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Two apocalyptic challenges for society: Mitigating the effects of anthropogenic biodiversity loss & climate change . . .

•Habitat Conservation •In , and more broadly, Indonesia and Southeast Asia (the tropics?) mitigating biodiversity loss and global climate change is largely about preserving forests, or rather, stemming habitat destruction. . . . with one of the world’s most unusual

: like “Gremlins”, tarsiers have (at least) two very dispositons •Cute little with ‘neotenized’ facial features •Horrifying alpha predator of the forest understory What are tarsiers and why study them? What Are Tarsiers? What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia —Primates Suborder— Superfamily—Tarsioidea Family—Tarsiidae Genus— What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder— Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini? Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini? Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Prosimii? Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Prosimii Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes? Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Tarsiiformes Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates What Are Tarsiers?

primates

“using likelihood ratio tests [one topology] cannot be shown to be significantly preferred to alternative topologies” (Yoder, 2003) based on results from the largest study of primate DNA sequence at that time (Murphy et al., 2001) What Are Tarsiers?

primates What Are Tarsiers? A textbook case of “long branch attraction” primates

adapted from Yoder (2003) What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini* Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius

*probably, but less confidently resolved with molecular systematics than are other major clades. What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius What Are Tarsiers?

primates

Kingdom—Animalia Phylum—Chordata Class—Mammalia Order—Primates Suborder—Haplorhini Superfamily—Tarsioidea Family—Tarsiidae Genus—Tarsius ? What Are Tarsiers? One genus or three genera?

Shekelle 2008 Groves & Shekelle 2010 Variation among Genera

Western Philippine Tarsier Eastern Tarsier Cephalapachus Carlito Tarsius Least social Tolerates sociality in captivity Social in wild Least vocal More vocal Highly vocal, “duets” 80 chromosoms 80 chromosome 46 chromosomes What Are Tarsiers?

primates What Are Tarsiers? primates What Are Tarsiers? primates primates small-bodied What Are Tarsiers? primates primates small-bodied

Smallest primate outside of Madagascar

Adult Tarsier Body Weight Range: 59-150g Means: 59-125g 33g infant What Are Tarsiers? primates primates small-bodied What Are Tarsiers? primates small-bodied What Are Tarsiers? primates small-bodied obligate faunivores What Are Tarsiers? primates small-bodied obligate faunivores Kay's threshold ≈ 500 g approximate body weight that separates primarily insectivorous from non- insectivorous primates

Tarsiers are obligate faunivores specializing in large-bodied arthropods What Are Tarsiers? primates small-bodied obligate faunivores

Tarsiers diet is unique among primates 100% live-caught prey 0% plant matter What Are Tarsiers? primates small-bodied obligate faunivores What Are Tarsiers? primates small-bodied obligate faunivores What Are Tarsiers? primates small-bodied obligate faunivores nocturnal What Are Tarsiers? primates small-bodied obligate faunivores nocturnal Tarsier Anatomy

eyeball: 2030-2140 mm3 brain: 2400-4500 mm3 What Are Tarsiers? primates small-bodied obligate faunivores nocturnal

Largest relative eyesize among

Each eyeball is approximately equal in size its brain What Are Tarsiers? primates small-bodied obligate faunivores nocturnal What Are Tarsiers? primates small-bodied obligate faunivores nocturnal What Are Tarsiers? primates small-bodied obligate faunivores nocturnal What Are Tarsiers? primates small-bodied obligate faunivores nocturnal vertical clingers and leapers What Are Tarsiers? primates small-bodied obligate faunivores nocturnal vertical clingers and leapers = VCL What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL What Are Tarsiers? primates small-bodied vertical clinging obligate faunivores nocturnal VCL What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied leaping obligate faunivores nocturnal VCL

Low intermembral index (forelimb/hindlimb) correlates with VCL Tarsiers have the lowest IM index among primates What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual newborn weight ≈ 20-33% mother’s weight linear measurements ≈ 50% mother’s largest neonatal weight among primates What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual newborn weight ≈ 20-33% mother’s weight linear measurements ≈ 50% mother’s largest neonatal weight among primates and yet infants are transported by mouth! What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual births are singletons but mother has 2-3 pairs of mammae What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual gestation length ≈ 6 months longer than in Macaca What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual head can rotate 180º in either direction, giving 360º mobility no other can do this What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual Western and Philippine tarsiers have 80 chromosomes The highest among mammals What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual Western and Philippine tarsiers have 80 chromosomes The highest among mammals

and yet Eastern Tarsiers have only 46! What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual First primate documented to use ultrasound and to hear ultrasound to >80 kHz What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied The world’s tarsier-ologists don’t meet in a phone booth, but we could What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied personal observation What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied personal observation What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied tarsiers are cool! What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied tarsiers are cool! What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual and they’re cool! relatively unstudied What Are Tarsiers? primates small-bodied obligate faunivores nocturnal VCL unusual relatively unstudied Tarsiers Are Very Old essentially a third, nearly independent lineage of primates

Phylogram of Tarsier Evolutionary History (based on nDNA, Meireles et al. 2003) Primates are found throughout the tropics, but extant tarsiers are found only in insular southeast Asia and Distribution of Tarsiers Southeast Asia: Last Glacial Maximum Bird et al. (2005) Tarsier Distribution & Last Glacial Maximum Bird et al. (2005) Tarsier Distribution & Last Glacial Maximum Bird et al. (2005) Tarsier Distribution & Last Glacial Maximum Bird et al. (2005) Paleohabitat & Last Glacial Maximum Bird et al. (2005) Paleohabitat & Last Glacial Maximum Bird et al. (2005) A little bit newer, but nevertheless similar . . .

Canon et al. (2009)

105 Three genera are each endemic to a distinct biogeographic region.

What about Sulawesi?

106 Hybrid Biogeographic Hypothesis for Sulawesi Based on Biological and Geological Data (Shekelle & Leksono, 2004)

l Tarsiers are immigrants to Sulawesi, with a distribution first shaped by dispersal throughout the proto-Sulawesi archipelago, and subsequently re-shaped by Pleistocene vicariance events.

Pre-Pleistocene Dispersal (1.8-20 mya): data of Hall (2001) principally concerns Miocene and Pliocene tectonic activity of the proto-Sulawesi archipelago.

Pleistocene Vicariance (0-1.8 mya): data of Evans et al. (2003) principally concerns Pleistocene and post-Pleistocene evolution of . Biogeographic and conservation conclusions:

1. How many taxa of tarsiers? within Niemitz’s (1984) T. spectrum spectrum alone are 16 known acoustic forms. 2. Why are there so many? dispersal throughout the proto-Sulawesi archipelago during the Miocene and Pliocene, followed by range fragmentation in the Pleistocene. 3. What are the implications of this? a taxonomic explosion of tarsiers indicates a conservation crisis for Sulawesi, where biodiversity may be underestimated by as much as an order of magnitude. Conservation •Two main objectives •Identifying priorities •Implementation

 The first step is a challenging, but rewarding scientific endeavor.  The latter is a frustrating and almost entirely futile expenditure of effort in places I have worked.

•Sulawesi is subdivided into 16+ subregions, cryptic species

•Biodiversity underestimated by an order of magnitude

Results from the Indonesian CAMP, 2001 Siau Island Tarsier, Tarsius tumpara. New Species, Critically Endangered “Top 25 Most Endangered Primates” 2006-2008, 2008-2010, 2010-2012

Shekelle and Salim, 2009 (Oryx) Shekelle et al. (2008) Tarsius tumpara: A New Tarsier Species from Siau Island, North Sulawesi

Adapted from . . . and the locals eat them! What else?

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