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Home , Bose Basin, Cleaver (Stone Age tool), Duboisia santeng, Flores, Hexaprotodon, Hominization

Evolutionary 24:185–213 (2015)

ARTICLE

Early Hominin in Island Southeast

ROY LARICK AND L. CIOCHON

Island covers ’s tropical expanse of continental shelf Solo Basin, has since produced more and active subduction zones. Cutting between island landmasses, ’s than 80 erectus cranial and Line separates Sunda and the Eastern Island Arc (the Arc) into distinct tectonic dental . The and Tri- and faunal provinces. West of the line, on Sunda, Island yields many fossils nil fossils have thick cranial vaults of . East of the line, on the Arc, Island provides one skele- and cranial capacities of 840 to 1,059 ton and isolated remains of . Luzon Island in the cc.3 A much later set of Solo Basin has another hominin. preserves early hominin archeology. This Homo erectus fossils, from Ngandong insular divergence sets up a unique regional context for early hominin dispersal, and related sites, have cranial isolation, and . The evidence is reviewed across three cli- capacities reaching 1,250 cc. mate periods. Patterns are discussed in relation to the pulse of global -level In 2003, at , on Flores, shifts, as as regional geo-tectonics, catastrophes, dispersal, and east of Wallace’s Line, Homo flore- paleogenomics. Several patterns imply evolutionary processes typical of oceanic siensis was defined on the basis of islands. Early hominins apparently responded to changing island conditions for one nearly complete skeleton and a million-and-a-half , likely becoming extinct during the period in which fragmentary remains of several indi- Homo sapiens colonized the . viduals.4,5 Compared to Sunda Homo erectus, the fossils from Liang Bua have a small cranial capacity In 1859, Alfred Russell Wallace Sunda and the Arc. Wallace’s Line 3 identified two faunal provinces represents a series of sea-channel (417 cm ). In relation to most Pleis- within Island Southeast Asia (ISEA), barriers to the dispersal of large tocene early hominins, the Liang between them. The provin- Bua skeleton is short (1.06 m) and ces are based primarily on the conti- has primitive wrists and large feet, 6 Roy Larick owns Bluestone Heights, an nental origin of large terrestrial as well as a late age (60 ka). Other environmental education and consulting mammals (Fig. 1, Box 1).1 West of members of the Liang Bua vertebrate firm in Cleveland, OH. He is a fauna share similar insular charac- archeologist who has done field work Wallace’s Line, mammalian 6 covering , , and Asia. He is a have Eurasian origins; east of the ters (Box 2). When compared with founding member of the Iowa- line, “Wallacean” mammals and related species on other ISEA land- Java Project. Email: [email protected] other vertebrates show a mixture of masses, the Liang Bua fauna show Russell Ciochon is Professor of Anthro- Eurasian and Australian origins. pology and Director of Museum Studies signs of isolation on Flores for a sig- at the University of Iowa, Iowa City. He is Similar, if less provincial differentia- nificant part of the Pleistocene.7 a paleoanthropologist with field work tion can be observed for some spe- In 2007, fragmentary fossils were projects throughout Asia. He also is a cies of fish, insects, and birds. The founding member of the Iowa-Bandung collected from Callao on the Java Project. Larick and Ciochon have line should pose a factor for ISEA island of Luzon, in the Philippines.8 collaborated on paleoanthropological early hominin dispersal, isolation, projects in , , and Indone- Because of its small dimensions and sia. Email: [email protected] and extinction. gracile morphology, a complete meta- tarsal resembles those in small- EARLY HOMININ bodied early hominins, including and Homo floresiensis. Key words: Sunda; ; island rule; BIOGEOGRAPHY IN ISLAND Homo erectus; Homo floresiensis; Marine Isotope Its date, 66.7 ka, is close to that of Stages (MIS); Java; Flores; Luzon; Sulawesi; SOUTHEAST ASIA Liang Bua. In 2014, an upper In 1891, Eugene Dubois’ named row of archaic character was Pithecanthropus erectus (now Homo recovered, as were additional small 9 VC 2015 Wiley Periodicals, Inc. erectus) based on a calotte and limb bones. The new Callao finds DOI: 10.1002/evan.21460 suggest a possible third group to the Published online in Wiley Online Library found at , in the Solo Basin of (wileyonlinelibrary.com). eastern Java.2 Sangiran, also in the ISEA Pleistocene hominin population 186 Larick and Ciochon ARTICLE

Figure 1. Southeast Asia: sedimentary basins and catastrophe origin points of relevance to early hominin . Terrestrial sur- face extends to 100 m below current sea level. We followed ’s modification of Wallace’s Line208 as illustrated in Cooper and Stringer204. Boxes show areas detailed in Figure 3. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] and a second colonization east of numerous and widespread. Archeol- three sections corresponding to these Wallace’s Line. ogy therefore fleshes out early homi- events. A period framework makes Early hominins are now definitively nin biogeography. The historical for some repetition in presenting situated on Sunda and at two wide- trend of archeological research par- sites with long stratigraphic sequen- spread points on the Arc. Both Arc allels that of fossils. Recent research ces. Nevertheless, parsing regional fossil hominins have the small size provides a richer comparative base developments by period helps to and specialized skeletal traits seen in (Box 3). Stone are incorporated identify the effects of climate change insular evolutionary contexts and, to into the review when an excavated and several regional environmental a lesser extent, in the earliest Homo stratigraphic sequence contains catastrophes (Box 5). Figure 2 erectus from Eurasia (). The fauna and artifacts within a recog- presents the overall scheme. Each Arc hominins diverge greatly from all nized geological level and when a period section has a synoptic table known Sunda forms. After more than sequence-long sampling strategy has for the relevant events, sites, lithics, a century of accumulating fragmen- consistently produced Pleistocene and fauna. Table 1, for example, tary evidence, ISEA early hominin ages. presents the earlier Pleistocene biogeography is now a significant Using dated Marine Isotope Stages benchmarks. research topic. Here we review rele- (MIS) global events and a long chro- vant evidence from Sunda and the nology for regional occurrences, we OLDUVAI SUBCHRON, OR Arc within a framework of Pleisto- can begin to comprehend ISEA bio- EARLIER PLEISTOCENE cene climate change, our primary geographic events in their global goal being to evaluate potential roles contexts (Fig. 2, Box 4). Three global During the earlier Pleistocene for well-known insular evolutionary events can be tied to crucial regional (2.6-0.9 Ma), a 41-kyr orbital cycle processes in ISEA early hominin evo- biogeographic transitions: the Oldu- drove global climate. This periodic- lution (Box 2). vai paleomagnetic event, the Late ity, based on ’s orbital obliquity, While ISEA early hominin fossils Revolution (aka exemplifies one of three such orbital are few and spatially isolated, Mid-Pleistocene Revolution), and the patterns, known as Milankovitch Pleistocene-age stone artifacts are Mid-Brunhes Event. This review has cycles.10 During this time, glacial- ARTICLE Early Hominin Biogeography in Island Southeast Asia 187

Figure 2. ISEA Pleistocene and site correlation. Even-numbered MIS represent cooler phases (Northern hemispheric glacials); odd-numbered stages represent warmer phases (Northern hemispheric interglacials). MIS ages are drawn from Bowen and Sikes,272 Berger et al.,109 and Willoughby.273 Ages before MIS 19 are approximate. Since early hominin fossils are known only from sites in Java and Liang Bua, Flores, artifacts serve as the evidence of early hominins at other sites. FAD 5 first appearance datum; LAD 5 last appearance datum. Except where noted, site or event placement marks earliest occurrence. Blue lines indicate first-occurrence pre- sumed hominin continuous occupation in each regional site sequence unless otherwise noted. Sites with named formations and/or geologic designations: Lower Lahar, Lower Lahar Unit (Sangiran Formation); Sangiran, Sangiran Formation; Bapang, Bapang Forma- tion; Ngebung, Bapang Formation; Song Terus, karst cave; Ngandong, 20 m of ; Wolo Sege, Tangi Talo, , Boa Lesa, and Kobatuwa, Ola Kile Formation; Liang Bua, karst cave; S Enrile Q, Awidon Mesa Formation; Cagayan, Ilagan and Awidon Mesa Formations; Arubo, Sierra Madre foothills; Callao, karst cave; Cabenge, Walanae Basin fill. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.]

interglacial cycles were relatively the size and distribution of Sunda mammals arrived on an emergent short and had low temperature landmasses. Sunda.19–21 amplitude.11,12 Toward the middle of glacial ice advanced significantly this phase, the Olduvai Subchron between about 1.8 and 1.74 Ma (MIS marks a brief geomagnetic pole 62, 60, and 58), about 1.56 Ma (MIS Sunda reversal event, 1.98-1.79 Ma.13 Soon 52), between 1.24 and 1.1 Ma (MIS Sangiran, Java () after Olduvai, increases in 36, 34, and 30), and about 0.9 Ma intensity were recorded in the terres- (MIS 22).16 During these stages, sea- Much of the Solo Basin lies at trial contexts of the level lowstands opened emergent about 78 S latitude and sits above the () and Heqin (China) landmasses throughout ISEA.17,18 It Indonesian subduction zone (Fig. basins.14,15 Glacial-interglacial was, apparently, just after the Oldu- 3A). Mammalian fossils are pre- cycling had an increased effect on vai event that Pleistocene Eurasian served within a 500,000- 188 Larick and Ciochon ARTICLE

Box 1. Geomorphologic Setting

ISEA encompasses Sunda and Luzon in the north, southward create Wallace’s line are the - the Arc. ISEA is the result of the through Sulawesi and the Maluku oro Strait, Sulu and Celebes , Indian and Pacific oceanic plates Islands and, in the extreme , to and the Makassar and subducting under the Eurasian ter- Flores, , Timor, and related Straits. Trending east-west are the restrial plate. Regional topography small islands. Much of the Arc lies Flores and Banda Seas and, south ranges from broad coastal plain in , the geographic and eco- of Flores, the Sea. and continental shelf to volcanic logical transition zone between Regional tectonic events may islands, plateaus, and deep sea Sunda and Sahul ( and have been involved with specific trenches (Figs. 1 and 8). ).208 During the and short-lived dispersal pathways. Sunda: Much of the Indonesian and Pleistocene, especially during Sartono implicated small-scale Archipelago lies on the Sunda periods of glacio-eustatic sea-level uplift in relation to dispersal corri- shelf, a vast, now mostly sub- drawdown, large mammals, includ- dors along the Palawan, Sulu, San- merged southward extension of the ing , evidently penetrated gihe, and Selayar archipelagos (Fig. Eurasian continental plate.206,207 current marine barriers between 8).85 While this particular model is The continental shelf extends Sunda and Wallacea.209,210 Fossil now dated, small-scale tectonic southward from the present South- and archeological evidence indicates effects could be approached in new east Asia mainland toward Java that early hominins traversed cur- terms. Within the area covering the and the Indian (8.18 S). rent straits numerous times during Seas of Java, Banda, and Molucca Here, the term “Sunda shelf” is the Pleistocene. (Fig. 8), there are numerous plate- reserved for the continental projec- Eustasy, Tectonics, Volcanism: lets with active convergent and tion. while “Sunda” refers to the Glacio-eustatic sea level changes of divergent boundaries.212 Such pla- shelf’s island landmasses. The term 125 m are documented for the telets are subject to forming pop- “Sunda” updates the nineteenth- Pleistocene.211 Glacial eustasy up blocks.213 This process could century names of “” and repeatedly redistributed habitable have enhanced the formation of “Sunda Land.” Today, Sunda takes island landmasses and interisland lowstand land bridges. Regional in the current large landmasses of connections. Glacial period sea- tephra catastrophes could rear- the Malay Peninsula, , level lowstands maximized condi- range local habitats. The Sunda , and Java, as well as tions for regional dispersals. Inter- sedimentary record shows frequent smaller islands such as . glacial highstands separated early volcanic eruptions along plate mar- Eastern Island Arc: East of hominin groups and set up condi- gins after the Olduvai Subchron. Sunda, the sea bottom is complex tions for insular endemism. Several Truly large emissions could extend with trenches and ranges. The subduction zones have made for local landmasses seaward, provid- landmasses of this area constitute deep sea barriers of full Pleistocene ing larger lowland areas and, in the Arc, which stretches from duration. Trending north-south to some cases, small land bridges.214

sequence (1.53-0.98 Ma) of aggrad- mandibular fragment reflects terres- mudstones, and thin tuffs overlies the ing volcaniclastic sediments.19 Cur- trial conditions in the higher Lower Lahar Unit. These represent rently, more than 80 fossil fragments reaches.20 Freshwater mollusks indi- slow-moving streams draining nearby of Homo erectus are known from cate swamps or shallow lakes in the low volcanic highlands into shallow Sangiran (Fig. 4). The area has three lower reaches.22 Sangiran itself lakes. Such watercourses intermit- primary sedimentary deposits: the reflects near-shore marine environ- tently flooded lake margins and Lower Lahar Unit, the Sangiran For- ments transformed into estuarine marshes, transforming coastal lagoons mation, and the Bapang Formation. and marsh settings. Shortly there- into inland lakes. Occasional volcanic after, glacio-eustatic sea level regres- eruptions deposited thin blankets of Lower Lahar Unit. As early as 1.90 sions exposed lahar-infilled lagoons ash. Lake-edge and marsh environ- Ma, a nearby volcanic cone, pro- and near-shore environments to cre- ments supported sedges, , water- duced a massive lahar- debris ate terrestrial habitats.23 Since nei- tolerant grasses, and trees.24,25 The flow. Sangiran represents a relatively ther hominin fossils nor stone associated fauna comprised aquatic distal point of the flow, where the artifacts were incorporated, it is and semi-aquatic vertebrates such lahar entered marine conditions on assumed that the Lower Lahar Unit pygmy hippo (), croco- Sunda’s south coast. Incorporated predates the arrival of Homo erectus. dile (Crocodylus), and tortoise (Geo- fossils indicate that the lahar pushed chelone), as well as turtle and fish through numerous environments Sangiran Formation. A sequence of (fragmentary remains).21,26–28 Wet before arriving at Sangiran. A cervid dark-colored lacustrine siltstones, grasslands with scattered shrubs ARTICLE Early Hominin Biogeography in Island Southeast Asia 189

Figure 3. ISEA early hominin sites: A, central/eastern Java; B, central Flores; C, northern Luzon; D, southwest Sulawesi. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] occupied slightly higher landscapes. upper reaches of the formation date to ments, clamshell cutting tools best Still higher, better-drained parts of the between 1.66 and 1.57 Ma.23 replicated the Bukuran bovid bone landscape supported a community of At Bukuran, apparent technologi- cut marks.31 sedges, grasses, ferns, and scattered cal cutmarks have been observed on trees.29 Fauna included stegodon bovid bones from legacy collections. Bapang Formation. Between 1.6 and (Stegodon elephantoides), cervids (Cer- Two specimens show marks outside 1.5 Ma, volcanic cones grew north- vus zwanni, C. hippelaphus,andCer- the range of natural causes. While west and southeast of Sangiran. vus sp. indet.) and one small bovid stone tools are absent from the San- Larger, more powerful streams began (Duboisia santeng), as well as Homo giran Formation, molluscan shells scouring and infilling the local low- erectus.19,26,27,30 The hominin-bearing are abundant. In a series of experi- lands.23 A stark fluvial erosion 190 Larick and Ciochon ARTICLE

large bovids ( palaeokerabau, sively studied. Old endemic species Bibos palaesondaicus), stegodon include Duboisia santeng and Axis (Stegodon trigonocephalus), and pri- lydekkeri, as well as Stegodon trigono- mates (Homo erectus, Macaca sp. cephalus, Bubalus palaeokerabau, indet., Trachypithecus cf. aura- Bibos palaesondaicus, Sus brachygna- tus).27,32,33 After about 0.9 Ma, Poh- thus, and Panthera tigris trinilen- jajar Formation fluvial deposits sis.26,44 The assemblage has been covered the Bapang sequence with a compared with that of the Bapang higher proportion of air-fall tuffs, formation Grenzbank Zone, having fluvially reworked ash fall, and lahar an age, using the short chronology, deposits (Fig. 2). The Pohjajar has of around 1.0 Ma.26 The long chro- not yielded Homo erectus remains at nology places the Grenzbank Zone at Sangiran or elsewhere. 1.5 Ma (Box 4).19 In any event, com- parison with the Sangiran sequence Trinil, Java is of limited value. In its middle course, the Solo Using Trinil and other legacy fau- River makes an abrupt northward nal collections, Storm modeled the Figure 4. Homo erectus cranium (S 17): turn to cut through the Kendeng ecological role of ISEA Homo erec- 45 Bapang Formation below the Middle Tuff, Hills (Fig. 3A). In 1891, Eugene tus. The number of identified Sangiran, Solo Basin, Java. Cranial 274 Dubois made the discovery specimens and the minimum num- capacity: 1,004 cc. (Photograph: R. Cio- ber of individuals reflect trophic lev- chon). [Color figure can be viewed in the of a calotte and femur on the Solo online issue, which is available at wileyonli- River bank at Trinil. Dubois named els of primary and secondary nelibrary.com.] the collection deposit the Hauptkno- consumers. Further, the numbers of chenschicht ( bone layer) of Tri- remains of Homo erectus, at sites 34,35 surface often marks the contact nil. Much later, four more such as Sangiran, resemble those of between the Sangiran and Bapang femora were collected in the same large carnivores. These numbers sug- area.36–38 The Trinil skullcap lies gest that Homo erectus functioned as Formations. This contact represents a 45 period of net sediment removal from within the range of morphological a carnivorous omnivore. the Sangiran area.29 Immediately variation for the large collection of A recent analysis of curated Haupt- above the contact, the Grenzbank Sangiran specimens. Together, the knochenschicht materialsisprovoca- Zone has poorly sorted coarse sedi- Sangiran and Trinil crania should tive. Results suggest a significantly ments. Heavy clasts, including verte- represent the earlier Sunda Homo younger age for the site and that brate fossils, can be highly erectus paleodeme. Homo erectus made complex use of 40 39 fragmented and indicate multiple The stratigraphic relationship freshwater shellfish. Ar/ Ar and reworking. Hominin fossils are found among the fossils has always been luminescence dating methods have throughout the Bapang. However, in questioned, especially between the been applied to sediment adhering to comparison with higher reaches of calotte and the original Femur I.39,40 freshwater mollusk shells (Pseudodon) 46 the formation, the Grenzbank lacks Recently, all five Trinil femora have in the surviving faunal collection. hominin skeletal elements of rela- been morphologically compared and TheresultsareindicativeoftheMid- tively low density, such as calottes, their structural and density charac- Brunhes Event (480 ka). As the new and retains only the denser mandibu- teristics evaluated by computed age analysis depends on materials lar and maxillary fragments and tomography.41 Femur I is anatomi- removed from stratigraphic context, it teeth. 40Ar/39Ar radiometric analysis cally more modern and less fossil- is difficult to make a definitive judg- suggests that the deposit began accu- ized than Femora II-V. Femur I is ment. The dates and behavioral mulating more than 1.5 Ma and con- apparently younger than the calotte, hypothesis are presented in the Mid- tinued to 0.9 Ma,19 thus recording while Femora II-V may be more Brunhes Event section. climate cycles in the range of MIS 47- closely related in time to the Homo 23. erectus calotte.41 Eastern Island Arc The Bapang sequence contains Trinil has seen little field investiga- numerous paleosols. These devel- tion since 1891. The stratigraphy Soa Basin, Flores (Indonesia) oped on riverine landscapes, repre- remains unimproved and contention The Soa Basin covers 200 km2 of senting riparian forest, savanna, and persists about the age of Hauptkno- west-central Flores and contains open woodland environments.29 chenschicht. The main bone layer is two volcaniclastic sedimentary Paleosol morphology and carbon iso- now considered an overbank deposit units. At its base, the Ola Kile For- tope values indicate a long-term shift built up during repeated floods. It mation consists of andesitic brec- toward longer annual dry seasons.29 may contain materials of different cias and volcanic mudflows with The more open habitat supported origins and ages.42,43 The Dubois minor interbedded tuffaceous silt- carnivores (Panthera), (Sus bra- fauna collection, curated at the Reiks stones, sandstones, and lava flows.47 chygnatus), cervids (Axis lydekkeri), Museum in Leiden, has been inten- From near the top of the formation, ARTICLE Early Hominin Biogeography in Island Southeast Asia 191

Box 2. Insular Evolutionary Process

Oceanic islands are isolated set- size217 and positively correlated This is because large-bodied indi- tings in which with its degree of isolation from viduals use more resources, so and can intensify and the mainland.218 The smaller and island natural selection favors thus accelerate evolutionary rates. more isolated an island, the more smaller individuals which, over Immigrating island populations are significant the role of island rule. time, produce smaller-bodied popu- smaller and less genetically diverse Island dwarfing has been observed lations.221 Smaller body size aids in than the mainland source popula- in a wide variety of both living and maintaining relatively large popula- tion, resulting in a founder effect. fossil mammals. There are well- tions on island resource bases.222 Islands have lower biodiversity known cases of Pleistocene probosci- Pleistocene ISEA demonstrates than do mainland areas, which dean dwarfing relating to Pleistocene numerous cases of island endem- results in species expanding and sea-level fluctuations. These include ism. At least five dwarfed species of shifting ecological niches from on several Mediterranean Stegodon emerged in the region. their mainland counterparts. More- islands, mammoths on the Califor- Van Valen’s island rule is directly over, island size is linked with fau- nia Channel Islands, and applicable in the large and small nal turnover, with smaller islands in Island Southeast Asia. forms observed in the Liang Bua having higher extinction rates.215 The strong hypothesis for island vertebrate fauna.6,216 These include These conditions force microevolu- dwarfing revolves around popula- giant tortoise (Geochelone), giant tionary changes that may lead to tion size versus food availability.219 (Spelaeomys or Hooijeromis macroevolutionary changes, includ- In general, island landmasses offer nusatenggara), a very small stego- ing . Van Valen’s “island relatively reduced nutritional don (Stegodon florensis insularis), rule” observes that in long-term resources to relatively small popu- and the dwarfed hominin Homo island contexts, large-bodied mam- lations. Islands also tend to have floresiensis (Box 6). As a dwarf mals tend to become smaller and fewer large predators. In general hominin, Liang Bua Homo flore- smaller ones bigger.216 The effects response, mammals have fewer siensis fits well within its of the island rule tend to be inver- young, while patterns of body size ISEA ecological and evolutionary sely proportional to an island’s are much accelerated.220 context.6

a fission-track analysis provides a to 0.90 6 0. 07 Ma.48 Dating analyses large mammals encountered on minimum age of 1.86 6 0.12 Ma. continue at Tangi Talo, with new entering Flores.47 Above Ola Kile, the Ola Bula For- results based on 40Ar/39Ar eruption mation comprises 100 m of volcanic age suggesting 1.3 Ma.49 Walanae Basin, South Sulawesi and fluvio-lacustrine deposits. Like (Indonesia) the Solo Basin Bapang Formation, Wolo Sege. This archeological site the Ola Bula is highly volcaniclastic lies in the Ola Bula basal tuff interval, Cabenge. The Walanae Basin lies on and indicates young riverine habi- just above the Ola Kile breccias. The Sulawesi’s southwest peninsula (Fig. tats and the presence of lakes. Two sedimentary environment is highly 3D). In the late 1940s, H. R. van Soa Basin sites, one archeological volcaniclastic and represents a stage Heekeren collected surface stone and one paleontological, reveal before the fluvial-lacustrine land- artifacts and fauna near the village Flores sedimentary and faunal envi- scapes had truly developed. Wolo of Cabenge (formerly Tjabenge). The ronments toward the end of the Sege has in-situ stone artifacts, Walanae fauna included giant Early Pleistocene (Fig. 3B). including some -like imple- tortoise (Testudo morgae), pygmy ments50 (Fig. 5). There is no in-situ (Archidiskodon celebensis), Tangi Talo. This paleontological fauna. Overlying the layers is stegodon (Stegodon sompoensis), locale lies low in the Ola Bula strati- an ignimbrite with an 40Ar/39Ar erup- giant (Celebochoerus heekereni), graphic sequence. Fauna include tion date of 1.02 6 0.02 Ma.47 and dwarf buffalo ( depressicor- dwarf Stegodon (Stegodon sondaari), The Soa Basin’s Ola Bula deposits nis).51 Although the presence of giant tortoise (Geochelone), and provide the oldest record of Pleisto- Archidiskodon suggested an earlier dragon (Varanus komodoen- cene fauna and stone tools east of Pleistocene age, Van Heekeren linked sis). There are no signs of butchering Sunda. The Wolo Sege artifacts indi- the Tjabenge stone and no stone artifacts. The Tangi cate that hominins arrived on Flores with this fauna.52 He cautioned, Talo small-bodied S. sondaari is the well before 1.0 Ma. The young vol- nevertheless, that the Walanae Archi- earliest stegodon on Flores. An over- canic environment represents the diskodon could represent a Middle lying tuff of volcanic debris is dated conditions that hominins and other Pleistocene descendant of a long- 192 Larick and Ciochon ARTICLE

lived endemic island form. Questions arose about the association of the industry with the fauna and the age of each.53–56 With new research, the trends Walanae terraces are now inter- larger N hemishere ice volumes significant glaciation/ draw-down significant glaciation/ draw-down Dmanisi H. erectus begin 41-kyr obliquity cycle: glacial cycles of short duration and low temperature amplitude preted as the upper part of a normal basin fill sequence uplifted during the .32 New work at nearby Talepu suggests that some artifact horizons have an early Pleis- Stegodon tocene age.57 S. sompoensis The Walanae basin volcaniclastic gravel series displays clasts of yellow and red . These flake and other highly colored, fine-

(Tjabenge) grained siliceous rocks are the raw materials for the Tjabenge Indus- try.52 Tjabenge flakes are relatively

sites lithics small and thick, with signs of having Cabenge small been struck in all directions from irregular cores. Tool types include points, concave scrapers, core and keeled scrapers, endscraper picks, Stegodon sondaari and chopping tools. Van Heekeren noted similarities between the Tja- cores radial benge and Sangiran flake industries, including the use of small, highly colored, fine-grain raw materials and fm Talo sites lithics Wolo Sege Tangi the production of irregular cores.

eruptions He concluded that these and other similar flake industries were pro- duced by a single species of homi- nin52 (Fig. 5).

Stegodon LATE EARLY PLEISTOCENE

S. elephantoides REVOLUTION OR LATE EARLY- S. trigonocephalus S. trigonocephalus TABLE 1. Early Pleistocene Synoptic Chart EARLY MIDDLE PLEISTOCENE

Java Flores Sulawesi The MIS 24-22 complex (1.0-0.9 Ma) marks a shift in glacial-

(Sangiran) interglacial forcing from the 41-kyr obliquity-based cycle to a 100-kyr eccentricity-based cycle.15 The MIS 24-22 complex has thus been called sites lithics

Bapang small flake the late Early Pleistocene Revolution Lower Lahar(EPR) Ola Kile or the Mid-Pleistocene Revolu- tion.58–61 The full period of transi- ka 990 Soa age tion, 1.25 Ma to 700 ka, has been termed the Mid-Pleistocene Transi- tion.62–64 We call it the “late Early- early Middle Pleistocene.” After EPR, glacial periods became Jaramillo eruptions subchron longer and more evenly distributed.

ka events Longer glacials served to increase end  global ice volume, thereby pushing

ka sea-level lowstands as much as 50 m  start lower than during the earlier Pleisto- cene.62 Short but warm interglacials MIS 24 975 920 EPR / 31 1070 Trinil 66 1980 1790 Olduvai 62-52 1700 1550 Sangiran 36 1200 3746-38 1400 1240 1250 47 1450 Sangiran decreased ice volume quickly. The ARTICLE Early Hominin Biogeography in Island Southeast Asia 193

Box 3. Stone and Shell Tool

Pleistocene ISEA Gesher Benot Ya’aqov (GBY) in sequences represent a assemblages are of three types. .73,74 The LFA chaine operatoire manifestation of the Acheulean arrives at simple yet effective cutting cultural formation. 1. Flake Industries. Early sur- tools in a small number of well- 3. Radial-Core Reduction. This face collecting at several locations structured steps. LFA assemblages is present on Flores, particularly have led to this category, in which large (>10 cm) flakes, either at Mata Menge and Liang Bua. flake and cobble tools are generally raw or minimally retouched as uni- Lithic resources were brought to ad hoc in flake production and facial tools and simple Acheulean- thesiteintheformofflake bifacial reduction. Flake industries type bifaces (convergent tip) and blanks. The Acheulean-like ele- have relatively few biface and cleavers (broad tip). The LFA is ments of Wolo Sege are not pres- -type tools.54,55,75 Flake associated with coarse-grain - entatMataMenge,wherethe industries show neither chronologi- ite and related materials rather than more complex tools show a reduc- cal development nor regional fine-grain . Kombewa flake tion sequence based on centripetal differentiation. types are common as cleaver blanks. or radial removals on the blanks.4 Of relevance are the flake indus- As a cultural force, the LFA A radial reduction sequence is tries named after their modern set- appeared by 1 Ma at found again at Liang Bua, 50 km tlements of Cabenge (Sulawesi), and spread throughout Africa and to the west and 800 kyr later. Kota Tampan (Malay Peninsula), beyond. While timing is imprecise, ISEA has few Pleistocene arche- and Ngandong and Pacitan (Java). the LFA arrived in the Levant, Ibe- ological sites in which complex Questions of artifact age and verac- ria, and by 600 ka. After stone-tool technology is associated ity plagued the early surface finds. 500 ka, only sub-Saharan Africa with diverse occupation debris. Late twentieth-century examina- holds sure LFA sites.73,74 Recently, Several explanations have been tions of terrace structure and theLFAprofilehasbeenapplied offered. Early hominins main- assemblage technology generally to assemblages from Ngebung tained a vegetarian diet that did ascribed these surface finds to the (Sangiran, Java) and Gunung not require complex stone late Pleistocene.53–55,75 Recent Sewu. The attribution was based tools.224 Early hominins used research, nevertheless, has recov- on typological grounds and a per- more readily available nonlithic ered stone tools in association with ceived between the Pinjor raw material, such as bamboo225 fauna and, sometimes, datable fauna (associated with LFA sites and wood.226 Since the late 1990s, material. Generally speaking, the in India) and the “Stegodon-Homo evidence has mounted to support earliest well-dated archeological erectus fauna” of Sunda.223 Bifaces the use shellfish for food and assemblages on both sides of Wal- and cleavers from the Cagayan shells for tools. Shell archeology lace’s Line have ages greater than and Arubo basin sites (Luzon) has been published for three sites: 900 ka. This pattern holds for have also been termed Acheulean Bukuran and Trinil on Java, and Java,66 Flores,47 and Sulawesi.53 and even Large-Flake Acheu- To’os on Timor.31,46,87,227 Individ- The earliest stone-tool horizon on lean.103,104 Similar tools from ually, each site has marginal evi- Luzon is about 800 ka.97,101 Wolo Sege (Soa Basin, Flores) are dence of shell use. When pooled, a 2. Large-Flake Acheulean also called Acheulean.78 The ques- hypothesis for early hominin (LFA). This category is defined by tion remains of whether or not shellfish use can nevertheless be the biface- and cleaver-rich site of Acheulean-like tools and reduction entertained.

MIS 23 interglacial is therefore lowstands likely made for extensive tation trails off rapidly thereafter. implicated in flooding insular land- dispersal corridors. Likewise, early The EPR marks the last known masses.65 At Heqin (China), 0.92 Ma Middle Pleistocene interglacial high- occurrence of Homo erectus at San- marks the transition to a long period stands led to insular isolation and giran. The early Middle Pleistocene (0.92 to 0.13 Ma) during which the endemic evolution. saw two region-wide catastrophes: Indian Summer Monsoon (ISM) was The early Middle Pleistocene may the Australasian tektite impact and structured by both southern high also be associated with generally the oldest Toba Tuff super-eruption and northern low pressure. During lower sedimentation rates. In the (Box 5). this period, ISM maxima coincided basins of Soa (Flores) and Cagayan Two areas, Sangiran and Soa, with Northern Hemisphere ice vol- (Luzon), the period immediately sur- have sedimentary sequences that ume minima (interglacials) and Ant- rounding the EPR is well represented extend from the Early to the Middle temperature maxima.15 Early by open-air archeological and faunal Pleistocene. Earlier, we presented Middle Pleistocene glacial sea-level sites, but evidence of hominin habi- Early Pleistocene evidence. We now 194 Larick and Ciochon ARTICLE

Figure 5. ISEA Early Pleistocene lithic artifacts. Ngebung 2, Sangiran, Solo Basin, Java: A, retouched large flake; B, polyhedron (courtesy of F. Semah).66 Cagayan Basin, Luzon, Philippines (photograph: R. Ciochon). Arubo, Luzon, Philippines: bifacially retouched large flake (cour- tesy of E. Z. Dizon).97 Wolo Sege, Soa Basin, Flores: small flakes (courtesy of A. Brumm). Cabenge, Walanae Basin, Sulawesi: small flake tools (modified after H. R. van Heekeren).52 [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] address the Middle Pleistocene Sunda type sites. The terrain is a dissected record for Sangrian and Soa, as well escarpment ranging from as other areas. Table 2 shows the Ngebung (Sangiran) terraces atop the Pohjajar Formation events, sites, lithics, and fauna that On Sangiran’s northwest flank, the down through the Bapang to the San- were present during the early Middle Ngebung Hills lie some 3 km NNW of giran Formation. From 1989 to 1994, Pleistocene. the Sangiran and Bapang Formation a French-Indonesian team excavated ARTICLE

TABLE 2. Mid-Pleistocene Transition Synoptic Chart start end age Java Flores Luzon/Bose MIS ka ka events ka sites lithics Stegodon sites lithics Stegodon sites lithics Stegodon trends 11 420 360 MBE 420 ISEA shift to cave occupation/ preservation 12 480 420 13 520 480 Gunung tropical Sewu uplift rainforest New & karst Guinea & formation Australia 14 580 520 15 630 580 in following interglacials: shift to more modern forests with reduced

diversity 195 Asia Southeast Island in Biogeography Hominin Early 16 680 630 Plateau min forest 19 800 680 Brunhes- 778 Arubo LFA? Matuyama 20 800 800 Kobatua radial Cagayan LFA S. luzonensis cores 21 Oldest 803 Boa Lesa radial Bose bifaces Toba/ cores Tektites 22 880 870 Mata radial S. florensis Tibet Plateau Menge cores frigid glacial 23 880 EPR 880 Ngebung? LFA? S. trigonocephalus 24 975 920 Jaramillo 990 Hequing shift to cold 196 Larick and Ciochon ARTICLE

Box 4. Long Chronology

During 125 years of ISEA paleonto- chronology.’ 236–238 Alternatively, the 60 ka),184 and Luzon (Homo sp logical research, an early hominin first widespread radiometric analy- indet, 67 ka).156 chronology has been difficult to ses, based on counting fission tracks, Marine Isotope Stages resolve. The most productive yielded young and highly scattered TheglobalMISprovideacali- research area has been Sangiran, in dates.21,239–241 Since the 1980s, a brated environmental framework , Indonesia. With a thick growing number of 40Ar/39Ar dates relating to basic ISEA habitat volcaniclastic sedimentary sequence and continually refined sampling change. The MIS system uses oxygen presenting at many locales, Sangiran schemes have generally shown San- isotope levels to indicate alternating has produced a great majority of the giran’s hominin-bearing sequence to warm and cool periods during the region’s Homo erectus fossils, archeo- have begun more than 1.5 Ma, possi- Quaternary or Pleistocene. logical sites, and chronological inter- blyasmuchas1.86Ma.19,20 Similar Isotope data are drawn from pollen pretations. It is worth summarizing chronological debates have been and foraminifera remains in sea- the evolution of chronological frame- seen for Arc locales, including Flores, bottom sediments. Deep-sea core works at Sangiran. Luzon, and Sulawesi. sediments also preserve paleomag- A micro-paleontological Plio-Pleis- The long chronology suggests netic reversals that can aid in estab- tocene (current early Pleistocene) that early hominins arrived in lishing chronology. ISEA deep-sea chronology was established for San- Sunda (Sangiran) in the form of cores also contain air-fall tephra giran well before WWII.228–230 After Homo erectus before 1.5 Ma. An from volcanic eruptions and micro- the war, a reinterpretation of the evi- early form arrived east of Wallace’s tektites from the Australasian Tektite dence suggested that the fossils Line, to Flores, about 1.1 Ma. In Strewnfield (Box 5). When subjected belonged to, in current terms, the the Late Pleistocene, the probable to radiometric age analysis, these late Early-early Middle Pleistocene ultimate survivors of these early horizons can be used to refine (1.0 to 0.4 Ma).84,231–235 However, dispersals are found on Java regional chronology. new microfossil analyses done in the (Ngandong Homo erectus, 125 strengthened the initial ‘long ka),141 Flores (Homo floresiensis, at Ngebung 2, where several meters of sent migrants contemporary with the collections, such as the Sangiran and Bapang riverine sands and gravels initial Homo erectus dispersal. Most nearby Pacitanian industries.53–55,75 overlie the Grenzbank Zone.66 Five of these foraged in open settings but, It is possible that some or all of the stratigraphic ensembles yielded stone for rest, depended on closed habi- stone artifacts at Ngebung are sur- artifacts and a highly fragmented tats. The fauna may therefore reflect face intrusions from a later period. fauna.66 40Ar/39Ar dates on tuffaceous a forest fringe setting.70 layer heavy minerals produced ages of Ngebung stone artifacts feature 800 ka.67 Our own analysis gave an large flakes (including Kombewa Eastern Island Arc 68 age of 900 ka. type), polyhedrons, cores, cleavers, Soa Basin, Flores (Indonesia) The 2,0001 fossil bones and teeth and hammer stones with little lithic include more than 800 fos- (Fig. 5).71 Spheroids are About 0.9 Ma, local volcanic erup- sils assigned to species: large bovids recorded from an erosion zone in a tions contributed heavily to Ola Bula comprise 45%; smaller cervids, 18%; lower stratigraphic unit (ensemble basin infill. The earlier volcanic tuff- and stegodons 8%.69 Endemic spe- A); these could be more recent intru- dominated facies grades upwards cies include stegodon (Stegodon trig- sions.66 Ngebung flakes, tools, and into a sandy interval dominated by onocephalus) and bovid (Duboisia spheroids are made on peb- fluvio-lacustrine tuffaceous sand 47 santeng). Mainland forms include bles and cobbles,72 likely quarried layers. At this time, the earlier pygmy hippo (Hexaprotodon sivalen- from Bapang-era erosion surfaces in Pleistocene dwarf stegodon (Stego- sis), buffalo (Bubalus paleokerabau), the Lower Lahar Unit.23 don sondaari) of Tangi Talo is not rhino (Rhinoceros sondaicus), and The Ngebung assemblage has two represented, but appears to have pig (Sus brachygnathus). possible origins (Box 3). On one been replaced by a large-bodied form Bone surfaces are highly eroded. , the biface and cleaver ele- (Stegodon florensis).76,77 S. florensis There are no cutmarks present, but ments recall the Large Flake Acheu- likely dispersed from a larger land- bovid long bones appear to have lean (LFA).73,74 Should these be true mass during MIS 22, the first been intentionally broken and LFA artifacts, Ngebung would be extreme glacial period of the Pleisto- defleshed. Some Stegodon tusks are among the oldest such sites outside cene. The accompanying sea-level broken as if to create ivory flakes.69 of Africa.71 On the other hand, the lowstand certainly opened dispersal The large mainland forms, Bubalus, Ngebung assemblage resembles ISEA routes from Sunda to the Arc and Rhinoceros, and Sus, should repre- flake industries defined on surface across the Arc. In such conditions, ARTICLE Early Hominin Biogeography in Island Southeast Asia 197

Box 5. Pleistocene Catastrophes

Three region-wide catastrophes are Bose, the Australasian event may found in undersea deposits across the visible in the ISEA early hominin have caused widespread, devastating , as well as the Arabian archeological record. The Australa- forest fires which early hominins and .255 YTT dates to sian Tektite Impact Event produced adapted to through the use of bifacial about 73,500 years 6 3,000 years148 the widespread Australasian Tektite stone tools.247 The association of or 73,000 years 6 4,000 years ago.149 Strewnfield. ’s TonleSap, stone tools and tektites in Bose is, YTT has been implicated in initiating the largest freshwater lake in South- nevertheless, in debate.251,252 the last glacial cycle due to its coinci- east Asia, may represent the impact The Toba caldera of northern dence with not only ice buildup dur- crater (Fig. 1).242 AgefortheAustra- Sumatra is earth’s largest ing this time,256 but population lasian Impact is estimated at 0.8 Ma, Pleistocene-age volcanic complex. bottlenecks of in west- based on 40Ar/39Ar and fission-track Two of numerous eruptions pro- ern Eurasia and Homo sapiens in analysis of individual tektites. 243–247 duced the Oldest Toba Tuff (OTT) Africa.257,258 However, the arguments Corroborating evidence comes from and the Youngest Toba Tuff (YTT). are weak. YTT traces have not been the position of microtektites in deep- New research suggests that OTT rep- identified at ISEA fossil hominin sea cores, always below indications resents a total tephra volume of 2,300 sites. The Middle Solo sites and Liang for the Brunhes-Matuyama geomag- cubic km,253 which is comparable to Bua appear to predate the event and netic reversal of 0.78 Ma.248–250 Aus- the better-known YTT total volume of the Callao stratigraphy apparently tralasian tektites are known in 2,800 cubic km.149 The Australasian postdates YTT. Given the present associationwithearlyhomininarti- Strewnfield and OTT are closely state of knowledge, it seems that facts and mammalian fauna in vari- related in time. At ODP site 758, across ISEA the YTT event did not ous contexts, including the Bose northwestofToba,Australasian generate significant bottlenecks or Basin, Guangxi, China, and the Caga- microtektites and OTT are associated among mammalian yanBasin,Luzon,Philippines.In in ash layers D and E.254 YTT is populations.259–261 larger-bodied stegodons could have blanks (Box 3). The Acheulean-like cal proximity. Nevertheless, age and dispersed to formerly small land- elements of Wolo Sege are not pres- relationship remain unclear. masses where dwarf species could ent at Mata Menge. have been overwhelmed. The Soa Basin Ola Bula formation, Talau basin (Timor). Pleistocene from Wolo Sege to Mata Menge, has fluvial conglomerates are exposed Mata Menge, Boa Lesa, and Koba- ISEA’s most complete archeological along the Talau River border tuwa. These three stratified sites lie record. Wolo Sege indicates hominin between Indonesian Timor and East in the sandy interval above Wolo Sege arrival at more than 1.0 Ma. The Timor. During the 1950s and 1960s, 47 and Tangi Talo (Fig. 3B). Mata change in fauna between Wolo Sege localities east and south of Atambua Menge has a basal date of 880 ka. and Mata Menge demonstrates an (Indonesian Timor) produced in situ 79,82,84 Boa Lesa and Kobatuwa lie up-section EPR-related faunal turnover at about fossils and surface stone tools. from Wolo Sege and Tangi Talo. A 900 ka. After this turnover, the Taxa included Stegodon timorensis series of tuffs runs through the entire Soa Basin shows phylogenetic conti- (pygmy stegodon), Geochelone (giant sequence. Fission track dates for cap- nuity in large-bodied until tortoise), and Varanus komodoensis 80,81,83,86 ping sediments place them at 700 ka. the arrival of Homo sapiens at (). During Among the three sites, major faunal 12 ka.47 the 1990s, fauna and artifacts were elements include large-bodied stego- associated at two more sites, but don (Stegodon florensis), Komodo Southern Wallacea Outer Arc accounts remained preliminary.87 In dragon (Varanus komodoensis), giant (Indonesia) 2015, at Raebia, east of Atambua, rat (Hooijeromys nusatenggara), and the Iowa-Bandung project located giant tortoise (Geochelone). The large South of Flores, the Savu Basin stegodon, giant tortoise and possible stegodon on Flores is somewhat subduction trench ranges to more stone artifacts in relation to bracket- smaller than the large form on Java.77 than 3,000 m in depth (Fig. 8). ing tuffaceous lenses. Nearby, at With a total of 507 stone tools, Sumba and Timor lie on the basin’s Sadi laun, stone artifact clusters Mata Menge has the largest early southern arc. At the lowest Pleisto- were excavated from deflated terrace hominin archeological assemblage in cene sea-level lowstands, overwater deposits. ISEA.78 Lithic resources were channels of 20–40 km separated brought to the site in the form of these islands from Flores. Both Noelbaki (Timor). At Timor Island’s flake blanks. The more complex tools islands have sites at which Paleo- west end, near , Paleolithic show a reduction sequence based on lithic stone artifacts and Pleistocene stone artifacts appear in coarse flu- centripetal or radial removals on the vertebrate fossils are found in physi- vial gravels cut through by modern 198 Larick and Ciochon ARTICLE watercourses. In 1978, Soejono sur- Pleistocene flake tool assemblages fit wear analyses suggest curation and veyed at Noelbaki and Noeltarus, within the Large Flake Acheulean.97 reuse.103,104 The Arubo complex age reporting choppers, chopping tools, remains unknown, but is likely to be flakes, blades, and proto-hand- Cagayan Basin. At Luzon’s northern correlative with the LFA features at made from large flakes.88 In 2015, at end, the Cagayan Basin is a 250 x 80 the Cagayan sites. thesamelocalities,theIowa-Bandung km subduction zone feature with 10 The Stegodon-artifact-tektite (Box project differentiated a laminar indus- vertical km of sedimentary infill (Fig. 5) association at Awidon Mesa For- try with limited patina from large 3C). The upward-coarsening volcani- mation is crucial for understanding flake tools with heavy patina. Contex- clastics reflect the tectonic and vol- Luzon early hominin arrival (Fig. 5). tual materials are in analysis. canic evolution of the adjacent This association suggests that early Cordillera Central volcanic arc. The hominins arrived at Luzon by 800 Watambuka and Lewapaku upper 900 m comprises two forma- ka, probably bringing LFA technol- (Sumba). On Sumba’s northeast tions of interbedded fluvial and pyro- ogy. It remains to be seen whether shore, Watumbaku lies in current estu- clastic deposits, the Ilagan (lower or not the Luzon dispersal relates to arine contexts. In 1978, a stegodon left 500 m) and the Awidon Mesa (upper the Soa Basin Middle Pleistocene mandible (Stegodon sumbaensis)was 400 m). The sequence consists of faunal turnover. It must be noted recovered without context.89,90 In four depositional environments: that in the Cagayan and Arubo 2015, the Iowa-Bandung project found meandering stream, braided stream, Basins, the Middle Pleistocene stone lahar and pyroclastic flow, and ash- technology is not easily differenti- two retouched flakes in deposi- 98 tional contexts indicating lower fall deposits. Cagayan has scores of ated from the Tabonian industries of (Pleistocene) sea level. In 2012, at localities with stone tools and, in the late Upper Pleistocene found on Lewapaku, 30 km inland, the Wollongong- some cases, tektites and Pleistocene Palawan Island, which are associated 99 94 Bandung Geological Museum project fauna. Field work has recently with Homo sapiens. found, in situ, tusk fragments, a tooth been resumed in the Cagayan Basin (Varanus komodoensis), a giant where the contemporaneous occur- MID-BRUNHES EVENT OR LATE murine (?Hooijeromys), and a rence of stone tools, tektites, and MIDDLE AND LATE PLEISTOCENE bird long bone. A loose comparison Pleistocene faunas has been 100 MIS 12/11 (480-360 ka) began a was made with the fauna of Tangi confirmed. new pattern within the 100-kyr cli- Talo.91 Enrile Southern Quarry. An impor- mate cycle. Glacial phases became Luzon (Philippines) tant Awidon Mesa formation locality long (70-90 ka) and very cold, while is Southern Enrile Quarry, near interglacials became short (10-30 ka) The Philippine archipelago has Penablanca.~ Here, a Danish- and warmer.105 This transition, the early hominin evidence northern- Australian team is bracketing the Mid-Brunhes Event (MBE), began most on the Arc and farthest from fossil and artifact-bearing level using the four large-amplitude glacial- Sunda. Deep sea channels currently 40Ar/39Ar on volcanic elements and interglacial cycles that have struc- separate the archipelago from other luminescence on low-temperature tured global climate to the pres- ISEA landmass systems, suggesting sediments. Preliminary 40Ar/39Ar ent.106 With MBE, earth’s climate the presence of dispersal barriers ages range from the late Early and became more orderly, predictable, throughout the Early and Middle Middle Pleistocene to 0.4 Ma as the and extreme.107 Pleistocene. secure youngest age.101 It is reasona- MBE had significant global conse- Paleoanthropological research has ble to entertain an Early Pleistocene quences. MIS 12 exhibited severe cool- a long history on Luzon. During the Revolution (EPR) time frame for the ing effects.108,109 MIS 11 marked the 1950s, Paleolithic flake tools and artifact-fauna-tektite association. longest, warmest interglacial, with sea extinct fauna were found together in level rising to 20 m above the present the Cagayan Basin, and probable dis- Arubo Basin. Approximately 300 highstand.110 The MBE 12/11 complex 92 persal routes were hypothesized. km south of Cagayan, Arubo is a probably aided in both dispersing and Australasian tektites were also dis- complex of open sites in the Sierra isolating ISEA large mammals. It is 93 covered at several locations. During Madre foothills of Central Luzon.102 worth noting that in western Eurasia, the 1970s, researchers at the A morphologically heterogeneous the appearance of Neandertal traits is National Museum of the Philippines lithic assemblage has been collected correlated with the onset of MIS 11.111 surveyed many more Cagayan co- here, primarily from sites out of geo- Extreme interglacial sea level high- occurrences of stegodon and arti- logical context.97 The site complex stands also came during MIS 9 and 5e. facts.94 More recently, fossils of sev- lies close to a chert boulder deposit, On mainland China, in conjunc- eral mammal families have been which served as the raw material tion with MBE, the Hulu and found in archeological contexts, source. The stone tool assemblages Dangge cave flowstones record the including small stegodon (S. luzonen- include the Large Flake Acheulean beginning of monsoon rainfall. The sis), and the tektite association has (LFA) hallmarks of bifaces, cleavers, flowstones record the dry been confirmed.95,96 Recent research flake cores, retouched flakes, and directly associated with the com- also suggests that the Luzon Middle choppers (Fig. 5). Microscopic use- mencement of massive periodic ARTICLE Early Hominin Biogeography in Island Southeast Asia 199

North Atlantic glacier ice calving known as Heinrich events.112,113 These events produced cold winters in Europe and dry monsoons in China.114,115 Another consequence of Heinrich events was the southward

trends shift of the Tropical Rain Belt.116 The sequence of events, stretching halfway around the globe, indicates the swiftness of glacial climate

Holocene MIS 5-2: ITCZ placementMonsoon. for Dry SE conditions throughout ISEA Last Interglacial; high sea levels Liang Bua sedimentation drying in New GuineaAustralia & Song Terus sedimentation high interglacial sea levels severe cooling 100-kyr obliquity cycle: cold glacials (70-90 ka); warm interglacials (10-30 ka) change. The Mid-Brunhes Event marked yet another significant change in regional erosion and sedimentation patterns. Some limestone Stegodon S. florensis S. insularis massifs developed karst landscapes. Central Java’s Southern Mountains Miocene coral beds (Gunung Sewu) provide an example. At Punung III in radial cores Gunung Sewu, a U-series age of 492 6 38 ka for the lower flowstone pro- vides a minimum age for uplift.117 sites lithics Liang Bua This determination corroborates other Middle Pleistocene uplift esti- mates relating karst development in the Southern Mountains.118–120 Early hominins found shelter in these newly opened . The Song

Stegodon Terus cave fluvial stage documents the accumulation of stone artifacts S. trigonocephalus S. trigonocephalus before 300 ka. Liang Bua opened for infilling about 195 ka and shows evi- dence of hominin habitation shortly thereafter. The evolution of is not yet clear. The Mid-Late Pleistocene bound- Java & [ ] Flores flakes (Pacitanian?) ary (MIS 5e, 130-120 ka) was a glob- TABLE 3. Mid-Brunhes Event Synoptic Chart ally important climatic event, represented in Paleolithic sites across northwestern Eurasia. How-

sites lithics ever, sedimentary sequences in

Terus Southeast Asia do not seem to regis- ter the Last Interglacial in significant ways. One Late Pleistocene catastro- 17 73 [Lenggong] flakes ka phe, the Youngest Toba Eruption, is age not identifiable in sedimentary sequences throughout ISEA (Box 5). Table 3 shows the events, sites, lithics, and fauna present during the Toba Mid-Brunhes Event, late Middle eruptions Pleistocene, and Late Pleistocene. ka events end  Sunda Trinil, Java

ka Recent observations from Dubois’  start historical faunal collection at the Museum in Leiden46 MIS 112 2 28 125e Flores 6 13078 120 190 2509 300 130 10 190 11 340 250 360 420 300 340 360 MBE 420 Song 36228 4 74 62 Youngest 5 120 74 Ngandong flakes suggest that Homo erectus may have 200 Larick and Ciochon ARTICLE occupied Trinil during the Middle ied chronology may have merged in Pleistocene, displaying advanced cog- this context.121 Given the poorly nition and cultural behavior. Within known of the site and the the collection, the Pseudodon shell primitive morphology of the Trinil assemblage represents adult large- calotte, we hesitate to grant sized specimens (80-120 mm in advanced symbolic behavior to the length) from varied riverine settings. group represented by the calotte. This suggests that individual shell- fish were selected for consumption Gunung Sewu, Java based on large size. One shell valve shows signs of modification by The Southern Mountains of East retouch, possibly resulting from use Java lie on the Indian Ocean coast, as a cutting or scraping tool. Another 100-150 km south of the Solo Basin. shell displays a zigzag pattern of The area has been of geological on the central part of the interest since the 1830s and of arche- left valve. The marks are attributed ological interest since the 1930s.124 to the engraving of meaningful In 1935, von Koenigswald and Twee- abstract patterns on an attractive die collected a range of flake and substrate.121 In other words, Trinil pebble tools from the Baksoka riv- hominins are thought to have inten- erbed, later described as the Pacita- tionally marked the shell. nian industry.125 Since the 1990s, 40 39 Luminescence and Ar/ Ar analy- Gunung Sewu research has focused sis on the shells’ adhering sediment on cave and habitations give a maximum of 0.55 Ma and a and the use of local lithic resources, 46 minimum of 0.44 Ma, or about including chert, jasper, limestone, half the age estimate based on analy- and meta-limestone.124 Gunung sis of the associated mammalian Sewu stone sources may have pro- 44 fauna. This new age would seem to vided the raw materials for Sangiran pull Trinil away from the Sangiran sites such as Ngebung. fossils and an early Middle Pleisto- Figure 6. Two bifaces from the Baksoko val- cene Ngebung cultural affiliation. Song Terus. Most Gunung Sewu ley, Gunung Sewu, Java (modified after Should the shell marking prove fac- caves have yielded Late Pleistocene G.-H. Bartstra).125 tual, does it indicate cognitive abil- and deposits, but one, ities expectable in the later Java sites Song Terus, features an important indicative of waning Sunda Homo in the Song Terus karst and the Mid- Middle Pleistocene infill. In 1953, erectus populations.126 dle Solo Terraces? On date alone, it Soejono and Basuki excavated fauna Other Gunung Sewu caves, such is tempting to draw a comparison and an archeological industry involv- as Song Klepek and Braholo, aid in with mid-Middle Pleistocene early ing stone, bone, and shell.52 In the understanding late Homo erectus hominin dentognathic fossils from late 1990s, an Indonesian-French and landscape use, Hexian, eastern China, and Penghu, team excavated more than 15 m of and may eventually shed light on the Taiwan. Trinil at a mid-Middle Pleis- stratigraphy in two units.126 The tocene age may represent a time in terrace lithic industries such as the lower unit consists of flood alluvium which Homo erectus diverged in sev- Pacitanian and Sangiran Flake from the 12-m terrace. The layers eral regional contexts. Industry. The deep Gunung Sewu contain fauna (rhinoceros, tapir, and caves may be especially important The Dubois collection zigzag shell 85,126 could represent part of a regional cervid) and flake tools. Com- for understanding hominin adapta- trend toward early hominin cultural bined U/Th-ESR analysis indicates tions during MIS 6 (150-130 ka), diversification. Nevertheless, the Tri- that the lowest archeological levels when the Southeast Asian mainland 127 nil shell marking has a complexity arrived 300 ka. The flake tools was exceptionally cold and dry. At normally associated with early Late may relate to classical Pacitanian this time, the equatorial insular 126 Pleistocene Homo sapiens in lithics (Fig. 6). The fluvially province may have served as an early Africa.122,123 It is possible that diver- derived archeological deposits hominin refuge. increase in density after 180 ka. By sifying Middle Pleistocene Homo Middle Solo Terraces, Java erectus had similar cognitive ability, 80 ka, the local stream at Song Terus but the find is unique. The problem had entrenched below the cave As the Solo River cut through the remains that the Trinil Hauptkno- entrance and a more typical cave Kendeng Hills, it entrenched within chenschicht is a complex deposit, infilling took over. Archeological Pleistocene terraces. In 1931, Geologi- apparently a mixed sequence of remains include and fauna, cal Survey of the Netherlands geolo- flood-borne overbank accumulations. but few lithics. This seems to repre- gist Carel ter Haar discovered a bone Bones, shells, and sediments of var- sent sparse inhabitation and may be bed near the base of the Solo River ARTICLE Early Hominin Biogeography in Island Southeast Asia 201

TABLE 4. Sambungmacan-Ngawi Calvaria Sites Village Location Date ID Poloyo – 1973 Sm 1 calvaria Poloyo – 1973 Sm 2 tibia fragment Mlale-Cemeng 4 km upstream from Poloyo 1977 Sm 3 calvaria Mlale-Cemeng 4 km upstream from Poloyo 2001 Sm 4 calvaria Selopuro 6 km downstream from Trinil 1987 Ngawi 1 calvaria

20-m terrace. Ngandong sat on the and are more specifically identified The Ngandong hominins range cutbank of an acute river bend within by the three villages nearest their from cranial vault fragments to a the Kendeng Hills. From 1931 to find spots (Table 4). nearly complete calvaria retaining 1933, the Survey excavated the bone delicate ethmoid and sphenoid struc- bed within a volcaniclastic sand stra- Middle Solo cranial morphol- tures.134 The Ngandong calvaria are tum.128–130 Ngandong produced 12 ogy. Compared to Sunda Early larger and more robustly built than “” cranial remains. Pleistocene Homo erectus,theSam- the Ngawi-Sambungmacan series Since the early 1970s, four bungmacan-Ngawi specimens show (Fig. 7). The lateral walls of the Ngandong-like calvaria have been derived features, including a relatively braincase are more vertically ori- found 15–20 km upstream from horizontal supraorbital torus that ented, helping cranial capacity range Ngandong, a few kilometers thickens laterally, continuous supra- upward to 1,250 cc. Raised temporal upstream from the Trinil locality. meatal/supramastoid crests, and a lines on the cranial vault give a puffy The calvaria are chance finds in the well-defined occipital torus and supe- appearance to the vault when viewed eroding Solo River bank; the sedi- rior nuchal line.131 Cranial walls are from the front. The strongly devel- mentary details of their provenance relatively tall and the coronal profile is oped supraorbital torus does not are as yet unknown. As a series, the “roundedly tent-shaped.”131 Cranial form a continuous bar, but rather fossils are generally called by the capacity is increased.131–133 The series meets in the glabellar region in a dis- names of their administrative munic- exhibits minor postmortem distortions tinct depression. The pronounced ipalities, Sambungmacan–Ngawi, or surface erosion. occipital torus has greater rearward projection.131

Ngandong . The 1930s excavations produced 25,000 verte- brate fossils, including at least 10 terrestrial mammals: buffalo (Buba- lus), cattle (Bibos), deer (Cervus palaeojavanicus), (Hexaprotodon), leopard (Panthera pardus), muntjac (Muntiacus), pig (Sus terhaari), rhinoceros (R. sondai- cus), stegodon (S. trigonocephalus), and tiger (Panthera tigris). Bovids represent more than half of the assemblage.135 Broken and disarticu- lated elements greatly outnumbered well-preserved specimens and par- tially articulated skeletons. No cut marks have been reported and few verifiable artifacts appeared in excavation. Recent excavations have isolated the extent of the 1930s excavation and located the bone bed stratum within. The Homo erectus specimens can now be placed surely within the original facies C of the basal fossilif- erous horizon.130,134,136 Bone bed sedimentary dynamics are also under Figure 7. Homo erectus calvaria (Solo V, Ng 6): Ngandong, Middle Solo Valley, investigation. The deposit, compris- Java, Cranial capacity: 1,251 cc.275 Courtesy of the American Museum of Natural His- ing poorly sorted, high-energy fluvial tory, New York. sand and gravel, shows hyper- 202 Larick and Ciochon ARTICLE

TABLE 5. Middle Solo Terraces Historical Ages Analyses Year Methodology Age 137 1939 Von Koenigswald attributes the Ngandong 1930s excavation fauna Upper Pleistocene 1988 U-series on Ngandong bone fragments 165 ka129 1996 ESR/U-series on bovid teeth from the Ngandong 1930s excavation area 53-27 ka138 2007 luminescence and U-series on Punung fauna breccias* 128 6 15 to 118 6 3ka117 2008 gamma-ray spectrography on Ngandong/Sambungmacan hominin fossils 70-40 ka139 2011 40Ar/39Ar incremental heating & ESR/U-series on Ngandong and Jigar fauna 546.6 6 12 ka140 ESR/U-series min age: 143120/-17 ka140 2014 red thermoluminescence (red TL) on Ngandong bone bed fluvial sediments 130-102 ka141 concentrated flow features typical of Lenggong Valley, Perak of the river exposing, then invading a lahar event. The volcaniclastic (Peninsular Malaysia) the karst system. The river now lies source appears to be an andesitic 30 m below and 200 m distant from cone located about 50 km away.134 Since the 1930s, several Lenggong the cave.5 With this new information, a Valley localities have produced sim- ple stone tool assemblages collec- Liang Bua. The cave is 14 km north taphonomic hypothesis can be 143 sketched, its events including tively known as the Tampanian. of and 25 km from the north upstream multi-species aggregation, As surface finds, the Tampanian was coast. Stone artifacts began accumu- thought to be an early hominin tech- lating at about 190 ka. After 100 ka, mass death and relatively speedy car- 144 cass decomposition, and mass flow nology similar to the Pacitanian. channel erosion created relief within of osseous elements downstream to The site of Kota Tampan is one of the soft debris. Remnant areas of concentration and burial at Ngan- several localities that show a tuffa- higher ground later became a focus dong. The aggregation may have ceous stratigraphy without faunal for hominin habitation from 74–61 been caused by drought or volcanic preservation. Evidence is now con- ka.152 There is no direct indication clusive that the site lies within the of YTT in the cave sediments. The eruption; mass death may be related 145–147 to localized ash fall. During lahar Youngest Toba Tuff (YTT). Liang Bua archeofauna sequence YTT dates to about 73,500 years 6 ranges from 95 ka to the pres- transport, the carcasses were disar- 148 3,000 years or 73,000 years 6 ent.153,155 Recently, a depositional ticulated and individual bones bro- 149 ken, but surfaces and edges were not 4,000 years ago (Box 5). Other hiatus has been identified between eroded.134 excavated Lenggong sites are strati- 60 and 17 ka.153 graphically younger. One much The Liang Bua archeofauna con- Middle Solo Terraces Ages. The age younger site includes a Homo sapi- sists of unfossilized but well- 150 of the 20 m terrace sediment constit- ens skeleton. preserved mammal, bird, reptile, and uents is yet to be resolved. The his- The Lenggong Valley sites and the mollusk remains. The pre- and post- torical sequence of age analyses is Tampanian are difficult to place in hiatus assemblages are distinct in presented in Table 5.137–141 biogeographic context. The species representation. For the early The oldest dates push these fossils YTT date puts Kota Tampan within sequence, that associated with Homo back into the Middle Pleistocene. the realm of possibility as an early floresiensis, large vertebrates include The youngest results (53 ka – 23 hominin site; however, the stone giant tortoise (Geochelone), three ka)138 straddle the 47 ka arrival technology associated with the site giant rat species (Papagomys, Spelae- threshold for Homo sapiens to ISEA could represent either early homi- omys, or Hooijeromis), and Komodo and Sahul. The youngest dates sug- nins or modern . The YTT dragon (Varanus komodoensis). A gest that Homo erectus survived the date has been used as evidence of a similar faunal association is seen at arrival of Homo sapiens for a signifi- pre-YTT eastward dispersal of Homo the Soa Basin sites about 700,000 151 cant time. It is reasonable, neverthe- sapiens. years earlier. However, Liang Bua less, to reject the earliest and latest uniquely preserves a very small steg- ages as anomalous. The bulk of the Eastern Island Arc odon (Stegodon florensis insularis), ages suggest that the hominins at thought to be the dwarfed descend- Wae Racang Karst, Flores Middle Solo terraces generally relate ant of the large-bodied EPR-arrival, to the Last Interglacial (MIS 5e), fall- On northwestern Flores, the Man- Stegodon florensis, as well as a ing just before, during, or just after. garri limestone massif (Miocene age) diminutive hominin (Homo floresien- The general consensus is that the extends more than 500 m above sea sis).153 The origin and unique mor- Middle Solo terraces represent the level (Fig. 3B). The local karst system phology of Homo floresiensis has latest form of ISEA Homo erectus.140 began developing about 600 ka as been the subject of numerous inter- The question is open as to whether the Wae Racang River incised more pretations (Box 6). or not arriving Homo sapiens than 100 m into the massif. Five Excluding the hominin fossils, the encountered Homo erectus of the river terraces record this evolution. Liang Bua fauna is characterized by Ngandong type.142 The Liang Bua cave site is the result phylogenetic continuity and low ARTICLE Early Hominin Biogeography in Island Southeast Asia 203

Box 6. Homo floresiensis Skeleton

The Liang Bua skeleton (LB1), joint263;shortenedfemurandtibia; A decade after discovery, the evo- the smallest known fossil hominin, trapezoid, scaphoid, and capitate lutionary processes responsible for expresses a mosaic of primitive and wrist bones that are primitive and the small size and unique morphol- derived features that are absent in resemble those of Homo habilis ogy of the LB specimens are still other early hominin groups.262 (OH8)266; and a foot exhibiting debated.265 An extreme interpreta- Fragmentary remains have been primitive features including long tion relates LB1 to a pathologically recovered from at least nine individ- lateral toes, a short hallux, and the dwarfed Homo sapiens individual. uals.263,264 Our description of Homo absence of a well-defined medial Jacob and colleagues suggest that floresiensis is based on partial skele- longitudinal arch, resulting in flat LB1 represents a “pygmoid austral- ton LB1 and mandible LB6. LB1 feet.267 The only relevant wrist ele- omelanesian” with developmental exhibits a marked reductive trend in ment known for Homo erectus in abnormalities.179 Other hypothe- its facial skeleton, with extremely either Africa or Asia is one damaged sized conditions include Laron syn- small overall cranial size, a primi- lunate from .268 It is drome,269 cretinism,270 and Down tive low and anteriorly narrow vault therefore impossible to compare the syndrome.271 However, LB1 mor- shape with thin cranial bones, a rel- LB1 wrist morphology with that of phology does not definitively reflect atively prognathic face, and small Homo erectus. It is distinctly possi- any known systemic . Cur- teeth.184 Nevertheless, the mandi- ble that the wrist morphology of rently, the LB1 stratigraphic context bles of Homo floresiensis (LB1 and early Homo erectus resembled that is being reevaluated.50 A probable LB6) are buttressed, as seen in the of Homo habilis (OH8), and thus older last occurrence for Homo flor- holotype of Homo habilis (OH7).265 could have been a likely precursor esiensis at Liang Bua, in excess of The postcranial skeleton exhibits to Homo floresiensis.Thisismore 60 ka, would preclude the patholog- some primitive features, including plausible than a proposed transcon- ical Homo sapiens hypothesis. flared hipbones; short collarbone; tinental migration of Homo habilis. forwardly positioned shoulder species richness, as well as by rela- Homo sapiens arrived in ISEA, Mijares began excavating at the tively large and small body sizes. likely including Flores, by 47 ka. At Callao Cave opening.156 In 2007, a These are features of “impoverished Liang Bua, nevertheless, modern Philippine-Australian partnership and disharmonic insular faunas” typ- humans appear after deposition expanded the work. The excavation ical of isolated oceanic islands.6 resumes at 12 ka. The Homo sapi- encountered a rich fauna including Within this context, the Homo flore- ens archeofauna sequence has fresh- native brown deer (Cervus mariannus; siensis skeleton exhibits small stat- water mollusks, including Thiaridae 90% of the identifiable bone frag- ure, a small brain, relatively long (Thiara granifera and Melanoides ments), Philippine warty pig (Sus arms, robust lower limbs, and long tuberculata) and Neritidae (Neritina philippensis), and an unspecified feet. In sum, Homo floresiensis skele- pulligera, Neritodryas cornea, Nerito- extinct bovid. Element representation tal features resemble those of small- dryas dubia, Septaria porcellana, and and fragmentation for the cervids bodied taxa on Flores and across the Clithon squarrosus).155 Modern indicated that both whole and partial Arc. Small bodies helped these forms humans also introduced a range of carcasses were brought into the cave remain successful as long-term mem- exotic animals to the island, includ- for further processing. Some bones 6,7 bers of insular Arc faunas (Box 2). ing the Sulawesi warty pig (Sus cele- show cut marks, but no stone tools 8 The early Liang Bua stone technol- bensis) and the Eurasian pig (Sus were found. ogy shows a radial or centripetal scrofa), long-tailed macaque (Macaca Near the base of the excavated area, reduction sequence similar to that fascicularis), Javanese porcupine Layer 14 yielded a carbonized breccia seen at Mata Menge during the Mid- with relatively dense fauna. Among 154 (Hystrix javanica), and masked palm dle Pleistocene. The stone tools civet (Paguma larvata).155 Only the fragmentary hominin limb bones, the are most closely associated with the breccias contained a complete third Komodo dragon (Varanus komodoen- butchered stegodon remains, primar- metatarsal. Two cervid teeth from sis), an opportunistic predator and ily dental and skeletal elements of Layer 14 were dated. U-series abla- scavenger, is represented throughout juveniles.155 Continuity in the stone tion on one tooth produced a mini- the archeofauna sequence. artifact technology, Mata Menge to mum age estimate of 66.7 ka. ESR on Liang Bua (800 kyr), strengthens the the other tooth corroborated the Cagayan Basin, Luzon image of Flores as an isolated early result.8 In 2014, further excavation in hominin territory.78,154 Archaeo- Callao Cave. The Cagayan’s eastern Layer 14 produced more hand and faunal continuity seems to corrobo- flank has a significant Miocene karst foot bones and a series of upper teeth. rate the trend. massif (Fig. 3C). In 2003, Armand The dentition is of archaic character.9 204 Larick and Ciochon ARTICLE

The initial interpretation of the be significantly older. This is a major overwater to the Arc, Eurasian fauna Callao metatarsal went in two direc- change in the archeological column. could have arrived from various tions. The small dimensions and grac- The most economical interpretation Sunda sources via current Java or ile morphology were linked to small- is that at 41 ka H. sapiens replaced Borneo.32,167 bodied Homo sapiens, including mod- an earlier population of archaic Alfred Wallace was the first to dis- ern Philippine Negrito populations. hominins.161 The cave environments cuss overwater fauna movements Alternatively, the fossil drew compari- of the Maros karst may represent the with waif dispersal among the more son with small early hominin species best preservation of levels indicating common means.168,169 Waifs are such as Homo habilis and Homo flore- the arrival of H. sapiens and the individuals or small groups swept siensis. The new Callao finds corrobo- extinction of an as-yet undetermined away from one environment and rate the early hominin comparison early hominin population. transported by water or air currents and strengthen the hypothesis for a to a new territory.170 Some terres- second colonization east of Wallace’s DISCUSSION trial forms, such as stegodon, giant Line. It is tempting to link the origins tortoise, and Komodo dragon could With basic space and time data in of these finds with the EPR-age Caga- float or swim following the prevail- hand, we can turn to basic issues of yan Basin dispersal involving homi- ing currents. Hominins more likely ISEA early hominin biogeography. nins, large-bodied stegodons, giant rafted, either accidently or purpose- Two assumptions guide us. First, , and other insular species. The fully.7 Overwater waif dispersal ren- early hominins arrived at ISEA as Australasian Tektite Impact age sug- ders unlikely any subsequent generalized hunter-scavengers inte- gests that such an event would date exchange with the source popula- grated within mainland Eurasian 171 to at least 800 ka. 162 mammalian faunas. As such, hom- tion. Such long-term isolation on small islands is the basis for insular Maros Karst, South Sulawesi inins directly responded to the move- ments of nonhuman fauna members. evolution. Some 80 km southwest of the Second, within insular contexts, Isolation Walanae sedimentary basin lies the mammalian faunas were particularly Maros limestone karst system. Here, sensitive to environmental processes, With the late Early Pleistocene local stream courses follow intersect- including dispersal, isolation, vicar- Transition (MIS 23 900 ka), the ing joints to form plateau-like hill iance, relict survivorship, and genetic glacial-interglacial pulse became lon- masses. In areas of maximum pla- drift. During a million-and-a-half ger and more extreme. Early Middle teau dissection, steep-sided towers years of shifting ISEA habitats, early Pleistocene interglacial high sea lev- 157 are isolated on alluvial plains. The hominins responded to many insular els may have been the first to isolate towers have many caves and rock opportunities and constraints. Here, hominin groups across ISEA, espe- shelters, some of which have prehis- we address current issues relevant to cially in the Arc. The Solo, Soa, and toric archeology. the dispersal and isolation of early Cagayan basins show major faunal Recent excavations in several caves hominins and their extinction in turnovers during MIS 23-20. EPR have exposed cultural levels extend- relation to the arrival of Homo also marks the last fossil evidence of ing back more than 35 ka. Stone sapiens. Early Pleistocene Homo erectus as technology is based on small bipolar defined at Trinil and Sangiran. With cores that can resemble very small Dispersal the Mid-Brunhes Event (MIS 11 bifaces. Archeofaunas have small Overwater Transit. Even at maxi- 450 ka), regional karst systems mammals, including monkeys mum sea-level lowstands, deep-water began to develop. By MIS 9 (300 (Macaca sp.), bear cuscus (Ailurops channels separated Sunda and East- ka), caves were accumulating early sp.), and Celebes warty pigs (Sus cel- ern Island Arc landmasses. Heading hominin living debris and becoming ebensis). Freshwater gastropods east from Java, the Lombok and an important record of fossil and (Tylomelania perfecta), fish, and birds Komodo Straits ranged from 20 to archeological evidence. By the Last 158,159 are also represented. - 35 km at maximum lowstands.163 Interglacial (MIS 5e 125 ka), ISEA based is now dated to 39.9 Farther north, between Borneo and early hominin groups were appa- ka.160 Sulawesi, the was rently isolated in at least three areas. At one Maros cave, an upper never less than 40-km wide. Sulawesi Two lines of evidence illuminate archeological complex ranges back and Flores are separated by a mini- some details of Middle and Late to 41 ka. Well below, a lower mum of 60 km of sea with cross- Pleistocene isolation. assemblage is quite different. The currents (Fig. 8). It is unlikely that a stone technology is based on mini- dry-land connection ever existed Stegodontidae. The extinct family mally reduced cores and retouched across the Mindoro Strait to connect Stegodontidae comprised Asian rela- cobbles yielding flakes and debitage. Borneo with Luzon via Palawan.164 tives of mammoths and modern ele- The fauna has and Stegodon Presently, strong north-to-south phants. The fossil diversity of and no smaller species. The levels flow-through currents assist in iso- stegodon species is greatest near contain no . The deposits lating populations on either side of Yunnan, in southern China, their are older than 53 kyr ago and may Wallace’s Line.165,166 In crossing presumed area of origin. Pliocene ARTICLE Early Hominin Biogeography in Island Southeast Asia 205

Figure 8. ISEA early hominin sites in bathymetric and geographic context including all geographic and a site names used in this paper. We followed Huxley’s modification of Wallace’s Line208 as illustrated in Cooper and Stringer204. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] 206 Larick and Ciochon ARTICLE

TABLE 6. Pleistocene ISEA Stegodon-Artifact Co-occrrence Chart

Region Island Site Geological context Technology Age Reference Sunda Java Bukuran Sangiran Fm shell S. elephantoides 1.6 Ma 19,32 Arca Flores T. Talo1W. Sege Ola Bula Fm flake S. sondaari 1.3 Ma 48 Arc Sulawesi Cabenge Walanae Basin flake S. sompoensis Early Pleist 48 Arc Luzon numerous Cagayan Basin LFA S. luzonensis 0.8 Ma 96 Arc Luzon numerous Arubo Basin LFA S. luzonensis 0.8 Ma 96 Arc Timor Atambua Weaiwa Fm flake S. timorensis n/a 79,83 Arc Timor Motaoan Weaiwa Fm flake S. timorensis n/a 87 aEastern Island Arc fossils are found across southern can–Ngawi and Trinil (if later), did morphology, the LB1 skeleton seems China and . Pleistocene fossils not decrease in size, nor is there to show signs of insular evolutionary are known in North China, South- indication of endemic dwarfing in development. We can imagine that east Asia, and India.172,173 In Pleisto- Sunda stegodon. Homo floresiensis is the insular result cene ISEA, Stegodon became a There are several associations of overwater dispersal of either of dominant faunal element in large- of stegodon fossils with early hominin two forms, a small-bodied arrival and small-bodied forms.85,174,175 artifacts, especially east of Wallace’s that became further specialized or a Large-bodied Stegodon trigonoce- Line. Minimally, such co-occurrences larger-bodied arrival that became phalus is first documented in the suggest that on the smaller Arc land- dwarfed. middle Sangiran Formation (c. 1.6 masses, early hominins and stegodon- At discovery, LB1’s remarkably Ma) of Java.27 Presumably from tids frequented similar habitats. small (417 cc) caused some Sunda, and with the aid of sea-level Maximally, as may be the case at researchers to argue that LB1 was a lowstands, Stegodon dispersed Liang Bua, the association indicates microcephalic modern human178 or a throughout ISEA. Earlier Pleistocene that early hominins scavenged or pygmoid Australomelanesian modern dwarfs appeared on Flores (S. son- hunted these proboscideans. Loosely human with developmental abnormal- daari), Sulawesi (S. sompoensis), speaking, the stegodon-artifact co- ities179 (Box 6). There is now convinc- Timor (S. timorensis), Sumba (S. occurrence sequence appears to ing evidence that LB1 followed an sumbaensis), and Mindanau (S. reflect early hominin dispersal across insular evolutionary path for reduced mindanensis).32 Later in the Pleisto- ISEA. The dispersal and isolation of brain size demonstrated in other cene, on Flores, a medium-bodied S. Pleistocene ISEA stegodon may there- mammalian lines. Three recent stud- florensis probably gave rise to the fore serve as a heuristic model for ies bear directly on Homo floresiensis insular development among ISEA Late Pleistocene small-bodied S. flor- brain size: island-dwarfed hippos; early hominins. Table 6 summarizes ensis insularis. The latter is known foxes, mice, and humans; and callitri- this evidence. from the upper levels of Liang Bua, chids (marmosets and tamarins). With EPR (MIS 24-22, 1.0-0.9 where dwarf hominins may have Weston and Lister180 compared scal- Ma), glacial period sea-level low- hunted or scavenged this and other ing models for dwarfed hippos and stands and aridity became more dwarfed forms.155 their mainland ancestors. Dwarfed extreme. For large mammals, includ- Dwarfing among ISEA Stegodonti- species have significantly smaller ing hominins and stegodon, aridity dae began during the earlier Pleisto- brains, in relation to cranial size, than may have prompted dispersal while cene and continued throughout the predicted from scaling mainland sea-level lowstands enabled it. EPR epoch.32 The great preponderance of thus provided the first real means to forms. Observing brain size reduction dwarfed forms is found east of Wal- in multiple mammalian lines, leave Sunda for the Arc and to set 181 lace’s Line. Here, stegodon size up Arc islands up first as glacial Schauber and Falk concluded that reduction may be related to the period refugia, then as interglacial the Homo floresiensis brain could be increasingly high interglacial high- period bottlenecks. A similar EPR proportionally dwarfed from a larger- stands commencing with the EPR. forcing effect, to provide for early bodied ancestor having similar rela- Body size reduction may in part hominins and elephants (Elephas tive brain size. Montgomery and 182 explain the long-term success of and Mammuthus) dispersing in tan- Mundy correlated callitrichid brain select large mammals, including dem, has been suggested for south- size reduction with a slowdown of the 176 stegodon and hominins in the Arc. ern Europe.60,61,177 prenatal growth rate. Based on the West of Wallace’s Line, on Sunda, callitrichid example, the brain size of there is no evidence of size reduction Homo floresiensis insularity. Among Homo floresiensis may have been sub- among hominins. Between the EPR the larger vertebrates, small-bodied ject to selection pressure at early (MIS 23 900 ka) and the Last Inter- species have relatively more success stages of development. glacial (MIS 5e 125 ka), the Java in overwater dispersal and better The LB1 wrist has been compared hominins, as indicated by cranial survival potential on resource-limited with that of Homo habilis,known and dental fossils at Sambungma- islands.170 With its small, specialized exclusively from (Box 6). ARTICLE Early Hominin Biogeography in Island Southeast Asia 207

Figure 9. Hypothetical ISEA hominin lineages based on fossil, paleogenomic, and archeological evidence: yellow 5 Homo sapiens; red 5 Homo neanderthalensis; blue 5 Homo erectus and derived lines. On the Homo erectus line, archeological evidence provides divergence ages for Flores, Sulawesi, and Luzon early hominin lineages, presumably from Java origins. show genomic introgressions to dispersing Homo sapiens from Eurasian Neandertals and, possibly, from ISEA Homo erectus-derived lines. Since is considered a descendant of Homo erectus, we assume that the paleogenomic evidence carried by Homo erectus is similar to that recorded recently at Sima de los Huesos in .198 [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.]

Late Pliocene or earliest Pleistocene conditions. If a larger-bodied Dma- Modern Humans ISEA dispersal of a small African hom- nisi hominin arrived at Flores in the The arrival date for Homo sapiens inin has thus been suggested.183 Two same time frame, island dwarfing is at ISEA and Sahul (New Guinea and lines of evidence weaken the Africa implicated. In island contexts, ungu- Australia) is coalescing on a thresh- connection. First, LB1 has advanced lates can develop significantly short- old of 47 ka.189 ISEA’s two well- craniodental features not seen in any ened limb bones, shortened dated early Homo sapiens sites fall African early hominin.184,185 Second, metapodials, and stiffer joints, some- 187 on or near the threshold: 47 ka at there is no fossil evidence across main- times resulting in fused elements.  land Eurasia for a Homo habilis dis- Moreover, small-bodied hominins the (Palawan, Philip- pines)97 and 42 ka at Niah Cave persal out of Africa. ( and Homo) tend to 190 Alternatively, Homo floresiensis have short, -like lower limbs as a (Borneo, Indonesia). Alternatively, represents a diminutive Eurasian function of body size scaling.188 Wadjak (Java, Indonesia), once con- arrival. Here, the Dmanisi fossils Currently, we conclude that LB1 sidered the earliest ISEA modern (Republic of , southwest Eur- can best be seen as the product of human, has been radiometrically 191 asia) reflect an Early Pleistocene (1.8 insular effects on a representative of dated to 37–28 ka. Regarding Ma) paleodeme186 as a potential the Dmanisi paleodeme. Likely path- Sahul, a much larger area with many ISEA donor. Dmanisi Homo erectus ways include the specialization of more archeological sites, age deter- features include a small brain (546– small-bodied arrival or the dwarfing minations fall into a 47-40-ka inter- 189 780 cc capacity) and small body of a larger-bodied arrival. Either val. The 47-ka arrival threshold (145-166 cm height, 40-50 kg way, LB1’s small and specialized suggests dispersal out of Africa weight).3 LB1 cranial capacity (417 skeletal morphology reflects three toward the onset of MIS 3 (59 ka), cc) is not significantly smaller than features of Pleistocene Flores fauna: when renewed warmth and wetness that of Dmanisi skull 5 (D4500) (546 phylogenetic continuity, low species made the normally arid areas of cc). A small-bodied representative of richness, and disharmony. “All three northwest Africa and Arabia more the Dmanisi paleodeme could have aspects stem from the isolated posi- habitable than during the preceding dispersed eastward across Wallace’s tion of the island and have resulted MIS 4.192,193 Such conditions may Line at 1.3-1.0 Ma. Once on Flores, in the distinct morphological charac- have initiated the primary H. sapiens its wrist and foot specialized to local teristics of the Flores fauna.”6 out-of-Africa dispersal.194–196 208 Larick and Ciochon ARTICLE

At present, the latest ISEA early CONCLUSION Liang Bua skeletal morphology likely hominin fossil occurrences predate represents development in relation to Among all paleoanthro- the 47 ka H. sapiens arrival thresh- low rates of resource availability and pological areas, ISEA is unique in its old: Ngandong Homo erectus at predator stress. The diversification and great range of maritime environ- 143-130 ka,140,141,189 Liang Bua dwarfing of ISEA Stegodontidae pro- ments. Pleistocene glacial eustasy Homo floresiensis at 60 ka,50 and vides a relevant large mammalian fossil  gave the region a fast-changing char- the Callao hominin at 66.7 ka.8 In analogue. Liang Bua archeology indi- acter during the early hominin sum, current evidence does not sug- period. The current geography repre- cates social and technological organi- gest significant overlap between sents an extreme sea-level highstand zation within which a very small early and modern hominins. with near maximum marine cover human could consume large verte- and terrestrial isolation. Such high- brates. Parallels may be drawn with Paleogenomics. Paleoge- stands typified relatively short peri- Late Pleistocene western Eurasia, nomics provides an emerging line of ods of the Pleistocene. The generally where Homo neanderthalensis showed evidence for understanding the rela- longer glacial lowstands, ranging specializations in skeletal morphology tionship between ISEA early and mod- from 20 m to 125 m below pres- and stone technology. The Last Inter- ern hominins during the Late ent sea level, produced more contin- glacial may have isolated ISEA early Pleistocene (Fig. 9). Denisovan uous terrestrial exposures and the hominin and stegodon populations to have been identified in widely situated means for mammalian dispersal. their evolutionary limits. The early Eurasian Pleistocene contexts: extreme Nevertheless, highstands were criti- hominin period apparently ended with southwest and relatively early (Sima de cal for isolating large mammalian the arrival of Homo sapiens. los Huesos, 430 ka) and far northeast fauna, including early hominins. and much later (Denisova, 60 197–199 The spatial context for ISEA early ACKNOWLEDGMENTS ka). The Sima mitochondrial hominin biogeography centers on This long-term project was funded sequence is identified as Denisovan Wallace’s Line. For more than two by the L.S.B. Leakey Foundation (4 and is distinct from that of Neander- million years, glacial eustasy, tec- 197,199 grants), the Wenner- Gren Foundation tals. The Denisovan mtDNA tonic uplift, and volcaniclastic depo- for Anthropological Research (2 shares, for at least one million sition have structured and grants), and the National years, a last common ancestor with the restructured dispersal routes and Foundation (2 grants). Continuing sup- leading to Homo sapiens and province habitats east and west of 197 port has also come from the Human Neandertals. The nuclear sequence the line. On Sunda, fossil evidence Evolution Research Fund at the Uni- shows a closer relationship to Nean- suggests that early Homo erectus 200 versity of Iowa Foundation. New fund- dertals. The Denisovan genome is arrived from mainland Eurasia dur- ing from the Center for Global and thus associated with several Eurasian ing a low sea-level stand before 1.6 201 Regional Environmental Research early hominin lineages. The genome Ma (Solo Basin, Sangiran Forma- (CGRER) at The University of Iowa also shows evidence of another, even tion). On the Arc, stone tools indi- 197,202 older hominin lineage. cate that hominins arrived at the Soa and the Ann and Gordon Getty Foun- In modern human populations, (Flores) and Walanae (South Sula- dation is being used to continue and Denisovan alleles appear in two wesi) basins well before 1.0 Ma and expand this research. For research widely spaced areas of Eurasia: to the Cagayan and Arubo basins assistance, we thank K. Lindsay Eaves, 203 Tibet and east of Wallace’s Line, (Luzon) probably by 800 ka. Natalie O’Shea, Kiran Patel, Toby Ava- specifically in the Philippines and The evidence of interglacial isola- los, Chloe Daniel, and Madeleine Hoof- Sahul islands of New Guinea and tion is more subtle. By 300 ka, Sunda nagle. Gregg Gunnell and Jonathan 204,205 Australia, and in . groups inhabited caves and used flake Bloch carefully reviewed the manu- Given this far eastern distribution, tools to process rhinoceros, tapir, and script. For our many years of field col- Cooper and Stringer argue that Deni- cervids (Gunung Sewu karst). Liang laboration, we gratefully acknowledge sovan alleles introgressed into Homo Bua cave opened for hominin habita- Yahdi Zalm, Yan Rizal, and A. Aswan sapiens populations east of Wallace’s tion by 195 ka, when the Flores (Institute of Technology-Bandung, Line.204 Cooper and Stringer propose hominin used an advanced flake tool Java), E. Arthur Bettis Ill (University of Homo heidelbergensis as the east-dis- industry in pursuit of the islands’ Iowa), Gregg Gunnell (Duke Univer- persing Denisovan-carrying early endemic large fauna (Wae Racang sity), Frank Huffman (University of hominin. However, with no fossil karst). Most of the later evidence Texas, Austin), and John-Paul Zonne- evidence for this taxon in ISEA, the postdates the Last Interglacial (MIS veld (University of Alberta). issue remains open. The ISEA Late 5e). 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