AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 57:2–32 (2013) Great Ape Skeletal Collections: Making the Most of Scarce and Irreplaceable Resources in the Digital Age 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 con- tent 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 eval- uation 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 mor- phological 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,