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08 Early Primate Evolution Paper No. : 14 Human Origin and Evolution Module : 08 Early Primate Evolution Development Team Principal Investigator Prof. Anup Kumar Kapoor Department of Anthropology, University of Delhi Dr. Satwanti Kapoor (Retd Professor) Paper Coordinator Department of Anthropology, University of Delhi Mr. Vijit Deepani & Prof. A.K. Kapoor Content Writer Department of Anthropology, University of Delhi Prof. R.K. Pathak Content Reviewer Department of Anthropology, Panjab University, Chandigarh 1 Early Primate Evolution Anthropology Description of Module Subject Name Anthropology Paper Name Human Origin and Evolution Module Name/Title Early Primate Evolution Module Id 08 Contents: Fossil Primates: Introduction Theories of primate origin Primates: Pre- Pleistocene Period a. Palaeocene epoch b. Eocene epoch c. Oligocene epoch d. Miocene – Pliocene epoch Summary Learning outcomes: The learner will be able to develop: an understanding about fossil primates and theories of primate origin. an insight about the extinct primate types of Pre-Pleistocene Period. 2 Early Primate Evolution Anthropology Fossil Primates: Introduction In modern time, the living primates are graded in four principle domains – Prosimian, Monkey, Ape and man. On the basis of examination of fossil evidences, it has been established that all the living primates have evolved and ‘adaptively radiated’ from a common ancestor. Fossil primates exhibit a palaeontological record of evolutionary processes that occurred over the last 65 to 80 million years. Crucial evidence of intermediate forms, that bridge the gap between extinct and extant taxa, is yielded by the macroevolutionary study of the primate fossil evidences. The paleoanthropologists often utilize the comparative anatomical method to develop insight about morphological adaptations in fossil primates. Crucial insight into the diet and behaviour of fossil primates is exhibited by comparing their dentition and skeletal traits with those of living primates. The examination of dental fossil material (such as teeth) could yield crucial information about the climate conditions of the period when the animal existed, if the geological structure in which the teeth were found is known. The primate fossil records and evidences provide crucial information on evolutionary trend of primate diversity over tens of millions of years. The geological time scale provides crucial evidences related to appearance and dominance of ancestral primate forms. The Cenozoic Era is characterized by the end of the “Age of the Dinosaurs” and the beginning of the “Age of the Mammals”. This geological era witnessed significant climate and topographic changes on earth and held utmost importance in relation to primate origins and evolution. There were considerable geological and climatic changes during the Cenozoic era. The Cenozoic is further ramified into seven epochs - Palaeocene, Eocene, Oligocene, Miocene, Pliocene, Pleistocene, and Holocene. During the Cenozoic era, primates appeared and adaptively radiated into all the various species living today. It has been emphasized that earliest primates of Palaeocene epoch were small prosimians (like tree shrews). During the Eocene epoch, lemurs and tarsiers of various kinds were distributed across North America and parts of Europe and Asia. But gradually with time, their number declined. In subsequent periods, in the new world, prosimian forms lead to development of the platyrrhine monkeys and in the Old World, these forms eventually evolved and led to emergence of catarrhine moneys, apes and man. 3 Early Primate Evolution Anthropology Fig I: Geological time scale (Source: Lambert, 1985) 4 Early Primate Evolution Anthropology ©Encyclopedia of Life Support Systems (EOLSS) Fig II: A possible hypothesis of relationships among extant and extinct primates. (Source: Masters, 2009) PP CENE = Plio-Pleistocene epoch; dotted lines and question marks represent inferred relationships and uncertainties respectively. Theories of primate origin Scientists have proposed several theories in relation to primate origin and evolution, and the following three theories have gained recognition and acceptance – arboreality, predation, and ecology (Lehman, 2009). Fred Szalay (1972) proposed the arboreal theory of primate origins and emphasized that primate origins represent an adaptive radiation of new species from early arboreal ancestral mammals. A change in dietary pattern from insectivory to herbivory was the impulse for this evolutionary shift. In order to ease out locomotion and movement in the complex web of flexible tree branches, grasping 5 Early Primate Evolution Anthropology hands and feet also evolved in these early mammals. Although the theory throws light upon modification in dentition and limb morphology, it could not explain certain aspects of the primate visual system. Mart Cartmill (1992) developed an alternative idea on primate origins, which is known as the visual predation theory. Cartmill emphasized that primate origins can be traced back to visual adaptations and modifications for hunting prey in arboreal dwellings. He further opined that arboreal hunting lead to evolution of grasping hands and nails which could facilitate primates to capture and hold prey in small and terminal branches. But the detailed ecological examination of insectivorous primates showed significant flaws in Cartmill’s theory. Wright et al. (2003) suggested that nocturnal insectivores are more likely to hunt using smell and sound rather than relying upon visual system and Garber (1980) suggested that tamarins, characterized with claw-like nails, exhibit adaptation in both hunting for insects in terminal branches and clinging to tree trunks to feed on exudates. These observations led, in part, to Robert Sussman’s (1991) more ecologically based idea on primate origins, called the angiosperm co-evolution theory. He suggested that the prime impulse behind primate origins and adaptations was the roughly coeval evolution of angiosperm plants. The evolution of angiosperm plants formulated a new set of ecological niche for animals. The former lured animals to disperse their seeds and in turn provided tasty and nutritious fruit to them. Primates were one of the principle taxonomic groups that evolved to take advantage of this scenario. But it was emphasized that the field studies of living mammals similar to those that led to the first primates did not provide crucial evidence in support of either Sussman’s or Cartmill’s theories singly but provided support for aspects of both theories. Pre-Pleistocene epoch In order to develop understanding about early primate evolution, primate fossil remains from the different strata of pre-Pleistocene period deposits are discussed and dealt with. Paleocene primates The Paleocene epoch was characterized by warm climatic conditions with savannas and woodlands dominating the geographic domain in North America and Europe (Swindler, 2004). The mountainous regions of North America and France in Europe yielded fossil remains of several varieties of the 6 Early Primate Evolution Anthropology primate – like taxa and most of the remains consisted of teeth and fragments of Jaw. The deposit of the middle Paleocene epoch yielded the fossil evidence of existence of the earliest primates (Das, 2008). The transition from Cretaceous to Palaeocene epoch was noted by appearance of small and primitive insectivorous mammal called Purgatorius ceratops. The late Cretaceous deposit yielded a single lower molar tooth which is linked to Purgatorius ceratops (Valen and Sloan, 1965). This genus along with other species of primitive primates is grouped under the infraorder Plesiadapiformes (Clemens, 1974). Plesiadapiformes encompasses three major families namely Carpolestidae, Phenacolemuridae and Plesiadapidae. On the basis of fossil evidences it is emphasized that most of these genera existed in North America and a few in Europe. The Family Carpolestidae is defined by certain typical fossil evidences such as jaws, both maxillae and mandibles, and teeth. Phenacolemuridae group represents the earliest of the primate forms but there is a lack of fossil evidences related to this group. Among the three groups, Plesiadapidae represent the best known family. The fossil remains of Plesiadapidae were constituted by a complete fossil skull and other parts of the skeletons. The members of this family were devoid of stereoscopic vision. Swindler (2004) suggested that members of Plesiadapiformes were characterized by an auditory bulla formed by the petrosal portion of the temporal bone (a hallmark of living primates) and a dental formula (in most species) of 2.1.3.3, for both upper and lower jaws, indicating a reduction from the basic eutherian 3.1.4.3 dental formula. Several scientific investigation have emphasized that Plesiadapis are unlikely to be members of the Primate Order. Plesiadapis - a prominent archaic under Plesiadapiformes - possessed primitive mammalian features such as claws instead of nails, rodent-like jaws and teeth, eyes at the side of the head, greatly enlarged incisors, a long snout and no postorbital bar (Fig III). 7 Early Primate Evolution Anthropology Fig III: Reconstruction of Plesiadapis – a prominent archaic under Plesiadapiformes (Source: Lambert, 1987) The Plesiadapiformes is traditionally different from Prosimii and Anthropoidea and it can be argued that the former are not ancestral to modern primates but represent an early offshoot from the primate family tree that does not lead to modern primates (Sellers,
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