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Journal of 47 (2004) 25e63 www.elsevier.com/locate/jhevol

A for late prehistoric

a,) a,b c David A. Burney , Lida Pigott Burney , Laurie R. Godfrey , William L. Jungersd, Steven M. Goodmane, Henry T. Wrightf, A.J. Timothy Jullg

aDepartment of Biological Sciences, Fordham University, Bronx NY 10458, USA bLouis Calder Center Biological Field Station, Fordham University, P.O. Box 887, Armonk NY 10504, USA cDepartment of Anthropology, University of Massachusetts-Amherst, Amherst MA 01003, USA dDepartment of Anatomical Sciences, Stony Brook University, Stony Brook NY 11794, USA eField Museum of Natural , 1400 S. Roosevelt Rd., Chicago, IL 60605, USA fMuseum of Anthropology, University of Michigan, Ann Arbor, MI 48109, USA gNSF Arizona AMS Facility, University of Arizona, Tucson AZ 85721, USA

Received 12 December 2003; accepted 24 May 2004

Abstract

A database has been assembled with 278 determinations for Madagascar. Materials 14C dated include pretreated sediments and plant macrofossils from cores and excavations throughout the island, and bones, teeth, or eggshells of most of the extinct megafaunal taxa, including the giant , hippopotami, and . Additional measurements come from uranium- dates on and thermoluminescence dating of pottery. Changes documented include late climatic events and, in the late , the apparently human- caused transformation of the environment. Multiple lines of evidence point to the earliest human presence at ca. 2300 14C yr BP (350 cal yr BC). A decline in megafauna, inferred from a drastic decrease in spores of the coprophilous fungus Sporormiella spp. in sediments at 1720 G 40 14C yr BP (230e410 cal yr AD), is followed by large increases in particles in sediment cores, beginning in the SW part of the island, and spreading to other coasts and the interior over the next millennium. The record of human occupation is initially sparse, but shows large human populations throughout the island by the beginning of the Second Millennium AD. Dating of the ‘‘’’ megafauna, including pygmy hippos, elephant , giant tortoises, and large lemurs, demonstrates that most if not all the extinct taxa were still on the island when humans arrived. Many taxa overlapped chronologically with humans for a millennium or more. The extinct lemurs stenognathus, Pachylemur insignis, pithecoides, and Daubentonia robusta, and the elephant birds spp. and Mullerornis spp., were still present near the end of the First Millennium AD. ingens,

) Corresponding author: D.A. Burney, Department of Biological Sciences, Fordham University, Bronx NY 10458 USA, Tel.: 718.817.3674; Fax: 718.817.3645. E-mail addresses: [email protected] (D.A. Burney), [email protected] (L.P. Burney), [email protected] (L.R. Godfrey), [email protected] (W.L. Jungers), sgoodman@fieldmuseum.org (S.M. Goodman), [email protected] (H.T. Wright), [email protected] (A.J.T. Jull).

0047-2484/$ - see front matter Ó 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2004.05.005 26 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 edwardsi, and sp. (cf. edwardsi) may have survived until the middle of the Second Millennium A.D. One specimen of Hippopotamus of unknown provenance dates to the period of European colonization. Ó 2004 Elsevier Ltd. All rights reserved.

Keywords: Madagascar; ; ; dating; paleoecology; megafauna; 14C; human settlement

Introduction Holocene is apparently less complete (see Burney, 1999). Less than two thousand separate our It has been known for well over a century that world from that of the extinct giant lemurs from Madagascar’s biotic and human are quite Madagascar. This is one of the last great habitable distinct from those of other nearby lands, notably landmasses to be colonized by humans and the Africa. Madagascar is a biogeographic entity un- only place on the planet in the late Quaternary to like any other on earth. It has many features of have a strepsirrhine-rich megafauna. , soil, and climate that make it a sort of The sequence of natural and human events in ‘‘Africa in miniature,’’ yet its long isolation in the late prehistoric Madagascar is also interesting from western Indian Ocean gives it an almost entirely a global perspective because this is the last place independent -track through the Cenozoic. In a on earth in which a diverse mammalian megafauna land without native ruminants, canids, or anthro- was extinguished prehistorically. The impact of poids, Nature has forged a unique ecological human colonization on continents and most other and evolutionary configuration in which at least large islands occurred much earlier. The relatively 17 of extinct lemurs, all larger than fresh record of paleoecological change in Mada- and some as large as , shared a diverse gascar deserves focused exploration through an floristic landscape with many species of smaller intensive program of and other strepsirrhines, as well as the largest birds that ever techniques for measuring time on relevant scales. lived, and a grazing maintained by giant The age range of dates from individual sites tortoises and three species of Hippopotamus varies markedly. The majority of sites (Godfrey and Jungers, 2002; also see articles in containing extinct megafauna that have been dated Goodman and Benstead, 2003). are late Holocene. A few open sites in the high- What happened next was also unprecedented: lands and caves at widely scattered locations Iron Age people found their way to Madagascar contain . Both cores and and settled in this strange new land, long after excavations have provided dates concerning the most of the habitable world except , earliest evidence for humans and subsequent Hawaii, and a few other remote islands had changes in the biota. A few regions of the island, been colonized by prehistoric humans. Almost notably portions of the humid eastern lowlands, are nothing is known about how or why they came to not well represented by prehistoric sites of any type. Madagascar from distant parts of the Indian In the west and southwest of the island, on the other Ocean, how they lived in the beginning, or even hand, two fortuitous sites, Belo-sur-Mer (Ankili- how they blended with subsequent waves of behandry Site) and Ambolisatra/Andolonomby, colonists to become the uniquely Malagasy people have yielded paleontological, archaeological, and (Dewar and Wright, 1993). We do not even know paleoecological information in an integrated con- for sure whether the earliest visitors to the island text with stratigraphic resolution of the key period were Asians or Africans. of transformation since human arrival (Burney, In an effort to better understand the timing 1999; Burney et al., 2003). Anjohibe Cave in of these remarkable transformations, we report the northwest and Ampasambazimba in the central here new Accelerator (AMS) highlands have also yielded a multiplicity of 14C dates for events and taxa, and present for dated evidence, but their chronology for the late the first time a compilation of all published and D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 27 unpublished absolute-age determinations for Ma- dates and other bone dates from the early liter- dagascar known to us. It is hoped that, with careful ature not specifying treatment received the mini- attention to the limitations of each pretreatment mum score (1). In addition, a scale factor has been protocol, material dated, and dating method, we included for assessing the age specificity of pre- may generate at least the broad outlines of the treated bulk sediment dates from cores. For island’s biotic history over the last few tens of details of the rating system, see Footnote 2 in the millennia, focusing particularly on refining the Appendix. In general, we mention in the text any chronometry of certain key events in the wake of caveats concerning the dating evidence presented the island’s relatively recent invasion by our own for a particular event. species. Radiocarbon dates have been calibrated using INTCAL98 (Stuiver et al., 1998). Some dates from the literature were originally reported with earlier Methods calibration sets that differ slightly from the range given here. Dates are reported as conventional The chronological inferences presented here are radiocarbon years ( generally corrected for isotopic drawn from a corpus of 278 dates from sites fractionation) G 1s ( yr BP), followed by the throughout the island (Figure 1; Appendix). These calibrated range at 2s (cal yr AD, BC, or BP). were determined on samples from caves and open Many dates from the literature prior to the 1980s paleontological sites, as well as additional dates on have probably not been corrected for isotopic sediment cores for paleoecological studies, and fractionation, and some later papers do not indi- dates on material from archaeological sites. Sub- cate whether this has been done or not. However, stances radiocarbon dated include bones, teeth, since most dates are on material that generally fecal pellets, and eggshells of vertebrates and pre- gives a d13C close to the standard, correction of treated sediments and plant macrofossils from these dates would change them only very slightly. cores. Also reported are 11 U-series determina- tions on speleothems and 16 thermoluminescence measurements on pottery. See individual citations Results and discussion and the ‘‘Comments’’ section in the Appendix for methods specific to individual dates. Climate and vegetation change before humans A numerical rating system for dating reliability is adapted from Mead and Meltzer (1984), with Although Madagascar has undoubtedly experi- modifications based on more recent information enced many profound shifts in climate and vegeta- and the requirements of the present study. For tion over the Cenozoic, details only begin to emerge instance, eggshell is available in many sites in in the dated records of the latest Pleistocene (Table Madagascar, and is rated highly in our scheme (6 1). Some of the earliest dated evidence for climate points) because experimental studies (Long et al., and vegetation change in Madagascar comes from 1983) suggest that this is a dating material rela- Lake Tritrivakely in the highlands (Burney, 1987a). tively free of -reservoir and contamination Subsequent longer cores to 40 m from this crater problems. Bone pretreatment issues, as lake extend beyond the range (ca. 40 kyr) of the discussed in Stafford et al. (1991) and Hedges and radiocarbon method (Gasse et al., 1994; Gasse and Van Klinken (1992), are reflected in our rating Van Campo, 1998, 2001; Williamson et al., 1998). system. A high score (5) is awarded to dates based At this 1778 m ASL site, the crater’s sediments on collagen purified by alkali pretreatment or record approximately 150 kyr of pollen and diatom gelatinization, or inspected visually after acid evidence, based on comparison to isotopic varia- pretreatment and confirmed to yield a large amount tions in ice cores from Antarctica. of pure white collagen. Bone collagen dates from Evidence from this and other sites in the high- material with poor collagen yield or humate dis- lands (reviewed in Burney, 1996a; Burney, 1997a; coloration received a low score (3). Whole-bone Straka, 1996) shows that the typical full-glacial 28 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 29 : Figure 1. Location of dated sites in Madagascar. A few are omitted for which specific location is unknown. KEY: 1. Imerina sites, including archaeological sites of Amboatany, Ambohidahy, Ambohinanjakana, Ambohitrikanjaka, Angavobe, Ankadivory, Fanongoavana, and coring site of Anosizato 2. Alaotra ( paleoecological) 3. Ambohitratroandro ( paleoecological) 4. Ambolisatra/Andolonomby (integrated) 5. Amboromena/Andonaham Pasipotsy (Andringitra) ( paleoecological) 6. Ampanihy/Ampoza ( paleontological) 7. Amparihibe ( paleoecological) 8. Amparihingidro ( paleontological) 9. Ampasambazimba (integrated) 10. Anakao ( paleontological) 11. Anavoha ( paleontological) 12. Andavambatobe (archaeological) 13. Andrafiabe/Antsiroandoha, Ankarana ( paleontological) 14. Andrahomana ( paleontological/archaeological) 15. Andraikiba/Andraikibakely/Miangola ( paleoecological)/Masinandraina ( paleontological) 16. Andranosoa (archaeological) 17. Anjohibe/Anjohikely/Lavakasaka (integrated) 18. Ankazoabo ( paleontological) 19. Ankilitelo ( paleontological) 20. Antsetsindrano (archaeological) 21. Antsirabe ( paleontological) 22. Baie des Galions ( geological) 23. Belo-sur-Mer (integrated) 24. Bemafandry ( paleontological) 25. Benavony ( paleoecological) 26. Bevoha ( paleontological) 27. Irodo (archaeological/paleontological) 28. Itampolo ( paleontological/archaeological) 29. Itasy/Kavitaha ( paleoecological) 30. Komango ( paleoecological) 31. Lakaton’i Anja (archaeological) 32. Lamboharana ( paleontological/archaeological) 33. Lohavohitra, Vonizonga (archaeological) 34. Mahery ( paleontological) 35. Mahilaka (archaeological) 36. Manambovo River Mouth/Talaky (archaeological/paleontological) 37. Mananjary ( paleontological) 38. Maroaloke ( paleontological) 39. Matsabory Ampozalana ( paleoecological) 40. Mitoho ( paleontological) 41. Mitsinjo ( paleoecological) 42. Nosy Mangabe (archaeological) 43. Ranobe ( paleoecological) 44. Sandrakatsy (archaeological) 45. Sarodrano (archaeological) 46. Taolambiby ( paleontological/archaeological) 47. Torotorofotsy ( paleoecological) 48. Tritrivakely ( paleoecological) 49. Tsiandroina ( paleontological) 50. Tsirave ( paleontological) 51. Tsiroanomandidy ( paleoecological) 52. Vohimasina/Vohitrarivo (archaeological) 30 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

Table 1 Chronological highlights of late prehistoric Madagascar Time Event >40e20 kyr BP Full glacial conditions, with domination of central highlands by ericoid bush. Mean temperatures >4(C colder than today. Maximum dryness recorded at LGM. 17,000 cal yr BP Deglaciation marked by warming conditions, beginning with partial replacement of ericoid bush by savanna and woodland in highlands. 9800 cal yr BP Holocene-type vegetation, probably a mosaic of wooded grassland and woodland, has replaced Pleistocene vegetation in the highlands. ca. 5000 cal yr BP Sea level reaches approximate present height. ca. 4000 cal yr BP Climatic desiccation in SW, continuing through rest of Holocene. Woodland vegetation on increase in central highlands due to higher effective moisture and lower temperatures. 2300 cal yr BP (350 cal yr BC) First evidence for humans, in form of modified bones of extinct megafauna and pollen of introduced Cannabis/Humulus. 230e410 cal yr AD Drastic decline of coprophilous Sporormiella fungus spores in sediments, due to reduced megafaunal densities. Abrupt increases in charcoal particles follow in SW sites, signaling increased human impact on the local landscape. Charcoal increase comes later in other regions, as humans spread up the west coast and into the interior. 780e1010 cal yr AD Earliest indirect evidence for proliferation, based on Sporormiella resurgence in NW. ca. 1000e1400 cal yr AD Charcoal particle evidence suggests deforestation was underway in the lowland rain forests. Islamized Indian Ocean traders active on coasts. Peak development of the entrepot of Mahilaka on the NW coast, followed by collapse. Sporormiella resurgence in highlands suggests pastoralism has reached this area. 1500 AD Portuguese explorer Diogo Dias lands on N shore of Madagascar. vegetation, down to 1000 m or less, was ericoid have been dated to as much as 40.1 kyr BP using bushland with composites and grasses. The con- uranium-series dating on speleothems containing figuration was quite different from today, with microfossils (Burney et al., 1997). They show a much of the high interior dominated by vegetation Medemia palm savanna vegetation in this region that is now found primarily on isolated summits north of Mahajanga quite similar to the vegetation above 1900 m. At (LGM) at the site today, with only minor change through around 20,000 years ago, lowland forest would the LGM and into the Holocene. The late Pleisto- probably have been restricted to refugia along the cene vegetation of Madagascar’s arid SW is not east coast and in the northwest of the island, with known, as published pollen evidence for this much of the high interior simply too dry and region reaches back only five millennia (Burney, cold (O4(C lower mean annual temperature than 1993a). However, the highly endemic taxonomic today; Gasse and Van Campo, 1998). A much status of this region’s flora would support the idea lower atmospheric CO2 concentration could also of long persistence of these ecosystems, although have contributed to this vegetation pattern (Street- wetter episodes such as in the mid-Holocene and Perrot et al., 1997). In any case, the biogeographic previous interglacials might have driven some of implications of this major difference between Pleis- the xerophytic species into refugia periodically, tocene glacial and Holocene interglacial Mada- promoting (Burney, 1993a; gascar is profound, as montane species isolated in Burney, 1996a). various high regions of the island today would have Throughout Madagascar, there is a variety of probably been prevalent over a much larger area, evidence suggesting that LGM desiccation was and many forest taxa may have been found only in widespread. Lake Alaotra, a large shallow lake in a very restricted area of the more humid lowlands humid eastern Madagascar, was almost certainly (Burney, 1996a; Goodman and Ganzhorn, 2004). greatly reduced in area if not completely dry ca. On the west coast, a few pollen spectra from 20 kyr BP (Reyes, 1993; Burney, 1997a). Lake the seasonally dry area around Anjohibe Cave Tritrivakely in the central highlands registers a D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 31 dry phase at this time, as does the bog known as Pollen of tree genera such as Celtis rise at this time, Matsabory Ampozalana on the Mt. d’Ambre in perhaps as higher elevation savannas were being the far north (Burney, 1987b). colonized by more trees, at least along water- At Tritrivakely, Ericaceae pollen drops from courses (Burney, 1987a; Gasse and Van Campo, 86% at 16,900 cal yr BP to 39% at 16,500 cal yr 1998). BP (Gasse and Van Campo, 1998), signaling the Spores of the coprophilous fungus Sporormiella, onset of warming postglacial conditions. Romani a proxy for megafaunal density, were higher in the et al. (2002) present evidence that glacial or perigla- late Holocene (up to 15% of pollen sum + spores) cial conditions were present on the Andringitra in semiarid coastal savannas than in any other Massif at LGM, and sedimentological changes habitats for which the spore has been measured. In prior to 11,580 G 145 yr BP (14,030e13,150 cal yr the woodland/grassland mosaic of the central BP) show that warming conditions had freed Lake highlands, spore-inferred megafaunal densities Amboromena (2400 m elevation) from the ice by appear to have been moderate. In humid forests this time. and montane habitats, the spore occurs only as At Tritrivakely, a brief return to cooler condi- a trace before humans (Burney et al., 2003). The tions is noted at 16,000e15,100 cal yr BP (Gasse spore occurrences may primarily reflect densities of and Van Campo, 1998). Later at this site, pollen hippos and other non- megafaunal taxa. evidence shows the nearly complete replacement, Whatever the ecological preferences of the ex- by 9800 cal yr BP, of ericoid heath vegetation with tinct taxa, one fact is becoming increasingly clear pollen of plants typical of wooded grassland or from the growing corpus of dates: major climate a grassland/woodland mosaic (Burney, 1987a; and vegetation changes of the late Pleistocene and Gasse and Van Campo, 1998). prehuman Holocene were survived by most, if not Outside these palynological studies, remarkably all, the known megafauna. Despite strong evidence little is known about the late Pleistocene biota, for range shrinkage in the late Holocene for some except for a few dates on extinct from three lemurs with very limited distributions today sites. The density of data increases greatly for (Vuillaume-Randriamanantena et al., 1985; Jungers megafaunas and their environments in mid- et al., 1995; Simons et al., 1995; Godfrey et al., Holocene , probably owing to more favorable 1999), no prehuman disappearances, climate-driven conditions for fossil site formation along the pre- or otherwise, have been detected in the record for sent coastline. Along the west coast, for instance, these . Of the nine genera of extinct lemurs many lakes and swamps apparently formed only dated, only radofilai, for which only one after postglacial sea level had approached its date has been obtained, is not securely dated to the modern height, probably ca. five millennia ago. human period. Although some sites in the SW show Whether sea level exceeded present heights at least increasing desiccation, and perhaps local faunal briefly after this time is not known for certain (see declines, around two millennia ago (Mahe´ and Burney, 1993a). Sourdat, 1972; Burney, 1993a; Goodman and In any case, wetter conditions than today Rakotozafy, 1997), pollen records show that even prevailed along the semiarid SW coast in the more severe climate change has occurred many mid-Holocene, with the expansion of dry forest at times before on the island, with no discernible effect the expense of palm savanna and xerophytic com- on megafaunal extinction patterns. munities. Evidence for desiccation appears after ca. 4,000 cal yr BP in this region. From this time Human arrival and palynological signals until human arrival, the Andolonomby pollen record (Burney, 1993a) shows that palm savanna Many lines of evidence converge on the notion was expanding locally. that humans first settled in Madagascar about Meanwhile, in the central highlands, pollen evi- two millennia ago: 1) first occurrence of human- dence shows that cooler conditions had returned modified bones of extinct animals, 2) first occur- by 3500 cal yr BP, with higher effective moisture. rence of pollen of introduced Cannabis/Humulus 32 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

( probably the former; see Merlin, 2003), 3) sudden The very earliest date for a human presence, large increases in microscopic charcoal particles 2325 G 43 yr BP (2366e2315 cal yr BP), comes from above background values, and 4) an increase in the SW interior site of Taolambiby. It is from ruderal pollen and other vegetational disturbance a of the lemur Palaeopropithecus ingens indicators. These key events have all been dated to with cut marks that suggest flesh removal with within three centuries of 2000 14C yr BP. Studies of a sharp object (Perez et al., 2003). This is approx- the Malagasy language also show a separation imately coeval with the earliest occurrence of from its closest surviving linguistic relatives in the Cannabis/Humulus pollen at Tritrivakely in the highlands of Borneo about two millennia ago central highlands, at an interpolated age of 2200 (Dahl, 1951). Divergence could have begun before cal yr BP (Burney, 1987a; Gasse and Van Campo, proto-Malagasy speakers departed from Indone- 1998). A human-modified femur of extinct Hippo- sia, of course. If people had arrived many centuries potamus from Ambolisatra on the SW coast yielded earlier, they would have predated the advent of the an age of 1970 G 90 yr BP (60 BCe130 cal yr AD) Iron Age. No plausible evidence has been found (MacPhee and Burney, 1991). Thus, a conservative for a stone-age culture on Madagascar (Dewar, estimate for the earliest human presence on 1984). This is merely a negative-evidence argu- Madagascar is ca. 2300 cal yr BP (350 cal yr BC). ment, but the persistence of stone artifacts in the This date is marked as a point of reference in Fig. 2 archaeological record is much better than iron, yet and subsequent figures. no stone tools except sinkers and musket-flints In the Andolonomby core from the vicinity of have been found. the SW fossil site of Ambolisatra, a plant

Figure 2. Radiocarbon dates for key events in the human prehistory of Madagascar, plotted as 2s calibrated ranges. D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 33 macrofossil date on the layer of sediments showing elephant birds would have been formidable prey. the first marked decrease of Sporormiella, a proxy Consequently, their population densities may have for the presence of megafauna (Burney et al., reached high levels under favorable conditions 2003), yielded 1720 G 40 yr BP (230e410 cal yr of climate and vegetation, so that edible forage AD). A similar spore decline, less securely dated, within reach may have cycled through the stom- was noted at Belo-sur-Mer, another important achs of megaherbivores too quickly for a large megafaunal site, in sediments 10 cm above a layer standing crop of plant litter to accumulate, thus dated 1990 G 50 yr BP (100 cal yr BCe110 cal yr imposing a limit on the flammability of the land- AD) on acid-treated organic bulk sediments. scape under ordinary circumstances. It would appear that humans interacted with Dating of charcoal spikes, in which values megafauna, although the evidence for this is scant, increase at least one or two orders of magnitude as it is in many other places experiencing compar- above background, has permitted some inferences able extinction events around the globe (Martin, regarding the rate of transformation of the re- 1984). If Sporormiella is indeed a reliable proxy for cently human-dominated landscape. This change megafaunal density, then humans had a noticeable in the fire ecology of the island shows several impact within a few centuries of arrival. However, interesting patterns. First, it is time-transgressive, the numerous taxa that yield much more recent showing up nearly two millennia ago in the south- terminal ages (see below) show that megafaunal west, then spreading up the west coast to wetter decline and megafaunal extinction were not coeval, climates, appearing in the central highlands about and in fact the two may be separated by at least 1400 yr BP at Lake Kavitaha (Burney, 1987c) and one millennium for many taxa. within a century or two later at the higher Lake Although causes for the decline and extinction of Tritrivakely site (Burney, 1987a). Charcoal increase the megafauna have long been debated (reviewed in does not appear at Benavony, a humid interior Burney, 1999), fire has often been invoked as lowland site in the Sambirano rain forest of the a major factor in the transformation of Madagas- northwest, until 710 G 110 yr BP (1050e1430 cal car’s landscapes. Radiocarbon dating of sedimen- yr AD). This conclusion is based on an acidealkali tary charcoal particles is problematic due to treated bulk sediment date that may well be slightly inherent-age and old-carbon reservoir effects (Bur- too old, since some older carbon released by the ney et al., 2003). The stratigraphy of Ambolisatra/ fires may have been incorporated. Andolonomby and Belo-sur-Mer, however, show It is possible that Madagascar was first settled that the Sporormiella-inferred decline of megafauna in the semi-arid southwest, and that over the next slightly predates the concomitant rise in micro- approximately seven centuries people gradually scopic charcoal particles. In both profiles, ca. 50 cm settled the other coasts and the central highlands. of sediment separates the two events, although they Humid forests of the low interior may have been are apparently too close in time for good discrim- the last areas settled (or at least burned), perhaps ination by 14C measurement (Burney et al., 2003). because of limitations posed by such human Fire incidence may have increased at these sites, diseases as plague, malaria, or dysentery (Burney, and perhaps throughout Madagascar, due to the 1999). decline of such large herbivores as hippos, ratites, Another interesting pattern in the charcoal and tortoises. The loss of grazers and browsers results is for charcoal to increase rapidly at the could have resulted in excessive plant litter accu- presumed time of the advent of local human mulation, leading to devastating fires. Before the activity, remain high for a few centuries, then advent of humans, adults of these taxa may have gradually decline to moderate values. This may be a lacked predators other than perhaps the largest global trend on remote islands in the wake of human crocodiles (Brochu, 2000). Juveniles of these spe- arrival, but it is not known certainly whether this cies and large lemurs may have also been vulner- trend is driven by ecological, microclimatic, or able to the largest viverrids (Goodman et al., socioeconomic dynamics, or some combination 2004), but the cow-sized adult hippos and half-ton (Burney et al., 1994; Burney, 1997b). 34 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

It is important to keep in mind that people did with the pottery on the deflated surface. The not introduce fire to Madagascar. Sediments of charcoal could well have been from a natural brush mesic sites of the center, north, and northwest fire. The site has since been destroyed by a cyclone. contain large amounts of particulate carbon even Northeast Madagascar has yielded many occu- tens of millennia before human arrival (Burney, pation sites of the latest First Millennium and 1987b; Matsumoto and Burney, 1994). The pri- early Second Millennium (reviewed in Dewar and mary effect of humans on the pre-existing fire Wright, 1993). A stratified occupation site on the ecology was to further increase burning, at least shelf island of Nosy Mangabe contained a layer for a while, and to spread fire to environments that with a date of 1250 G 60 yr BP (660e990 cal yr are naturally too wet or too dry to support fre- AD) on charcoal associated with coarse ceramic quent fires before the arrival of humans. Very dry and chlorite schist sherds and iron slag. ecosystems often lack sufficient ground litter to Within a few centuries of the beginning of the support a natural fire ecology. Second Millennium, evidence for human occu- pation is far more common in many parts of Madagascar. For example, in the small Gorge Population growth and ecological transformation d’Andavakoera (Dewar and Rakotovololona, 1992; Dewar and Wright, 1993), there are more It is worrisome that there are no dated occupa- than 40 sites of mostly twelfth-thirteenth century tion sites during the first half millennium of sig- age, based on the occurrence of imported ceramics nificant indirect evidence for a human presenced of a known age range. These sites contain few including bones cut in an apparently fresh state; remains of domesticates, but many bones and introduced pollen, disturbance indicators, and shells of marine and terrestrial . Among the particulate charcoal in cores; and Sporormiella primate and tortoise remains are a few bones of decline. It is typical of many lands, however, that extinct species. These are ‘‘generally in a fragmen- indirect evidence for humans predates the earliest tary state, but obvious cut and butchery marks are definite occupation sites, since a founding pop- rare’’ (Dewar and Wright, 1993: 436). ulation may have considerable impact on the envi- Along the northeast coast in the vicinity of ronment while their living sites are still few in Irodo there are a number of sites that appear on number and difficult to detect archaeologically (see the basis of imported pottery to range in age from Burney et al., 1994). slightly earlier than the twelfth to the fourteenth The oldest direct evidence for occupation, pro- century, with possible smaller occupations a few bably temporary encampments for exploiting forest centuries earlier (Ve´ rin, 1975; Dewar, 1984). A few resources, comes from the Gorge d’Andavakoera dates have been obtained, but all present inter- on the north tip of the island (Dewar and pretive problems (Dewar and Wright, 1993). A Rakotovololona, 1992; Dewar and Wright, 1993). fragment of Aepyornis eggshell from one of these A cave in this valley, Lakaton’i Anja, contained sites that dates to 1150 G 90 yr BP (680e1030 cal several levels of faunal materials mixed with yr AD) confirms that the survived until then, pottery sherds. Upper levels include sgraffiato but it is not possible to determine whether its wares of the twelfth-fourteenth century. Below presence in the site was directly associated with this level there are similar deposits lacking im- human activity (Battistini and Ve´ rin, 1966). ported sherds. Two charcoal dates yielded 1680 G Settlements in the southernmost forests of the 65 yr BP (230e530 cal yr AD) and 1300 G 80 yr east coast are of similar age (Rakotoarisoa et al., BP (620e900 cal yr AD). 1993), as are sites such as Andranosoa, where Another possible early site, Sarodrano on the pastoralists and foragers occupied a large area southwest coast, yielded a charcoal date of 1460 G around the confluence of two rivers in the extreme 90 yr BP (410e710 cal yr AD). Battistini and Ve´ rin south of the island, and the nearby coastal site of (1971) were not certain that the charcoal was Talaky (reviewed in Dewar and Wright, 1993). At derived from human activity and contemporaneous Andranosoa, which yielded dates as early as D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 35

920 G 90 yr BP (1020e1210 cal yr AD) and as late and increased charcoal in sediments, was un- as 730 G 90 yr BP (1160e1410 cal yr AD), a pit derway by 960 G 90 yr BP (900e1210 cal yr AD; contained large quantities of bones of sheep/goat Burney, 1987c). This is in good agreement with and Tenrec ecaudatus and only a few cattle evidence from Fiekena ceramic assemblage sites remains. A carapace fragment from an extinct such as Ankadivory. People with a different pottery giant tortoise and hippo teeth were excavated from and other cultural distinctions from the an underlying terrace deposit nearby (Rasamuel, coasts were well-established in the area by the 1984). thirteenth and fourteenth centuries, based on ra- Although dated evidence for earliest introduc- diocarbon and thermoluminescence dates, and con- tion of livestock to Madagascar is not available, firmed by some imported pottery, mostly of Far indirect evidence in the form of a resurgence of Eastern types (Wright et al., 1992; Rakotovolo- Sporormiella probably can be used to assess cattle lona, 1994). arrival in Madagascar, as it has in North The first European known to have visited America (Davis, 1987). At the Amparihibe crater Madagascar was Diogo Dias, a Portuguese ex- lake on the shelf island of Nosy Be, a plant plorer, in 1500 AD. History records that, over the macrofossil date of 1130 G 50 yr BP (779e1020 next three centuries, Portuguese, Dutch, French, cal yr AD) on the Sporormiella increase in and English colonial ventures were attempted, and sediments is consistent with the settlement along pirates used the many remote harbors as bases this coast by Islamized Indian Ocean traders. A from which to launch their mischief (Brown, similar signal from Lake Kavitaha in the high- 1979). Some of these European colonists and lands at 960 G 90 yr BP (900e1210 cal yr AD) adventurers, with their new technology, religions, suggests cattle may have spread to the people of and languages, were absorbed into the indigenous the interior highlands within two centuries or less. setting, contributing to the unique Malagasy cul- Pollen data from sites throughout the island tural configuration of today. Population growth (reviewed in Burney, 1997a) show that, during and land conversion have been particularly this period around the beginning of the second rapid since the mid-twentieth century (Green and millennium AD, woody vegetation was declining Sussmann, 1990), with the result that many of the and grasslands were on the increase, most likely surviving biotic communities are under severe through human agency. Urban-scale human threat from human pressures. population aggregations were occurring in some areas by this time, notably at the walled city of Mahilaka on the northwest coast, which thrived Last of the megafauna between the eleventh and fourteenth centuries (Radimilahy, 1997). This city of several thousand Intensified radiocarbon dating of apparent late people declined suddenly near the end of the occurrences of extinct giant lemurs, elephant birds, fourteenth century for unknown reasons. It has and hippos has brought into sharp focus the been suggested that plague, malaria, dysentery, or protracted nature of the subfossil . The other diseases that were rampant at this time in ‘‘extinction window,’’ defined as the duration of Eurasia and Africa might have limited human the period from earliest human evidence to last expansion in the more humid parts of Madagas- occurrence of extinct megafauna, shows an over- car (Burney, 1999), although no direct evidence lap between humans and some of these lost crea- has been found to support this idea. tures of up to two millennia. The extinct lemurs Evidence for very early settlements in the now Hadropithecus stenognathus, Pachylemur insignis, heavily populated central highlands has proven Mesopropithecus pithecoides, and Daubentonia elusive. Pollen and charcoal data (Burney, 1987c) robusta, and the elephant birds Aepyornis and point to local human impacts beginning in the late Mullerornis, were still present near the end of the First Millennium. Definite settlement activity, with First Millennium AD (Figure 3; Table 2). Dating pollen of agriculturally associated ruderal plants suggests that Archaeolemur sp. (cf. edwardsi), 36 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

Figure 3. A. Dated late occurrences for eight genera of extinct lemurs, plotted as 2s calibrated ranges. B. Dated late occurrences for extinct non-primate vertebrates, plotted as 2s calibrated ranges. D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 37

Megaladapis edwardsi, and Palaeopropithecus in- instance, and the extent of evidence for butchery gens survived at least until 830 G 60 yr BP (1040e has been slim, although this body of evidence con- 1290 cal yr AD), 630 G 50 yr BP (1280e1420 cal tinues to grow as more sites come to light (reviewed yr AD), and 510 G 80 yr BP (1300e1620 cal yr in Burney, 1999; also see Perez et al., 2003). AD), respectively. Burney (1999) advocated the use of a conceptual It is therefore not surprising that the French matrix approach to test the competing hypotheses colonial governor, Etienne de Flacourt (1661), for the extinctions. By tabulating all proposed reported eyewitness accounts of beasts in the hypotheses according to their expectations regard- remote interior that could have been giant lemurs, ing the extinction rate, its geographic pattern, and elephant birds, and hippopotami. Hippos in the inferred processes involved, it is possible to particular have cropped up in other colonial evaluate chronometric data for consistency with accounts from the eastern, central, and northwest- the unique combination of predictions for each ern parts of the island as late as the ninenteenth hypothesis. Direct evidence for the process by and early twentieth centuries (Godfrey, 1986; which the extinction occurred, which is regrettably Burney and Ramilisonina, 1998). Dates on a large scarce, may thus be augmented by looking at the hippo of unknown provenance (an unlabeled landscape-level parameters of time and space as specimen at the Universite´ d’Antananarivo that displayed by radiocarbon-dated occurrences. may have been collected at the eastern site of Table 3 provides an updated version of this Mananjarydor perhaps brought over from analysis of hypotheses. Although each of these has Africa) were as recent as 99 G 36 yr BP and been proposed for Madagascar, the degree to 213 G 40 (combined calibrated range 1639e1950 which the expectations are articulated in the cal yr AD). Ethnographic data collected in the literature varies. All of these have likewise been Belo-sur-Mer region include putative eyewitness proposed for other regions with late prehistoric accounts of a hippolike as recently as 1976 megafaunal extinctions, notably North America (Burney and Ramilisonina, 1998). (Burney and Robinson, in review). Hypotheses H1, H3, and H5 each predict that the time period of overlap between humans and Implications of chronological refinement extinct megafauna at a given site with suitable for extinction models stratigraphic and temporal resolution should be very short, probably a century or less (essentially The general outlines of human activity and the 95% confidence interval for AMS 14C dates in biotic change are summarized in Figures 2 and 3, this time-frame), because the entire extinction event although many questions about cause and effect in should advance as a rapid wave (‘‘Blitzkrieg,’’ H3) Madagascar’s transformation are left unanswered or occur simultaneously throughout the island by the present state of chronometric analysis. The (‘‘Great Fire,’’ H1, and ‘‘Hypervirulent Disease,’’ earliest humans on Madagascar saw the remark- H5). Two other hypotheses (‘‘Great Drought,’’ H2, able megafauna, and the coincidence of their and ‘‘Synergy,’’ H6) allow for a slower process temporal overlap is in itself strong circumstantial (102e103 yr), with regional and mosaic patterns, evidence for a human hand in the disappearances. respectively. Hypothesis H4 is not explicit as to This inference is rendered more compelling by the how long the process of environmental degradation apparent lack of extinctions in the wake of and postulated extinctions caused by cattle and documented major natural vegetation changes in other introduced herbivores should take at a given the late Pleistocene and Holocene. location or island-wide, but comparisons to histor- The argument over causes for the extinctions and ically recorded biological invasions on other islands rapid transformation of the island have generally (Atkinson, 1989; Burney, 1996b; Vitousek et al., centered around the lack of direct evidence for 1996) would suggest that population build-up of a human role (see Dewar, 1984). Human occupa- feral herbivores would be moderately rapid (101 yr tion sites from the earliest period are unknown, for locally, 102 yr island-wide). 38 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

Table 2 14C dates for of Madagascar (see Appendix for sources and details) Taxon Age Calibrated Site Comments BP G 1s range at 2s Archaeoindris fontoynontii 2291 G 55 2362e2149 BP Ampasambazimba -sized ‘‘sloth lemur.’’ Dates show that the largest of all extinct lemurs was extant when humans arrived. Archaeoindris fontoynontii 2402 G 45 2711e2338 BP Ampasambazimba cf. Archaeolemur 830 G 60 AD 1047e1280 Anjohikely Latest dated occurrence of ‘‘monkey lemur’’ ; from fecal pellet associated with juvenile skeleton. Archaeolemur sp. 1020 G 50 AD 990e1180 Antsiroandoha Archaeolemur cf. majori 1370 G 40 AD 650e780 Belo-sur-Mer Archaeolemur sp. 1550 G 50 AD 435e650 Belo-sur-Mer Archaeolemur majori 1650 G 50 AD 370e575 Belo-sur-Mer Archaeolemur majori 2050 G 60 BC 165eAD 120 Belo-sur-Mer Archaeolemur cf. edwardsi 2060 G 70 BC 332eAD 110 Anjohikely Archaeolemur cf. edwardsi 7790 G 70 8740e8410 BP Anjohibe Babakotia radofilai 4400 G 60 5290e4840 BP Antsiroandoha This medium-sized ‘‘sloth lemur’’ is the only extinct genus not yet dated to the human period. Daubentonia robusta 1065 G 40 AD 891e1027 Beloha Date on the giant aye-aye shows its late survival.

Hadropithecus stenognathus 1413 G 80 AD 444e772 Belo-sur-Mer Latest dated occurrence for this ‘‘monkey lemur.’’ Hadropithecus stenognathus 2194 G 70 2344e1998 BP Tsirave Hadropithecus stenognathus 6724 G 54 7660e7490 BP Andrahomana

Hapalemur simus 4560 G 70 5460e4980 BP Andrafiabe Date shows that the extant bamboo lemur had a range in the mid-Holocene that extended to the northernmost part of the island. Megaladapis edwardsi 630 G 50 AD 1280e1420 Ankilitelo Latest dated occurrence for this large-bodied koala lemur. Megaladapis grandidieri 1035 G 50 AD 900e1040 Ampasambazimba Presumably whole-bone radiometric date may not be reliable. Megaladapis edwardsi 1277 G 35 AD 666e816 Beloha Megaladapis sp. 1591 G 60 AD 339e609 Site unknown Megaladapis sp. 1815 G 60 AD 27e412 Bevoha Megaladapis madagascariensis 2140 G 50 2320e1990 BP Belo-sur-Mer Megaladapis madagascariensis 2713 G 44 2870e2760 BP Taolambiby Megaladapis sp. 4566 G 35 5436e5059 BP Andrahomana Megaladapis grandidieri/ 12,760 G 70 15,670e14,380 BP Andrafiabe madagascariensis Megaladapis grandidieri/ 26,150 G 400 Antsiroandoha Oldest date for a lemur; poor collagen madagascariensis. quality suggests this may be a minimum age. Mesopropithecus pithecoides 1410 G 40 AD 570e679 Ampasambazimba Latest dated occurrence for this genus of ‘‘sloth lemurs,’’ smallest of the extinct lemur taxa. Mesopropithecus globiceps 1694 G 40 AD 245e429 Tsiandroina Mesopropithecus globiceps 1866 G 45 AD 58e247 Belo-sur-Mer Mesopropithecus globiceps 2148 G 35 BC 354e60 Ankazoabo Pachylemur insignis 1220 G 50 AD 715e985 Belo-sur-Mer Latest dated occurrence for this large relative of the ruffed lemur. D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 39

Pachylemur insignis 2000 G 50 BC 45eAD 145 Belo-sur-Mer Palaeopropithecus ingens 510 G 80 AD 1300e1620 Ankilitelo Latest dated occurrence for this large, extremely suspensory ‘‘sloth lemur.’’ Calibrated range overlaps the period of European contact. Palaeopropithecus ingens. 1269 G 80 AD 640e946 Ankazoabo Palaeopropithecus ingens 2008 G 90 2292e1729 BP Belo-sur-Mer Palaeopropithecus ingens 2285 G 40 2352e2157 BP Ampoza Palaeopropithecus ingens 2325 G 43 2366e2345 BP Taolambiby Cut marks on bone interpreted as evidence for butchery. Propithecus verreauxi Modern Taolambiby Extant with cut marks on bone interpreted as evidence for recent butchery. These marks are very similar to those described by Perez et al. (2003) for extinct lemurs at the same site.

Inferring processes, as opposed to rates and either a long coexistence between domesticated patterns, in the extinctions (see last column of herbivores and native herbivores, or the disap- Table 3) has required an indirect approach, taking pearance of the natives before the appearance of care to avoid the logical trap of negative evidence. introduced species in the fossil record, would Hypotheses H1 through H5 each postulate a single refute H4, although Dewar (1984, 1997) and cause that would presumably be sufficient to Burney (1993a,b, 1999) have suggested that bi- produce the rates and patterns observed. Detecting ological invasions may be more important as actual cause and effect in the fossil record, a contributing cause rather than the exclusive cause however, is challenging and prone to circular in the decline of native species. Recent Sporor- logic. Showing, for instance, that there is archae- miella results (Burney et al., 2003), in combination ological evidence for human hunting of the with the large number of late-surviving megafau- megafauna in no way proves that hunting caused nal species documented here, suggest that, in fact, the extinctions. Ironically, under a strict Blitzkrieg the major decline of native animals may have come regime (Martin, 1984; also see Grayson, 1984), well before livestock proliferated. Livestock do abundant evidence for overhunting would not be overlap with the persistent taxa that survived into an expectation, as the extinction wave would pass the Second Millennium AD, such as Megaladapis any given point too rapidly to leave an extensive edwardsi, Palaeopropithecus ingens, Archaeolemur archaeological record of the event. Direct refuta- sp. (cf. edwardsi), and Daubentonia robusta among tion is thus probably out of the question, although the large lemurs, as well as the hippos and perhaps one must ask how much evidence for hunting the ratites, so that competition between native and would be necessary to show that overhunting was introduced herbivores, or more diffuse effects from not a major factor (i.e., that hunting was sustain- the landscape modifications associated with pas- able). toralism, may have played a role in finishing off If a rich site like Ankilibehandry (Belo- some species. sur-Mer), which probably contains fossils from Synergy (H6), like all combinatorial hypothe- the entire period of the ‘‘extinction window’’ ses, poses special problems for refutation, owing to (Burney, 1999), also shows that some of the the multifactorial nature of the inferred causes. megafauna sustained hunting pressure and other However, the integrated site analysis approach has human impacts for nearly a millennium, then offered a partial solution. With subphreatic Blitzkrieg and Hypervirulent Disease models have excavation and integrated site analysis (Burney, been refuted. Similar logic would suggest that 1999; Burney and Robinson, in review), it has been 40 D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63

Table 3 Extinction hypotheses proposed for Madagascar No. Type Source Rate Pattern Process H1 ‘‘Great Fire’’ Humbert (1927) Rapid Simultaneous Landscape transformed by throughout introduction of fire by humans; extinctions followed as a result of forest loss. H2 ‘‘Great Drought’’ Mahe´ and Slow Southern Climatic desiccation, extinctions due Sourdat (1972) region only to spread of semiarid conditions. H3 ‘‘Blitzkrieg’’ Martin (1984) Rapid Wave across island First-contact overkill of naı¨ ve fauna, beginning at coastal settlements and moving inland. H4 ‘‘Biological Invasion’’ Dewar (1984) Moderately Multiple waves Introduced herbivores disrupt natural rapid across island vegetation and compete with native herbivores. H5 ‘‘Hypervirulent Disease’’ MacPhee and Very rapid Panzootic Unknown pathogen(s) introduced by humans, Marx (1997) disease pattern spreading quickly; lethal to wide array of , and possibly some birds and reptiles. H6 ‘‘Synergy’’ Burney (1999) Very slow Mosaic Full array of human impacts played a role, but to differing extents in various regions; factors interacting to multiply effects; some amplification by background climate change.

to some extent possible to tease apart the evidence Therefore it is difficult at this to specify in the sediments for fire (H1), climate change (H2), what such an event would look like in the fossil overhunting (H3), and biological invasion (H4). If record, except that one would predict by these ‘‘event horizons’’ specified by the unicausal reference to known virulent diseases that an hypotheses H1-H4 occurred at different times, as essentially instantaneous die-off would occur at the radiocarbon evidence indicates, and all the first human contact. Instead, the radiocarbon extinctions correlate with a particular one, then evidence suggests that humans, and presumably synergy is unlikely to be important. If, on the other any diseases they might have brought, over- hand, as has been suggested here, these four lapped with many of the extinct taxa for at least events, or some combination of a subset, coincide one, and in some cases nearly two, millennia. temporally with disappearance of some megafau- The rather sudden decline in megafaunal densi- nal species, the case is strengthened (but, of course, ties within a half millennium of human arrival, not ‘‘proved’’dsee Popper, 1969) for a likely as inferred from the Sporormiella results, prob- enhancement of the roles of individual factors ably supports models that suggest rapid (but through synergistic interaction. For instance, only partial) overkill by humans, followed by anthropogenic deforestation may make hunting a protracted array of human impacts working less sustainable; biological invasions may involve against smaller surviving populations. To opt introduced predators or herbivore competitors instead for a disease explanation requires that that exacerbate the effects of human hunting one must argue for an unknown disease or and/or deforestation. diseases having a different epidemiology from No disease mechanism of the type specified by any known disease, knocking down the bulk of MacPhee and Marx (1997), lethally infecting the population but somehow preventing rapid a wide range of mammals, birds, and reptiles recovery of the immunized survivors to previous and transported by humans or their introduced numbers. commensals (H5), has been documented for There is considerable likelihood that the data Madagascar or any other part of the world. presented here will not fully resolve the scientific D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 41 controversy over the cause of these extinctions. fragmentation, and local extirpation. The late Building and carefully dating the large, precise, occurrences of some of the taxa documented are multiproxy data sets needed in order to address also consistent with the notion that the ‘‘extinc- some of the questions surrounding the landscape tion window’’ is unfortunately not yet closed. transformations and extinctions is more likely to Today in Madagascar, the same negative synergies provide resolution, or at least clarification, than of resource overutilization, fire-mediated vegeta- any conceivable alternative. A detailed mechanism tion change, and biological invasion continue to for the Synergy Hypothesis was proposed in swirl about, pulling smaller lemur species and Burney et al., (2003). Sporormiella results, in a host of other organisms into the extinction combination with charcoal particle analysis and vortex. precision dating of both sediments and bones of extinct megafauna, suggest additional details con- cerning Madagascar at the brink of human trans- formation. What is novel in these data sets is evidence Acknowledgements suggesting that the late prehistoric extinctions in Madagascar were not a single, brief event with This research was supported by NSF BCS- a simple unicausal explanation, but rather a more 0129185 (to DAB, LRG, and WLJ), BCS-0237388 complex and drawn-out process in which human- (to LRG), and EAR-0115488 (to AJTJ), and perturbed systems underwent a series of drastic conducted under collaborative agreements with phase transitions. By combining the dated evi- the De´ partement de Pale´ ontologie et d’Anthropo- dence from , , and paly- logie Biologique, Universite´ d’Antananarivo, and nology, it is reasonable to hypothesize that: 1) the the Acade´ mie Malgache. For dates first reported megafauna was probably concentrated in wooded here, we thank the curators and caretakers of the savannas, a plant community type rare in Mada- following collections from which dating samples gascar today; 2) the large biomass were collected: Laboratoire de Pale´ ontologie des decreased sharply within a few centuries of first Verte´ bre´ s, Universite´ d’Antananarivo (UA; Ar- evidence for humans in the locality; 3) immediately mand Rasoamiaramanana, Gise` le Randria); Aca- following this ecological collapse, a new fire regime de´ mie Malgache, Parc Tsimbazaza, Antananarivo, emerged that was likely a response to plant Madagascar (AM; C. Rabenoro, L. Rakotozafy); biomass buildup following decline of the grazer- Natural History, Oxford, United Kingdom (OX- browser regime; 4) the vegetation was transformed UM; Thomas Kemp, Malgosia Nowak-Kemp); to less productive systems (depauperate steppe the Swedish Museum of Natural History (S. grasslands and closed forest) with feedbacks to the Steunes); and the University of Massachusetts local climate system and the simpler, reorganized Anthropological Primate and Comparative Col- biota. These ideas are consistent with changes lection. Alan Walker donated specimens for documented for other areas, and help explain the dating, and Anne Yoder provided unpublished extent of the great transformations observed for dates. Nancy Beavan of the Rafter Radiocarbon Madagascar ( for details see Burney, 1999; Burney Laboratory (New Zealand) supplied advice on et al., 2003). pretreatment protocols and interpretation of Whatever the specific reconstruction of mega- results. Darden Hood of Beta Analytic, Inc. faunal paleoenvironments, the dating evidence (Miami, FL, USA) assisted with calibration of presented is also consistent with the hypothesis that many older dates. Paul S. Martin gave useful the extinction event here, as perhaps elsewhere, is advice on dating priorities and interpretations. not a single event but a stepwise cascade of ex- Mirya Ramarolahy, Lydia Raharivony, Ramiliso- tinctions, as the synergistic combination of human nina, Toussaint Rakotondrazafy, and Natalie impacts and nonlinear natural responses creates Vasey assisted with collecting and subsampling a mosaic of environmental change, population materials for dating. 42

Appendix Catalog of age determinations for Madagascar Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Alaotra sediment/ 5190 G 110 BP BP 6250e5670 61e70 cm b-45005 ÿ16.30 17(25# S, Acid/alkali/ Reyes, 1993 3 + 2 Large lake radiometric (4x) ALO-2A + 48(30# E acid 2B 752 m Alaotra sediment/ 6830 G 110 BP BP 7860e7480 91e105 cm b-45854 ÿ18.10 17(25# S, Acid/alkali/acid Reyes, 1993 3 + 2 Large lake radiometric ALO-3C, 48(30# E

3D,3E 752 m 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Alaotra sediment/ 29,690 G 510 BP 191e200 cm b-31514 ÿ17.90 17(25# S, Acid/alkali/acid Reyes, 1993 3 + 2 Large lake radiometric (4x) ALO-4A 48(30# E 752 m Ambila-Lemaitso sediment/ not given AD 1030e1260 100e150 cm KI-3398 Straka, 2001 3? + 1 radiometric Ambinanibe, / 700 G 100 BP AD 1160e1420 Sondage I, LY-3911 25(04#S. Rakotoarisoa 5+2 Anosy radiometric Layer 4 46(57#E et al., 1993 5m Amboatany, potsherd/TL 1670 G 65 AD Operation D, DurTL-50-11(a) 18(46# S, Wright et al., TL Imerina Layer 4c AS 47(32# E 1992 Archaeological Amboatany, potsherd/TL 1480 G 100 AD Operation D, DurTL-50-11(b) 18(46# S, Wright et al., TL Imerina Layer 4c AS 47(32#E 1992 Archaeological Ambohidahy, Imerina potsherd/TL 1520 G 100 AD Sondage 1, DurTL-77-6 AS 18(46# S Wright et al., TL Archaeological Layer 2 47(31 #E 1992 Ambohidahy, Imerina potsherd/TL 1500 G 100 AD Sondage 1, DurTL-77-3 AS 18(46# S Wright et al., TL Archaeological Layer 2 47(31 #E 1992 Ambohidahy, Imerina potsherd/TL 1420 G 110 AD Sondage 1, DurTL-77-5 AS 18(46# S Wright et al., TL Archaeological Layer 2 47(31 #E 1992 Ambohidahy, Imerina potsherd/TL 1100 G 190 AD Sondage 1, DurTL-77-4 AS 18(46# S Exceptionally Wright et al., TL Archaeological Layer 2 47(31 #E early for human 1992 occupation on the

central highlands e 63 Ambohinanajakana, potsherd/TL 1780 G 50 AD Operation C, DurTL-77-2 AS 18(44# S Wright et al., TL Imerina Layers 9-10 47(35 #E 1992 Archaeological Ambohinanajakana, potsherd/TL 1720 G 50 AD Operation C, DurTL-77-1 AS 18(44# S Wright et al., TL Imerina Archaeological Layers 9-10 47(35 #E 1992 Ambohinanajakana, potsherd/TL 1650 G 70 AD Operation C, DurTL-50-14 AS 18(44# S, Wright et al., TL Imerina Archaeological Layer 2b 47(35# E 1992 Ambohinanjakana, charcoal/ 221 G 61 BP AD 1520e1950 Operation C, SMU-1522 18(44# S, Wright et al., 6+2 Imerina Archaeological radiometric Layer 7 47(35#E 1992 Ambohitratoandro sediment/ 3720 G 50 BP BP 4230e3910 160 cm b-28591 ÿ21.50 18(54# S, Acid/alkali/acid Burney, 3+2 Wetland radiometric AMB-160 46(39# E 1997a,b Ambohitrikanjaka, charcoal/ 380 G 90 BP AD 1410e1670 Site 1, Gif-5154 18(51# S, Wright et al., 6+2 Imerina Archaeological radiometric Excavation 2 47(38 #E 1992 Layer 3 Ambolisatra/Andolonomby plant 1720 G 40 BP AD 230e410 150 cm b-167657 ÿ26.10 23(04# S, Sporormiella Burney et al., 5+3 Shallow pond macrofossil/AMS ALB-150 43(45# E? decline 2003 2m (megafaunal collapse) Acid/ alkali/acid Ambolisatra/Andolonomby sediment/ 1890 G 90 BP BC 60eAD 90e100 cm b-24345 ÿ24.10 23(04# S, Acid/alkali/acid Burney, 1993a 3 + 2 Shallow pond radiometric (4x) 350 ALB-1A,B 43(45# E 2m Ambolisatra/Andolonomby Hippopotamus 1970 G 50 BP BC 60eAD MNIP MAD AA-2895 23(03# S, Human-modified MacPhee and 5+3 Shallow pond femur 130 1711 43(45#E? megafaunal bone Burney, 1991 (human-modified)/ AMS G e ( # Ambolisatra/Andolonomby bone/ 2915 170 BP BP 3470 2740 1.8 m AND- b-23149 ÿ24.50 23 04 S, Burney, 1993a 5 +3 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Shallow pond radiometric (4x) 87-2A,2B 43(45# E 2m Ambolisatra/Andolonomby sediment/ 4920 G 90 BP BP 5900e5470 306e326 cm b-24469 ÿ26.50 23(04# S, Acid/alkali/acid Burney, 1993a 3 + 2 Shallow pond radiometric ALB-2A,B,C,D 43(45# E (6-25) 2m Ambolisatra/Andolonomby bone/radiometric 4965 G 70 BP BP 5900e5590 2.6 m AND- bÿ23148 ÿ24.60 23(04# S, Burney, 1993a 5 + 3 Shallow pond 87-1 43(45# E 2m Amboromena Etched sediment 11,580 G 145 BP BP 14030e 165 cm Romani et al., 3+3 depression lake 13150 2002 Amboulitsate Paleolake/ Geochelone abrupta 750 G 370 BP AD 580e1960 Muse´ um BM-2125 ÿ20.90 23(04# S, Late occurrence Burleigh and 5+3 Wetland bony carapace/ National d’His- 43(45# E of extinct taxon Arnold, 1986 radiometric toire Naturelle 2m Collagen (Pale´ ontologie), extraction Paris. 1884-30 Ampamalora Aepyornis eggshell 1970 G 90 BP BC 190eAD not given not given Late occurrence Marden, 1967 6+2 (9 pieces 240 of extinct taxon (p.487) Sauer, collectively dated) 1972 (p.12) radiometric Ampangandiria, Astimo potsherd/TL 1490 G 55 AD Feature 1, 20(45#S, This paper TL Archaeological Sounding I 44(02#E Ampanihy unidentifiable bone 2430 G 100 BP BP 2760e2190 Hela-3854 22(19.8# S, Jernvall et al., 5? + 2 fragments not given 44(42.7# E 2003 660m e 63 Amparihibe Deep plant macrofossil/ 1130 G 50 BP AD 780e1010 156 cm AHB- b-67392 ÿ21.10 13(18# S, Sporormiella in- Burney, 1999 5 + 3 crater lake AMS 156 ETH-11460 48(13# E crease (livestock) Acid/alkali/acid Amparihibe Deep sediment/AMS 1660 G 90 BP AD 150e600 275 cm AHB- bÿ62509 13(18# S, Acid/alkali/acid This paper 3 + 3 crater lake 3:10-11 CAMS-6424 48(13# E Amparihibe Deep sediment/AMS 1780 G 70 BP AD 80e410 193 cm AHB- b-62508 13(18# S, Acid/alkali/acid This paper 3 + 3 crater lake 2:15-16 CAMS-6423 48(13# E Date rejected old carbon contamination Amparihibe Deep sediment/AMS 1820 G 60 BP AD 70e370 338 cm AHB- b-55059 13(18# S, Acid/alkali/acid This paper 3 + 3 crater lake 4-74 CAMS-3542 48(13# E Amparihingidro wood/radiometric 2850 G 200 BP BP 3460e2470 220 cm Gif-? Mahe´ and 3+2

Sourdat, 1972 43 (continued on next page) Appendix (continued) 44 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13 C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D.

Ampasambazimba (Itasy) ArchaeoindrIs 2291 G 55 BP BP 2362e2149 AM 6239 (type NZA-18519 ÿ23.50 18(54# S, Collagen extrac- This paper 5 + 3 Paleolake/Wetland fontoynontii crown specimen) R-28331/1 46(44# E tion with gelatini- of upper left p4/ DB -03-1 1200m sation AMS Ampasambazimba Megaladapis 1035 G 50 BP AD 900e1040 not recorded Pta-739 18(54 S, Tattersall, 1973 1? + 3 Paleolake/Wetland grandidieri tibia/ BM M 9927 46(44# E radiometric 1200 m Ampasambazimba Mesopropithecus 1410 G 40 BP AD 570e679 MP12, UA3926 NZA-18523 ÿ22.09 18(54# S, Collagen extrac- This paper 5 + 3 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Paleolake/Wetland pithecoides DB-03-8 R-28331/6 46(44# E tion with alkali proximal left 1200m femur/AMS Ampasambazimba Archaeoindris 2402 G 45 BP BP 2711e2338 AI4 DB-03-4 NZA-18520 ÿ23.40 18(54# S, Collagen extrac- This paper 5 + 3 Paleolake/Wetland fontoynontii R-28331/3 46(44# E tion with alkali proximal end of 1200m femur/AMS Ampasambazimba wood 4725 G 180 BP BP 5890e4870 unit 1, trench B GX-9989 18(54# S, MacPhee et al., 5+2 Paleolake/Wetland ca 1m 46(44# E 1985 1200m Ampasambazimba wood 7540 G 200 BP BP 8760e7960 unit 1,trench G GX-9836 18(54# S, MacPhee et al., 5+2 Paleolake/Wetland ca 1.8m 46(44# E 1985 1200m Ampasambazimba wood 8245 G 215 BP BP 9580e8580 unit 1, trench GX-9835 18(54# S, MacPhee et al., 5+2 Paleolake/Wetland G, ca 2.9m 46(44# E 1985 1200m Ampasambazimba Alopochen 22,550 G 170 BP UA-360 b-63116 18(54# S, Goodman, 5+3 Paleolake/Wetland sirabensis Bone 46(44# E 1999 fragments/AMS 1200m Ampoza Wetland/ Hypogeomys 1350 G 60 BP AD 620e770 b-72676 22(19# S, Goodman and 5+3 floodplain antimena 44(44# E Rakotondravony, 570m 1996 Ampoza Wetland/ unidentifiable 1705 G 55 BP AD 225e435 65 cm Hela-157 22(18.9#S, Jernvall et al., 5? + 2

floodplain bone fragments 44(42.3#E 2003 e not given 570m 63 Ampoza Wetland/ unidentifiable 1810 G 60 BP AD 70e380 110 cm Hela-156 22(18.9#S, Jernvall et al., 5? + 2 floodplain bone fragments 44(42.3#E 2003 not given 570m Ampoza Wetland/ unidentifiable 1850 G 55 BP AD 550e325 top Hela-158 22(18.9#S, Jernvall et al., 5? + 2 floodplain bone fragments 44(42.3#E 2003 not given 570m Ampoza Wetland/ Bones 1910 G 120 BP BC 190eAD GaK-2309 22(19# S, Goodman says Mahe´ and 1? + 3? floodplain 400 44(44# E this is giant Sourdat, 1972 tortoise Ampoza Wetland/ Geochelone 2035 G 35 BP BC 160eAD 50 B.M.N.H. (P) BM-1628 ÿ27.30 22(19# S, Late occurrence Burleigh and 5? + 3 floodplain abrupta R5890 44(44# E of extinct taxon Arnold, 1986 (cf. gigantea) 570 m Collagen / extraction radiometric Ampoza Wetland/floodplain Palaeopropithecus 2285 G 40 BP BP 2350e AMO-2 NZA-16997 ÿ20.60 22(19# S, Late occurrence This paper 5 + 3 cf. ingens/AMS 2160 R-28139/8 44(44# E of extinct taxon 570m Collagen extrac- tion with gelatini- sation Ampoza Wetland/floodplain Hippopotamus/ 2370 G 50 BP BP 2700e GrA-11626 22(19# S, Late occurrence This paper 5 + 3 AMS 2330 44(44# E of extinct taxon

570m 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Ampoza Wetland/floodplain Hippopotamus 2517 G 40 BP BP 2747e AMO-1 NZA-16996 ÿ21.57 22(19# S, Collagen extrac- This paper 5 + 3 lemerlei 2372 R-28139/7 44(44# E tion with gelatini- AMS 570m sation Ampoza Wetland/floodplain Hippopotamus/ 2760 G 60 BP BP 2980e2760 Pta-7728 22(19# S, This paper 5 + 3 AMS 44(44# E 570m Anakao, south of Tulear Aepyornis eggshell 2375 G100 BP BP 2740e2150 QC-971 23(39#S, Late occurrence Dewar, 1984 6+3 Coastal dunes 43(39#E of extinct taxon (p. 582) Anakao, south of Tulear Aepyornis eggshell 2775 G 95 BP BP 3160e2740 QC-970 23(39#S, Dewar, 1984 6+3 Coastal dunes 43(39#E (p. 582) Anakao, south of Tulear Aepyornis eggshell 3960 G 150 BP BP 4840e3980 QC-972 23(39#S, Dewar, 1984 6+ 3 Coastal dunes 43(39#E (p. 582) Anavoha Coastal dunes wood/radiometric 1954 G 110 BP BC 200eAD 3- 77 cm GaK-1654 Mahe´ and 5+2 30 Sourdat, 1972 Andavambatobe Cave charcoal/ 580 G 100 BP AD 1260e Sq 1, layer 8 b-55092 363.0- 42 cm. In Wright Wright et al., 6+2 radiometric 1480 77 cmbd 1125.7 (L) et al., 1996 1996 HFJ-92-10 Andonaham Pasipotsy Small sediment 7060 G 110 BP BP 8050e 170 cm 19(04# S, Romani et al., 3+3 glacial cirque 7670 46(17# E 2002 2120 m Andrafiabe Cave Hapalemur simus/ 4560 G 70 BP BP 5460e4980 b-62841 Simons et al., 5+3 AMS CAMS-6775 1995 Andrafiabe Cave Megaladapis/AMS 12,760 G 70 BP BP 15670e b-61703 Simons et al., 5+3 14380 CAMS-6394 1995 e 63 Andrahomana Cave Hypogeomys 4440 G 60 BP BP 5640e b-73370 25(11#55$ Goodman and 5+3 australis/AMS 4420 S, 46(37#- Rakotondravony, 59$E10m 1996 Andrahomana Cave Hadropithecus 6724 G 54 BP BP 7660e not given AA-45963 ÿ9.10 25(11#55$ This paper 5 + 3 stenognathus 7490 BJ-HS-1 T-16044A S, 46(37#- humeral fragment/ 59$E10m AMS Andrahomana Cave Hypogeomys 1536 G 35 BP AD 428e618 AHA-I, NZA-18996 ÿ20.00 25(11#55$ Collagen extrac- This paper 5 + 3 australis tooth/ Sq C,D 1,2 R-28421/1 S, 46(37#- tion with gelatini- AMS layer 1 59$E10m sation AHA-03-01 (continued on next page) 45 Appendix (continued) 46 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Andrahomana Cave Megaladapis sp. 4566 G 35 BP BP 5436e AHA-03-7 NZA-18997 ÿ20.10 25(11#55$ Old spoil NE of This paper 5 + 3 infant femur/AMS 5059 R-28421/7 S, 46(37#- AHA-G Collagen 59$E extraction with gelatinisation Andraikiba Deep crater lake sediment/ 1200 G 70 BP AD 670e990 76e90 cm ck b-23486 ÿ24.50 19(53# S, Acid This paper 3 + 2 radiometric depth 46(57# E AND-2A,B,C 1520 m Andraikiba Deep crater lake sediment/ 2310 G 100 BP BP 2720e2100 327e341 cm b-22837 ÿ25.90 19(52# S, Acid Burney, 1988 3 + 2 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. radiometric (2x) AND-4 A,B,C 46(57# E 1520 m Andraikiba Deep crater lake sediment/ 2320 G 70 BP BP 2690e2150 219e233 cm b-23487 ÿ25.90 19(52# S, Acid This paper 3 + 2 radiometric AND 3,A,B,C 46(57# E 1520 m Andraikibakely Pond sediment/ 6990 G 130 BP BP 8030e7590 ca. 110 cm b-67393 ÿ20.60 19(52# S, This paper 3 + 2 radiometric AKK-2A,B 46(58# E (extended) 1520m Andrakibaikely Pond sediment/ 27,970 G 540 BP ca. 381e390 cm b-31513 ÿ18.30 19(52# S, This paper 3 + 2 radiometric AKK-4 46(58# E 1520m Andrakibaikely Pond sediment/ 31,400 G 630 BP 786e822 cm b-14852 ÿ21.33 19(52# S, This paper 3 + 2 radiometric AKK-9 II 46(58# E A,B,C,D 1520m Andranosoa Archaeological charcoal/ 220 G 70 BP AD 1500e1950 iron- b-8094 24(46#S, Emphoux, 1981 6 + 2 radiometric feature south 45(35#E end of site 240m Andranosoa Archaeological charcoal/ 730 G 90 BP AD 1160e1410 undesignated Gif-4570 24(46#S, Early Emphoux, 1979 6+2 radiometric pit 45(23#E archaeological cited in Dewar, 240m site 1984 (p. 582) Andranosoa Archaeological charcoal/ 920 G 90 BP AD 970e1280 ‘‘fosse Gif-4571 24(46#S, Early Emphoux, 1979 6+2 radiometric d’odure’’ 45(23#E archaeological cited in Dewar, 240m site 1984 (p. 582)

Andranosoa Archaeological potsherd/TL 950 G 210 AD pit 1.02 mbs DurTL-75 4AS 24(46#S, Emphoux TL e 45(35#E unpublished 63 240m Angavobe, Imerina charcoal/ 330 G 80 BP AD 1430e1800 Sondage II, GaK-1058 18(55# S, Wright et al., 6+2 Archaeological radiometric Pocket I 47(44#E 1992 1780m Anjohibe Cave Eidolon dupreanum 330 G 70 BP AD 1430e1800 surface (19-8) b-56770 15(32.55# S, Burney et al., 5+3 postcranial bones/ CAMS-4255 46(53.17#E 1997 AMS 100 m Anjohibe Cave / 371 G 50 BP 28e38 mm from 15(32.55# S, Brook et al., 1999 U U-series top MA2 46(53.17#E 100 m Anjohibe Cave speleothem/ 1122 G 43 BP 84e90 mm from 15(32.55# S, Brook et al., 1999 U U-series top MA2 46(53.17#E 100 m Anjohibe Cave speleothem/ 3412 G 14 BP base MA 1 15(32.55# S, Brook et al., U U-series 46(53.17#E 1999 100 m Anjohibe Cave Hippopotamus cf. 3730 G 70 BP BP 4280e3880 Sq E3, level 2 b-55093 CAMS- 15(32.55# S, Burney et al., 5+3 lemerlei rib/AMS (19-5) HFJ-92- 3562 46(53.17#E 1997 11 100 m Anjohibe Cave speleothem/ !4.0 kyr BP 0-3 cm 15(32.55# S, Burney et al., U U-series ANJ-92-3 46(53.17#E 1997 100 m Anjohibe Cave Hippopotamus cf. 5300 G 60 BP BP 6270e5920 surface (19-8) b-64961 CAMS- ÿ31.80 15(32.55# S, Burney et al., 5+3 lemerlei / HFJ-93-2-B 8648 46(53.17#E 1997 AMS 100 m G e ( # Anjohibe Cave speleothem/ 6.52 2.64 kyr 0 2 cm ANJ- 15 32.55 S, Outermost layer Burney et al., U 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. U-series BP 92-2 46(53.17#E of collapsed 1997 100 m column Anjohibe Cave sediment/AMS 7040 G 70 BP BP 7970e 100 cm ANJ- b-60947 CAMS- 15(32.55#S, Burney et al., 3+3 7700 AM-2-100 5645 46(53.17#E 1997 100 m Anjohibe Cave sediment/AMS 7080 G 80 BP BP 8020e 248e252 cm b-58395 CAMS- 15(32.55#S, Burney et al., 3+3 7720 ANJ-AM-2-250 4628 46(53.17#E 1997 Anjohibe Cave speleothem/ 7340 G 46 BP base MA2-6 15(32.55#S, Brook et al., U U-series 46(53.17#E 1999 100 m Anjohibe Cave Archaeolemur cf. 7790 G 70 BP BP 8740e surface (19-8) b-64960 CAMS- ÿ27.50 15(32.55#S, Collagen extrac- Burney et al., 5+3 edwardsi pelvis/ 8410 HFJ-93-1-A 8647 46(53.17#E tion with alkali 1997 AMS 100 m Anjohibe Cave speleothem/ 22.88 G 4.48 kyr 0e2cm 15(32.55#S, Burney et al., U U-series BP ANJ-92-4 46(53.17#E 1997 100 m Anjohibe Cave speleothem/ 37.55 G 4.67 kyr ANJ-92-5 15(32.55#S, Date rejected due Burney et al., U U-series BP 46(53.17#E to ratio 1997 100 m Anjohibe Cave speleothem/ 40.11 G 7.21 kyr 100 cm 15(32.55#S, Center of col- Burney et al., U U-series BP ANJ-92-2 46(53.17#E lapsed column 1997 100 m Anjohibe Cave speleothem/ 69.6 G 7.3 kyr NCC-1 #3 15(32.55#S, Directly below This paper U U-series BP 46(53.17#E bone breccia in e 63 100 m collapsed-cave deposit Anjohibe Cave speleothem/ 86.8 G 7.8 kyr NCC-1 #4 15(32.55#S, Laminated sta- This paper U U-series BP 46(53.17#E lagmite on floor 100 m of collapsed- cave feature Anjohikely Cave cf. Archaeolemur 830 G 60 BP AD 1040e site 1-1 b-55060 15(33# S, Late occurrence Burney et al., 5+3 dung/AMS 1290 HFJ-92-13A CAMS-3543 46(53# E of extinct taxon 1997 Acid washes Anjohikely Cave Archaeolemur cf. 2060 G 70 BP BC 350eAD 80 site 31-1 b-55095 15(33# S, Late occurrence Burney et al., 5+3 edwardsi small HFJ-92-16 CAMS -3709 46(53# E of extinct taxon 1997 pieces of bone/ Collagen extrac-

AMS tion with alkali 47 (continued on next page) Appendix (continued) 48 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Ankadivory, Imerina charcoal/ 523 G 80 BP AD 1290e Sq. P 108/bV- b-23989 18(46# S Wright et al., 6+2 Archaeological radiometric 1500 Pit 3 47(34# E 1992 Ankadivory, Imerina charcoal/ 810 G 100 BP AD 1020e Sq. E 116/ SMU-2632 18(46# S Early archaeolo- Wright et al., 6+2 Archaeological radiometric 1390 O,Layer XVI 47(34# E gical site 1992 Ankadivory, Imerina charcoal/ 810 G 185 BP AD 880e Sq. E 116/ SMU-2077 18(46# S, Early archaeolo- Wright et al., 6+2 Archaeological radiometric 1440 O,Layer VII 47(34# E gical site 1992 Ankadivory, Imerina charcoal/ 880 G 50 BP AD 1030e Sq. P 108/b-Pit SMU-2361 18(46# S, Early archaeolo- Wright et al., 6+2 Archaeological radiometric 1260 10 47(34# E gical site 1992 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Ankadivory, Imerina pottery/TL 1295 G 140 AD Sq E 116/O, DurTL 97-3AS 18(46# S Early archaeolo- Wright et al., TL Archaeological Layer VII 47(34# E gical site 1992 Ankazoabo Cave Palaeopropithecus pr- 1269 G 80 BP AD 640e946 AM 6184 #11 NZA-10059 ÿ19.00 24(33# S, Late occurrence This paper 5 + 3 emolar/AMS R-24649/1 43(56# E of extinct taxon Collagen extrac- tion with gelatini- sation Ankazoabo Cave Mesopropithecus 2148 G 35 BP BC 354e60 AM 6531, UA NZA-18998 ÿ15.80 24(33# S, Univ. of Anta- This paper 5 + 3 globiceps tooth/ 4853 DB -03-5 R-28421/9 43(56# E narivo collection AMS Collagen extrac- tion with gelatini- sation Ankazoaraka sediment/ not given BP 3364e 260e280 cm KI-3223 9 m In Straka, 2001 Straka, 1993 3? + 2 radiometric 3049 this date is given as BC 1400 to 1040 yr Ankazomiranga sediment/ not given AD 1520e 10e20 cm KI-3854 290 kms Straka, 2001 3? + 2 radiometric 1670 of Tana 1330 m Ankazomiranga sediment/ not given BP 3479e 20e30 cm KI-3854 290 kms of Straka, 2001 3? + 2 radiometric 3259 Tana 1330 m Ankazomiranga sediment/ 12,200 G 90 BP BP 15,340e 120e140 cm KI-3455 290 kms of Straka, 2001 3? + 2 radiometric 13,840 Tana 1330 m

Ankazomiranga sediment/ 19,450 G 280 BP 160e180 cm KI-3854.17 290 kms of Straka, 2001 3? + 2 e radiometric Tana 1330 m 63

Ankazomiranga sediment/ 28,850 G 720 BP 180e200 cm KI-3854.19 290 kms Straka, 2001 3? + 2 radiometric of Tana 1330 m Ankilitelo Cave Palaeopropithecus in- 510 G 80 BP AD 1300e not given b-22(54.819#S, Late occurrence Simons, 1997 5 + 3 gens AMS 1620 43(52.610#E of extinct taxon Collagen extrac- tion with alkali Ankilitelo Cave Megaladapis 630 G 50 BP AD 1280e not given b-22(54.819#S, Late occurrence Simons, 1997 5 + 3 edwardsi AMS 1420 43(52.610#E of extinct taxon Collagen extrac- tion with alkali Anosizato Wetland sediment/ 9020 G 120 BP BP 10430e 1.45 m AZ-47 b-28592 ÿ21.20 18(55# S, Rafamantanantsoa,3+2 radiometric 9760 47(30# E 1991 1260 m Antsetsindrano, Vakinan- charcoal/ 250 G 60 BP AD 1490e Shelter 7, Pit 2 b-14168 20(07#S, Wright et al., 6+2 karatra Archaeological radiometric 1950 Layer IIIC 47(00# E 1992 0.20e0.30m 2100m Antsetsindrano, Vakinan- charcoal/ 430 G 60 BP AD 1410e Shelter 2, Pit 3 b-14169 20(07# S, Wright et al., 6+2 karatra Archaeological radiometric 1630 Layer IIIC 47(00# E 1992 0.66e0.73m 2100m Antsetsindrano, Vakin’ charcoal/ 360 G 60 BP AD 1430e Shelter 7, Pit 2 b-14167 20(07# S, Wright et al., 6+2 ankaratra Archaeological radiometric 1660 Feature 3 47(00# E 1992 0.32e0.39 m 2100m G e ( # Antsirabe Paleolake/Wetland Numida meleagris bo-90 60 BP AD 1660 UA-322 b-63120 19 52 S, Goodman, 1999 5 + 25 3 (2004) 47 Evolution Human of Journal / al. et Burney D.A. ne > 1960 47(02# E Antsirabe Paleolake/Wetland Alopochen sirabensis 17,100 G 100 BPBP 20,910e UA-119 b-63118 19(52# S, Goodman, 1999 5 + 3 bone/AMS 19,820 47(02# E Antsirabe Paleolake/Wetland Centrornis majori bo- 17,370 G 90 BP BP 21200e UA-214 b-63119 19(52# S, Goodman, 1999 5 + 3 ne/AMS 20150 47(02# E Antsirabe Paleolake/Wetland Alopochen sirabensis 19,250 G 110 BPBP 23470e UA-142 b-63117 19(52# S, Goodman, 1999 5 + 3 bone/AMS 22240 47(02# E Antsiroandoha Cave Archaeolemur sp. 1020 G 50 BP AD 910e b-60797 CAMS- 12(54# S, Late occurrence Simons et al., 5+3 fibula/AMS 1150 5484 49(05# E of extinct taxon 1995 Collagen extrac- tion with alkali Antsiroandoha Cave Babakotia radofilai/ 4400 G 60 BP BP 5290e b-60798 CAMS- Collagen extrac- Simons et al., 5+3 AMS bone 4840 5485 tion with alkali 1995 Antsiroandoha Cave Megaladapis sp./ 26,150 G 400 BP b-30982 ETH-52- poor collagen Simons et al., 1+3 AMS 55 yield Collagen 1995 extraction with alkali Baie des Galions Wave-cut no- shell/radiometric 2250 G 420 BP BC 900eAD 1e1.4 m asl GaK-277 25(30# S, collected by Kigoshi et al., 2+2 tch 1010 46(30#E Battistini 1964 1-1.4m Bassin de Bealanana sediment/ not given BP 2779e 230e250 cm KI-3612 18(5# S, Straka, 2001 3? + 2 radiometric 2739 47(20# E 1500m Behavoha Coastal dunes Geochelone carapace/ 2160 G 110 BP BC 410e GaK-1658 Late occurrence Mahe´ and Sourdat,1? + 3 e 63 radiometric AD 70 of extinct taxon 1972 Beloha Daubentonia robusta 1065 G 40 BP AD 891e UA 3890, Dr5 NZA-18524 ÿ16.30 20(08# S, Collagen extrac- This paper 5 + 3 left humerus/ 1027 (1932) DB-03- R-28331/7 47(01#E tion with alkali AMS 11 Beloha Alopochen sirabensis/ 1380 G 90 BP AD 530e b-63675 20(08# S, Late occurrence Goodman and Ra- 5+3 AMS 860 47(01#E of extinct taxon kotozafy, 1997 Beloha Megaladapis 1277 G 35 BP AD 666e UA 4595, AM NZA-18999 ÿ20.40 20(08# S, University of An-This paper 5 + 3 edwardsi tooth/ \816 6565 DB-03-21 R-28421/10 47(01#E tananarivo collec- AMS tion Collagen extraction with gelatinisation Belo-sur-Mer Ankilibehandry palm leaf base/ 190 G 80 BP AD 1515e Sq. 3, 60e70 c- b-103347 ÿ26.60 20(44# S, Acid/alkali/acid This paper 5 + 1 ( # Interdunal swale radiometric 1950 m BSM-95-8 44 01 E 49 (continued on next page) Appendix (continued) 50 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Belo-sur-Mer Ankilibehan- Pachylemur insignis 1220 G 50 BP AD 680e960 Sqs. 3ÿ5 100+ b-90098 ÿ20.60 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale ulna/AMS cm BSM-95-5 44(01# E of extinct taxon 2m Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- cf. Mullerornis right 1280 G 60 BP AD 650e890 Sq. 3, 50e70 c- b-103349 ÿ24.90 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale tibiotarsus/AMS m BSM-95-10 44(01# E of extinct taxon 2m Collagen extrac- tion with alkali ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Belo-sur-Mer Ankilibehan- Archaeolemur cf. 1370 G 40 BP AD 620e700 Sq. 5, Level 3 b-90095 ÿ19.90 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale majori jaw/AMS BSM-95-2 44(01# E of extinct taxon 2m Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Hadropithecus ste- 1413 G 80 BP AD 444e772 1934 C. Lam- NZA-12582 ÿ8.40 20(44# S, Late occurrence This paper 5 + 3 dry Interdunal swale nognathus AMS berton BSM- R-26341/1 44(01# E of extinct taxon 00-1 AM 6083 Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Archaeolemur sp. 1550 G 50 BP AD 410e620 Sq.4, Level 2 b-90096 ÿ20.50 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale Fibula/AMS BSM-95-3 44(01# E of extinct taxon 2m Collagen extrac- tion with gelatini- sation Belo-sur-Mer Ankilibehan- Archaeolemur 1650 G 50 BP AD 260e530 Sq. 5, Level 1 b-90094 ÿ19.90 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale majori jaw/AMS BSM-95-1 44(01# E of extinct taxon 2m Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Aepyornis cf. max- 1830 G 60 BP AD 60e350 Sq. 5, level 3 4- b-90099 ÿ17.30 20(44# S, Late occurrence Burney, 1999 6 + 3 dry Interdunal swale imus eggshell/ 5 cm BSM-95-6 44(01# E of extinct taxon. radiometric 2m Charcoal in- (extended) crease, probably anthropogenic Acid etching

Belo-sur-Mer Ankilibehan- Mesopropithecus 1866 G 45 BP AD 58e247 MP22, UA3932 NZA-18522 ÿ20.30 20(44# S, Collagen extrac- This paper 5 + 3 e dry Interdunal swale globiceps left femo- (1934) DB-03-7 R-28331/5 44(01# E tion with alkali 63 ral shaft/AMS 2m Belo-sur-Mer Ankilibehan- sediment at base of 1990 G 50 BP BC 100eAD Sq. 5, 105 cm b-103348 ÿ28.00 20(44# S, 10 cm below Sp- Burney, 1999 3 + 2 dry Interdunal swale bone layer/AMS 110 BSM-95-9 44(01# E orormiella decline 2m (megafaunal col- lapse) Acid wa- shes Belo-sur-Mer Ankilibehan- Pachylemur insignis 2000 G 50 BP BC 110eAD Sqs. 3-5 b-90097 ÿ20.20 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale ulna/AMS 100 100+ cm 44(01# E of extinct taxon BSM-95-4 2m Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Palaeopropithecus 2008 G 90 BP BP 2292e1729 1934 C. Lam- NZA-12583 ÿ21.00 20(44# S, Late occurrence This paper 5 + 3 dry Interdunal swale ingens/bone AMS berton BSM-0- R- 26341/2 44(01# E of extinct taxon 0-2 UA 1042 2m Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Archaeolemur ma- 2050 G 60 BP BC 200eAD 70 Sq.1, Layer 1 b-76612 CAMS- ÿ18.30 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale jori mandible/AMS BSM-1-1 16368 44(01# E of extinct taxon Collagen extrac- tion with alkali Belo-sur-Mer Ankilibehan- Megaladapis mada- 2140 G 50 BP BP 2320e1990 Sq.3, 105+ cm b-102567 ÿ22.40 20(44# S, Late occurrence Burney, 1999 5 + 3 dry Interdunal swale gascariensis talus/ BSM-95-7 44(01# E of extinct taxon AMS 2m Collagen extrac- tion with alkali G e ( # Bemafandry Coastal dunes wood adjacent to 1980 90 BP BC 190 AD 98 cm GaK-1656 24 59 S, Mahe´ and 5+2 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. dated Geochelone 230 44(07 #E Sourdat, 1972 carapace/AMS Bemafandry Coastal dunes Geochelone cara- 2060 G 150 BP BC 400eAD GaK-1655 not given Late occurrence Mahe´ and 1? + 3 pace/radiometric 250 of extinct taxon Sourdat, 1972 Benavony Wetland sediment/radiomet- 710 G 110 BP AD 1050e1430 197e201 cm b-81218 ÿ28.80 13(42# 48$ Charcoal in- Burney, 1999 3 + 3 ric (4x) Ben-2-3A,B S, 48(28# crease, probably 53$E anthropogenic Acid/alkali/acid Benavony Wetland sediment/AMS 2980 G 50 BP BP 3330e2980 383e387 cm b-103532 ÿ26.90 13(42# Acid washes This paper 3 + 3 Ben-2-5A 48$S, 48- (28#53$ E Benavony Wetland sediment/ 3790 G 50 BP BP 4350e3990 828e837 cm b-77549 ÿ30.10 13(42# Acid washes This paper 3 + 2 radiometric BEN-2-9A,B 48$S, 48- (28#53$ E Beropitika Archaeological charcoal/ 750 G 90 BP AD 1050e1400 Gif-4496 24(37#S, Early archaeolo- Emphoux, 1979 6+2 radiometric 45(35#E gical site cited in Dewar, 360m 1984 (p. 582) Bevoha Megaladapis tooth/ 1815 G 60 BP AD 27e412 AM 6479 #6 NZA-10060 R -2- ÿ20.20 23(04# S, Late occurrence of This paper 5 + 3 AMS 4649/2 43(45# E? extinct taxon Col- 2m lagen extraction with gelatinisation Fanongoavana, Imerina charcoal/radio 410 G 100 BP AD 1320e1660 Unit 15030, LY-3161 19(03# S, Wright et al., 6+2 Archaeological metric El.1624.05 47(51#E 1992 Fanongoavana, Imerina charcoal/ 450 G 120 BP AD 1290e1660 Unit 600-16600 LY-3162 19(03# S, Wright et al., 6+2 e 63 Archaeological radiometric El. 1625.15 47(51#E 1992 Fanongoavana, Imerina charcoal/ 450 G 100 BP AD 1300e1650 Unit 1000-15.0- Gif-547 19(03# S, Wright et al., 6+2 Archaeological radiometric 00 Sq. E5m 47(51#E 1992 Forest of Ambohitantely sediment/ not given BP 2479e2329 230e250 cm KI-3454 18(5# S, Straka, 2001 3? + 2 radiometric 47(20# E 1500m Ft. Dauphin, 50 km WSW Aepyornis eggshell/ 1000 G 150 BP AD 690e1290 unknown UCLA-1893 ÿ14.40 21(01# S, Late occurrence Berger et al., 6+3 radiometric 47(E of extinct taxon 1975 Ifanja sediment/ not given AD 1270e1430 260e280 cm KI- 3221.95 49(10 E, Straka, 2001 3? + 2 radiometric 18(50# S Ifanja sediment/ not given BC 370e40 500e520 cm KI-3221.51 S of Straka, 2001 3? + 2 radiometric Sanganoro

(continued on next page) 51 Appendix (continued) 52 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13 C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Irodo (Antanimenabe) charcoal/ 980 G 100 BP AD 880e1260 GaK-350b 12(38# S, Early archaeolo- Battistini and 6+2 Archaeological radiometric 49(31# E gical site Ve´ rin, 1967 Irodo (Antanimenabe) charcoal/ 1200 G 40 BP AD 710e960 GaK-380 12(38# S, Early archaeolo- Battistini and 6+2 Archaeological radiometric 49(31# E gical site Ve´ rin, 1967 Irodo (Tafiampatsa) dune Aepyornis eggshell/ 1150 G 90 BP AD 680e1030 GaK- 12(38# S, Early archaeolo- Mahe´ and Sour- 6+3 radiometric 49(31# E gical site dat, 1972 (p. 305) Irodo (Tafiantsirbeka) Gastropod shell/ 1090 G 90 BP AD 720e1160 GaK-692 12(38# S, Early archaeolo- Battistini and 2+2 Archaeological radiometric 49(31# E gical site Ve´ rin, 1967 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Itampolo Interdunal swale Hippopotamus sp./ 980 G 200 BP AD 660e1400 15 cm GaK-1506 24(41 S, Late occurrence Mahe´ and Sour- 1+3 radiometric 43(57#E of extinct taxon dat, 1972 Itampolo Interdunal swale Aepyornis tibiotar- 1880 G 70 BP BC 30eAD 1-A HFJ-92-12 b-67659 Human-modified Burney, 1999 5 + 2 sus (human-modi- 320 megafaunal bone fied) Collagen extrac- tion with alkali Itampolo Interdunal swale wood/radiometric 2290 G 90 BP BP 2700e2100 120 cm GaK-1652 24(41# S, Mahe´ and Sour- 5+2 43(57# E dat, 1972 Itasy Large lake sediment/ 2260 G 60 BP BP 2360e2130 400e420 cm KI-3160 19S, 46.3( Straka, 1993 3? + 2 radiometric E Itasy Large lake sediment/ 8165 G 90 BP BP 9410e8800 900e930 cm GRO-2804 not given DeWaard and 3? + 2 radiometric Straka, 1961 Kavitaha Lava-barrier lake sediment/ 670 G 80 BP AD 1220e1420 151e160 cm VI b-14856 ÿ18.28 19(02# S, Burney, 1987c 3 + 2 radiometric A,B 46(47# E 1210 m Kavitaha Lava-barrier lake sediment/ 710 G 100 BP AD 1160e1420 10 cm I (KAV- b-11729 ÿ19.31 19(02# S, Date rejected due Burney, 1987c 3 + 1 radiometric IA,B) 46(47# E to old carbon 1210 m contamination Kavitaha Lava-barrier lake sediment/ 960 G 90 BP AD 900e1260 220 cm (cen- b-15528 ÿ25.00 19(02# S, Sporormiella in- Burney, 1987c 3 + 3 radiometric tered) KAV-2A 46(47# E crease (livestock). 1210 m Acid washes Kavitaha Lava-barrier lake sediment/ 1400 G 80 BP AD 530e780 291e295 cm V b-14855 ÿ23.51 19(02# S, Charcoal in- Burney, 1987c 3 + 3 radiometric (KAV-3), VA 46(47# E crease, probably

1210 m anthropogenic e Kavitaha Lava-barrier lake sediment/ 1490 G 60 BP AD 430e660 341e350 cm II- b-11730 ÿ22.63 19(02# S, Burney, 1987c 3 + 2 63 radiometric A,B (KAV-3, 46(47# E 101-110) 1210 m Komango Lake macrofossil/AMS 1750 G 60 BP AD 130e420 KOM-8 (48 b-68068 CAMS- ÿ27.90 19(9# 4$S, Charcoal in- Burney, 1999 5 + 3 cm) KOM-8 10245 44(48#30$ crease, probably E anthropogenic Komango Lake sediment/AMS 3390 G 50 BP BP 3720e3480 KOM-9 (41- b-103350 ÿ28.10 19(9# 40 Acid washes This paper 3 + 3 46 cm) KOM- #S, 44(48# 9A 30$E Lakaton’i Anja Archaeologi- charcoal/ 1300 G 80 BP AD 620e900 b-18424 Not given Dewar and Wrig- 6+2 cal Rockshelter radiometric ht, 1993 (p.429) Lakaton’i Anja Archaeologi- charcoal/AMS 1680 G 65 BP AD 230e530 b-29946 ETH- Not given Early archaeolo- Dewar and Wrig- 6+2 cal, Rockshelter 5408 gical site ht, 1993 (p.429) Lamboharana Shallow pond Bones presumably 1220 G 80 BP AD 660e990 40 cm GaK-2310 22(11# S, Mahe´ and Sour- 1? + 2 megafaunal/radio- 43(16# E dat, 1972 metric Lamboharana Shallow pond Hippopotamus le- 1740 G 50 BP AD 155e415 MNIP MAD TO-1437 22(11# S, Human-modified MacPhee and 5+3 merlei femur 1709 43(16# E megafaunal bone Burney, 1991 (human-modified)/ Collagen extrac- AMS tion Lamboharana Shallow pond Hippopotamus 2020 G 300 BP BC 805eAD MNIP MAD TO-1438 22(11# S, Human-modified MacPhee and 5+3 femur (human- 640 1710 43(16# E megafaunal bone Burney, 1991 modified)/AMS Lamboharana Shallow pond bone/radiometric 2350 G 120 BP BP 2740e2100 60 cm GaK-2307 22(11# S, Mahe´ and Sour- 1? + 2 43(16# E dat, 1972 G ( #

Lamboharana Shallow pond Hippopotamus 3495 55 BP MNIP MAD AA-2894 22 11 S, Date rejected; sa- MacPhee and 5+1 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. femur (human- 1710 43(16# E mple redated to Burney, 1991 modified)/AMS 2020 BP G 300 Lavakasaka Cave cf. Mullerornis 2380 G 70 BP BP 2720e2320 sondage 1, layer b-55094 CAMS- 15(34#S, Late occurrence Burney et al., 6+3 eggshell/AMS 1 HFJ-92-15 3547 46(51#E of extinct taxon 1997 Lohavohitra, Vonizonga charcoal/ 90 G 30 BP AD 1680e1950 Zone 1, Loca- SMU-2075 19(38# S, Wright et al., 6+2 Archaeological radiometric tion 03 Strati- 47(17# E 1992 graphic Unit 25 Lohavohitra, Vonizonga charcoal/ >100 BP Sondage I, LY-3749 19(38# S, Wright et al., 6+2 Archaeological radiometric Level 7 47(17#E 1992 Lohavohitra, Vonizonga charcoal/ 230 G 60 BP AD 1510e1950 Zone 1, Loca- Gif-7470 19(38# S, Wright et al., 6+2 Archaeological radiometric tion 03 Strati- 47(17#E 1992 graphic Unit 29 Lohavohitra, Vonizonga charcoal/ 260 G 60 BP AD 1480e1950 Zone 1, Loca- Gif-7471 19(38# S, Wright et al., 6+2 Archaeological radiometric tion 01 Strati- 47(17#E 1992 graphic Unit 16 Lohavohitra, Vonizongo charcoal/ 270 G 45 BP AD 1510e1670 Sondage I, SMU-1520 19(38# S, Wright et al., 6+2 Archaeological radiometric Layer 4 47(17#E 1992 Mahery Shallow crater lake sediment/ 2950 G 70 BP BP 3340e2890 MAH-0 b-23488 ÿ35.50 12(26# S, Burney, 1988 3 + 2 radiometric (11-30 cm) 49(14# E MAH-0A,B 400 m Mahery Shallow crater lake sediment/ 3680 G 70BP BP 4230e3840 61-70 cm b-22618 ÿ32.40 12(26# S, This paper 3 + 2 radiometric MAH-1A 49(14# E 400 m Mahery Shallow crater lake sediment/ 4620 G 90 BP BP 5580e5040 MAH-2 b-24614 ÿ17.70 12(26# S, Burney, 1988 3 + 1 e 63 radiometric (21-50 cm) 49(14# E MAH-2A,B,C 400 m Mahilaka Archaeological Oryza/AMS 830 G 70 BP AD 1030e1290 I - - 3 ST-12097 13(45#S, Early archaeolo- Radimilahy, 1997 5 + 3 48(19#E5m gical site (rice) Mahilaka Archaeological charcoal/ 890 G 45 BP AD 1025e1250 VIII-A-6 Ua- 3802 13(45#S, Radimilahy, 1998 6 + 2 radiometric 48(19#E5m Mahilaka Archaeological charcoal/ 910 G 75 BP AD 995e1270 II-P-3-4 Ua- 3804 13(45#S, Radimilahy, 1998 6 + 2 radiometric 48(19#E5m Mahilaka Archaeological charcoal/ 930 G 90 BP AD 965e1275 VI-N-4 Ua -3803 13(45#S, Radimilahy, 1998 6 + 2 radiometric 48(19#E Mahilaka Archaeological charcoal/ 1030 G 65 BP AD 885e1160 X-LT-6 Ua -3805 13(45#S, 4- Radimilahy, 1998 6 + 2 radiometric 8(19#E5m

(continued on next page) 53 Appendix (continued) 54 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Mahilaka Archaeological charcoal/ 1120 G 160 BP AD 630e1250 I - - 3 St-12098 13(45#S, Radimilahy, 1998 6 + 2 radiometric 48(19#E Mahilaka Archaeological charcoal/ 1175 G 70 BP AD 685e1005 VII-G-4 Ua-3806 13(45#S, Radimilahy, 1998 6 + 2 radiometric 48(19#E5m Maliavola Archaeological 1140 G 80 BP AD 690e1030 24(59#S, Rakotoarisoa and 6+2 46(54#E Radimilahy, 2003 15m Maliavola, Anosy potsherd/TL 1475 G 100 AD Operation I, DurTL-77-8AS 24(59#S, Same context as Rakotoarisoa TL ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Archaeological Feature I 46(54#E SMU 2078 et al., 1993 15m Maliavola, Anosy charcoal/ 1144 G 29 BP AD 810e980 Operation I, SMU-2078 24(59#S, Same context as Rakotoarisoa 6+2 Archaeological radiometric Feature I 46(54#E DurTL 77 - 8AS et al., 1993 15m Manambovo River mouth Aepyornis 840 G 80 BP AD 1020e1290 no given 25(28# S, Early archaeolo- Battistini, Ve´ rin 6+2 Sandstone terrace shell and pottery/ 45(42E gical site and Raison, 1963 radiometric Mananjary Flood plain Hippopotamus 99 G 36 BP AD 1670e1950 not given AA-45965 T-160- ÿ8.00 21(14# S, Collagen extrac- This paper 5 + 3 laloumena AMS DB-HA-1 46A 48(20# E tion with gelatini- sation Mananjary Flood plain Hippopotamus 213 G 40 BP AD 1639e1945 DB-HA-1 (2) NZA-16993 ÿ8.69 21(14# S, Collagen extrac- This paper 5 + 3 laloumena AMS R- 28139/1 48(20# E tion with gelatini- sation Mananjary Flood plain Hippopotamus 2327 G 40 BP BP 2364e2212 DB-03-20 NZA-18525 ÿ17.20 21(14# S, Collagen extrac- This paper 5 + 3 laloumena AMS R -28331/10a 48(20# E tion with gelatini- sation Marais d’Ifanja Swamp/ sediment/ 4200 G 80 BP BP 4870e4520 780e800 cm GRO-2197 not given Near Itasy DeWaard and St- 3? + 2 Lake radiometric raka, 1961 Maranakely sediment >41,400 BP 360e370 cm KI-3576.25 not given Straka, 2001 3? + 2 radiometric Maroaloke Coastal dunes charcoal/not given 915 G 45 BP AD 1020e1210 OxA-5638 25(12# S, Early archaeolo- Parker Pearson 6+2 46(12# E gical site et al., 1995

Maroaloke Coastal dunes Aepyornis eggshell/ 1415 G 40 BP AD 570e670 OxA-5078 25(12# S, Late occurrence Parker Pearson 6+3 e not given 46(12# E of extinct taxon et al., 1995 63 Maroaloke Coastal dunes Aepyornis eggshell/ 2285 G 40 BP BP 2350e2160 OxA-5077 25(12# S, Late occurrence Parker Pearson 6+3 not given 46(12# E of extinct taxon et al., 1995 Masinandriana Large Plesiorycteropus 2154 G 40 BP BP 2309e2006 MAS-2 NZA-16994 ÿ18.69 Late occurrence This paper 5 + 3 Wetland pelvic fragment/ R-28139/3 of extinct taxon. AMS Did not produce sufficient gelatin for CN analysis. Collagen extrac- tion with gelatini- sation. Masinandriana Large Aepyornis sp. 4496 G 40 BP BP 5307e4974 MAS-5 NZA-16995 ÿ13.99 Collagen extrac- This paper 5 + 3 Wetland Tibiotarsus/AMS R-28139/6 tion with gelatini- sation Matsabory Ampozalana sediment/radio 18,860 G 240 BP BP 23310e 36e40 cm MA- b-17531 ÿ28.52 12(31# S, Acid washes Burney, 1987b 3 + 3 Wetland metric 21500 P-1 49(11# E 975 m Matsabory Ampozalana sediment/radio- 19,680 G 240 BP MAP-3 (26-30) b-17532 ÿ27.52 12(31# S, Acid washes Burney, 1987b 3 + 3 Wetland metric 49(11# E 975 m Matsabory Ampozalana sediment/radio- 35,090 G 800BP MAP-4 (31- b-16809 ÿ26.32 12(31# S, Acid washes Burney, 1987b 3 + 2 Wetland metric 70 cm) MAP- 49(11# E 4,A,B,C,D,E, 975 m F,G,H Matsabory Ampozalana sediment/radio- 35,750 G 1360 MAP-5 III (M- b-14853 ÿ25.89 12(31# S, Acid washes Burney, 1987b 3 + 2 Wetland metric BP AP-5) IIIA, II- 49(11# E

IB,IIIC,IIID 975 m 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Miangola Crater bog sediment/radio- 30,580 G 590 BP MIA-2A (25.5- b-22348 ÿ24.50 19(52# S, Acid washes This paper 3 + 3 metric 30 cm) MIA- 46(59# E 2A 1520 m Miangola Crater bog sediment/radio- 32,710 G 820 BP MIA-9 (51-70 - b-14851 ÿ24.92 19(52# S, Acid washes Burney, 1987b 3 + 2 metric cm) I (MIA-9) 46(59# E IA,IB 1520 m Mitoho Cave egg- 4030 G 70 BP BP 4810e4300 surface b-16442 ÿ13.25 24(03# S, Acid etched MacPhee, 1986 6 + 3 shell/radiometric 43(46# E Mitsinjo Flood plain lake sediment/radio- 470 G 90 BP AD 1300e1640 198e212 cm b-50275 ÿ24.80 16(02# S, Acid washes Matsumoto and 3+2 metric MIT-3A,B 45(51# E Burney, 1994 9m Mitsinjo Flood plain lake sediment/radio- 530 G 80 BP AD 1290e1490 90e110 cm b-48177 ÿ27.40 16(02# S, Acid washes Matsumoto and 3+2 metric MIT-2A 45(51# E Burney, 1994 9m Mitsinjo Flood plain lake sediment/radio- 890 G 80 BP AD 1000e1280 264e268 cm b-51797 ÿ29.40 16(02# S, Acid washes Matsumoto and 3+3 metric (4x) MIT-3.5A 45(51# E Burney, 1994 (3:81-85) Mitsinjo Flood plain lake wood,sediment 2400 G 90 BP 2740e2180 390e420 cm b-51799 ÿ27.30 16(02# S, Acid washes Matsumoto and 3+2 radiometric (4x) MIT-6W,A,B,C 45(51# E Burney, 1994 (6:41-70) 9m Mitsinjo Flood plain lake sediment/radio- 3160 G 210 BP BP 3870e2840 323e349 cm b-52365 ÿ26.70 16(02# S, Rejected due to Matsumoto and 3+1 metric (4x) MIT-5A,B,C,D 45(51# E old carbon con- Burney, 1994 (5:1-27) 9m tamination Mitsinjo Flood plain lake sediment/radio- 3420 G120 BP BP 3980e3390 480e500 cm b-48178 ÿ23.50 16(02# S, Acid washes Matsumoto and 3+2 e 63 metric MIT-7A 45(51# E Burney, 1994 9m Nosy Mangabe Archaeologi- pottery/TL AD 950 G 160 Dur91TL-15- 15(30# S, Early archaeolo- Dewar and Wrig- TL cal 2ASQi 49(46# E gical site ht, 1993 (p.439) Nosy Mangabe Archaeologi- charcoal/radio- 1250 G 60 BP AD 660e990 SMU-2501 15(30# S, Early archaeolo- Dewar and Wrig- 6+2 cal metric 49(46# E gical site ht, 1993 (p. 429) Ranobe Pond sediment/radio- 600 G 70 BP AD 1280e1440 51e70cm RAN- b-24423 ÿ26.30 23(01# S, This paper 3 + 2 metric 0-A,B,C,D 43(37# E 51-70 10 m (continued on next page) 55 Appendix (continued) 56 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Ranobe Pond sediment/ 1300 G 80 BP AD 620e900 RAN-2:81-100 b-22619 ÿ27.40 23(01# S, This paper 3 + 2 radiometric cm RAN-2A,B 43(37# E 10 m Ranobe Pond sediment/ 1360 G 80 BP AD 550e860 RAN-2:21-40 b-23489 ÿ26.60 23(01# S, Burney, 1997a,b 3 + 2 radiometric RAN-1A,B 43(37# E 10 m Sandrakatsy East charcoal/ 1140 G 60 BP AD 770e1010 Operation D, SMU-2076 16(09# S, Early archaeolo- Wright and 6+2 Archaeological radiometric Layer 3 49(46# E gical site Fanony, 1992 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Sandrakatsy East charcoal/ 1240 G 50 BP AD 670e900 Operation D, SMU-2359 16(09# S, Early archaeolo- Wright and 6+2 Archaeological radiometric Layer 3 49(46# E gical site Fanony, 1992 Sandrakatsy West charcoal/ 970 G 60 BP AD 980e1200 Operation E, SMU-2502 16(09# S, Early archaeolo- Wright and 6+2 Archaeological radiometric Layer 2 49(46# E gical site Fanony, 1992 Sarodrano Archaeological charcoal/ 160 G? BP Site 1 GaK-926 23(30# S, B and V believe Battistini and 6+1 radiometric 43(44# E that the modern- Ve´ rin, 1971 (p.58) looking ceramics and glass bead suggest the layer mixed with older natural charcoal. No # given for G Sarodrano Archaeological charcoal/ 210 G 80 BP AD 1490e1950 Site 2 GaK-927 23(30# S, Battistini and Ve´ - 6+1 radiometric 43(44# E rin, 1971 (p.58) Sarodrano Archaeological charcoal/ 535 G 80 BP AD 1290e1480 Site 1 GaK-1057 23(30# S, Battistini and Ve´ - 6+1 radiometric 43(44# E rin, 1971 (p. 58) Sarodrano Archaeological charcoal/ 1460 G 90 BP AD 410e710 Site 2 GaK-928 23(30# S, Early archaeolo- Battistini and Ve´ - 6+1 radiometric 43(44#E gical site? rin, 1971 (p.60) Southern Madagascar Aepyornis eggshell/ 2930 G 85 BP BP 3340e2860 not given 19(#S, Marden, 1967. 6+3 radiometric 47(#E (p.487) SW Madagascar near Nossi Geochelone grandi- 1250 G 50 BP AD 670e890 B.M.N.H. (Z) BM-1399 ÿ13.90 not given Late occurrence Burleigh and 5? + 3 Be´ dieri carapace 1984.1282 of extinct taxon Arnold, 1986 radiometric

Talaky Archaeological, charcoal/ 840 G 80 BP AD 1020e1290 Sq. C4 and C5 GaK-276 25(27# S, Early archaeolo- Battistini and 6+2 e Sandstone bench radiometric of 2nd site in B 45(43# E gical site Ve´ rin, 1967 63 zone 20 cmbs 3m Taolambiby seepage deposit Propithecus ver- modern not given AA-45962 23(40# S, This paper 5 + 3 reauxi humerus AW-T-66-1 44(25# E (cut marks)/AMS Taolambiby seepage deposit Tortoise bone/ 2290 G 90 BP BP 2700e2100 1.90 m GaK-1651 23(40# S, Late occurrence Mahe´ and Sour- 1? + 3 radiometric 44(25# E of extinct taxon dat, 1972 Taolambiby seepage deposit Palaeopropithecus 2325 G 43 BP BP 2366e2315 not given OX- AA-45960 ÿ20.20 23(40# S, Human-modified This paper 5 + 3 ingens radius (cut UM 14346j T16041A 44(25# E megafaunal bone; marks)/AMS Collagen extrac- tion with gelatini- sation Taolambiby seepage deposit Megaladaois grand- 2713 G 44 BP BP 2870e not given OX- AA-45961 ÿ18.90 23(40# S, This paper 5 + 3 idieri/madagascar- 2760 UM 5105 T-16042 44(25# E iensis 2nd , lower jaw/AMS Torotorofotsy Paleolake/ wood/radiometric 9390 G 100 BP BP 11070e 2.1m TTF-2.1 b-28590 ÿ29.50 18(55# S, Rakotondrazafy, 5+2 Wetland 10270 48(26# E 1992 956m Torotorofotsy Paleolake/ sediment/ 35,000 + 2500/ 200e220 cm KI-3222 18(55# S, Straka, 1993 3? + 2 Wetland radiometric - 1900 BP 48(26# E 956m Tritrivakely Crater lake plant macrofossil/ modern 15 cm LT1 H-616 19(47# S, Gasse et al., 5+3 AMS 46(55# E 1994 G e e ( # Tritrivakely Crater lake sediment/ 1240 100 BP AD 630 41 45cm b-15884 ÿ26.51 19 47 S, Charcoal in- Burney, 1987a 3 + 25 3 (2004) 47 Evolution Human of Journal / al. et Burney D.A. radiometric 1000 TRI-1A 46(55# E crease, probably 1778 m anthropogenic Tritrivakely Crater lake plant macrofossil/ 2840 G 60 BP BP 3140e 140 cm LT2/5 H-667 ÿ26.00 19(47# S, Gasse et al., 5+3 AMS 2790 46(55# E 1994 (p. 1517) 1778m Tritrivakely Crater lake sediment/ 3810 G 70 BP BP 4420e 151e155 cm T- b-15885 ÿ22.65 19(47# S, Burney, 1987a 3 + 3 radiometric 3980 RI-2A (61-65) 46(55# E 1778 m Tritrivakely Crater lake plant macrofossil/ 4340 G 90 BP BP 5290e 305 cm LT4 H-618 ÿ24.80 19(47# S, Gasse et al., 5+3 AMS 4650 46(55# E 1994 (p. 1517) 1778 m Tritrivakely Crater lake sediment/ 5730 G 70 BP BP 6670e 246e255 cm b-16807 ÿ24.31 19(47# S, Burney, 1987a 3 + 2 radiometric 6390 TRI-3A,B 46(55# E (56-65) 1778 m Tritrivakely Crater lake plant macrofossil/ 5840 G 80 BP BP 6800e 300 cm T-II/ 19(47# S, Williamson 5+3 AMS 6450 300 cm 46(55# E et al., 1998 1778 m Tritrivakely Crater lake plant macrofossil/ 6290 G 110 BP BP 7430e 340 cm LT4/5 H-669 ÿ22.80 19(47# S, Gasse et al., 5+3 AMS 6900 46(55# E 1994 (p. 1517) 1778 m Tritrivakely Crater lake plant macrofossil/ 8460 G 130 BP BP 9690e 350 cm LT4 H-711 ÿ21.11 19(47# S, Gasse et al., 5+3 AMS 9130 (40) 46(55# E 1994 (p. 1517) 1778 m Tritrivakely Crater lake sediment/ 8490 G 100 BP BP 9600e 326e335 cm b-16808 ÿ23.75 19(47# S, Burney, 1987a 3 + 2 e 63 radiometric 9290 TRI-4A,B (36- 46(55# E 45) 1778 m Tritrivakely Crater lake plant macrofossil/ 9920 G 90 BP BP 11650e 350 cm T-II/350 19(47# S, Williamson 5+3 AMS 11180 cm 46(55# E et al., 1998 1778 m Tritrivakely Crater lake sediment/ 10,470 G 110 BP BP 12910e 473e490 cm IV b-14854 ÿ18.31 19(47# S, Burney, 1987a 3 + 2 radiometric 11910 (TRI-5) IVA, 46(55# E IVB (83-100) 1778 m Tritrivakely Crater lake plant macrofossil/ 12,750 G 170 BP BP 15930e 379 cm LT4 H-712 ÿ18.83 19(47# S, Gasse et al., 5+3 AMS 14220 (69) 46(55# E 1994 1778m (continued on next page) 57 Appendix (continued) 58 Site name and Material/ Age G 1s Calibrated Provenance or Lab nr d13C Lat./long. Comments Source Rating site type dating range at 2s Accession # Elevation method Sample I.D. Tritrivakely Crater lake plant macrofossil/ 13,740 G 130 BP BP 17030e 411 cm T-II/ 19(47# S, Williamson 5+3 AMS 15980 411 cm 46(55# E et al., 1998 1778m Tritrivakely Crater lake plant macrofossil/ 15,070 G 170 BP BP 18700e 412 cm LT5 H-710 ÿ18.06 19(47# S, Gasse et al., 1994 5+3 AMS 17390 (19) 46(55# E (p. 1517) 1778 m Tritrivakely Crater lake charcoal/AMS 18,550 G 210 BP BP 22870e 450 cm T-II/ 19(47# S, Williamson et al., 6+2 21220 450 cm 46(55# E 1998 ..Bre ta./Junlo ua vlto 7(04 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. 1778 m Tritrivakely Crater lake plant macrofossil/ 19,210 G 180 BP BP 23590e 463 cm LT5/8 H-670 ÿ16.70 19(47# S, Gasse et al., 1994 5 + 3 AMS 22030 46(55# E 1778 m Tritrivakely Crater lake charcoal/AMS 22,700 G 290 BP 552 cm T-II/ 19(47# S, Williamson et al., 6+2 552 cm 46(55# E 1998 1778m Tritrivakely Crater lake sediment/AMS 24,010 G 330 BP 550 cm LT6 H-624 ÿ19.90 19(47# S, Gasse et al., 1994 3 + 3 46(55# E 1778 m Tritrivakely Crater lake sediment/AMS 28,290 G 500 BP 732 cm LT8 H-650 ÿ21.90 19(47# S, Gasse believes the Gasse et al., 1994 3 + 3 46(55# E sediment dates 1778 m may be slightly biased by con- tamination by organic detritus Tritrivakely Crater lake sediment/AMS 34,630 G 990 BP 904 cm LT10 H-626 ÿ10.40 19(47# S, Gasse et al., 1994 3 + 3 46(55# E 1778m Tritrivakely Crater lake sediment/AMS 34,720 G 980 BP 1165 cm LT14 H-619 ÿ24.50 19(47# S, Gasse et al., 1994 3 + 3 46(55# E 1778m Tritrivakely Crater lake plant macrofossil/ 36,200 G 120 BP 1300 cm LT16 H-617 ÿ25.70 19(47# S, Gasse et al., 1994 5 + 3

AMS 46(55# E e 1778 m 63 Tritrivakely Crater lake sediment/AMS 43,100 G 2700 1034 cm LT12 H-628 ÿ21.00 19(47# S, Date rejected Gasse et al., 1994 3 + 2 BP 46(55# E 1778 m Tsiandroina Mesopropithecus 1694 G 40 BP AD 245e429 MP13 DB- 03-6 NZA-18521 ÿ19.30 Late occurance of This paper 5 + 3 globiceps right dis- R-28331/4 extinct taxon; tal humerus/AMS Collagen extrac- tion with gelatini- sation Tsiandroina Wetland Coua primavea 1980 G 60 BP BC 110eAD b-63676 Late occurrence Goodman and 5+3 bone/AMS 130 of extinct taxon Rakotozafy, 1997 Tsiandrora Archaeological charcoal/ 950 G 80 BP AD 960e1260 Sondage I, LY 3912 25(03#S, Rakotoarisoa, 6? + 2 radiometric Level 10-12 46(54#E 1993 12m Tsirave Hadropithecus 2194 G 70 BP BP 2344e1998 not given NZA-15672 ÿ13.22 21(50# S, Late occurrence This paper 5 + 3 stenognathus tooth/ BJ-HS-2 R-26931/1 45(7# E of extinct taxon AMS Collagen extrac- tion with gelatini- sation Tsiroanomandidy sediment/ 1160 G 110 BP AD 660e1040 2.2e2.3 m b-28593 ÿ27.30 18(45# S, This is same as Rafamantanant- 3+2 Wetland radiometric KZ-26 46(3# E Kizato soa, 1991 Tulear (Belande) Aepyornis eggshell/ 5210 G140 BP BP 6290e5640 not given Sauer, 1972 6 + 3 Coastal dunes radiometric unknown Megaladapis tooth/ 1591 G 60 BP AD 339e609 AM 6527 #5 NZA-10102 ÿ20.40 Late occurrence of This paper 5 + 3 AMS R-24649/3 extinct taxon Col- lagen extraction

with gelatinisation 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. Vinaninony Wetland sediment/ 11,100 G 150 BP BP 13,750e 200e220 cm KI-3220.21 19.5( S, Straka, 1993 3? + 2 radiometric 12,660 47.2( E 1875 m Vinaninony Wetland sediment/ > 27,000 BP 600e620 cm KI-3220.61 19.5( S, In Straka, 2001 Straka, 1993 3? + 2 radiometric 47.2( E this date is given 1875 m as 27,000 BP Vohimasina, Vakin’ankara- charcoal/ 480 G 50 BP AD 1400e1480 Sondage 3, SMU-1519 20(08# S, Wright et al., 6+2 tra Archaeological radiometric Layer V 47(01# E 1992 Vohimasina, Vakin’ankara- charcoal/ 710 G 60 BP AD 1220e1400 Sondage 1, SMU-2630 20(8# S, Early archaeolo- Wright et al., 6+2 tra Archaeological radiometric Layer VI 47(01# E gical site 1992 Vohimasina, Vakin’ankara- potsherd/TL 1410 G 115 AD Sondage 1, DurTL-50-13 AS 20(08 #S, Wright et al., TL tra Archaeological Layer VI 47(01# E 1992 Vohimasina, Vakin’ankara- potsherd/TL 1515 G 95 AD Sondage 3, DurTL-50-9 AS 20(08 #S, Wright et al., TL tra Archaeological Layer V 47(01# E 1992 Vohitrarivo, Vakin’ankaratra charcoal/ 290 G 80 BP AD 1440e1950 not given b-24479 20(05# S, Wright et al., 6+2 radiometric 47(05# E 1992 Refer notes to the appendix for key to abbreviations for dating laboratories. Refer notes to appendix for rating system.

Notes to the appendix Key to abbreviations for dating laboratories: Abbreviation Laboratory Location e

AA NSF-Arizona AMS Facility USA 63 BM British Museum England ß Beta Analytic USA CAMS Center for Accelerator Mass Spectrometry USA Dur Durham University England ETH ETH/AMS Facility Switzerland GaK Gakushuin University Japan Gif Gif sur Yvette France GrA Groningen AMS The Netherlands GRO) Groningen The Netherlands GX Geochron Laboratories USA H LHGI, Orsay France Hela Helsinki Finland 59 (continued on next page) 60 Abbreviation Laboratory Location KI Kiel Germany LY University of Lyon France NZA New Zealand New Zealand OxA Oxford Radiocarbon Accelerator Unit England Pta Pretoria South Africa QC) Queens College USA R Rafter New Zealand SMU) Southern Methodist Univ. USA ST) Stockholm Sweden T Trondheim Norway TO IsoTrace Canada Ua Uppsala AMS Sweden 25 (2004) 47 Evolution Human of Journal / al. et Burney D.A. UCLA University of California Los Angles USA *Indicates laboratories that are closed or operating under a different code designation

Rating system: For each 14C determination, a score was assigned to provide a rough guide for assessing the reliability of the date based on the simple system proposed by Mead and Meltzer (1984) with modifications based on suggestions in Long et al. (1983); Stafford et al. (1991); and Hedges and Van Klinken (1992). Ratings are based on adding together a score from each section.

Section I. Material dated Score Faunal Eggshell 6 Collagen 5 Fecal pellet 5 Pretreated bone with poor collagen yield 3 Whole bone 1 Other organic material Charcoal 6 Plant macrofossil (wood, seeds) 5 Peat, organic mud, soil 3 Shell (freshwater/terrestrial/marine) 2 Section II. Strength of association with event of interest Strong (includes pretreated sediments spanning !5cm) 3 e

Unknown or medium (includes pretreated sediments 5-20 cm) 2 63 Weak (includes sediments >20 cm) 1 Dates scoring a total of 8-9 are likely to be the most reliable; 5-7, may have less certainty, depending on context; 2-4, generally unreliable. Question marks indicate insufficient information was available to determine score. D.A. Burney et al. / Journal of Human Evolution 47 (2004) 25e63 61

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