CA✩ Forum on Theory in Anthropology The Animal Connection and Human Evolution

A suite of unique physical and behavioral characteristics distinguishes Homo sapiens from other mammals. Three diagnostic human behaviors played key roles in human evolution: tool making, symbolic behavior and language, and the domestication of plants and animals. I focus here on a previously unrecognized fourth behavior, which I call the animal connection, that characterized the human lineage over the past 2.6 million years. I propose that the animal connection is the underlying link among the other key human behaviors and that it substantially influenced the evolution of humans. The evolution of any species can be understood by first identifying unique or diagnostic characteristics of the modern species and then tracing their development through time. Traits that are ancient and continuous and that provide causal explanations for significant attributes of the species in question are likely to have played crucial roles during evolution. Traits often considered diagnostic of humans and significant in their evolution are (1) making and using tools (Darwin 1871; The animal connection comprises an increasingly intimate and reciprocal set of interactions between animals and humans (i.e., members of the genus Homo) starting ∼2.6 million years ago (mya). The animal connection began with the exploitation and observation of animals by humans. Over time, regular social interactions were incorporated into the animal connection. This trait is expressed today in the widespread Pat Shipman is Professor of Biological Anthropology in the Department of Anthropology of Pennsylvania State University (University Park, Pennsylvania 16802, U.S.A. [[email protected]]). This paper was submitted 3 X 08 and accepted 23 VII 09. adoption, or cross-species alloparenting, of animals-including dingoes, possums, bandicoots, raccoons, deer, moose, bison, fruit bats, lizards, bears, tapir, monkeys, sloths, coatimundis, antelopes, zebra, tree kangaroos, rabbits, weasels, ferrets, rodents, and birds, cervids, felids, and canids of all types (Serpell 1989)-as members of the family. This adoption involves active nurturing and care; ethnographic studies and early travelers' accounts often report women breast-feeding young animals or prechewing food for them (e.g., Galton 1865; Tool Making and Tool Using Tool making and tool using are much more complex behaviors in humans than in other animals. Humans make a vast array of tools and have made tools for ∼2.6 million years (m.yr.; Early humans used stone tools repeatedly for carcass processing. Direct evidence of this function comes from cut marks, scraping marks, and percussion scars on bones from many different sites (e.g., Blumenschine 1995; Tool using in the broad sense is an extrasomatic adaptation (White 1959) of humans: a means by which humans evolved behaviorally without adapting physically. Tool use is a predominant, universal, and daily aspect of present human life, implying that the adaptive advantage of tools has been greater to humans than to apes. Among nonhuman mammals, tool use is less complex and less frequent. Modern chimpanzees are the most adept animal tool makers and produce the largest tool kit. Chimpanzees make and use nests; termiting or anting implements; leaf sponges; hammerstones and anvils for breaking nuts; and sticks to pick marrow from bones, to dig for underground storage organs (e.g., tubers), and to locate and extract bush babies from tree hollows (e.g., Captive but not wild bonobos make flaked stone implements The importance of tool use in chimpanzees varies. Chimpanzees at Mahale spend little time ant fishing (∼1%-2% of the daylight hours) and gain little food (0.6 g protein; Nishida and Haraiwa 1982). However, Taï chimps spend hours cracking nuts and gain as much as 3,000 calories/day in season. Excluding nut cracking, Taï chimpanzees use a tool only every second day Significantly, humans use tools for different purposes than do nonhuman primates. Ape tools are rarely involved in killing or obtaining meat (Stanford 1996; Symbolic Behavior Following others Language, specifically, is widely considered a primary hallmark of humanity (e.g., Language differs from other types of communication by its symbolic and reflective content, using arbitrary symbols (words) to convey meaning (Pinker and Jackendoff 2005). Language today involves syntax, grammar, lexical items (which refer to concrete objects, perceptible attributes, or actions), and disambiguators or grammatical items that permit humans to express complex relational or referential ideas and to speak of the past, the present, the future, and the hypothetical Wild animals demonstrably communicate with each other. Vervet monkeys give different alarm calls to signal aerial versus terrestrial predators (Cheney and Seyfarth 1990). The symbolic vocabulary is clearly shared by the audience (the social group). However, the topic is not "snake" but "danger-snake"; the call cannot be used to ask the color of the snake, for example. Animals in experimental situations are capable of fairly sophisticated communication. Educated apes use about 400 words, do not use syntax or grammar, and do not combine more than two or three words into sentences (Savage-Rumbaugh and Lewin 1994). One of the most linguistically adept nonhumans, Kanzi the bonobo, uses fewer than 400 words (lexigrams) but understands about 3,000 Domestication Central to understanding the Neolithic Revolution Domestication itself is a process in which selective breeding by humans produces genetic change in a target species (Bö-könyi 1989; Only after fixation of behavioral traits are the morphological changes selected for, making the onset of domestication difficult to recognize. At the heart of the process of taming a wild animal, bringing it into the household, and controlling its reproduction and genetic output over generations is the formation of an intimate, reciprocal, and individual relationship between a particular animal and a particular human (Clutton-Brock 1999). Forming a relationship with an animal is more akin to negotiating a partnership than to caretaking a plant The ease with which zoonoses spread from domestic animals to humans shows the biological intimacy of the animal connection. In the archaeological record, domestic plants are recognized by changes in seed morphology, by appearance in new geographic regions, and/or by genetic analysis. The apparatus of agriculture, such as ploughs, hoes, scythes, granaries, fields, fences, or irrigation ditches, leaves archaeologically visible traces. Other mammals are not known to domesticate plants or animals. The Animal Connection as a Diagnostic Trait Like the traits reviewed above, the animal connection is universal among humans, is capable of powerfully transforming behavior, and is absent or extremely rare among other species. Same-species alloparenting is reasonably common among nonhuman primates Numerous reports allege cross-species alloparenting of young humans by wild animals. Purported alloparents include wolf, bear, dog, gazelle, chimpanzee, vervet monkey, goat, sheep, cow, ostrich, jackal, panther, and leopard. Few written reports of feral children raised by other mammals The Hypothesis I hypothesize that as an ancient, diagnostic trait of the human lineage, the animal connection had a major influence on human evolution, genetics, and behavior. This hypothesis predicts that the fossil and archaeological record will include abundant evidence that (1) humans were intimately and persistently connected with animals, (2) human adaptive changes were causally linked to the animal connection, and (3) a meaningful adaptive advantage of the animal connection can be identified in each stage of human evolution. Below, I review the archaeological and fossil record in three broad chronological stages and discuss evidence related to my hypothesis. I focus on the following key adaptive transformations: the origin and evolution of tool making, the manipulation of symbols and origin of language, and the domestication of animals and plants

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