Sample size measured in this study.
\r\n\tIncreasingly, governments and development institutions are recognizing the importance of addressing social exclusion for sustainable development. As such, the book will examine the role of government and the contribution of international development partners in the protection and support of marginalized groups and communities. Additionally, the role, responsibility, and response of academia as a socially accountable partner will form part of the discourse.
\r\n\t
Mainland and island are two unique bodies of landmasses that hold not only the obvious different area dimension but also a timeline that portrayed dynamic changes on their geographical and ecological features. Southeast Asia that comprises mainland and the patches of island is a home for the primate species diversity with high rate of endemism and provinciality [1]. Since the emergence of primates in the region during Quaternary to recent, 13 genera have been taxonomically recognized:
Lying over a wide range of latitude and various sizes of islands, the Southeast Asian region is frequently subjected for the studies of primate insularity that involved spatial factors (e.g., island size, latitude, and island-mainland distance) [3, 4, 5, 10] and temporal factors (e.g., isolation duration and geological chronology) [5]. Insularity on primates is an interesting phenomenon that invites many reports, linking to their ecomorphological complex (body size and body shape) [6] and biodiversity changes [11].
\nIn many ecological aspects, mainland environment differs from island environment. In addition, large-sized island provides different ecological scenarios from small-sized island. Certain duration of isolation on a relatively small island may lead to limited resources, fewer predators, and reduced interspecific competition [12]. Although it is not impacted universally, the combinative geographical effects on island size and island isolation can promote gigantism in smaller insular mammal species and dwarfism in larger mammal species. It is widely known as island rule (=Foster’s rule) [6, 13, 14, 15, 16, 17, 18]. With the wide span of latitudinal range, primates inhabiting the Sunda Shelf region are also assumed to follow Bergmann’s rule, by testing the effect of latitudinal position to body size [3, 4, 5]. This study aims to elicit the validity of ecogeographical rules affected body size and biodiversity changes of primates around Sunda Shelf throughout the geological chronology, since their appearance in Quaternary until recent.
\nAmong mammal taxa, the record of body size shift has not been found spectacular in all primate species [19]. Before the Quaternary, the primate fossil records adapted to island rule are found in Madagascar and Caribbean islands. Strepsirrhine primates found in Madagascar (e.g.,
Hominine taxa represented by the
Gained with the fact that three primate genera (
Map showing two different generalized bathymetric levels from 40 and 120 m throughout Sunda shelf. Closed dash lines present the group of islands with relatively equal range of sea depth.
Sex group | \n\n | \n\n | \n\n | \n||
---|---|---|---|---|---|
\n | \n\n | \n\n | \n\n | \n\n | \n|
31 | \n9 | \n60 | \n20 | \n38 | \n|
22 | \n12 | \n39 | \n8 | \n43 | \n
Sample size measured in this study.
All specimens are housed in Lee Kong Chian Natural History Museum and museum Zoologicum Bogoriense Indonesia.
Abbreviation | \nDefinition | \n
---|---|
PRS | \nProsthion: anteroinferior point on projection of premaxilla between central incisors | \n
PRS2 | \nProsthion2: anteroinferiormost point on premaxilla, equivalent to prosthion but between central and lateral incisors | \n
PMS | \nThe point where premaxillary suture crosses alveolar margin | \n
MP3 | \nMesial P3: most mesial point on P3 alveolus, projected labially onto alveolar margin | \n
MM1 | \nMesial M1: contact points between P4 and M1, projected labially onto alveolar margin | \n
MM3 | \nMesial M3: contact point between M2 and M3, projected labially onto alveolar margin | \n
DM3 | \nDistal M3: posterior midpoint onto alveolar margin of M3 | \n
PMA | \nMost posterior point of maxillary alveolus on the maxilla palatine | \n
NSP | \nNasospinale: inferiormost midline point of piriform aperture | \n
WPA | \nPoint corresponding lo largest width of piriform aperture | \n
NPM | \nMeeting point of nasal and premaxilla on margin of piriform aperture | \n
RHI | \nRhinion: most anterior midline point on nasals | \n
PMN | \nPremaxillary maximum superior PMS where premaxillo-maxillary suture meets nasal bone or aperture | \n
NAS | \nNasion: midline point on fronto-nasal suture | \n
GLA | \nGlabella: most forward projecting midline point of frontals at the level of the supraorbital ridges | \n
BRG | \nBregma: junction of coronal and sagittal sutures, on sagittal crest if necessary | \n
INI | \nInion: most posterior point of cranium, when viewed in the Frankfurt horizontal, be it on sagittal/nuchal crest or not | \n
OPS | \nOpisthion: posterior most point of foramen magnum | \n
LOC | \nMost anterior point on the occipital condyle along the margin of the foramen magnum | \n
AOC | \nOccipital condyle along the margin of the foramen magnum between POC and AOC | \n
Abbreviation and definition used in this study [27].
Frontal (left) and lateral (right) views of the generalized M. fascicularis skull, showing 20 landmark positions used in the analysis. Number and position of landmark points are applied with the same procedure in all species measured.
The box and whisker plot diagrams (Figure 3) exhibit two distinction profiles between Hylobatidae and Cercopithecidae. Island populations of
Box and whisker diagram showing the variation of craniolateral centroid size (CS) among five non-human primate species in mainland and island group.
For the last 30 years, benefited by the advanced methodology of molecular biology, the expansion of studies on primates of Southeast Asia have resulted in the increased number of taxonomic diversification [28, 29, 30], which was previously mostly explained by the superficial character (e.g., pelage color, tail length, and behavior) on the living taxa [2, 5]. Mainland and large islands have been claimed to correspond to the higher taxonomic diversity than islands [31]. With the wide span of area, mainland and large islands have a great advantage to develop more topographic diversity, formed as geographic barriers (e.g., peak, valley, river), linking to high possibility to allopatric speciation [32].
\nPrincipal component analyses (PCA) on the craniolateral shape of the five species share similarities in the wider shape variance of all three insular species (Figure 4). The mixed category between large-sized island and small-sized island in this study (Table 3) may strongly correspond to the higher craniolateral morphology, by considering (i) each isolated small island with unique geographical-ecological condition and different degrees of isolation may contribute to the shape modification, furthermore to endemism [12]; (ii) large islands may lead to various shape modifications, generated by various topographic-diversity-derived habitat variations [32]. Reflecting the wide variance morphology on three insular genera of this study, insularity does not gain merely on taxonomic diversity; furthermore strong individual differentiation within population or intraspecific variation could also possibly generated.
\nPlots of principal component PC1–PC2 displaying the variance between mainland and island population among five species observed.
Genera | \nSpecies/subspecies | \nIsland | \nLatitude | \nIsland size (km2) | \nIsland size category [33] | \nmax. elevation (m) | \n
---|---|---|---|---|---|---|
CONTINENTAL ISLAND | \n||||||
Ponginae | \n\n | \nBorneo | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nSumatra (north) | \n2°–4°N | \n473,481 | \nLarge | \n3805 | \n
\n | \n | \nSumatra (north) | \n2°–4°N | \n473,481 | \nLarge | \n3805 | \n
Hylobatidae | \n\n | \nJava (west) | \n8°–10°N | \n128,300 | \nLarge | \n3676 | \n
\n | \n | \nBorneo (south) | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nBorneo (north) | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
Cercopithecinae | \n\n | \nKhram Yai | \n12.70°N | \n20,28 | \nSmall | \n219 | \n
\n | \n | \nCon Son | \n8.71°N | \n51,52 | \nSmall | \n560.8 | \n
\n | \n | \nRiau Islands | \n2.50°–3.13°N | \n106 | \nSmall | \n959 | \n
\n | \n | \nBawean | \n5.80°S | \n196,27 | \nSmall | \n655 | \n
\n | \n | \nKarimun Jawa | \n5.85°S | \n71,2 | \nSmall | \n506 | \n
Colobinae | \n\n | \nNatuna Besar | \n4°N | \n1720 | \nSmall | \n187 | \n
\n | \n | \nSumatra (north) | \n2°–4°N | \n473,481 | \nLarge | \n3805 | \n
\n | \n | \nBorneo | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nBorneo (north) | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nBorneo (northeast) | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nBorneo (east) | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
\n | \n | \nJava | \n8°–10°N | \n128,300 | \nLarge | \n3676 | \n
\n | \n | \nBorneo | \n8°N–2°S | \n743,330 | \nLarge | \n4095 | \n
Hylobatidae | \n\n | \nMentawai Islands | \n1.2°–3°S | \n268–4030 | \nSmall | \n384 | \n
Cercopithecinae | \n\n | \nSulawesi (southwest) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nSulawesi (southeast) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nSulawesi (central) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nSulawesi (northwest) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nSulawesi (north) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nSulawesi (northeast) | \n0.3°N–5.3°S | \n174,600 | \nLarge | \n3478 | \n
\n | \n | \nMentawai Islands | \n1.2°–3°S | \n268–4030 | \nSmall | \n384 | \n
\n | \n | \nMentawai Islands | \n1.2–3S | \n268–4030 | \nSmall | \n384 | \n
\n | \n | \nLittle Nicobar | \n7.32°N | \n140 | \nSmall | \n435 | \n
\n | \n | \nMaratua | \n2.25°N | \n22,8 | \nSmall | \n94.18 | \n
\n | \n | \nPalawan | \n9.70°N | \n14,650 | \nLarge | \n2086 | \n
\n | \n | \nLuzon | \n16.9°N | \n110,000 | \nLarge | \n2922 | \n
\n | \n | \nLasia | \n2.17°N | \n15,12 | \nSmall | \n69 | \n
\n | \n | \nSimeulue | \n2.65°N | \n2310 | \nSmall | \n567 | \n
Colobinae | \n\n | \nMentawai Islands | \n1.2–3°S | \n268–4030 | \nSmall | \n384 | \n
\n | \n | \nMentawai Islands | \n1.2–3°S | \n268–4030 | \nSmall | \n384 | \n
\n | \n | \nMentawai Islands | \n1.2–3°S | \n268–4030 | \nSmall | \n384 | \n
List of modern non-human primate species/subspecies native to islands with the latitudinal position.
The category of island refers to the indicator of small island category (<12,000 km2) [34].
The isolation process on an island may lead to enforce the possibility of extinction in certain species [30]. For example, in Java Island, with area span 138,000 km2, three primate species (
Genera | \nSpecimen | \nLocality | \nPleistocene | \nHolocene | \n||
---|---|---|---|---|---|---|
\n | \n | \n | Early | \nMiddle | \nLate | \n\n |
MAINLAND | \n||||||
Hominidae | \n\n | \nZhoukoudian Caves, China | \n\n | 0.6–0.4 | \n\n | \n |
\n | \n | \nZhoukoudian Caves, China | \n\n | 0.4–0.5 | \n\n | \n |
\n | \n | \nHad Pu Dai, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nTham Khuyen, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nLang Trang, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nMa U’Oi, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nThum Wiman Nakin, Thailand | \n\n | ● | \n\n | \n |
Ponginae | \n\n | \nGigantopithecus Cave, China | \n● | \n\n | \n | \n |
\n | \n | \nJianshi, China | \n● | \n\n | \n | \n |
\n | \n | \nBaikong, China | \n2.2 | \n\n | \n | \n |
\n | \n | \nJuyuan, China | \n1.8 | \n\n | \n | \n |
\n | \n | \nSanhe, China | \n1.2–1.6 | \n\n | \n | \n |
\n | \n | \nQueque, China | \n<0.7–1 | \n≤0.7–0.8 | \n\n | \n |
\n | \n | \nYangliang, China | \n● | \n\n | \n | \n |
\n | \n | \nHad Pu Dai, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nDaxin, China | \n\n | ● | \n\n | \n |
\n | \n | \nWuming, China | \n\n | ● | \n\n | \n |
\n | \n | \nBama, China | \n\n | ● | \n\n | \n |
\n | \n | \nTham Khuyen, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nTham Hai, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nHeijang, China | \n\n | ● | \n\n | \n |
\n | \n | \nShuangtan, China | \n\n | \n | ● | \n\n |
\n | \n | \nGigantopithecus Cave, China | \n● | \n\n | \n | \n |
\n | \n | \nBaikong, China | \n>2.2 | \n\n | \n | \n |
\n | \n | \nJuyuan, China | \n>1.8 | \n\n | \n | \n |
\n | \n | \nSanhe, China | \n1.2–1.6 | \n\n | \n | \n |
\n | \n | \nQueque, China | \n<0.7–1 | \n≤0.7–0.8 | \n\n | \n |
\n | \n | \nYangliang, China | \n● | \n\n | \n | \n |
\n | \n | \nHad Pu Dai, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nTham Khuyen, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nThum Wiman Nakin, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nDaxin, China | \n\n | ● | \n\n | \n |
\n | \n | \nHoshantung, China | \n\n | ● | \n\n | \n |
\n | \n | \nKoloshan, China | \n\n | ● | \n\n | \n |
\n | \n | \nBama, China | \n\n | ● | \n\n | \n |
\n | \n | \nTam Hang, Laos | \n\n | ● | \n\n | \n |
\n | \n | \nTham Khuyen, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nTham Hai, Vietnam | \n\n | ● | \n\n | \n |
\n | \n | \nPhnom Loang, Cambodia | \n\n | ● | \n\n | \n |
\n | \n | \nThum Wiman Nakin, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nKao Pah Nam | \n\n | ● | \n\n | \n |
\n | \n | \nThum Wiman Nakin, Thailand | \n\n | ● | \n\n | \n |
\n | \n | \nHei, China | \n\n | 0.3–0.38 | \n\n | \n |
\n | \n | \nHeijang, China | \n\n | ● | \n\n | \n |
\n | \n | \nTongzi, China | \n\n | \n | ● | \n\n |
\n | \n | \nKeo Leng, Vietnam | \n\n | \n | ● | \n\n |
\n | \n | \nHang Hum II, Vietnam | \n\n | \n | ● | \n\n |
\n | \n | \nShuangtan, China | \n\n | \n | ● | \n\n |
\n | \n | \nYixiantian, China | \n\n | \n | ● | \n\n |
\n | \n | \nGonglishan, China | \n\n | \n | ● | \n\n |
\n | \n | \nZhiren, China | \n\n | \n | ● | \n\n |
\n | \n | \nNongbashankou, China | \n\n | \n | ● | \n\n |
\n | \n | \nBaxian, China | \n\n | \n | ● | \n\n |
\n | \n | \nLoushan, China | \n\n | \n | ● | \n\n |
Hylobatidae | \n\n | \nBaikong, China | \n2.2 | \n\n | \n | \n |
\n | \n | \nJuyuan, China | \n1.8 | \n\n | \n | \n |
\n | \n | \nSanhe, China | \n1.2–1.6 | \n\n | \n | \n |
\n | \n | \nQueque, China | \n0.7–1 | \n≤0.7–0.8 | \n\n | \n |
\n | \n | \nHei, China | \n\n | 0.3–0.38 | \n\n | \n |
\n | \n | \nHeijang, China | \n\n | 0.4–0.32 | \n\n | \n |
\n | \n | \nYenchinkou, China, China | \n● | \n● | \n● | \n\n |
\n | \n | \nSzechwan, China | \n● | \n● | \n● | \n\n |
\n | \n | \nNiah Cave, Borneo, China | \n\n | \n | ● | \n\n |
\n | \n | \nShuangtan, China | \n\n | \n | ● | \n\n |
\n | \n | \nYixiantian, China | \n\n | \n | 0.1 | \n\n |
\n | \n | \nGonglishan, China | \n\n | \n | ● | \n\n |
\n | \n | \nZhiren, China | \n\n | \n | 0.11 | \n\n |
\n | \n | \nBaxian, China | \n\n | \n | ● | \n\n |
\n | \n | \nLoushan, China | \n\n | \n | \n | ● | \n
Cercopithecinae | \n\n | \nBaikong, China | \n2.2 | \n\n | \n | \n |
\n | \n | \nJuyuan, China | \n1.8 | \n\n | \n | \n |
\n | \n | \nSanhe, China | \n1.2–1.6 | \n\n | \n | \n |
\n | \n | \nQueque, China | \n<0.7–1 | \n≤0.7–0.8 | \n\n | \n |
\n | \n | \nYangliang, China | \n● | \n\n | \n | \n |
\n | \n | \nHei, China | \n\n | 0.3–0.38 | \n\n | \n |
\n | \n | \nHeijang, China | \n\n | 0.4–0.32 | \n\n | \n |
\n | \n | \nShuangtan, China | \n\n | \n | ● | \n\n |
\n | \n | \nYixiantian, China | \n\n | \n | 0.1 | \n\n |
\n | \n | \nGonglishan, China | \n\n | \n | ● | \n\n |
\n | \n | \nZhiren, China | \n\n | \n | 0.11 | \n\n |
\n | \n | \nNongbashankou, China | \n\n | \n | ● | \n\n |
\n | \n | \nBaxian, China | \n\n | \n | ● | \n\n |
\n | \n | \nLoushan, China | \n\n | \n | \n | ● | \n
Colobinae | \n\n | \nBaikong, China | \n2.2 | \n\n | \n | \n |
\n | \n | \nJuyuan, China | \n1.8 | \n\n | \n | \n |
\n | \n | \nSanhe, China | \n1.2–1.6 | \n\n | \n | \n |
\n | \n | \nQueque, China | \n<0.7–1 | \n≤0.7–0.8 | \n\n | \n |
\n | \n | \nHei, China | \n\n | 0.3–0.38 | \n\n | \n |
\n | \n | \nHeijang, China | \n\n | 0.4–0.32 | \n\n | \n |
\n | \n | \nShuangtan, China | \n\n | \n | ● | \n\n |
\n | \n | \nYixiantian, China | \n\n | \n | 0.1 | \n\n |
\n | \n | \nGonglishan, China | \n\n | \n | ● | \n\n |
\n | \n | \nZhiren, China | \n\n | \n | 0.11 | \n\n |
\n | \n | \nNongbashankou, China | \n\n | \n | ● | \n\n |
\n | \n | \nBaxian, China | \n\n | \n | ● | \n\n |
\n | \n | \nLoushan, China | \n\n | \n | \n | ● | \n
CONTINENTAL ISLAND | \n||||||
Hominidae | \n\n | \nSangiran, Java | \n0.99–1.5 | \n\n | \n | \n |
\n | \n | \nSangiran, Java | \n0.78–1.3 | \n\n | \n | \n |
\n | \n | \nSangiran, Java | \n1.2–0.98 | \n\n | \n | \n |
\n | \n | \nSangiran, Java | \n1.2–0.99 | \n\n | \n | \n |
\n | \n | \nSambungmacan, Java | \n≤0.78 | \n\n | \n | \n |
\n | \n | \nNgawi, Java | \n\n | ● | \n● | \n\n |
\n | \n | \nNgandong, Java | \n\n | ● | \n0.05–0.032 or 0.1 | \n\n |
\n | \n | \nPunung, Java | \n\n | \n | 0.0118 | \n● | \n
Pongidae | \n\n | \nSemedo, Java | \n? | \n? | \n\n | \n |
\n | \n | \nPunung, Java | \n\n | \n | 0.125 | \n\n |
\n | \n | \nLida Ayer, Sumatra | \n\n | \n | \n | ● | \n
Hylobatidae | \nHylobatidae [41] | \nTrinil, Java | \n\n | ● | \n● | \n\n |
\n | \n | \nPunung, Java | \n\n | \n | 0.0118 | \n● | \n
\n | \n | \nLida Ayer, Sumatra | \n\n | \n | \n | ● | \n
\n | \n | \nNiah Cave, Borneo | \n\n | \n | 0.04 | \n\n |
Cercopithecinae | \n\n | \nSangiran, Java | \n\n | \n | \n | \n |
\n | \n | \nPunung, Java | \n\n | \n | 0.0118 | \n0.008 | \n
\n | \n | \nSangiran, Java | \n1 | \n\n | \n | \n |
\n | \n | \nSangiran, Java | \n1 | \n\n | \n | \n |
\n | \n | \nCallao Cave, Luzon | \n\n | \n | 0.065 | \n\n |
\n | \n | \nIlle Cave, Palawan | \n\n | \n | ● | \n● | \n
Colobinae | \n\n | \nSangiran, Java | \n\n | ● | \n\n | \n |
\n | \n | \nPunung, Java | \n\n | \n | \n | 0.01 | \n
\n | \n | \nSangiran, Java | \n1.9 | \n\n | \n | \n |
OCEANIC ISLAND | \n||||||
Hominidae | \n\n | \nMata Menge, Flores | \n\n | 0.7 | \n\n | \n |
\n | \n | \nLiang Bua, Flores | \n\n | ● | \n0.06–0.1 | \n\n |
\n | \n | \nCallao Cave, Luzon | \n\n | \n | 0.06 | \n\n |
Cercopithecidae | \n\n | \nCallao Cave, Luzon | \n\n | \n | 0.065 | \n\n |
\n | \n | \nIlle Cave, Palawan | \n\n | \n | ● | \n● | \n
\n | \n | \nTimor Island | \n\n | \n | \n | 0.007 | \n
List of fossil/subfossils of primate species/subspecies discovered in archeological sites throughout Southeast Asia.
Southeast Asia with wide span of latitude ranging from 6°N to 14°S is split by the equator line, demanding at least two comprehensive separations that require thermoregulation connection from the equator to southern and northern latitudes. Mammals of mainland Southeast Asia have been subjected to describe body size variation following thermoregulation effect, widely termed as Bergmann’s rule [6]. Concluding that Bergmann’s rule may occur within a species, it predicts that population in warmer climates (commonly referred to lower latitudes) have smaller mean body size than conspecifics in colder climates (generally marked with higher latitude) [6]. Published accounts applying this ecogeographical rule on non-human primates has been intensively investigated in the widely distributed species in Southeast Asia:
Interestingly, anti-Bergmann’s rule appears north side of Kra Isthmus (the narrowest area differing Indochinese mainland and Malay Peninsula at 12.2°N) [4, 5]. Explanatory cause for this inversed Bergmann’s rule has not been uncovered. In response to this matter,
Although serious attempts to test Bergmann’s rule on insular non-human primates have increased, the result of the statistical analysis on the cranial size of southern pig-tailed macaque (
In the context of conservative classification on island area, primate insularity has been investigated into categorization of area size, e.g., small and large island, which was directly calculated by metric size of island [31]. This ecogeographical rule implemented exclusively on island, commonly known as Foster’s rule, proposes that population of large-bodied mammals on island tend to have a smaller mean body size than mainland population (dwarfism), while small-bodied mammals become larger (gigantism) [6]. One suggested that, in the scope of insularity on Southeast Asian mammals, the small island criterion is defined by the island size <12.000 km2 [34] (Table 3). Without providing the specific primate species group, one suggested that primates follow island rule [19]. However, a study tested in body length of
The most spectacular evidences of dwarfism on extinct Homininae taxa are
Among gibbons, diminutive body size has been presented by
Researchers have long endeavored to uncover the Foster’s rule in Southeast Asian archipelago [4, 5, 10], but most outcomes show no statistically significant results [11]. On exclusively
Mainland Southeast Asia contains the high variation of non-human primate species. Recent molecular biological studies revealed critical systematics of non-human primates (i.e.,
In Java, a chain of 38 mountains forming east–west spine with various slopes, illustrated by jagged highlands by alternating peaks and valleys, leads to classes of topographic diversity [35]. This phenomenon led the geographically separated populations to undergo allopatric speciation. According to the modern Javanese mammal fauna, the low topographic diversity in East Java resulted in less variation in endemic mammals than in the West and Central Java. This topographic profile is supported by the presence of two endemic non-human primate species/subspecies strictly occupying western Java forests;
Conversely, a higher endemic mammal species diversity was more visible in East Java during the Middle Pleistocene, in the stage of
With the numerous
Time by duration and particular period falls to the temporal scope of inhabitation of certain population on island is pronounced to impact body size evolution [12]. Higher duration of island isolation increases the chance for ecological release to influence functional characters (e.g., diet, locomotion, and bauplan) among species. The report on paleoinsular mammals has claimed that body size shift on island mammal species occurred when residence time reached more than 10,000 years [12]. While the evidences are prominently strong on terrestrial herbivores, including terrestrial primates (e.g.,
Typically expressed by the estimated dispersal chronology in Southeast Asian Archipelago, duration of island isolation shows the function of maximum sea depth separating island from mainland or neighboring large island, mainly in small-sized island. Some oceanic islands in the region (Simeulue, Lasia, Siberut, Sipora, North Pagai, South Pagai) remarked with bathymetric barrier more than 120 m (Figure 1) display clear effect of isolation than the shallow-water fringing island over Sunda Shelf. The shallow depth of Sunda Shelf sea floor (0–40 m) allows the emergence of exposed dry land that permits colonization, reversed colonization, or recolonization of the island which most commonly occur during the sea level drop during the Last Glacial Maximum (LGM), which reduces the optimum genetic isolation.
\nOn the level of subspecies, the long duration of island isolation appears to indicate the development of new intraspecific features in
According to the previous paleontological works on mammal evolution of Southeast Asia, there is no fossil evidence of primates before ca. 0.9 Ma in Java Island. The first colonization of primates to Java is estimated to occur at the end of Early Pleistocene, when Sunda Shelf fully emerged and then periodically entered Java via Siva-Malayan corridor route during Middle Pleistocene [33]. Along with the balanced mammal association, including
To date, there is no chronological and geographical comparative study demonstrating body size of non-human primates between fossils and recent on Java Island. It rather revealed the similarities on morphological characters in accordance with the attempt in determining species. So, it was difficult to answer whether Middle Pleistocene non-human primates of Java are the continuously highly adapted species until recent or the extinct species that disappeared in the Middle Pleistocene like other mammals (including
Late Pleistocene displays the rise of tropical rain forest non-human primates (
With limited connection to the diverse mainland fauna, isolated island promotes the poor taxonomic diversity and the imbalanced rate between herbivores and carnivores. Small island has been claimed to reduce the sympatric speciation than large island [31]. This condition drove a disharmonic inter- and intraspecific variation [12]. For instance, in severe ecological condition when food resources are limited in long duration, the large-bodied species tends to expand their territory where small-bodied species fails to compete and being enforced to undergo stronger dietary adaptation. This response to ecological condition led to a radiation into different size classes and morphotypes, which arrives to appear in the form such as anatomical modification (e.g., dental pattern, size, and shape of limb bone) causing genetic radiation [12].
\nIn most case, this disharmonic taxonomic diversity condition dropped the survivability. The heavily impoverished condition leads to some species to extinction, for example, in all Late Pliocene-Early Pleistocene (
The vegetation type of an area derives from mean temperature caused from latitudinal position, geographical topography, seasonality by monsoon, and geological sediments. During Quaternary, the fluctuating temperature prominently contributes to habitat changes. The ecological shift from tropical rainforest to more open environment in Early-Mid Holocene resulted in biodiversity loss in non-human primates; for example, it is shown by the disappearance of
\n
Prefigured by many geographic properties, bathymetric barrier presents to appear as the strongest casual effect in enforcing island isolation in Southeast Asian Archipelago, expressed by the high degree of endemism in level of species in oceanic islands (i.e.,
The duration of island isolation widens to promote the evolutionary results that yield the island ecological mechanism becoming intensified. The higher time cost on ecological factors such as selective pressures and predator avoidance could escalate the chance for anatomical feature to be modified. Although it is hard to know the absolute duration of island isolation, the relative isolation can be seen from the present bathymetry showing the predicted terminal time for body of water to cover the maximum depth that stop the connection from mainland to surrounding islands. Constituted by this concept, oceanic islands with high bathymetric barrier will definitely prolong the disconnection signal from mainland than continental islands.
When we control geographical and chronological isolation factors, the two main island ecosystem factors, faunal association and vegetation type, strongly contribute to the change of body size and shape, resulting in a higher island effect. Patterns impacted by this ecosystem factors are not the same in all islands. The imbalanced condition on fauna between the number of herbivores and carnivores and less interspecific faunal diversity could lead to the body size shift and anatomical modification. On primates, oceanic islands located near the equator covered with the densely tropical rain forest gave less likely island effect (e.g., Mentawai Island and Simeulue Island) than in oceanic island with drier and more open environment where resource is less abundantly available (e.g., Flores Island).
Latitudinal factor is clear to be seen in the mainland. While each island holds unique geographical properties directing to isolation (e.g., bathymetric barrier and island size), most Southeast Asian islands that are located around the equator with tropical weather resulting in major rain forest cover and short latitudinal range rather rise to contribute to more diverse body size and body shape longitudinally. Thus, Bergmann’s rule is seemingly irrelevant to be evaluated in such condition.
The primates of Sunda Shelf occupying the great number of islands scattered in large scale area did not perform any pattern in regard to correlation between body size and island size. Potential causal relation to island size is more manifested in the increasing taxonomic diversity. Large-sized islands throughout Sunda Shelf hold higher diversity in anatomical variation than in small-sized island. It is supposedly due to the combination of possible isolation-derived process by geographic or ecological barrier and the resiliency of relict species along many stages of period. This circumstance is conceivably reassured from the Quaternary through recent, for example, the high diversity of calvarium morphology seen in
Endemism featured on non-human primates in continental islands of Sunda Shelf mostly direct to the resilience of relict groups occupying the island, not necessarily in response to a long-term island isolation process. In the level of species, this premise is endorsed by the existence of a single taxon occupying large islands (e.g.,
The author is indebted to Maharadatun Kamsi, the curator of vertebrate collection of the Museum Zoologicum Bogoriense, Indonesian Academy of Sciences (LIPI), as well as Kelvin Lim, the collection manager of Lee Kong Chian of Natural History Museum, National University of Singapore. I also wish to thank Dr. Tsuyoshi Ito for helpful suggestion in biostatistics work and data analysis and Thomas Priyo Ertanto, who drew the map in this publication. This research was financially supported by The Asahi Glass Foundation to HI (ID: 2017-F-06). We further thank Mao Asami, Indra Sutisna, Akhmad Herdiyanto, Nanang Supriatna, and Kurnianingsih for support during sample collection and laboratory work.
\nThe authors declare no conflict of interest.
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