Great Ape Skeletal Collections: Making the Most of Scarce and Irreplaceable Resources in the Digital Age

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 57:2–32 (2013)

Adam D. Gordon,1* Emily Marcus,2,3 and Bernard Wood3

1Department of Anthropology, University at Albany, SUNY, Albany, NY 12222 2Honors Program, George Washington University, Washington, DC 20052 3Center for the Advanced Study of Hominid Paleobiology, Department of Anthropology, George Washington University, Washington, DC 20052

KEY WORDS hominoids; Powell-Cotton Museum; ontogeny; online database

ABSTRACT Information about primate genomes information, who has made use of it, and what types of has re-emphasized the importance of the great apes data have been collected. We present a protocol for col- (Pan, Gorilla,andPongo) as, for most purposes, the lecting information about each individual animal (e.g., appropriate comparators when generating hypotheses which bones are preserved, their condition, etc.) and about the most recent common ancestor of the hominins have made that information about the Powell-Cotton and panins, or the most recent common ancestor of the Collection freely available in an online relational hominin clade. Great ape skeletal collections are thus database (Human Origins Database, www.humanori- an important and irreplaceable resource for researchers ginsdatabase.org). As an illustration of the practical conducting these types of comparative analyses, yet the application of these data, we developed a tabular sum- integrity of these collections is threatened by unneces- mary of ontogenetic information about each individual sary use and their availability is threatened by ﬁnan- (see Appendices A and B). Collections like the Powell- cial pressures on the institutions in which the Cotton are irreplaceable sources of material regarding collections reside. We discuss the general history of the hard-tissue evidence and recent history of the clos- great ape skeletal collections, and in order to get a bet- est living relatives of modern humans. We end this con- ter sense of the utility and potential of these important tribution by suggesting ways that curators and the sources of data we assemble the equivalent of a biogra- researchers who use and rely on these reference collec- phy of the Powell-Cotton Collection. We explore the his- tions could work together to help preserve and protect tory of how this collection of chimpanzee and gorilla them so that future generations can use and beneﬁt skeletonswasaccumulated,howitcametoberecog- from these priceless resources. Am J Phys Anthropol nized as a potentially important source of comparative 57:2–32, 2013. VC 2013 Wiley Periodicals, Inc.

INTRODUCTION skeletal collection of great apes. Most studies were content to use genus level samples (e.g., chimpanzees, low- New molecular (e.g., Arnold et al., 2010; Perelman land gorillas, or orangutans). However, the results of et al., 2011; Prado-Martinez et al., 2013) and morpholog- fine-grained morphological and genetic analyses (e.g., ical (Diogo and Wood, 2011) evidence has confirmed the Groves, 2001; Pilbrow, 2006; Gonder et al., 2011; Prado- close relationship between modern humans and the Martinez et al., 2013) make it clear that given the mor- great apes, and the particularly close relationship phological variation that is distributed geographically between modern humans and chimpanzees and bonobos. among subspecies, when researchers assemble samples Great ape skeletal collections thus provide the core data of great apes for comparative analyses care needs to be required to study hominin fossils in a comparative evolu- taken to assemble skeletal samples from regions that tionary context. Unfortunately, these collections, which coincide with the known ranges of species or subspecies. are uncommon to begin with, are, even with the best Yet to our knowledge there is no single source of infor- curatorial oversight, slowly but surely degrading over mation about great ape skeletal collections that takes time. The collections are irreplaceable in at least two into account the important developments that have senses. First, the animals involved are endangered, and occurred since Groves’ (2001) important review. second many museum collections sample taxa from parts There is also a growing realization within the research of their range that are no longer occupied by living ani- community that we cannot take the continued mals. We suggest the time has come for a thorough evaluation of these critically important collections. This would include assembling a detailed inventory of each collection, devising and implementing ways of making Grant sponsors: G. Harold & Leila Y. Mathers Foundation (BW), Wenner-Gren Hunt Postdoctoral Fellowship (ADG). use of information that has already been collected, and making recommendations about standard practices that *Correspondence to: Adam Gordon, Department of Anthropology, will help preserve these precious collections for the CAS 237, University at Albany, 1400 Washington Avenue, Albany, future. NY 12222, USA. E-mail: [email protected] Not that long ago, when conducting comparative morphological research addressing questions about hominin DOI: 10.1002/ajpa.22391 evolution, it would have been deemed sufficient (even if Published online in Wiley Online Library not ideal) to collect hard-tissue data from the nearest (wileyonlinelibrary.com).

Ó 2013 WILEY PERIODICALS, INC. GREAT APE SKELETAL COLLECTIONS 3 availability of great ape skeletal collections for granted. ships shifted from gross morphology to the morphology We believe they are threatened by a combination of over- of molecules (e.g., Grunbaum,€ 1902; Nuttall, 1904). In use, museums diverting resources from collections to the early 1960s, Zuckerkandl et al. (1960) and Goodman what is perceived to be more important “basic” research, (1963) used hemoglobin and albumin, respectively, to and the parlous financial state of host institutions that investigate the relationships among higher primates and lack the prestige and funding of national museums. they both concluded that chimpanzees were more closely Thus, there is a pressing need for the community to related to modern humans than to gorillas. Others con- maximize the utility of these collections by minimizing curred (Sarich and Wilson, 1967) suggesting that 99% of redundant data collection. No one should be turned the amino-acid sequences of chimpanzee and modern away from examining the collections, but repeated and human proteins were identical (King and Wilson, 1975). unnecessary measurement inevitably damages speci- Initial attempts to compare the DNA of higher prima- mens and wastes the collective effort of the research tes used a method called DNA hybridization (e.g., Cac- community. That said, there is an argument that could cone and Powell, 1989), but once sequencing methods be made for deliberately repeating some measurements became available they rapidly replaced hybridization as in the interest of evaluating comparability across data the preferred method for generating hypotheses about sets collected by different researchers. We revisit this the relationships among living hominoids and the num- last point toward the end of this article. ber of sequence-based studies increased each year (see We are conscious that some paleoanthropologists may Bradley, 2008; Arnold et al., 2010; Perelman et al., 2011; not be aware of the history of research about the rela- for reviews). When these DNA differences were cali- tionships among the great apes and modern humans, so brated using what was then the best paleontological evi- because this is the core of the case for the importance of dence for the split between the apes and the Old World these collections we begin by briefly reviewing this. We Monkeys, it was predicted that the hypothetical ancestor then set out recently accumulated morphological and of modern humans and chimpanzees/bonobos lived genetic evidence about the taxonomy and biogeography between about 5 and 8 million years ago (Ma) (Bradley, of each of the great apes. To our knowledge there is no 2008). New estimates based on empirical data about gen- single source of information about great ape skeletal col- eration times (Langergraber et al., 2012) suggest that lections, so in the next section we briefly survey informa- the date is probably closer to 8 than to 5 Ma, although a tion published in print and available online regarding more recent analysis of a larger data set (Prado-Marti- the major collections. We then suggest what types of nez et al., 2013) suggests it is closer to 5 Ma, and this information it would be useful to know about a collection date may be further affected by recalibrations of the before visiting it. In the main section of this review, we molecular clock based on the newly discovered Oligocene present a “biography” of one of the more comprehensive catarrhine Rukwapithecus fleaglei, which has been collections; for various reasons the exemplar we have argued to be a basal hominoid (Stevens et al., 2013). chosen is the Powell-Cotton Collection. We set out its Whole genomes can now be sequenced with acceptable history, how it came to be recognized as a potentially levels of coverage, and in the last few years researchers important source of comparative information about the have published good draft sequences of the genomes of African apes, then we review what use has been made of the chimpanzee (Chimpanzee Sequencing and Analysis it, and what types of data were collected. We introduce Consortium, 2005), the orangutan (Locke et al., 2011), the Human Origins Database (www.humanoriginsdata- the gorilla (Scally et al., 2012), and the bonobo (Prufer€ base.org), a freely-available online relational database et al., 2012). Scally et al. (2012) sampled two Western we constructed as a pilot of the type of detailed informa- Lowland and one Eastern Lowland gorilla and showed tion we suggest should be available for all great ape col- that if you take all of the genome then the greatest num- lections. Finally, we present suggestions about how the ber of similarities are between modern humans and community might collectively move forward to help pre- chimpanzees. However, they also found that “in 30% of serve and protect great ape skeletal collections for future the genome, gorilla is closer to human and chimpanzee generations. than the latter are to each other” (ibid, p. 169), a phe- nomenon referred to as incomplete lineage sorting (ILS). RELATIONSHIPS AMONG The Prufer€ et al. (2012) study showed that bonobos and EXTANT HIGHER PRIMATES common chimpanzees are 99.7% alike, whereas 98.7% of the bonobo genome resembles that of modern humans. In an essay entitled On the Relations of Man to the The latter authors also found evidence of ILS in their Lower Animals that formed the middle section of a small study in that 3% of the modern human genome is more book entitled Evidence as to Man’s Place in Nature, Hux- closely related to bonobos or to common chimpanzees ley (1863) concluded that the phenetic differences than either of these taxa is related to the other; they between modern humans and the gorilla (and by infer- also suggest that 25% of all genes contain evidence of ence the chimpanzee) were less marked than the differ- ILS. That said, a recent comparative study of 79 great ences between the gorilla and the orangutan and ape genomes representing all six currently recognized gibbons. Most recent attempts to use gross morphologi- species has also emphasized the presence of genetically cal evidence to generate hypotheses about higher pri- distinct populations within each great ape species mate relationships have confirmed the close relationship (Prado-Martinez et al., 2013). between modern humans and the African apes (e.g., It has been clear for several decades now that phyloge- Gibbs et al., 2002; Diogo and Wood, 2011), although netic relationships must be taken into account in com- some still claim to see gross morphological arguments in parative studies (e.g., see Felsenstein, 1985; Harvey and favor a modern human–orangutan sister relationship Pagel, 1991; Garland et al., 1992; Martins and Hansen, (Grehan and Schwartz, 2009). 1997; Nunn, 2011), and thus the correct taxonomic iden- During the first half of the 20th century the focus of tification of specimens in comparative collections is the search for evidence about higher primate relation- important in the context of correct phylogenetic

American Journal of Physical Anthropology 4 A.D. GORDON ET AL. most western Pan isolate, the subspecies Pan troglodytes verus (also known as the Western chimpanzee) extends from Senegal in the west, through Liberia to the Ivory Coast in the east. Morin et al. (1994) used evidence from mitochondrial DNA (mtDNA) to suggest that this isolate should be recognized as a separate species, but few have adopted this suggestion. Gonder et al. (1997) suggested that the mtDNA of chimpanzees from western Nigeria and from localities along the Nigeria-Cameroon border is distinct from that of P. t. verus, and this interpretation was confirmed by a more recent study of microsatellites (Gonder et al., 2011). Groves (2001) suggested that the latter popula- Fig. 1. Current consensus of phylogenetic relationships tions should be recognized as a separate subspecies, P. among modern humans and great ape species, and current sub- troglodytes vellerosus, also called the Nigerian- species designations in their presumed phylogenetic position Cameroonian chimpanzee. However, for reasons of prior- (dotted lines indicate branches leading to subspecies). Branch ity these populations have since been renamed as P. lengths within genera are not necessarily proportional to time. troglodytes ellioti (Oates et al., 2009). Note that the division of the eastern chimpanzee into Pan trog- The range of the subspecies P. troglodytes troglodytes, lodytes schweinfurthii and P. t. marungensis is based on a mor- or the Central chimpanzee, extends from its northern phological distinction that is not supported by molecular evidence (Gonder et al., 2011). Phylogenetic relationships within boundary with P. t. ellioti in Northern Cameroon (this each genus are drawn from a variety of sources (Pan: Groves, approximates to the Sanaga River) to the Oubangui 2005; Gonder et al., 2011; Gorilla: Groves, 2001; Scally et al., River in the east and the Congo River to the east and 2012; Pongo: Brandon-Jones et al., 2004; Singleton et al., 2004; south. The Oubangui River separates the range of P. t. Locke et al., 2011); more recently, Prado-Martinez et al. (2013) troglodytes from that of the Eastern chimpanzee, P. trog- suggested that P. t. verus is sister to P. t. ellioti rather than sis- lodytes schweinfurthii, and the Congo River separates it ter to a clade including all other P. troglodytes subspecies. from the range of the bonobo (see above). The Eastern chimpanzee is found from the Oubangui placement. However, taxonomic information associated River eastwards into Western Uganda, Rwanda, and with individual museum specimens is often dated and Tanzania; its southern range extends to the Congo River. may lag behind contemporary consensus taxonomy. Cor- Although often recognized as a single subspecies, P. t. rect taxonomic identification is also inextricably bound schweinfurthii, Groves (2005) has argued that this group up with accurate biogeographic provenance of speci- should be divided into two subspecies on the basis of cra- mens, information that regrettably is often poorly docu- niometric variation, but Gonder et al. (2011) found no mented. In the next two section, we review the current genetic support for such a division. In Groves’ scheme, P. consensus on the taxonomy and geographic distribution t. schweinfurthii occupies the northwestern part of the of Pan, Gorilla, and Pongo (Fig. 1). range of the Eastern chimpanzee, including the Ituri district in the Democratic Republic of the Congo as well as north and west of there. P. troglodytes marungensis is GREAT APE TAXONOMY AND BIOGEOGRAPHY found in Uganda and in the Maniema district in the Pan DRC, as well as in the southeast extent of the range. Pilbrow (2006) provides a useful detailed review of the The present consensus (e.g., Groves, 2001; Prufer€ literature covering the molecular and morphological evi- et al., 2012) is that the genus Pan includes at least two dence for P. troglodytes. Her odontometric data are con- species: Pan paniscus, the bonobo, and Pan troglodytes, sistent with other morphological evidence (Braga, 1995; the common chimpanzee (Fig. 1). Uchida, 1996; Braga, 1998; Guy et al., 2003; Taylor and The range of P. paniscus is presently confined to the Groves, 2003; Lockwood et al., 2004) and with genetic region in the Democratic Republic of the Congo that lies evidence that all suggest that P. t. verus is the most dis- to the south of the Congo River. Prufer€ et al. (2012) tinctive subspecies of P. troglodytes. Her data also sup- found “no indication of preferential gene flow between port the genetic and other morphological evidence for bonobos and any of the chimpanzee groups tested,” and the distinctiveness of P. t. ellioti. they suggest that this is “consistent with the suggestion that the formation of the Congo River c.2.5–1.5 Ma cre- Gorilla ated a barrier to gene flow that allowed bonobos and chimpanzees to evolve different phenotypes over a rela- The present consensus (e.g., Groves, 2001; Pilbrow, tively short time” (ibid, p. 528). The time range of esti- 2010; Scally et al., 2012) is that the genus Gorilla mates for the separation of the two extant species includes two species: Gorilla gorilla, the Western gorilla, matches the range of 2.6–1.5 Ma cited by Langergraber and Gorilla beringei, the Eastern gorilla (Fig. 1). When et al. (2012), and it is also consistent with the older Scally et al. (2012) compared the genomes of the West- end of the slightly later range of 1.7–0.75 Ma based ern and Eastern lowland gorilla individuals, they esti- on genome-level comparisons (Prado-Martinez et al., mated “an average sequence divergence time 1.75 Ma, 2013). but with evidence for more recent genetic exchange and The geographical range of P. troglodytes extends from a population bottleneck in the eastern species” (ibid, p. Senegal and Guinea in the west to Uganda and Tanza- 169); this is also consistent with the results of Prado- nia in the east, but it is not continuous between these Martinez et al. (2013) (i.e., divergence 1.7–0.75 Ma). The two longitudinal extremes for there are no chimpanzees original dates proposed by Becquet and Przeworski in the relatively arid Dahomey Gap. The range of the (2007) and Thalmann et al. (2007) for the split, 1.29 Ma

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 5 and 2.13–1.2 Ma, respectively, have recently been recali- The names and distributions of the three proposed brated by Langergraber et al. (2012) to 1.8 and 3.01– subspecies of P. pygmaeus are as follows: P. p. pygmaeus: 1.69 Ma, respectively. Northwest Bornean Orangutan, known from Sarawak The Western gorilla, or G. gorilla, is divided into two (Malaysia) and Northwest Kalimantan (Indonesia); P. p. subspecies (Groves, 2001). The bigger of the two in wurmbii: Central Bornean orangutan, known from terms of geographical range and population size is G. Southwest and Central Kalimantan (Indonesia), and P. gorilla gorilla, whose range extends from Cameroon p. morio: Northeast Bornean orangutan, known from (south of the Sanaga River) southwards through Equato- East Kalimantan (Indonesia) and Sabah (Malaysia). rial Guinea, the Gabon and Congo down to the mouth of The review presented above uses the contemporary the Congo River. Eastward, the range extends into the names for the various species and subspecies, but many Congo and the Central African Republic close to the museum collections antedate contemporary usage. The Oubangui River. In the southern part of this range goril- junior synonyms of the current taxa may have been las are not known beyond the Congo River in the Demo- used in some of the older collections, so we list these in cratic Republic of the Congo. The second subspecies of Table 1. In Table 2 we list the names that have previ- the Western gorilla, G. gorilla diehli, or the Cross River ously been used for the countries we refer to above. gorilla, is a small isolated population that straddles the Maps delineating the boundaries between taxa can be Nigeria-Cameroon border in the upper Cross River found on the website of the IUCN Red List of Threat- highlands. ened Species (http://www.iucnredlist.org). The Eastern gorilla, or G. beringei, is also divided into Unfortunately, many collections do not provide either two subspecies (Groves, 2001). The larger of the two in current or obsolete taxonomic information that corre- terms of geographical range and population size is G. sponds to current subspecific or even specific designa- beringei graueri, the Eastern Lowland or Grauer gorilla, tions, but rather lump all gorillas as G. gorilla, all whose range extends east from the Lualaba River into common chimpanzees (as opposed to bonobos) as P. trog- the Mitumba Mountains and as far south as Lake Tan- lodytes, and all orangutans as P. pygmaeus. As a conse- ganyika. The second eastern gorilla subspecies, G. berin- quence, the information derived from field notes gei beringei, or the Mountain gorilla, is represented in associated with each specimen are of critical importance. two small, isolated populations. One, in the Virunga Vol- In the next section, we consider the acquisition and dis- canoes, straddles the border between Rwanda, Uganda, position of these collections in general, and the informa- and the Democratic Republic of the Congo; the other is tion about them that is available online. located in the Bwindi Impenetrable Forest in southwest- ern Uganda. MAJOR RESEARCH COLLECTIONS OF GREAT Pilbrow (2010) provides a useful detailed review of the APE SKELETONS: PHYSICAL COLLECTIONS literature covering the molecular and morphological evi- AND ONLINE PRESENCE dence for Gorilla. Her odontometric data are consistent with recognizing the distinctiveness of G. g. diehli (see The primary destinations of “natural history” material above) and they also support the distinctiveness of the collected throughout the 19th and 20th centuries were regional subpopulations of G. b. graueri in Tshiaberimu Europe and the United States, thus it is not surprising and Kahuzi-Biega. Long ago these two populations had that this is where most of the large collections of great been flagged as a separate subspecies, G. beringei rex- ape skeletal material are found. Some of these collec- pygmaeorum (Schwarz, 1927). tions were acquired through field trips designed for that express purpose by scientists with the backing of aca- Pongo demic institutions and/or museums. A case in point is the Asiatic Primate Expedition of the late 1930s, which The present consensus (e.g., Groves, 2001; Locke included the physical anthropologists Adolph Schultz, et al., 2011) is that the genus Pongo includes two spe- Clarence Ray Carpenter, and a young Sherwood Wash- cies: Pongo pygmaeus, the Bornean orangutan, and burn, and which was backed by the Carnegie Institution, Pongo abelii, the Sumatran orangutan (Fig. 1). Columbia University, and Harvard University ([Anony- The ancestors of orangutans most likely migrated mous], 1937). The specimens in this collection (which, in from the mainland to Sumatra and from there to Bor- addition to orangutans, also include hylobatids, mon- neo. Steiper (2006) is in favor of a deep (c.2 Ma) diver- keys, lorisiforms, a tarsier, and many other mammals gence and suggests that the “Bornean and Sumatran and birds; Coolidge et al., 1940) are associated with orangutans are independent lineages” that are “in an detailed field notes regarding collection locality, date col- early stage of speciation” (ibid, p. 520). However, when lected, and in some cases, accurate drawings of skeletal Locke et al. (2011) compared the genomes of the two spe- elements and measurements made by Adolph Schultz. cies they estimated the split time to be more recent, c.1 However, many other collections were acquired through Ma (ibid, Fig. 1, p. 530); this is also consistent with the purchases made from individual collectors or through results of Prado-Martinez et al. (2013). companies that specialized in the purchase and distribu- Warren et al. (2001), on the basis of variation in the tion of skeletal specimens. The field data associated with control region of the mtDNA, identified four distinct sub- such collections vary in quality: some individual collec- populations with particular regional diversity and geo- tors took careful notes (including geographic location, graphic clustering: (1) Southwest and Central collection date, body mass, linear measurements of the Kalimantan; (2) Northwest Kalimantan and Sarawak; body, etc.) that were sent along with specimens, while (3) Sabah; and (4) East Kalimantan. Brandon-Jones other specimens were purchased in markets in source et al. (2004) recognized two subspecies of the Bornean countries from local hunters who provided no other orangutan: P. p. pygmaeus and P. p. wurmbii.An information at all. Unfortunately, it is not that uncom- additional subspecies, P. p. morio, was recognized by mon for collections to list the “country” of origin for Singleton et al. (2004). some gorilla and chimpanzee specimens as “Africa.”

American Journal of Physical Anthropology 6 A.D. GORDON ET AL.

TABLE 1. Junior synonyms for the genus, species, and subspecies of Gorilla, Pan, and Pongo adapted (with the author’s permis- sion) from Groves (2001, 2005)

Genus Gorilla (I. Geoffroy, 1853) Gorilla gorilla gorilla (Savage, 1847) 1847 Troglodytes gorilla Savage. 1848 Troglodytes savagei Owen. 1855 Gorilla gina I. Geoffroy. 1856 Satyrus adrotes Mayer. Replacement for Troglodytes gorilla. 1856 Sat[yrus] africanus Mayer. Replacement for Troglodytes gorilla. 1862 Gorilla castaneiceps Slack. 1877 Gorilla may^ema Alix and Bouvier. 1903 Gorilla gigas Haeckel. 1905 Gorilla gorilla matschiei Rothschild. 1905 Gorilla jacobi Matschie. 1912 Gorilla gorilla schwarzi Fritze. 1914 Gorilla hansmeyeri Matschie. 1914 Gorilla zenkeri Matschie. 1927 G[orilla] uellensis Schouteden. 1927 Gorilla gorilla halli Rothschild. 1943 Gorilla (Pseudogorilla) ellioti Frechkop.

Gorilla gorilla diehli (Matschie, 1904) 1904 Gorilla diehli Matschie.

Gorilla beringei beringei (Matschie, 1903) 1903 Gorilla beringeri Matschie. 1905 Gorilla beringei Matschie. 1917 Gorilla beringei mikenensis Lonnberg.€

Gorilla beringei graueri (Matschie, 1914) 1908 Gorilla manyema Rothschild. Lapsus for may^ema Alix and Bouv- ier, 1877. 1914 Gorilla graueri Matschie. 1927 Gorilla gorilla rex-pygmaeorum Schwarz.

Genus Pan (Oken, 1816) 1812 Troglodytes E. Geoffroy. Troglodytes niger E. Geoffroy, 1812. 1816 Pan Oken. Pan africanus Oken, 1816. 1828 Theranthropus Brookes. Troglodytes niger E. Geoffroy. 1838 Anthropopithecus de Blainville. Simia troglodytes Blumenbach, 1799. 1841 Hylanthropus Gloger. Simia troglodytes Blumenbach, 1799. 1860 Pseudanthropus Reichenbach. Replacement for Troglodytes. 1866 Pongo Haeckel. Replacement for Troglodytes. Not of Lacepe `de, 1799 (Ponginae). 1866 Engeco Haeckel. Simia troglodytes Blumenbach, 1799. 1895 Anthropithecus Haeckel. Emendation of Anthropopithecus. 1905 Fsihego de Pauw. Fsihego ituriensis de Pauw, 1905. 1954 Bonobo Tratz and Heck. Pan satyrus paniscus Schwarz, 1929.

Pan troglodytes (Blumenbach, 1799)

Pan troglodytes troglodytes (Blumenbach, 1799) 1758 Simia satyrus Linnaeus (in part). Name suppressed by the Inter- national Commission on Zoological Nomenclature (1929), Opin- ion 114. 1792 Simia satyrus pongo Kerr. 1792 Simia satyrus jocko Kerr. 1799 Simia troglodytes Blumenbach. 1812 Troglodytes niger E. Geoffroy. 1816 Pan africanus Oken. 1831 Troglodytes leucoprymnus Lesson. 1840 Anthropopithecus pan Lesson. 1855 Troglodytes tschego Duvernoy. 1856 Satyrus lagaros Mayer. 1856 Satyrus chimpanse Mayer. 1860 Troglodytes calvus du Chaillu. 1860 Troglodytes kooloo-kamba du Chaillu. 1862 Troglodytes vellerosus Gray. 1866 Troglodytes aubryi Gratiolet and Alix. 1870 Pseudanthropus fuliginosus Schaufuss. 1876 Anthropopithecus angustimanus Brehm. Nomen nudum.

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 7

APPENDIXTABLE 1. Continued

Genus Gorilla (I. Geoffroy, 1853) 1895 Anthropopithecus fuscus Meyer. 1899 Troglodytes livingstonii Selenka. Nomen nudum. 1903 Anthropithecus mafuca Haeckel. Nomen nudum. 1905 Simia pygmaeus raripilosus Rothschild. 1914 Anthropopithecus reuteri Matschie. 1914 Anthropopithecus ochroleucus Matschie. 1919 Anthropopithecus schneideri Matschie. 1919 Anthropopithecus pusillus Matschie. 1932 Anthropopithecus heckii Koch.

Pan troglodytes verus (Schwarz, 1934) 1904 Simia chimpanse Matschie. Not of Mayer, 1856. 1934 Pan satyrus verus Schwarz.

Pan troglodytes ellioti (Gray, 1862) 1914 Anthropopithecus ellioti Matschie. 1914 Anthropopithecus oertzeni Matschie. 1919 Anthropopithecus papio Matschie.

Pan troglodytes schweinfurthii (Giglioli, 1872) 1872 Troglodytes schweinfurthii Giglioli. 1905 Fsihego ituriensis de Pauw. 1912 Simia (Anthropopithecus) nahani Matschie. 1912 Simia (Anthropopithecus) ituricus Matschie. 1912 Simia (Anthropopithecus) kooloo-kamba yambuyae Matschie. 1914 Anthropopithecus schubotzi Matschie. 1914 Anthropopithecus steindachneri Lorenz.

Pan troglodytes marungensis (Noack, 1887) 1887 Troglodytes niger var. marungensis Noack. 1899 Troglodytes livingstonii Selenka. Nomen nudum. 1912 Simia (Anthropopithecus) cottoni Matschie. 1912 Simia (Anthropopithecus) adolﬁ-friederici Matschie. 1914 Anthropopithecus purschei Matschie. 1914 Anthropopithecus pfeifferi Matschie. 1914 Anthropopithecus graueri Matschie. 1914 Anthropopithecus calvescens Matschie. 1914 Anthropopithecus castanomale Matschie.

Pan paniscus (Schwarz, 1929) 1929 Pan satyrus paniscus Schwarz.

Genus Pongo (Lacepe `de, 1799) Pongo pygmaeus pygmaeus (Linnaeus, 1760) 1760 Simia pygmaeus Linnaeus. 1896 Pithecus satyrus landakkensis Selenka. 1896 P[ithecus] satyrus batangtuensis Selenka. 1896 P[ithecus] satyrus dadappensis Selenka. 1896 P[ithecus] satyrus genepaiensis Selenka. 1896 P[ithecus] satyrus skalauensis Selenka. 1896 P[ithecus] satyrus rantaiensis Selenka. 1896 P[ithecus] tuakensis Selenka.

Pongo pygmaeus morio (Owen, 1837) 1837 Simia morio Owen. Borneo. 1853 Pithecus brookei Blyth. Sarawak. 1853 Pithecus owenii Blyth. Sarawak. 1855 Pithecus curtus Blyth. Sarawak.

Pongo abelii (Lesson, 1827) 1827 Pongo abelii Lesson. 1841 Simia gigantica Pearson. 1896 P[ithecus] satyrus deliensis Selenka. 1896 P[ithecus] satyrus langkatensis Selenka. 1896 P[ithecus] satyrus abongensis Selenka.

Current designations are presented with taxon name ﬁrst; junior synonyms are preceded by the date of publication. Depending on the date when specimens were accessioned in collections, these taxon names may have been used in place of the currently valid taxa. Note the prevailing use of Simia, Troglodytes, and Anthropopithecus as the genus name for chimpanzees, and Pithecus as the genus name of orangutans. Readers wanting further information about these taxa and their geographical distribution should con- sult Groves (2001).

American Journal of Physical Anthropology 8 A.D. GORDON ET AL.

TABLE 2. History of names of nations in which great apes are found

Contemporary country names Names used during the accumulation of Western primate collections Africa Cameroon German colony of Kamerun: 1884–1919 French Cameroun and British Cameroons: 1919–1961 Republic of Cameroon: 1961 to present Central African Republic Oubangi-Chari: 1903–1908 Part of Oubangi-Chari-Tchad in French Equatorial Africa: 1908–1920 Oubangi-Chari (part of French Equatorial Africa): 1920–1958 Central African Republic: 1968–1958 Central African Empire: 1976–1979 Central African Republic: 1979 to present Congo French Congo: 1882–1903 Middle Congo: 1903–1910 Congo (part of French Equatorial Africa): 1910–1960 Republic of the Congo: 1960 to present Cote^ d’Ivoire (Ivory Coast) Cote^ d’Ivoire: 15th century to 1986 (independence from France in 1960) Republique de Cote^ d’Ivoire: 1986 to present Democratic Republic of the Congo Congo Free State: 1885–1908 Belgian Congo: 1908–1960 Republic of the Congo: 1960–1964 Democratic Republic of the Congo: 1964–1971 Zaire: 1971–1997 Democratic Republic of the Congo: 1997 to present Equatorial Guinea Spanish Guinea: 1926–1968 Republic of Equatorial Guinea: 1968 to present Gabon Gabon (French colony): 1885–1910 Gabon (part of French Equatorial Africa): 1910–1960 Gabonese Republic: 1960 to present Guinea French Guinea: 1895–1958 Republic of Guinea: 1958 to present Liberia Referred to as the Grain or Pepper Coast prior to colonization in 1822 Liberia: 1822–1847 Republic of Liberia: 1847 to present Nigeria Uniﬁcation of Southern and Northern Nigeria: 1914 Colony and Protectorate of Nigeria: 1914–1960 Republic of Nigeria: 1960 to present Rwanda Rwanda (part of German East Africa): 1885–1919 part of Ruanda-Urundi (Belgian colony): 1919–1962 Republic of Rwanda: 1962 to present Senegal Part of French West Africa: 1895–1958 Part of Mali Federation: 1959–1960 Part of Senegambia: 1982–1989 Republic of Senegal: 1960 to present Tanzania Part of German East Africa: 1885–1919 Tanganyika (British East Africa): 1920–1961 Republic of Tanganyika: 1961–1964 Zanzibar and Pemba merged with Tanganyika to form the United Republic of Tanzania: 1964 United Republic of Tanzania: 1964 to present Uganda British East Africa or East Africa Protectorate: 1890–1962 Republic of Uganda: 1962 to present Asia Indonesia (includes Sumatra and part of Borneo) Republic of Indonesia: 1949 to present Malaysia (includes part of Borneo) Straits Settlements: 1826–1946 British North Borneo: 1882–1963 Malay States: 1895–1946 Federation of Malaya: 1948–1963 Sabah and Sarawak merged with Malaya: 1963 Malaysia: 1963 to present Brunei (part of Borneo) Sultanate of Brunei: 15th century to 1959 Nation of Brunei, the Abode of Peace: 1959 to present

The field data that were recorded have been pre- log books, or preserved as separate field notes in an served in a variety of ways, some having to do with the archive. This information may be variably recorded collectors and methods of field preparation, others hav- even within the same collection, and it is probably the ing to do with past and present curatorial practices of case that many researchers have visited a museum, col- the institutions in which they reside. Field data may be lected their data, and left, without ever knowing of the recorded on tags attached to skins or skeletal elements, existence of some important field data relevant to their written on skulls, recorded on box labels or in museum research.

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 9 Field data are also occasionally associated with the With the advent of the internet, which became available wrong specimen or cannot be attributed to a particular to the general public in the 1990s, the better-resourced specimen. When field-prepared materials arrived at museums began building websites, and now even small museums, in some cases they were incorrectly matched local museums have some web presence. However, most of with their field notes, and in others cases elements from these websites tend to be geared primarily to the public the same specimen were incorrectly labeled; one visitor rather than the research community. Even in those reviewer noted that he or she has had to match skeletal cases where some collection information is available online, specimens and skins by their bullet holes. It is also not the types of data and the manner in which they are pre- unheard of for sex to be recorded incorrectly. This is sented (e.g., specimen-level data versus species-level aggre- obvious in the case of adult gorillas and orangutans, but gates) vary from institution to institution, and information is much less so for chimpanzees or for juveniles of any that is key for researchers intending to use a skeletal col- species. In some cases curators recognized attribution lection (see below) is often lacking. errors at the time of preparation, resulting in cataloging We suggest that there are several categories of infor- practices that at least alert researchers to the problem mation that it would be useful to have before making (e.g., at the Royal Museum of Central Africa in Tervu- plans to visit one of the institutions housing a collection ren, Belgium, the long bones from several G. b. graueri of great apes. They include the following: individuals all received the same catalog number and are stored in the same box). Situations like this are frus- 1. Sex-specific counts of specimens identified to the trating, particularly when they involve rare taxa among lowest taxonomic level possible. In the past, the already rare great ape collections (as in the case researchers were often content to collect data at the above), but individual attribution of elements may be genus level, but as noted above, over the past few deca- resolvable in time through genetic sampling if adequate des taxonomic designations of Gorilla, Pan,andPongo financial resources can be found to devote to the have been revised in favor of finer-scale divisions, and problem. these divisions are thought to represent distinct line- Access to resources surely plays a role in determining ages with differences that may be relevant to evolution- whether specimen-level data from the field have been ary questions (e.g., Tocheri et al., 2011). Although migrated to digital formats and made available online. museums usually identify specimens to at least the spe- Some collections are in publicly-funded national muse- cies level, in many cases these designations were made ums (e.g., the Smithsonian, the Natural History when all gorillas and all orangutans were considered to Museum in London) or major national or state-level belong to a single species in their respective genera. research institutes (e.g., the Zoologische Staatssamm- 2. Geographical information for specimens identi- lung Munchen).€ Others are in municipal museums that fied to the smallest geographical region possible, are public–private partnerships (e.g., the American and preferably by latitude and longitude. Biogeo- Museum of Natural History, the Cleveland Museum of graphic provenance is critical information that is often Natural History), and some are in museums that are neglected both in terms of data made available in col- still in private hands (e.g., the Powell-Cotton Museum). lections and data used by researchers. Where taxonomic A consequence of these different circumstances is that designations are out of date, finer-scale geographic there is wide variation in the amount and type of infor- information can help identify the subspecies present in mation available online for each of these collections the area where a specimen was collected. Although in (Table 3). some cases the nation of origin may be helpful, in other Historically, obtaining detailed information regarding cases the same modern political entity is home to multi- great ape skeletal collections without visiting them has ple subspecies. Latitude and longitude information from been difficult. Researchers and students usually learned original field notes is not common in many cases, but it about the existence, scope, and content of the various is invaluable for maintaining accurate taxonomic desig- collections from acknowledgment sections in journal nations for those specimens that have it. In addition, articles, from their academic supervisor, or from other detailed geographic information allows for evaluation of researchers in the field. In our experience such informa- the contribution of biogeographic variation to overall tion is often vague. A few journal articles from the late morphological variation within a taxon, an important 1960s to the early 1980s provided summaries of counts consideration when making comparisons with fossil of specimens by species at some museums (Tappen, samples that may be drawn from geographically (and 1969; Almquist, 1973; Albrecht, 1982), but they do not temporally) distant portions of a taxon’s range. go beyond basic “skin, skull, skeleton” distinctions, nor 3. Counts of specimens by skeletal element. Many do they provide geographical information. The most museums provide information on the number of skulls, detailed published information about primate collections the number of sets of postcrania, and the number of began as a catalog of primates in the Natural History skins. But it would be more useful to have information Museum in London that was later expanded to include about the presence or absence of each skeletal element, the primates in all British collections. The first volume for in many cases only portions of the postcranium are focused on platyrrhines (Napier, 1976) with the volume preserved, and some, or all, of the teeth may be miss- on hominoids coming out fourteen years later (Jenkins, ing from a cranium or mandible. 1990). Individual specimens located in British museums 4. Information on the maturity of individual speci- are listed in the latter volume by taxon, plus, where mens. Museums occasionally provide information on available, information about locality and age (adult, the number of adult and non-adult specimens, juvenile, or infant), but there is no detailed information although the criteria by which maturity is assessed about the elements present nor about the condition of are usually not indicated. Ideally, each specimen the specimens. In any event, the hominoid volume is would be associated with a set of developmental vari- limited to collections housed in the UK, and because it is ables indicating relative maturity (e.g., epiphyseal out of print it is difficult to get access to a copy. fusion of various elements, dental eruption).

American Journal of Physical Anthropology TABLE 3. Summary of the information available online for major and commonly used great ape skeletal collections in the US and Europe

Other specimen information (e.g., Information on Information on the wild vs. captive, Information regard- Web address for collection Counts of specimens Geographical Counts of specimens maturity of individ- condition of speci- reproductive ing previously col- Pubs citing Collection information by taxon? information? by skeletal element? ual specimens? mens/ elements? condition)? lected data? the collection? Notes

AMNH http://entheros.amnh.org/db/ Yes (some to species, Variable: from no Skin/Skull/ Skeleton No Only “disarticulated” Occasionally data on Occasionally in log No Attached to each speci- emuwebamnh/index.php some to information to or “mounted” captive/wild book scans men page there is a subspecies) specific locality “multimedia” cate- within a province gory. Some photo- graphs of the specimens are available and a scanned image of their entry in the catalogue of specimens. BMNH http://www.nhm.ac.uk/ Yes (species only) Rarely Skin/Skull/ Skeleton Age categories No No None No Attached to many research-curation/ in some cases (undefined) specimens there is a scientific-resources/ low resolution collections/zoological- image of the collec- collections/zoology-specimen- tions register, but database/index.php any associated information is not legible. CMNH http://www.cmnh.org/site/ Yes (species only) Most identify nearest Postcranium/Skull Age categories No Occasionally data on None No Available as spread- ResearchandCollections/ town (present or (undefined) captive/wild, no sheet, not online PhysicalAnthropology/ absent) reproductive sta- database Collections/Hamann- tus data ToddCollection/ Databases.aspx FMNH http://emuweb. Yes (some to species, Variable: from no Skin/Skull/ Skeleton No Rarely in “collection Often data on cap- None No Collection notes field fieldmuseum.org/ some to information to and a few specific notes” for tive/wild, no with useful but non- mammals/Query.php subspecies) specific locality elements elements reproductive sta- standardized with latitude and tus data information longitude MCZ http://mczbase.mcz.harvard. Yes (some to species, Variable: from no Skin/Skull/ Skeleton Age categories (unde- Rarely in log book Usually data on cap- Measurements No Attached to many edu/SpecimenSearch.cfm some to information to and a few specific fined) but room remarks (attached tive/wild occasionally specimens there is subspecies) specific locality elements for remarks as media file to provided an image of the within a province record), general collections register condition some- as well as any times noted in associated data database such as standard measurements, information about specific elements, etc. MNHN http://www.mnhn.fr/museum/ No No No No No No None No The mammal collection foffice/science/science/ is currently being ColEtBd/collectionsMuseum/ entered into a collectionSci/ database FicheCollection.xsp? COLLECTION_ COLLECTION_ID5259& COLLECTION_ID5259 &idx563&nav5liste MNK http://www. No No No No No No None No The museum web site naturkundemuseum-berlin. includes a link to de/en/sammlungen/zoologie/ the GBIF portal (see mammalogie/ text), but appears to only include animal audio recording and fossil data. APPENDIXTABLE 3. Continued

MRAC http://www.africamuseum. Yes (subspecies) Country Skin/Skull/ Skeleton Age categories (unde- No Wild status can usu- None No Available as spread- be/research/biology/ fined), notes latest ally be inferred sheet, not online vertebrates/mammalogy/ erupted tooth for from locality data database collections each specimen NHMV http://www.nhm-wien.ac.at/ Yes (species only) Country No No No No None No Database provides tal- en/research/_zoology_ lies by prepartion vertebrates/mammal_ (Skin, Skull, Skele- collection/database ton, Dermoplast, Fluid) and sex, not individual specimen data NMNH http://collections.mnh.si. Yes (subspecies) Country, Relation to Skin/Skull/ Skeleton “Stage” and Condition category Usually data on cap- Measurements often Rarely Some specimens have edu/search/mammals/ town, Region and most ele- “Remarks” cate- for specimens tive/wild, provided attachments such as ments in many gory for different usually blank “Reproductive cranial X-rays, cases ages usually Condition” field pathology reports blank present in database but typically empty NSF http://sesam.senckenberg.de/ Yes (most to species, Country Skin/Skull/ Skeleton Age categories occa- No Wild status can be Measurements occa- No some to sionally provided inferred from sionally provided subspecies) (undefined) locality data PCM http://www.quexpark.co. No* No* No* No* No* No* None* No* *No institutional data- uk/education-and- base, but see text research/research/ natural-science.html RBINS https://darwin. Yes (subspecies) Country No Age (undefined) No Wild status can be None No naturalsciences. inferred from be/search/search locality data RMNH http://science.naturalis. No No No No No No None No No specific information nl/collections/ about collection collections-in- available online naturalis/ vertebrates UZH http://www.aim.uzh.ch/ Yes (subspecies) No Tallies for Skeleton/ Tallies for adults No No None No Not a database, tallies Research/Collection/ Crania/Postcra- (undefined) of totals not individ- Prim.pdf nia/Cadavers ual specimen data YPM http://peabody.yale. Yes (some to species, Country or no Skin/Skull/ Skeleton Rarely age descriptor No Occasional data on None No edu/collections/ some to information in “Other attrib- captive/wild in search-collections?mam subspecies) utes” field “Collected” field ZSM http://www.zsm.mwn. No No No No No No None No Only information about de/mam/ptypes.htm type specimens is available online; no information about the extensive orangutan collection is available

Collection abbreviations are as follows: AMNH: American Museum of Natural History; BMNH: British Museum, Natural History; CMNH: Cleveland Museum of Natural History; FMNH: Field Museum of Natural History, Chicago; MCZ: Museum of Comparative Zoology, Harvard University; MNHN: Museum National d’Histoire Naturelle, Paris; MNK: Museum fur€ Naturkunde, Berlin; MRAC: Musee Royal de l’Afrique Centrale; NHMV: Naturhistoriches Museum, Vienna; NMNH: National Museum of Natural History, Smithsonian Institution; NSF: Naturmuseum Senckenberg, Frankfurt; PCM: Powell-Cotton Museum, Birchington; RBINS: Royal Belgian Institute of Natural Sciences, Brussels; RMNH: Naturalis (formerly the Rijksmuseum van Natuurlijke History), Lei- den; UZH: University of Zurich; YPM: Yale Peabody Museum; ZSM: Zoologische Staatssammlung Munchen.€ 12 A.D. GORDON ET AL. 5. Information on the physical condition of speci- POWELL-COTTON COLLECTION mens/elements. Although an element may be pres- OF AFRICAN APES ent in the collection, it may be rendered unusable for a given study because of in vivo pathology, post mor- We focused on this collection because it is the one tem damage, or post mortem destructive sampling. ADG and BW know best and because we suspect it is Furthermore, researchers interested in conducting the great ape collection for which the most comprehen- destructive sampling could identify elements that sive contextual information is available. The collection of have undergone previous destructive sampling, great ape skeletons (and skins) is just one component of researchers interested in documenting the frequency a large collection of Asian and African mammals within or nature of in vivo pathologies would have informa- the Powell-Cotton Museum. tion on collections and individuals to focus on, etc. 6. Additional information related to specimen con- History of the Powell-Cotton collection dition at time of collection. There is a variety of other information related to specimens that is useful The Powell-Cotton Museum, which is located in Quex to know when building a study sample, including Park on the outskirts of the village of Birchington in information on whether individuals were pregnant or Northeastern Kent, England, was founded in 1896 to lactating at the time of capture, or whether they were provide a home for the collections generated by Major captive or wild-caught. See Borries et al. (2013) for an Percy Horace Gordon (PHG) Powell-Cotton (1866–1940) illustration of how whether or not one takes into (Cooke and Barton, 1957). Percy Powell-Cotton was a account such metadata can dramatically alter the pat- keen naturalist, collector, and hunter who between 1887 terns identified in primate life history analyses. and 1939 made 28 collecting trips to Africa and parts of 7. Information regarding previously collected Asia; twenty-two of the trips focused on the African con- data. Preferably this would be in the form of the data tinent (see map of expedition routes and area of cover- themselves and a record of who collected the data. age on p. 276 by Cooke and Barton, 1957). Percy Powell- Given that many standard measurements have been Cotton’s goal in Africa was to acquire representative collected multiple times for some collections, such specimens of all of the major game animals in that conti- information would make it possible to assess interob- nent, and the resulting mammal collection is substan- server reliability for those measurements. It would tial. The non-human primate collection is particularly also facilitate a comparison of “standard” measure- extensive and it has long been recognized to be one of ments that may be measured in slightly different the world’s largest skeletal collections of wild-shot West- ways by different researchers. In the absence of the ern gorillas (G. g. gorilla) and chimpanzees (mainly P. t . data themselves, it would still be useful to have infor- troglodytes, but some P. t. schweinfurthii and P. t. ellioti) mation about the type of data collected (e.g., linear ([Anonymous], 1940; Cooke and Barton, 1957; Tappen, distances, radiographs, 3D surface scans, 3D land- 1969; Almquist, 1973). mark data, CT scans) and contact information for the The ape specimens collected by Powell-Cotton came researcher(s) who collected those data. from countries that were, at the time of his various col- 8. Publications citing the collection. Ideally, this infor- lecting trips, called the Congo Free State, the French mation would also include the types of data published in Congo, and the British and French Cameroons (Table 2), the articles that make reference to the collection. but he also purchased specimens from collectors operat- ing in what is now Cameroon. Some specimens were We do not claim that the above is an exhaustive list of purchased from Kurt Zenkerman, a German plantation all of the categories and types of collection information owner who supplemented his income by collecting natu- that could potentially be provided online either publicly, ral history specimens that he sold to European muse- securely, or some combination thereof, but we believe ums. However, the majority of the great apes in the this list covers most of the information a researcher Powell-Cotton Collection (P-CC) were acquired by Fred would need when preparing a grant proposal or when G. Merfield, a British expatriate living in the Cameroons making plans for a research trip. It would allow whom Powell-Cotton met in 1926. These acquisitions researchers to precisely and accurately budget the time were made in the name of the Yaounde Zoological Trad- required for a visit, and it would enable them to make a ing Company, which was formed with Merfield as execu- better informed assessment of whether the data they tive director and Major Powell-Cotton and Mrs. Hannah require could be obtained from previous researchers, Brayton-Slater Powell-Cotton as co-directors, with the thus minimizing wear and tear on irreplaceable speci- purpose of supplying the Powell-Cotton Museum and mens (see below). other European museums with specimens of African As mentioned above, institutions differ in the amount mammals. and type of information they make available for Merfield made a point of establishing friendly relation- researchers. Most of the larger institutions provide some ships with the heads of many villages throughout parts information on several of the points listed above; online of what is now southern and central Cameroon, and information for smaller (and some larger) institutions is after doing so, he persuaded them to collect mammal minimal or non-existent. In the next sections, we specimens for him. Merfield also collected specimens describe a collection at a small private institution with himself and it is estimated that he personally shot c.10– limited financial resources, and then present an online 15% of the specimens he sent to the Powell-Cotton database built by ADG and BW to provide much of the Museum (Malcolm Harman, Personal Communication). information we refer to above for that collection. We Specimens (including apes, monkeys, and other mam- present this as the starting point of a discussion about mals) were numbered in five series of up to 1,000 each. standardizing specimen information across great ape Collection of specimens from the regions of Yaounde, skeletal collections to ensure the maximum utility and Batouri, Yabassi, and Lomie in Cameroon began in 1927 the maximum lifespan for these irreplaceable resources. and continued up to the beginning of WWII in 1939. All

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 13 of the material was sent to the Powell-Cotton Museum colleagues honored this agreement. Charles Oxnard where PHG Powell-Cotton also acted as a distribution recalls “Colin Groves coming with huge calipers to mea- agent. Specimens were sold to museums, to Sir Francis sure long bone lengths,” and in 1966 BW was warmly Collier, and to dealers such as Rowland Ward Ltd. and welcomed in Birmingham when he was working on talus Edward Gerrard & Sons. The arrangement was that the morphology (Day and Wood, 1968, 1969; Wood, 1973). Powell-Cotton Museum was allowed to keep one speci- Zuckerman left Birmingham in 1968 and not long there- men for every one sold on to museums or dealers; the after the ape postcrania were returned to the Powell- museum still has detailed records about all of the speci- Cotton Museum in surplus ammunition boxes. Zucker- mens that were sold. man almost certainly had access to these because from The existence and significance of the wider P-CC was 1960 he was Chief Scientific Adviser to the UK Ministry well known long before data from the great apes in the of Defence, and in 1964 he was appointed the first UK collection began to be used in the scientific literature. Government Chief Scientific Adviser. Publications on Major Powell-Cotton’s collecting expeditions were comparative great ape morphology in the context of reported on regularly in Nature and Science (e.g., [Anon- human evolution that used the P-CC began appearing in ymous], 1899a,b, 1903b–f, 1904a,b, 1905, 1906, 1907a), the late 1960s, and one or more such articles have and the sale and contribution of specimens to the British appeared nearly every year up to the present (Fig. 2; for Museum (now the Natural History Museum) and to col- a bibliography of works referencing the collection go to lections at other institutions such as the Royal College www.humanoriginsdatabase.org). of Surgeons of England were noted in the scientific press (e.g., [Anonymous] 1903, 1907b–d, 1909, 1928, 1934, Nature and taxonomy 1936a, b, 1938). Major Powell-Cotton was well known in natural history circles, and many species and subspecies The P-CC is one of the world’s largest collections of of non-hominoid mammals were given the eponymous Western gorillas (G. gorilla) and Central African chim- name “cottoni.” For example, Lydekker and Matschie panzees (P. t. troglodytes). Western gorillas are repre- note “it is a great pleasure to name this interesting new sented by 148 complete or nearly complete skeletons species after Major P.H.G. Powell-Cotton”, and in many (most with skins) as well as six sets of postcrania with- places in Lydekker’s (1913–1916) Catalogue of Ungulate out skulls, 49 skulls without postcrania, 16 crania with- Mammals in the British Museum it states “type in Major out mandibles, and three mandibles without crania. Powell-Cotton’s collection in Quex Park, Birchington, Central African chimpanzees are represented by 146 Kent” (Colin Groves, Personal Communication). Mam- complete or nearly complete skeletons (most with skins) malian comparative morphologists were certainly aware in addition to 27 skulls without postcrania and 11 par- of the importance of the broader P-CC by the early tial or complete crania without mandibles. There are 1950s, and it was at this time that it apparently came to also six complete skeletons and one unassociated skull the attention of primatologists and students of human that appear to have been collected from the range of P. t. evolution. ellioti, and four complete skeletons and two unassociated skulls of chimpanzees from the Ituri Forest (P. t. African ape component schweinfurthii). The P-CC provides an excellent ontogenetic sequence for both genera. The gorillas range in The first report of research that used data collected maturity from specimens with only a few deciduous from the great apes in the P-CC was by Ashton and teeth erupted to adults with a completely erupted denti- Zuckerman (1951). It is a reasonable inference that Solly tion and complete epiphyseal and cranial suture fusion. Zuckerman was among the first, if not the first, to recog- Chimpanzees expand this range by including a specimen nize and realize the potential for comparative research with no erupted teeth. of the African apes in the collection. During WWII the part of Kent in which Quex Park is situated was effec- tively one large military base with most people in the area involved in the war effort in one way or another, so by the end of the war the majority of the great ape skeletons in the P-CC had not yet been fully macerated. Some anatomical regions (e.g., hands, feet) were still articulated with remnants of flesh and ligaments adher- ing to them. Solly Zuckerman arranged for the postcranial material in the P-CC to be carefully macerated and curated at the Anatomy Department in Birmingham, where in 1939 he had been appointed to the Sands Cox Chair of Anatomy. The arduous and time-consuming work of maceration and curation was undertaken by Ernie Sims, Bill Pardoe, and Tom Spence, under the supervision of Eric Ashton (Charles Oxnard, Personal Communication). Apparently, a quid pro quo of the arrangement between Zuckerman and the Powell-Cotton Museum was that the material was to be kept in Bir- Fig. 2. Time versus cumulative number of research publica- mingham for a period so that it could be used by Zucker- tions in peer-reviewed journals in which the Powell-Cotton Col- man and his colleagues for their research. However, it lection is explicitly cited as a source of primate data. The was also part of the agreement that while in Birming- master table from which these data were extracted, which is ham the material would be made accessible to anyone based on a literature search, is available at the Human Origins else who wanted to work on it. Zuckerman and his Database, www.humanoriginsdatabase.org

American Journal of Physical Anthropology 14 A.D. GORDON ET AL.

TABLE 4. Counts of Pan troglodytes specimens that have initiated and completed each developmental marker (dental eruption or epiphyseal fusion)

Initiation Completion

Pan troglodytes Abbr. M F Total M F Total Tooth/epiphysis deciduous first incisor, maxillary dI1 56 89 145 56 89 145 deciduous second incisor, mandibular di2 56 89 145 56 89 145 deciduous first incisor, mandibular di1 56 88 144 56 88 144 deciduous second incisor, maxillary dI2 56 88 144 56 88 144 deciduous third premolar, mandibular dp3 56 88 144 56 88 144 deciduous third premolar, maxillary dP3 56 88 144 56 88 144 deciduous fourth premolar, mandibular dp4 56 88 144 56 88 144 deciduous fourth premolar, maxillary dP4 56 88 144 56 88 144 deciduous canine, mandibular dc 56 88 144 56 88 144 deciduous canine, maxillary dC 56 88 144 56 88 144 first molar, mandibular m1 52 84 136 51 84 135 first molar, maxillary M1 52 84 136 51 84 135 first incisor, mandibular i1 39 77 116 36 76 112 second molar, mandibular m2 40 75 115 32 73 105 second incisor, mandibular i2 37 77 114 32 73 105 first incisor, maxillary I1 37 75 112 34 74 108 second molar, maxillary M2 36 75 111 30 70 100 second incisor, maxillary I2 35 75 110 33 74 107 third premolar, maxillary P3 30 73 103 30 72 102 fourth premolar, maxillary P4 31 71 102 31 70 101 fourth premolar, mandibular p4 31 71 102 29 70 99 third premolar, mandibular p3 30 71 101 29 70 99 canine, maxillary C286997236891 canine, mandibular c276895236891 humerus, distal HD 26 69 95 21 66 87 humerus, epicondyle HE 24 67 91 20 65 85 pelvis (os coxa) OC 24 67 91 20 64 84 third molar, mandibular m3 24 66 90 22 64 86 ulna, proximal UP 23 66 89 20 65 85 third molar, maxillary M3 22 65 87 22 64 86 femur, lesser trochanter FeLT 21 66 87 20 64 84 femur, greater trochanter FeGT 21 65 86 20 62 82 femur, head FeH 20 65 85 20 61 81 radius, proximal RP 20 64 84 19 61 80 fibula, distal FiD 20 62 82 18 57 75 tibia, distal TD 20 61 81 18 57 75 tibia, proximal TP 20 61 81 17 49 66 fibula, proximal FiP 20 59 79 17 53 70 femur, distal FeD 19 60 79 18 50 68 humerus, proximal HP 19 58 77 16 51 67 radius, distal RD 19 58 77 17 48 65 ulna, distal UD 19 56 75 18 48 66

Counts are for specimens which have complete data for all developmental markers. Dental markers are italicized, postcranial markers are not. Developmental markers are given in order of decreasing number of specimens that have initiated that marker; i.e., developmental markers that initiate (begin eruption or begin fusion) earliest are presented ﬁrst, those that initiate latest are presented last. Some markers appear to develop out of sequence; i.e., they initiate later than other markers but also complete earlier than those markers. The markers that are completed for a higher number of specimens than markers that are initiated for a higher number of specimens (roughly translating to markers that complete earlier than markers which initiated earlier) are shown in bold. For example, more chimpanzee specimens have at least initiated eruption of the second mandibular incisor (i2) than have initiated eruption of the ﬁrst maxillary incisor (I1) (i.e., i2 begins to erupt earlier on average than I1). However, eruption is complete for I1 in more specimens than is the case for i2, indicating that despite the earlier initiation of i2 eruption, I1 tends to be fully erupted earlier on average than i2.

Additional features of the African apes in the P-CC restricted geographical area, providing a large sample in make it a special resource. First, a large number of indi- one collection that records local geographic variation. viduals have most of the postcranial skeleton preserved, Fourth, many have linear measurements of the cadaver, including most with the hand and foot bones—the latter and some have information about body mass. thanks to the careful preparation undertaken in Birming- ham. The hyoid is also preserved for many individuals. THE POWELL-COTTON COLLECTION DATABASE Second, the greater part of the collection is backed up by ﬁeld notes that include the date of collection, the name of For nearly all of the mammal specimens in the P-CC, the locality, and its latitude and longitude in degrees and basic locality information, specimen sex, and general minutes. Third, most of the specimens are from a preparation information (i.e., skin, skull, and/or

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 15

TABLE 5. Counts of Gorilla gorilla gorilla specimens that have initiated and completed each developmental marker (dental eruption or epiphyseal fusion)

Initiation Completion

Gorilla gorilla gorilla Abbr. M F Total M F Total Tooth/epiphysis deciduous first incisor, mandibular di1 71 74 145 71 74 145 deciduous first incisor, maxillary dI1 71 74 145 71 74 145 deciduous second incisor, mandibular di2 71 74 145 71 74 145 deciduous second incisor, maxillary dI2 71 74 145 71 74 145 deciduous third premolar, mandibular dp3 71 74 145 70 74 144 deciduous third premolar, maxillary dP3 71 74 145 70 74 144 deciduous fourth premolar, mandibular dp4 70 74 144 68 72 140 deciduous canine, mandibular dc 69 74 143 68 72 140 deciduous canine, maxillary dC 70 73 143 68 72 140 deciduous fourth premolar, maxillary dP4 70 73 143 68 72 140 first molar, mandibular m1 63 69 132 62 66 128 first molar, maxillary M1 61 70 131 61 66 127 first incisor, mandibular i1 57 61 118 54 61 115 first incisor, maxillary I1 54 61 115 53 56 109 second incisor, mandibular i2 54 60 114 54 58 112 second incisor, maxillary I2 54 60 114 53 53 106 second molar, maxillary M2 53 58 111 53 49 102 second molar, mandibular m2 53 57 110 53 54 107 third premolar, maxillary P3 53 50 103 50 49 99 fourth premolar, maxillary P4 52 51 103 50 47 97 fourth premolar, mandibular p4 52 50 102 49 47 96 humerus, distal HD 49 49 98 38 45 83 third premolar, mandibular p3 50 47 97 49 47 96 canine, mandibular c484694414283 canine, maxillary C494594394382 third molar, mandibular m3 49 44 93 45 37 82 pelvis (os coxa) OC 47 46 93 37 42 79 third molar, maxillary M3 49 44 93 43 36 79 ulna, proximal UP 44 47 91 37 44 81 humerus, epicondyle HE 43 45 88 37 44 81 femur, lesser trochanter FeLT 40 46 86 34 43 77 femur, head FeH 40 45 85 33 40 73 femur, greater trochanter FeGT 38 45 83 33 40 73 radius, proximal RP 35 43 78 32 38 70 tibia, distal TD 33 39 72 29 35 64 fibula, distal FiD 33 39 72 28 34 62 femur, distal FeD 31 38 69 27 35 62 tibia, proximal TP 31 38 69 27 31 58 fibula, proximal FiP 30 38 68 28 33 61 radius, distal RD 29 35 64 26 29 55 humerus, proximal HP 29 35 64 26 27 53 ulna, distal UD 29 35 64 24 27 51

Counts are for specimens which have complete data for all developmental markers. Formatting and abbreviations follow Table 4. skeleton) are available upon request from the museum incorporated into an online relational database and made in spreadsheet format. Presently, Ph.D. student Jaimie available as specimen datasheets through the Human Morris of Canterbury Christ Church University is con- Origins Database (www.humanoriginsdatabase.org). verting these locality data to a digital format and enter- These data are searchable online in a variety of ways, ing them into ArcGIS, with the eventual goal of allowing users to identify whether, given a set of selec- providing these data to researchers. However, until tion criteria, a suitable sample is available within the P- recently, one would have to visit the collection to get any CC. In addition, users can identify the individual speci- more specific information regarding the number of indi- mens that meet those criteria. In all cases, the results of viduals preserving specific elements, the number of a database search can be exported from the Human Ori- adults vs. juveniles, or whether specimens were damaged. gins Database as a text file and then opened in a spread- In an attempt to make these important data available, sheet program such as Excel. for each African ape specimen ADG conducted a system- Using the various searches, a researcher anywhere in atic inventory of all skeletal elements (down to individual the world with a computer with internet access can iden- teeth and bones) in which the presence or absence was tify specific samples from the P-CC that would be appro- recorded along with notes regarding any in vivo priate for a particular study. For example, if one were pathologies, presence of connective tissue on bones that interested in studying a sex-balanced sample of right tali may obscure some measurements, and post mortem dam- from adult P. t. troglodytes, the Human Origins Database age. Developmental stage was also recorded for some ele- could be used to generate a list of all specimens at the P- ments (as detailed further below). These data were then CC that could be included in such a study.

American Journal of Physical Anthropology 16 A.D. GORDON ET AL.

Fig. 3. Developmental markers organized by decreasing number of individual specimens that have initiated that marker (i.e., dental eruption or epiphyseal fusion has begun). Dotted lines track the number of individuals that have initiated each marker and solid lines track completion of each marker. Represented in gray is the difference between the number of individuals that have completed eruption of the maxillary third molar (generally the last tooth to fully erupt in both species in this data set) and the number of individuals that have completely fused the distal ulna (generally the last epiphysis to completely fuse in both species in this data set).

Developmental sequence and ontogenetic series Unfused (U). Epiphysis is distinct from diaphysis. There is no bony connection, but there may be other con- As an example of what can be achieved with the types nective tissue between the epiphysis and diaphysis. of data we have recorded on the P-CC, we present below Partially fused (P). Epiphysis and diaphysis are con- an analysis of the sequence of developmental markers in nected by some bone, but the growth plate is still P. troglodytes and G. gorilla based on information present. recorded in the Human Origins Database. We also pro- Fully fused (F). Epiphysis and diaphysis are continu- vide the resulting lists for both species of all specimens ous. A line may mark the previous site of the growth in the collection sorted by relative maturity along with plate. each specimen’s developmental data (Appendices A and The degree of fusion of the ilium, ischium, and pubis B). The annotated inventory of African ape skeletal was scored as follows: material in the P-CC includes data about (A) the degree Unfused (U). Ilium distinct from ischium and pubis. of fusion of sixteen long bone epiphyses, (B) the eruption Ischium and pubis may, or may not, be fused. status of all of the teeth, and (C) the general state of Partially fused (P). Ilium and ischiopubis are connected fusion of all of the main cranial sutures (Tables 4 and 5 by some bone, but fusion is not complete. Fully fused (F). The components of the pelvic bone are and see below). These annotated inventories plus the completely fused with no gaps between bones. growth data have been incorporated into the online database, both as searchable values in the relational database and as a series of data sheets provided for each Dental eruption. The deciduous teeth are referred to specimen in the collection. by their developmental origin rather than by their morphology (i.e., we refer to the deciduous second “molar” as the deciduous fourth premolar). This allows for a scoring Epiphyseal fusion. For all of the long bones in the system that refers to the eruption stage at a particular collection, epiphyseal fusion at each epiphysis was tooth position rather than the eruption stage of a speciﬁc scored as follows: tooth in the dental succession.

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 17 Deciduous unerupted (dU). Deciduous tooth is in the in the collection, which are primarily Central chimpan- crypt, or if partially emerged, the crown does not project zees (P. t. troglodytes) (Tables 4 and 5). above the alveolar margin. Regarding relative dental and postcranial maturity, Deciduous partially erupted (dP). Deciduous tooth c.17% of all chimpanzee specimens (24 of 145) and c.19% crown projects above the alveolar margin, but it is not in of all gorilla specimens (28 of 145) were dentally but not full occlusion with occluding dentition. Or, if the corre- postcranially mature (as approximated by the gray boxes sponding tooth in the other jaw is not fully erupted, the in Fig. 3) at the time of death. The extent of the problem crown does not project above the alveolar margin at the of assuming that dentally mature specimens are “adult” level of other fully erupted teeth. becomes evident when the specimens referred to above Deciduous erupted (D). Deciduous tooth crown projects are considered as a percentage of dentally mature speci- above the alveolar margin. mens. In the chimpanzee sample c.28% (24 of 86) of den- Unerupted (U) (for teeth without deciduous antecedent- tally mature specimens are not skeletally mature, and in s). Tooth is in crypt, or if partially emerged, the crown the gorilla sample c.38% (28 of 74) of dentally mature does not project above the alveolar margin. specimens are not skeletally mature. Only two skeletally Partially erupted (P). Tooth crown projects above alveo- mature gorillas had not completed eruption of their third lar margin, but it is not in full occlusion with occluding molars; all of the skeletally mature chimpanzees were dentition. Or, if the corresponding tooth in the other jaw also dentally mature. Thus, in both taxa approximately is not fully erupted, the crown does not project above the one-quarter to one-third of dentally mature individuals alveolar margin at the level of other fully erupted teeth. are unlikely to have achieved their full adult size. For Fully erupted (F). Tooth crown is in full occlusion, or if example, facial growth has been shown to continue past the corresponding tooth in the other jaw is not fully dental maturity at different sex-specific rates in great erupted, the crown projects above the alveolar margin at apes and other primates; thus, depending on the criteria the level of other fully erupted teeth. used to define adulthood, this will affect estimates of mean adult facial size and sexual dimorphism (Wang et al., 2007; Balolia et al., 2013). For some studies the Cranial fusion. Cranial sutures were evaluated as difference between dental and skeletal maturity may not follows: be a concern, whereas for others it might represent a Unfused (U). All cranial sutures are open. significant source of error due to additional variation in Partially fused (P). At least one cranial suture is par- the traits under consideration. tially fused. It is reasonable to assume that other skeletal collec- Fully fused (F). Complete fusion of the sutures of the tions have similar proportions of gorillas and chimpan- neurocranium and face (i.e., not including sutures of the zees that are dentally, but not skeletally, mature. basicranium). Therefore, we extend the caution that a quarter or more In order to assess the sequence of developmental events of African ape specimens in museum collections that are recorded in this data set and to assess the relative matu- identified as adult based on the eruption of the third rity of each specimen, left and right scores on all dental molar may not be fully skeletally mature, and that this and postcranial variables were reduced to a single score may be an important consideration for some investiga- for each individual. If the scores differed between anti- tions. Similar cautions may apply to collections of other meres, the more advanced developmental stage was cho- higher primate taxa, and we are exploring how this may sen (but both sides are reported independently in the relate to subtle and not-so-subtle differences in life his- online database). Because fusion of the cranial sutures tory within and among great ape species. occurs after dental and postcranial maturity in all of the In addition to identifying the sequence of developmen- crania inspected, cranial data were not included in this tal events, we also identified the relative maturity of analysis (although they are included in the Appendices A each chimpanzee and gorilla in the P-CC collection. For and B; see below). The number of specimens in the P-CC each complete specimen, we calculated a “maturity of known sex with complete information regarding dental score,” which was calculated as the proportion of the eruption and postcranial epiphyseal fusion is the same permanent dentition that is completely erupted and the for chimpanzees and gorillas (P. troglodytes N 5 145, G. proportion of the postcranium that is completely fused. gorilla N 5 145). For each epiphysis, counts were taken of See King (2004) for a similar approach applied to 16 the number of complete individuals in each species that non-human primate species. For each specimen, the A) had begun the process of fusion at that epiphysis and number of permanent teeth that are completely erupted B) were completely fused at that epiphysis. Likewise, for (on one side) were added to the number of epiphyses each tooth (deciduous and permanent), counts were taken that are completely fused (on one side), as well as 0.5 of the number of complete individuals in each species that A) had at least begun to erupt that tooth, and B) times the number of teeth that are partially erupted and had fully erupted that tooth. These counts were then 0.5 times the number of epiphyses that are partially sorted in descending order (Tables 4 and 5, Fig. 3). In fused. The resulting value was divided by the total num- general, most developmental events were completed in ber of developmental markers exclusive of cranial suture the same order in which they initiated; the exceptions are fusion (i.e., 32) to produce a score between zero and one. shown in bold in Tables 4 and 5. The sequence of dental In Appendices A and B of this article, we list all of the development identified here for P. troglodytes and G. complete chimpanzee and gorillas specimens sorted by gorilla is generally similar to the patterns reported by maturity score. In the case of specimens with no perma- earlier workers (e.g., Schultz, 1935; Dean and Wood, nent dentition, specimens are ordered by the number 1981; Bolter and Zihlman, 2011). There is little published and type of deciduous teeth present. Specimens in which information available on the combined relative timing of fusion has occurred at all sutures except those of the dental eruption and epiphyseal fusion for African apes basicranium (which are rarely fused in this collection) (but see Zihlman et al., 2007). We report these data for are always completely mature both postcranially and both Western gorillas (G. g. gorilla) and all chimpanzees dentally, thus they are placed later in the sequence than

American Journal of Physical Anthropology 18 A.D. GORDON ET AL. otherwise fully mature specimens for which cranial temporal effort to identify the study sample on site. fusion is not complete. We estimated the maturity score Developing databases such as the one we have for specimens that are damaged or incomplete by match- assembled for the Powell-Cotton Collection would be a ing them as closely as possible with complete specimens. great help to researchers. Individuals with fully fused crania that lack postcranial A second point relates to the ultimately ephemeral evidence are listed last because all of the complete speci- nature of the collections themselves. Higher primate mens in our sample with fused cranial sutures were skeletons are vulnerable to damage, and given enough postcranially mature at the time of death. wear and tear over time, they will eventually disinte- Thus, in addition to the adult skeletons, the P-CC pre- grate beyond the point of yielding useful morphological serves a remarkable ontogenetic series of Western goril- information. The Powell-Cotton Collection is a case in las and Central chimpanzees. For the gorillas (all of point. When BW compared observations he made in which appear to have been collected within the range of 1973 with those made three years ago, several speci- G. g. gorilla), among the specimens that preserve all mens are now missing teeth that were present in 1973, developmental markers there are 47 fully adult complete and dimensions that could be measured then cannot be or nearly complete skeletons, and 101 complete or nearly measured now. Although there will always be a need to complete skeletons in various stages of development. In maintain access to the original specimens for specialist addition, there are many skulls without associated post- examination, the original specimens could be afforded a crania, and some postcrania without associated skulls. substantial degree of protection if most of the informa- With respect to the latter, at least one of the “missing” tion needed by most of the people who presently exam- gorilla skulls is in the collection of the Hunterian ine them was made available in the form of an electronic Museum at the Royal College of Surgeons of England relational database, or at the very least if those original (postcranial specimen P-CC M21 is from the same indi- data were published in ways that identified data with vidual as the skull specimen RCSOM/A 64.21). For the individual specimens (e.g., through online supplemental chimpanzees (including three subspecies, but nearly all tables linked to journal articles, or through other perma- of which are P. t. troglodytes), among the specimens that nently maintained online data depositories such as preserve all developmental markers there are 62 fully Dryad [http://datadryad.org]). adult complete skeletons and 84 complete skeletons in It is widely recognized that fossils are irreplaceable, various stages of development, again in addition to sev- but this is also the case for museum collections of extant higher primates. Many of these collections include the eral skulls without associated postcrania. skeletons of animals collected in locations where wild higher primates have locally been driven to extinction. RECOMMENDATIONS FOR MOVING FORWARD These important comparative collections are irreplaceable, and there is an urgent need for the raw, non- The temporal and paleoecological contexts of the homi- aggregated data obtained from them to be placed in the nin fossil record are important for its interpretation, but public domain so that the collections can be preserved the primary data used to investigate human evolutionary for future generations. With that in mind, we encourage history are morphological. Morphological evidence is the morphologists to publish their raw data with their anal- currency of hominin paleobiology; it is the raw material yses whenever possible, and we encourage journal edi- for all analyses, be they taxonomic, phylogenetic, func- tors to accommodate such efforts whenever they can. tional, or developmental. These morphological data are So what specifically can be done to address these analogous to sequence data in molecular biology, or to points, and what has already been done? In April 2007, seismic records and temperature proxy data from piston a workshop designed to address these questions in paleo- cores in the earth sciences. Yet, any testable hypotheses anthropology in general was organized by Eric Delson, about taxonomy, evolutionary relationships, and functional Stephen Frost, Will Harcourt-Smith, and Christopher morphology are dependent on having substantial collec- Norris, and was co-funded by the Wenner-Gren Founda- tions of extant higher primates for comparison. Some tion and the National Science Foundation (a summary of museums provide helpful information available online the workshop can be found in the work by Delson et al., about their collections (e.g., the Smithsonian Mammal col- 2007). An international group of participants (including lections database [http://collections.nmnh.si.edu/search/ ADG and BW) brought together researchers involved in mammals/] often includes in a notes field a list of major database creation and museum representatives to initi- elements present for individual great ape specimens), but ate discussion on issues such as database standards, for most of the collections it is difficult for researchers to data standards, and museum involvement in these ini- assess how much of a seemingly large collection will be of tiatives. The workshop brought to light many different any use for their particular research purposes. perspectives on these issues, and here we outline specific The ability to deliver specimen- and element-specific proposals for great ape skeletal collections that follow information through a relational database that allows from our thoughts on those discussions. users to search through that information in a variety of Regarding the first point, over the past decade an ways has many implications for our field, and below we international standard has emerged for organismal biol- draw attention to two of them. One of the realities we ogy. The Darwin Core (http://rs.tdwg.org/dwc/) is a data- face as a discipline is that financial resources for com- base standard for biodiversity information that was parative morphological analyses are decreasing and are originally developed in the late 1990s and was last unlikely to increase in the foreseeable future. In such an updated and ratified in October 2009. It sets out specific economic climate (and indeed, even in a climate where database field names for data related to specimen identi- research money is plentiful), it is particularly useful to fication, taxonomic and biogeographic information, and be able to identify the specimens that meet study crite- many other types of metadata. The adoption of such a ria before arriving at a museum so that researcher’s standard by collections/research databases has allowed time and money can be put to maximum use, as opposed for the development of web portals, which are online to spending a significant portion of that financial and aggregators of information derived from simultaneous

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 19 queries of multiple online databases using a common set The measurement flexibility within the Darwin Core of user-specified search terms. Such searches work leads into a discussion of the second issue raised above: because the various databases either use the standard standards for the data collected (as opposed to database data fields specified by the Darwin Core, or they have structure). Although comparative primate morphologists fields that can be translated to match the standards. ostensibly share a common set of “standard” measure- Examples of web portals in disciplines related to paleo- ments (e.g., buccolingual molar width or femoral length), anthropology are the Global Biodiversity Information in practice two researchers may use slightly different Facility (GBIF, http://data.gbif.org) the Mammal Net- definitions of the “same” measurement, or may even col- worked Information System (MaNIS, http://manisne- lect the same measurement using the same definition in t.org/), and the Paleontology Portal (http:// two slightly different ways. This is why it is important www.paleoportal.org/). These websites and others like to spend time in a manuscript describing measurement them typically have additional content, but the portal techniques, and why one of us devoted 27 pages of the portion of the site allows the user to build a query that monograph on the Koobi Fora cranial remains to is sent to some or all of a predetermined set of databases describing the 394 measurements used (Wood, 1991). for which the portal has a communication protocol in The recognition of differences in the protocols used for place. For example, the Paleontology Portal can query “standard” measurements tends to elicit one of two differ- from among several private and university-affiliated nat- ent responses from researchers who would like to use a ural history museums as well as MorphoBank. combined dataset of standard measurements collected by Web portals provide the ability to query many collec- two or more people. The first is to conclude that the tions at once and thus have the potential to aggregate, uncertainty regarding measurement differences degrades for example, all information on world-wide holdings of P. data quality for combined datasets to the point that they t. schweinfurthii into the results of a single query. Indi- are unreliable and should not be used, while the second is vidual apes collected from the same time and place are to recognize the problem but suspect that inter-observer often distributed among several museums, and it would bias will have minimal effect (e.g., Bailey et al., 2004) on be useful to know where those specimens are. For exam- the results of a particular analysis and hope for the best. ple, the populations represented by the Merfield mate- We would like to suggest a middle way, in which com- rial in the P-CC are also distributed across Europe and parison of repeated measurements on a common set of the US through the sale of half of the specimens Mer- specimens by different researchers can be used to deter- field collected, and the Asiatic Primate Expedition mate- mine whether a combined dataset is problematic, or not, rial is split between the Museum of Comparative in a given situation. This suggestion relies on access to Zoology at Harvard and the Department of Anthropology researchers’ measurements for individual specimens, as at the University of Zurich. In addition, in some cases well as specific information regarding how measure- elements from single individuals may be split between ments were collected. As a discipline we lack a single different collections. As mentioned above, at least one of source of definitions for common measurements used by the skulls that are missing from some African ape skele- a majority of practitioners in the field. We are not pro- tons in the P-CC is in the Royal College of Surgeons in posing the prescription of a limited set of “correct” meas- London; it also appears that some limb elements missing urements that should be taken on skeletal material, but from bonobo skeletons at the Royal Museum of Central rather the identification and description of the multiple Africa in Tervuren, Belgium probably ended up in Bel- ways “standard” measurement techniques are inter- gian university collections during the partitioning of preted so that researchers can explicitly identify the those samples in the mid-20th century (Wim Wendelin, technique they used when sharing their measurements Personal Communication). Queries to a web portal such with others. We develop this suggestion further below. as the GBIF that aggregates all of the information from The final issue raised above is that of museum great ape collections would allow researchers to identify involvement, and it is a critically important issue. First, where the components of these various populations cur- there is a considerable investment of trained-personnel rently reside, with one large caveat: web portals are reli- time that must go into a skeletal element-level anno- ant on both the existence of databases containing the tated inventory such as the one ADG and BW undertook information desired and standard database fields from with the P-CC, as well as into the development of a which the information can be queried. Leaving aside for database, translation into database fields of information the moment the issue of the existence of a database for a from what are often fading or crumbling hand-written specific collection and focusing on database standards, field notes, and other associated activities associated the Darwin Core already has such standards for meta- with compiling a database along the lines of what we data such as subspecies name and collection locality for have described here. Typically this investment must individual specimens, but does not specify a set of stand- come from the museum or institution curating the collec- ard osteometric measurements. tion, and as mentioned above, many institutions are The Darwin Core retains a great deal of flexibility already limited in their ability to meet whatever they regarding measurements derived from specimens. With- consider to be core functions of the institution. A chal- out going into too much detail on database structure, lenge we must face as a discipline is how to meet these rather than defining fields with specific names such as goals without impinging on museum resources for those “right maxillary canine height” or “femoral head super- collections that cannot prioritize these efforts. This chal- oinferior diameter,” the Darwin Core specifies fields such lenge could be met by external research groups taking as “measurementID,” “measurementMethod,” on components of the work (as ADG and BW have done “measurementType,” and “measurementValue.” This in the case of the P-CC), by raising funds to support in- allows database developers to name and define their house efforts, or by other mechanisms. own sets of measurements that can either be queried Second, we must recognize that museums housing specifically by measurement name or returned as part of great ape skeletal material vary in their access policies, a set of all measurements for an individual specimen. data sharing policies, whether or not they charge bench

American Journal of Physical Anthropology 20 A.D. GORDON ET AL. fees, etc. Some museums will be comfortable making researchers avoid collections that do not have such their collections information available online to the gen- resources available online. What we would suggest is eral public for free, while others may prefer to limit that developing measurement standards, migrating data access to bona fide researchers and/or to individuals or to the web where feasible, and encouraging the open institutions that pay for access to the information. There sharing of data will improve the utility and increase the are many factors that go into each museum’s existing lifespan of these essential and irreplaceable resources. policies, including (but not limited to) their statutory remit, concerns regarding specimen preservation over the long term, funds required to maintain collections, ACKNOWLEDGMENTS and maintaining institutional control over collection We would like to thank the Trustees of the Powell-Cotton data. Developing databases such as we describe here Museum and Malcolm Harman for granting generous will ameliorate some of these concerns and aggravate access to the African ape specimens and records under their others, with the consequence that it may not be clear to care, and for their warm reception to the efforts to highlight museum curators and administrators whether making the exceptional nature of the Powell-Cotton Collection. In such information available would be a net gain from particular, Malcolm Harman has been very helpful over the their perspective. Thus, it is imperative that museum years in acquainting us with the history of the collection representatives be party to discussions and decisions and its connection to collections at other institutions. We made regarding establishing data sharing standards for thank David Pilbeam for arranging library access for Emily data derived from great ape skeletal collections. This Marcus. Madison Evans researched the previous names of brings us to our closing suggestion. countries in which African great apes are found. We would also like to thank Trudy Turner and Chris Vinyard for their A congress to identify measurement and editorial guidance and the anonymous reviewers whose collections information standards comments have greatly improved the content and structure of this manuscript. The Human Origins Database and our Over the past few years there have been several work- work on the Powell-Cotton Collection was made possible shops and symposia at annual meetings that have through a grant to BW from the G. Harold & Leila Y. Math- brought together representatives from various research ers Foundation. ADG was partially supported in the writing groups to talk about the databases they have developed of this article by a Wenner-Gren Hunt Postdoctoral Fellow- for their various projects. We applaud these initiatives ship. Finally, ADG would like to extend a special thank you to get research database developers talking to each other to his good friends Wendy and Rob Teale, whose hospitality about common issues, but we also think that it is time and charm is well known to all who have stayed with them to try something different and more specific. over the years while working at the Powell-Cotton Museum. We propose that the time has come to bring together comparative morphologists working on great ape skeletal LITERATURE CITED material (and some who work on primates in general), regardless of whether they are involved in database proj- [Anonymous]. 1899a. (Notes). Nature 60:621–625. ects, and the curators who oversee those collections and [Anonymous]. 1899b. (Scientific notes and news). Science 10: who make policy about great ape collections. The pur- 741–743. pose of the meeting would be two-fold: first, to identify [Anonymous]. 1903a. (Natural history notes). Nature 68:92. and define a set of standard measurements typically col- [Anonymous]. 1903b. (Notes). Nature 68:458–461. lected on skeletal material, along with a set of variants [Anonymous]. 1903c. (Notes). Nature 67:302–306. with different definitions for each measurement, and [Anonymous]. 1903d. (Scientific notes and news). Science 18: 636–640. second, to identify specific concerns and limitations that [Anonymous]. 1903e. (Scientific notes and news). Science 18:93–95. those responsible for the collections have regarding mak- [Anonymous]. 1903f. (Scientific notes and news). Science 17: ing such information available on a searchable online 1019–1024. database. Such a meeting could be broken into working [Anonymous]. 1904a. (Notes). Nature 70:630–634. groups focusing on specific types of measurements (e.g., [Anonymous]. 1904b. (Scientific notes and news). Science 20: craniofacial, dental, axial, long bones, manual, and pedal 693–695. elements) and specific collection concerns (e.g., intellec- [Anonymous]. 1905. (Scientific notes and news). Science 21:638–639. tual property issues, funding concerns, impact on [Anonymous]. 1906. (Notes). Nature 74:578. research visit rates). While ambitious in scope, we [Anonymous]. 1907a. (Notes). Nature 75:443–447. believe that such a meeting would be an important step [Anonymous]. 1907b. (Notes). Nature 75:374–378. [Anonymous]. 1907c. The Royal Society conversazione. Nature that would A) greatly improve the utility of information 76:57–61. collected over the past several decades by the various [Anonymous]. 1907d. (Scientific notes and news). Science 26: practitioners in our field, B) help preserve the collections 921–928. and ensure their utility in the years to come, and C) pro- [Anonymous]. 1909. (Scientific notes and news). Science 29:808–812. vide a template for applying the same principles to col- [Anonymous]. 1928. (News and views). Nature 121:720–725. lections of fossil material. [Anonymous]. 1934. Gifts to the British Natural History In closing, we recognize that because of the vagaries Museum. Science 79:49. of the data collection protocols used in the field some- [Anonymous]. 1936a. (Recent acquisitions at the Natural His- times a century or more ago, for some collections not all tory Museum). Nature 137:182–183. of the types of data we would wish to have will be avail- [Anonymous]. 1936b. (Recent acquisitions at the Natural His- tory Museum). Nature 137:736. able in any form, digital or otherwise. We also recognize [Anonymous]. 1937. The Asiatic primate expedition. Science 85: that for a variety of reasons, some of which we have 11–12. touched on above, for some collections it may be a long [Anonymous]. 1938. British Museum (Natural History): acquisi- time before those data that do exist become available tions. Nature 142:204. online, and we certainly would not suggest that [Anonymous]. 1940. (Correspondence). Nature 146:223–224.

American Journal of Physical Anthropology GREAT APE SKELETAL COLLECTIONS 21

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American Journal of Physical Anthropology APPENDIX A.

Specimen Sex Maturity score m1 M1 i1 m2 i2 I1 M2 I2 P3 P4 p4 p3 C c m3 M3 HD HE OC UP FeLT FeGT FeH RP FiD TD TP FiP FeD HP RD UD Cranium CAM I 202 U 0.000 U U dU U dU dU U dU dU dU dU dU dU dU U U U U U U U U U U U U U U U U U U U M 465 F 0.000 U U dU U dU D U D dU dU dU dU dU dU U U U U U U U U U U U U U U U U U U U M 781 F 0.000 UUDUDDUDDDDDdPdPUU U M 475 series 2 F 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 475 series 1 M 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U babe M 556 F 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U CAM II 314 M 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 152 M 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 173 M 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 235 F 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 276 F 0.000 U U D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 888 M 0.016 UUDPDDUDDDDDDdPUU U N1 F? 0.016 UPDUDDUDDDDDDDUU U RA P KLTLCOLLECTIONS SKELETAL APE GREAT FC 70 F 0.031 P PDUDDUDDDDDDDUU U M 777 M 0.031 P P D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U CONGO 371 M 0.063 F FDUDDUDDDDDDDUU U M03 F 0.063 F FDUDDUDDDDDDDUU U M 133 F 0.063 F FDUDDUDDDDDDDUU U M 182 M 0.063 F FDUDDUDDDDDDDUU U M 300 F 0.063 F FDUDDUDDDDDDDUU U M 403 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 657 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 665 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 674 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 754 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 930 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U CAM I 222 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 250 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 369 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U

mrcnJunlo hsclAnthropology Physical of Journal American M 397 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 451 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 60 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 744 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 911 F 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 93 M 0.063 F F D U D D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 876 M 0.078 F F D U D D U D D D D D P D U U U U U U U U U U U U U U U U U U U CONGO 214 F 0.078 F F D U D D P D D D D D D D U U U U U U U U U U U U U U U U U U U M 27 series 2 U 0.094 F FDPDDPDDDDDDDUU U CAM I 93 F 0.094 F F P U P D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 507 F 0.109 F F F U P D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 675 M 0.109 F F P U P D U P D D D D D D U U U U U U U U U U U U U U U U U U U M 358 M 0.109 F F P P P D U D D D D D D D U U U U U U U U U U U U U U U U U U U M 259 M 0.109 F F D P D P P D D D D D D D U U U U U U U U U U U U U U U U U U U M 453 M 0.125 F F F F U D U D D D D D D D U U U U U U U U U U U U U U U U U U U CONGO 195 M 0.125 F F D P D P U U D D P P D D U U U U U U U U U U U U U U U U U U U M 363 M 0.125 F F P P D P P D D D D D D D U U U U U U U U U U U U U U U U U U U M 94 M 0.141 F F F P P D P D D D D D D D U U U U U U U U U U U U U U U U U U U M 881 F 0.172 F F F P P P U F D D D D D D U U U U U U U U U U U U U U U U U U U CAM I 118 M 0.203 F F F P P F P F D D D D D D U U U U U U U U U U U U U U U U U U U 23 mrcnJunlo hsclAnthropology Physical of Journal American 24 APPENDIX A. Continued

Specimen Sex Maturity score m1 M1 i1 m2 i2 I1 M2 I2 P3 P4 p4 p3 C c m3 M3 HD HE OC UP FeLT FeGT FeH RP FiD TD TP FiP FeD HP RD UD Cranium M 746 M 0.234 F F F P F F P F D D P D D D U U U U U U U U U U U U U U U U U U U M 801 M 0.234 F F F F F F U F D D D U D D U U P U U U U U U U U U U U U U U U U M 145 F 0.234 F F F F F F P F D D D D D D U U U U U U U U U U U U U U U U U U U M 635 M 0.250 F F F F P F F P D F D D D D U U U U U U U U U U U U U U U U U U U M 891 F 0.266 FFFFPFFPPFDDDDUU U M 274 F 0.281 F F F F P F P P P P F P D D U U U U U U U U U U U U U U U U U U U M 454 F 0.281 F F F F F F F F F U D D D D U U U U U U U U U U U U U U U U U U U M 805 F 0.297 F F F P F F P F F F D D D D U U P U U U U U U U U U U U U U U U U M 382 F 0.313 F F F F F F P F F U P F D D U U U U U U U U U U U U U U U U U U U M 636 M 0.313 F F F F F F F F F F D U D D U U U U U U U U U U U U U U U U U U U M 170 M 0.344 F F F P F F P F F F F F D D U U U U U U U U U U U U U U U U U U U M 573 F 0.359 FFFFFFFFPFFFDDUU U M 455 M 0.375 F F F F F F F F F F F F D D U U U U U U U U U U U U U U U U U U U M 371 F 0.391 F F F F F F F F F F F F P D U U U U U U U U U U U U U U U U U U U M 52 M 0.406 F F F F F F F F F F F F P P U U U U U U U U U U U U U U U U U U U ZVI 35 M 0.438 F F F F F F F F F F F F F F U U U U U U U U U U U U U U U U U U U M 707 M 0.438 F F F F F F F F F F F F P P U U P P U U U U U U U U U U U U U U U M 719 M 0.484 F F F F F F F F F F F F P P P U P P P P U U U U U U U U U U U U U

CAM I 217 F 0.484 F F F F F F F F F F F F F F U U F U P U U U U U U U U U U U U U U AL. ET GORDON A.D. CONGO 372 M 0.516 F F F F F F F F F F F F P P P U P P P P P P U U U U U U U U U U U M 886 F 0.516 FFFFFFFFFFFFFFPU U CONGO 370 F 0.516 F F F F F F F F F F F F F F P U P P P U P U U U U U U U U U U U U M 278 M 0.531 F F F F F F F F F F F F F F F F P U P U U U U U U U U U U U U U U M 181 F 0.547 F F F F F F F F F F F F F F F F P P U P U U U U U U U U U U U U U M 347 M 0.578 F F F F F F F F F F F F F F F F F P P P U U U U U U U U U U U U P M 449 F 0.641 F F F F F F F F F F F F F F U U F F F F F P P U P U U U U U U U U M 702 F 0.688 F F F F F F F F F F F F F F F F F F P F F P P P U U U U U U U U U M 275 F 0.688 F F F F F F F F F F F F F F P P F F F F F F P P U U U U U U U U U CAM II 15 F 0.703 FFFFFFFFFFFFFFFP U M 506 series 3 F 0.703 F F F F F F F F F F F F F F F F F F F F P P P P U U P U U U U U U ZVII 26 F 0.781 F F F F F F F F F F F F F F F F F F F F F F F F P P U U U U U U U M 144 M 0.797 F F F F F F F F F F F F F F F F F F F F F F F P P P P P U U U U U CAM I 205 F 0.813 F F F F F F F F F F F F F F F F F F F F F F F F P P P U P U U U U ZIX 51 F 0.828 F F F F F F F F F F F F F F F F F F F F F F F F P P P P P U U U P CAM I 216 F 0.859 F F F F F F F F F F F F F F F F F F F F F F F F P P P P P P P U P M 984 M 0.875 F F F F F F F F F F F F F F F F F F F F F F F F P P P P P P P P P M 655 F 0.891 F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P P U U ZV 83 F 0.906 F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P P P U M 664 F 0.906 F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P P P P M 967 F 0.906 F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P P P P M 743 F 0.922 F F F F F F F F F F F F F F F F F F F F F F F F F F P F P P P P U M 873 F 0.922 F F F F F F F F F F F F F F F F F F F F F F F F F F P F P P P P U M 401 M 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F P P F P F F U M 105 F 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F P F F F P P P ZVI 34 M 0.969 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P F U CAM I 203 F 0.969 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P U M 450 F 0.969 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P M 184 F 0.984 F F F F F F F F F F F F F F F F F F F F F F F F F F P F F F F F P M 676 F 0.984 F F F F F F F F F F F F F F F F F F F F F F F F F F F F P F F F P CAM I 204 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F U APPENDIX A. Continued

Specimen Sex Maturity score m1 M1 i1 m2 i2 I1 M2 I2 P3 P4 p4 p3 C c m3 M3 HD HE OC UP FeLT FeGT FeH RP FiD TD TP FiP FeD HP RD UD Cranium CAM I 228 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F U M 254 series 3 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F U M 475 series 1 ADULT F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F U M 800 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F U M 169 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P ZIX 49 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 207 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 219 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P FC 100 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 158 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 254 series 2 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 277 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 299 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 352 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 474 series 2 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P COLLECTIONS SKELETAL APE GREAT M 576 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 742 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 789 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 78 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 986 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 988 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P ZVII 23 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P ZVII 24 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P ZVII 25 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P ZVIII 10 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 279 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 424 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 504 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 670 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 273 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P

mrcnJunlo hsclAnthropology Physical of Journal American FC 116 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 13 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM I 147 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM I 206 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM I 218 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM II 301 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM II 62 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 148 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 155 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 171 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 172 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 186 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 234 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 249 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 272 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 348 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 425 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 440 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 467 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 498 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F 25 mrcnJunlo hsclAnthropology Physical of Journal American 26

APPENDIX A. Continued

Specimen Sex Maturity score m1 M1 i1 m2 i2 I1 M2 I2 P3 P4 p4 p3 C c m3 M3 HD HE OC UP FeLT FeGT FeH RP FiD TD TP FiP FeD HP RD UD Cranium M 501 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 506 series 2 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 650 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 677 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 706 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 712 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 720 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 724 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 745 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F ZIX 52 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M86 F 1.000 FFFFFFFFFFFFFF F F F FF F F F F F FFF F FF F F

CAM 236 M F F F F F F F F F F F F F F F F U AL. ET GORDON A.D. M134 F FFFFFFFFFFFFFFFF U M574 F FFFFFFFFFFFFFFFF U CAM 200 M F F F F F F F F P CAM II 326 F F F F F F F P M803 F FFFFFFFFFFFFFFFF P M792 F FFFFFFFFFFFFFFFF P CAM 11 F F F F F F F F F F CAM 26 F? F F F F F F F F F CAM 74 M F F F F F F F F F CAM II 9 F F F F F F F F F F M02 F F F F F F F F F F CAM 199 M F F F F F F F F F F F F F F F F F CAM 237 F F F F F F F F F F F F F F F F F F CAM 238 F F F F F F F F F F F F F F F F F F CAM M.0. M F F F F F F F F F F F F F F F F F PCMM01 F FFFFFFFFFFFFFFFF F PCM M 491 F F F F F F F F F F F F F F F F F F M814 F FFFFFFFFFFFFFFFF F Z VIII 0.2 M F F F F F F F F F F F F F F F F F M165 F FFFFF F FFF FFF F FC 181 U F F F F F F F F M04 F FFFFFFFFFFFFFFFF F CONGO 259 M FFFFFFFFFFFFFFFF CAM 201 U F P U U CAM 67 U CAMEROONS 1929 U U GREAT APE SKELETAL COLLECTIONS 27

APPENDIX B postcrania are listed after all other specimens because Developmental information for Gorilla gorilla gorilla long bone epiphyses may remain unfused or partially specimens at the Powell-Cotton Museum. Specimens are fused after full dental eruption. However, postcranial listed in increasing developmental age as determined by fusion precedes cranial suture fusion in all cases for degree of dental eruption and epiphyseal fusion. Devel- which complete skeletons are available, so completely opmental marker abbreviations (column headings) follow fused crania presumably belong to individuals that were Table 5 and are ordered within the dentition and post- also postcranially mature. Finally, three isolated mandi- cranium by their developmental sequence as shown in bles are listed at the very end of the list of specimens. A Table 5; developmental data abbreviations as given in color version is available at the Human Origins Data- the text. Calculation of maturity score is described in base, www.humanoriginsdatabase.org the text. Skulls with fully erupted dentition but without

American Journal of Physical Anthropology mrcnJunlo hsclAnthropology Physical of Journal American APPENDIX B. 28

Specimen Sex Maturity score m1 M1 i1 I1 i2 I2 M2 m2 P3 P4 p4 p3 c C m3 M3 HD OC UP HE FeLT FeH FeGT RP TD FD FeD TP FiP RD HP UD Cranium M 780 M ? U U U U U U U U U U U U U U U U M No number - M? ? UUUU U U U UUU U UUUUU babe M 476 M 0.000 U U D D D D U U dP dP dU dP dU dP U U U U U U U U U U U U U U U U U U U M 756 M 0.000 U U D D D D U U D dU dP D dU dU U U U U U U U U U U U U U U U U U U U M 887 F 0.000 U U D D D D U U D dU dP D dP dU U U U U U U U U U U U U U U U U U U U FC 129 M 0.000 U U D D D D U U D dP dP D dP dP U U U U U U U U U U U U U U U U U U U CAM I 43 F 0.000 U U D D D D U U D dP dP D dP dP U U U U U U U U U U U U U U U U U U U M 32 series 2 M 0.000 U U D D D D U U D D D D D D U U U CAM II 331 M 0.000 U U D D D D U U D D D D D D U U U M 631 M 0.000 U U D D D D U U D D D D D D U U U M 532 F 0.000 U U D D D D U U D D D D D D U U U CAM I 224 M 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 117 F 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 497 M 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 690 series 1 M 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 141 M 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 409 F 0.000 U U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U

M 99 M 0.016 P U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U AL. ET GORDON A.D. M 333 F 0.016 U P D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 471 M 0.031 F U D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U FC 124 F 0.031 P P D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 880 F 0.031 P P D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 29 F 0.031 P P D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 137 M 0.031 U U D D D D U U D D D D D D U U U F U U U U U U U U U U U U U U U M 000 (M) M 0.047 F PDDDDUUDDDDDDUU U FC 122 M 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 34 M 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 625 F 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 828 F 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 868 F 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 98 F 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 169 M 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 760 F 0.063 F F D D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 487 M 0.063 F FDDDDUUDDDDDDUU U M 000 series 1 F 0.063 F FDDDDUUDDDDDDUU U M 319 F 0.063 F FDDDDUUDDDDDDUU U M33 M 0.063 F FDDDDUUDDDDDDUU U M 000 series 2 F 0.063 F FDDDDUUDDDDDDUU U M 698 M 0.078 F F P D D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 119 series 3 M 0.078 F F P D D D U U D D D D D D U U U U U U U U U U U U U U U U U U P M 463 M 0.078 F F D D D D U U P D D D D D U U U U U U U U U U U U U U U U U U U M 855 F 0.078 F F D P D D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 450 M 0.125 F F F D F D U U D D D D D D U U U U U U U U U U U U U U U U U U U M 847 series 2 M 0.141 F F P P D P U U D D D D D D U U U F U U U U U U U U U U U U U U U CAM I 110 F 0.141 F F F P D P U U D D D D D D U U P U U U U U U U U U U U U U U U U FC 151 F 0.156 F F F P P P P U D D D D D D U U U U U U U U U U U U U U U U U U U FC 147 F 0.156 F F F P P P P U D D D D D D U U U U U U U U U U U U U U U U U U U CAM I 109 F 0.172 F F F P F P U P D D D D D D U U U U U U U U U U U U U U U U U U U CAM I 111 F 0.203 F F F F F P P P D D D D D D U U U U U U U U U U U U U U U U U U U APPENDIX B. Continued

Specimen Sex Maturity score m1 M1 i1 I1 i2 I2 M2 m2 P3 P4 p4 p3 c C m3 M3 HD OC UP HE FeLT FeH FeGT RP TD FD FeD TP FiP RD HP UD Cranium M 667 F 0.210 FFFFF PFDDDDDDUUUUUU U U U UUU U UUUUU P M 841 F 0.234 F F F F F F P F D D D D D D U U U U U U U U U U U U U U U U U U U M 865 F 0.234 F F F F F F P F D D D D D D U U U U U U U U U U U U U U U U U U U M 160 F 0.250 F F F F F F P P D D D D D D U U U U F U U U U U U U U U U U U U U M 32 series 1 F 0.250 F FFFFFF FDDDDDDUU U M 875 series 2 F 0.266 F F F F F P F F D P P D D D U U U U U U U U U U U U U U U U U U U M 691 F 0.266 F F F F F F F F D D D D D D U U P U U U U U U U U U U U U U U U U M 674 M 0.281 F F F F F F F F D D P P D D U U U U U U U U U U U U U U U U U U U M 22 M 0.281 F F F F F F F F P P D D D D U U U U U U U U U U U U U U U U U U U CAM I 108 M 0.297 F F F F F F F F P P P D D D U U U U U U U U U U U U U U U U U U U M 985 F 0.313 F F F F F F P F F F P D D D U U U U U U U U U U U U U U U U U U P M 180 F 0.313 F F F F F F F F F P P D D D U U U U U U U U U U U U U U U U U U U CAM II 315 F 0.313 F F DFDF F F F F F F D D U U P M 689 M 0.344 F F F F F F F F F F F D D D U U U U U U U U U U U U U U U U U U U M 387 F 0.375 F F F F F F F F F F F F D D U U U U U U U U U U U U U U U U U U U COLLECTIONS SKELETAL APE GREAT M II 1 F 0.406 F F F F F P P F P P D D D D U U F F P P P P P P U U U U U U U U P CAM I 14 F 0.406 F F F F F F P F F P F F P D U U P U P U P U U U U U U U U U U U U M 109 M 0.453 F F F F F F F F F F F F P P P P P U U U U U U U U U U U U U U U U CAM I 107 M 0.469 F F F F F F F F F F F F D P P P P P P U U U U U U U U U U U U U P FC 114 F 0.469 F F F F F F F F F F F F P P P P P P U U U U U U U U U U U U U U U M 862 M 0.469 F F F F F F F F F F F F P P P P P P U U U U U U U U U U U U U U P M 791 F 0.469 F FFFFFF FFFFFPPP P P M03 F 0.484 FFFFFFFFFFFFDFPF P M 462 M 0.484 F F F F F F F F F F P F P P F F P U U P U U U U U U U U U U U U P M 717 M 0.500 F F F F F F F F F F F F P P F F P P U U U U U U U U U U U U U U U M 505 M 0.500 F F F F F F F F F F F F P P P P P P F U U U U U U U U U U U U U P M 464 M 0.516 F F F F F F F F F F F F P P F P P P U P P U U U U U U U U U U U P M 342 M 0.547 F F F F F F F F F F F F F F F F P U P P U U U U U U U U U U U U P M 241 M 0.563 F F F F F F F F F F F F P P F F P U P U P P P U U U P U U U U U P FC 115 M 0.578 F F F F F F F F F F F F F F F F P P P P U P U U U U U U U U U U U

mrcnJunlo hsclAnthropology Physical of Journal American M 206 M 0.578 F F F F F F F F F F F F F F F F F P P P U U U U U U U U U U U U P M 204 M 0.594 F F F F F F F F F F F F F F F F P P P P P P U U U U U U U U U U P CAM I 208 M 0.641 F F F F F F F F F F F F F F F F F P P F P P P U U U U U U U U U P M 460 F 0.641 F F F F F F F F F F F F F F F F F P P F P P P U U U U U U U U U P M 878 F 0.641 F F F F F F F F F F F F F F P P F P F F F P P U U U U U U U U U P CAM I 97 F 0.641 F FFFFFF FFFFFPDUU P Z V 91 F 0.656 F FFFF F FFFFFFFP P P CAM I 198 F 0.672 F FFFFFF FFPFFFFUU P M 57 F 0.672 F F F F F F F F F F F F F F P P F F F F F P P P U U U U U U U U P FC 133 M 0.672 F F F F F F F F F F F F F F F F F P F F F P P U U U U U U U U U P CAM I 41 M 0.688 F F F F F F F F F F F F F P F F F F F F P P P P P U U U U U U U P Z II 63 F 0.688 F F F F F F F F F F F F F F F P F F F F F P P P U U U U U U U U P M 847 series 1 F 0.703 F F F F F F F F F F F F F F P P F P F F F F F P P U U U U U U U P M 857 F 0.734 F F F F F F F F F F F F P P U U F F F F F F F F P P P P P U U U P Z III 31 M 0.766 F F F F F F F F F F F F F F F F F F F F F F F P U P U P U U U U P M 854 M 0.781 F F F F F F F F F F F F F P F P F F F F F F F P P P P P P U U U P CAM I 44 F 0.781 F F F F F F F F F F F F F F F F F F F F F F F F U P U U P U U U P Z VI 30 M 0.781 F F F F F F F F F F F F F F F F F F F F F F F F P P U U U U U U P M 20 M 0.797 F F F F F F F F F F F F F F F F F F F F P P P F U U U U U F F F P CAM I 231 M 0.797 F FFFFFF FFFFFFFF F F F F F FP P P P UUU P 29 mrcnJunlo hsclAnthropology Physical of Journal American 30 APPENDIX B. Continued

Specimen Sex Maturity score m1 M1 i1 I1 i2 I2 M2 m2 P3 P4 p4 p3 c C m3 M3 HD OC UP HE FeLT FeH FeGT RP TD FD FeD TP FiP RD HP UD Cranium CAM I 48 M 0.813 F F F F F F F F F F F F F F F F F F F F F F F F P P U P P U U U P M 688 F 0.813 F F F F F F F F F F F F F F F F F F F F F F F F P P P P U U U U P M 119 series 1 M 0.813 F F F F F F F F F F F F F F F F F F F F F F F F F P P U U U U U P M21 M 0.828 PFPP F F F UFFFFFUUU M 729 series 2 F 0.828 F F F F F F F F F F F F F F F F F F F F F F F F P P P P P U U U P MII23 U 0.875 FFFF F F F PFPPP PP FC 112 M 0.906 F F F F F F F F F F F F P F P P P M 696 F 0.906 F F F F F F F F F F F F P F F F F F F F F F F F F F F P P P P P P M 35 F 0.922 F F F F F F F F F F F F F F P P F F F F F F F F F P F P P F F F P M 470 F 0.922 F F F F F F F F F F F F F F F F F F F F F F F P F F F P F P P P P Z I 30 M 0.938 FFFF F F F FF FF PPP M 136 F 0.938 F F F F F F F F F F F F F F F F F F F F F F F F F F F P F P P P P CAM I 47 M 0.938 F F F F F F F F F F F F F F F F F F F F F F F F F F P F F P P P P CAM I 105 M 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P CAM I 149 F 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P CAM I 230 M 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P CAM I 42 F 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P FC 146 F 0.953 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P P

CAM I 150 F 0.969 F F F F F F F F F F F F F F P P F F F F F F F F F F F F F F F F P AL. ET GORDON A.D. M 798 F 0.969 F F F F F F F F F F F F F F P P F F F F F F F F F F F F F F F F P M 174 M 0.969 F F F F F F F F F F F F F F F F F F F F F F F F F F F P F F F P P M 716 F 0.969 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P P M 856 F 0.984 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P Z II 64 M 0.984 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P P M 799 F 0.984 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P F P M 183 M 1.000 FFFF F F F FFFFFFFFF CAM I 95 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 98 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 372 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 490 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 690 series 2 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 96 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P Z VI 33 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P Z VI 32 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 134 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 96 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 139 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 58 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 687 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 755 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 835 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 840 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 877 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 879 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 89 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 902 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M 95 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M I 29 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P Z II 65 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 99 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P APPENDIX B. Continued

Specimen Sex Maturity score m1 M1 i1 I1 i2 I2 M2 m2 P3 P4 p4 p3 c C m3 M3 HD OC UP HE FeLT FeH FeGT RP TD FD FeD TP FiP RD HP UD Cranium M 461 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P M II 2 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F P CAM I 229 M 1.000 F FFFFFF FFFFFFFF F F F F F F F F FF F F F F F F CAM I 106 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F CAM I 139 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F FC 123 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F FC 130 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 135 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 138 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 150 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 264 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 300 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 329 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 340 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 720 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F COLLECTIONS SKELETAL APE GREAT M 729 series 1 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 766 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 786 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 875 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 932 F 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M 962 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F M I 28 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F Z I 17 M 1.000 F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F FC 215 F F F F F F F F F F F F F F F F F P M11 F F FFFFFF FFFFFFFF F P FC 206 M F F F F F F F F P FC 207 M F F F F F F F F P M04 F F F F F F F F F P FC 225 M ? ? ? ? ? ? ? ? P FC 216 M F F F F F F F F F F F F F F F ? P mrcnJunlo hsclAnthropology Physical of Journal American FC 195 M F F F F F F F F F F F F F F F F P FC 196 M F F F F F F F F F F F F F F F F P M 177 F F F F F F F F F F F F F F F F F P M 281 F F F F F F F F F F F F F F F F F P M 283 M F F F F F F F F F F F F F F F F P M 502 M F F F F F F F F F F F F F F F F P MII3 M F FFFFFF FFFFFFFF F P CAM II 282 F F F F F F F F F P FC 224 F F F F F F F F F F F F F F F F F P FC 113 M F F F F F F F F P FC111 M F FFFFFF FFFFFFFF F P M01 M F FFFFFF FFFFFFFF F P M02 M F FFFFFF FFFFFFFF F P M211 M F FFFFFF FFFFFFFF F P M 256 F F F F F F F F F F F F F F F F F P M 474 M F F F F F F F F F F F F F F F F P CAM I 128 M F F F F F F F F P FC 197 F F F F F F F F F P FC 208 F F F F F F F F F P 31 mrcnJunlo hsclAnthropology Physical of Journal American 32

APPENDIX B. Continued

Specimen Sex Maturity score m1 M1 i1 I1 i2 I2 M2 m2 P3 P4 p4 p3 c C m3 M3 HD OC UP HE FeLT FeH FeGT RP TD FD FeD TP FiP RD HP UD Cranium FC 219 F F F F F F F F F P M 349 F F F F F F F F F P FC 120 F F F F F F F F F F F F F F F F F P M2 F F FFFFFF FFFFFFFF F P

CAM II 323 M F F F F F F F F F F F F F F F F F AL. ET GORDON A.D. FC 154 F F F F F F F F F F F F F F F F F F FC 159 M F F F F F F F F F F F F F F F F F FC 162 M F F F F F F F F F F F F F F F F F FC 163 M F F F F F F F F F F F F F F F F F M10 M F FFFFFF FFFFFFFF F F M 212 M F F F F F F F F F F F F F F F F F M 288 M F F F F F F F F F F F F F F F F F M31 M F FFFFFF FFFFFFFF F F M 503 M F F F F F F F F F F F F F F F F F M 801 F F F F F F F F F F F F F F F F F F M 808 F F F F F F F F F F F F F F F F F F CAM I 83 M F F F F F F F F F CAM II 278 M F F F F F F F F F FC 217 F F F F F F F F F F FC 218 F F F F F F F F F F FC 226 F F F F F F F F F F M No number (F) F F F F F F F F P M 320 M F F F F F F F F M No number (M) M F F F F F F F F