonomy of the range from VU to Critically Endangered (CR) (Fig. 7, Table
latter two genera, and we encourage more fieldwork in order 3). Four of the sixteen taxa hypothesized by Shekelle and
to answer this question (see Brandon-Jones et al. 2004). Leksono (2004) are VU, seven are EN, one is CR, and four
are DD (Fig. 7, Table 3).
Conservation Gursky et al. (2008), following previous work by Supri-
atna et al. (2001) and Shekelle and Leksono (2004) estimated
One of the most significant outcomes of 25 years of sus- that the Togean Island tarsier would be Endangered [EN],
tained research on the alpha taxonomy by one of us (MS) and that the Togean Islands were a unique area of endemism
and subsequently by colleagues (for example, Merker and in Sulawesi, in that they possess native tarsiers, lack native
Groves 2006; Merker et al. 2009, 2010; Driller et al. 2015) monkeys, and appear to be geologically separated from the
has been to identify the critical conservation hotspots in the Sulawesi mainland by an ocean barrier that exceeds 120 m
Sulawesian region (i.e. Sulawesi plus nearby islands), of in depth—believed to be deeper than the lowest ocean levels
the Biodiversity Hotspot of Wallacea (Fig. 7). Reclassify- during Pleistocene glacial maxima. Each of these assertions
ing a single widespread species into numerous allopatric and are hypotheses for further testing. The broader implication
parapatric cryptic sibling species has the secondary effect is that the Togean Islands possess a largely endemic biota of
of clarifying conservation priorities (Bickford et al. 2007). taxa that do not disperse easily across water barriers.

Table 3. Conservation status for 16 Indonesian tarsier taxa and populations (listed in Brandon-Jones et al., 2004) show the impact of 25 years of sustained research
on the alpha taxonomy of Sulawesian tarsiers of the genus Tarsius toward priority areas in the Wallacea Biodiversity Hotspot.

*These populations still remain classified with T. tarsier, but are implausibly disjunct, and are hypothesized to be three separate taxa.
**These populations were classified with T. tarsier (=spectrum) in 2000.

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 New species of tarsier on the Togean Islands

Figure 7. Mapping the priority areas hotspots in the Wallacea Biodiversity Hotspot: a 25-year effort. (L) 2000 and prior, (R) present based upon combined sources
(Supriatna et al. 2001; Gursky et al. 2008; IUCN Red List). Green = LR/NT, Yellow = VU, Orange = EN, Red = CR. Note: distribution of T. pumilus is a conjectural
representation of regions between 1800–2200 m above sea level in the species’ polygon.

Acknowledgments
facilitated by a grant from the Margot Marsh Biodiversity
We are deeply saddened by the passing of our colleague Foundation and a private donation by the Tjiasmanto Family.
and coauthor, Colin Groves, on November 30, 2017. Colin
was described by Jane Goodall as the world’s greatest pri- Literature Cited
mate taxonomist, and while we concur, that description only
scratches the surface of his expertise. A recent bibliogra- Bickford D., D. J. Lohman, N. S. Sodhi, P. K. L. Ng, R. Meier,
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description, Didermocerus sumatrensis harrissoni Groves, cies as a window on diversity and conservation. Trends
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Groves authored the description of two new families, five D. J. Melnick, J. C. Morales, M. Shekelle and C.-B.
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and was supported by the National Science Foundation under duet songs of Sulawesi tarsiers: evidence for new cryptic
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ogy, Indonesian Institute of Sciences, Bogor, Indonesia. Revised: 7 July 2019
Shekelle M., C. Groves, S. Gursky, I. Neri-Arboleda and A Published: 28 December 2019
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Schulze and H. Fitch-Snyder (eds.), pp.71–84. Research In another sad twist to this work, on 12 April 2019, as
Center for Biology, Indonesian Institute of Sciences, we were in the final editing of this manuscript, Eslie Arteban
Bogor, Indonesia. Tamalagi, known as Ecil, passed away suddenly at the age
Shekelle, M., R. Meier, I. Wahyu, Wirdateti and N. Ting. 2010. of 47. Ecil was a field assistant for virtually every tarsier
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J. E. Janecka, C. A. Fisher and W. J. Murphy. 2012. Mac- this work to his memory. Photo from Ecil’s Facebook page.
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