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The evolutionary history that led to Eocene-and-later primates of modern aspect (Euprimates) has been uncertain. We describe a skeleton of Paleocene plesiadapiform Carpolestes simpsoni that includes most of the skull and many postcranial bones. Phylogenetic analyses indicate that Carpolestidae are closely related to Euprimates. C. simpsoni had long fingers and an opposable hallux with a nail. It lacked orbital convergence and an ankle specialized for leaping. We infer that the ancestor of Euprimates was primitively an arboreal grasper adapted for terminal branch feeding rather than a specialized leaper or visually directed predator.

Paleontological evidence indicates that the evolutionary diversification of mammals early in the Cenozoic era was characterized by an adaptive radiation of distal limb structures. Likewise, neontological data show that morphological variation in distal limb integumentary appendages (e.g., nails, hooves, and claws) can be observed not only among distantly related mammalian taxa but also among closely related species within the same clade. Comparative analysis of nail, claw, and hoof morphogenesis reveals relatively subtle differences in mesenchymal and epithelial patterning underlying these adult differences in distal limb appendage morphology. Furthermore, studies of regulatory gene expression during vertebrate claw development demonstrate that many of the signaling molecules involved in patterning ectodermal derivatives such as teeth, hair, and feathers are also involved in organizing mammalian distal limb appendages. For example, Bmp4 signaling plays an important role during the recruitment of mesenchymal cells into the condensations forming the terminal phalanges, whereas Msx2 affects the length of nails and claws by suppressing proliferation of germinal epidermal cells. Evolutionary changes in the form of distal integumentary appendages may therefore result from changes in gene expression during formation of mesenchymal condensations (Bmp4, posterior Hox genes), induction of the claw fold and germinal matrix (shh), and/or proliferation of epidermal cells in the claw matrix (Msx1, Msx2). The prevalence of convergences and parallelisms in nail and claw structure among mammals underscores the existence of multiple morphogenetic pathways for evolutionary change in distal limb appendages.

The issue of whether nails or claws were present on the digits of the last common ancestor of living primates is central to the understanding of the ecological context in which the order originated. Two lines of evidence are available, the shape (claw, nail, toilet-claw) and the histological structure (one or two horny strata). Here we review the existing data regarding the shape and histological structure of cheirideal appendages in primates and present new information from a wide range of living primates. We demonstrate the presence of a typical toilet-claw in Daubentonia madagascariensis and discuss its consequences, since the alleged lack of such structures in this species has long obscured the issue. The general view that primate nails, with the exception of those in New World primates, consist of only one layer is disproved by the presence of two distinct strata in the nails of the feet of three out of seven catarrhine species examined, as well as in Lemur catta. The combined new and old data indicate that the last common ancestor of the extant primates had lost the typical mammalian claws of its ancestors and developed nails on all pedal digits except digit II, which bore a toilet-claw. All nails as well as the toilet-claw originally consisted of two layers. We present a new hypothesis regarding the adaptational significance of these changes., (Copyright 1999 Academic Press.)

Primates of the subfamily Callitrichinae (Callimico, Callithrix, Leontopithecus, and Saguinus) are small-bodied New World monkeys (105-700 g) possessing clawlike nails on all manual and pedal digits excluding the hallux. Specialized nails in these genera serve a critical function in feeding by enabling tamarins and marmosets to cling to trunks and other large vertical supports while exploiting food resources. Within the subfamily, there is evidence of at least four distinct large-branch feeding patterns. These include (1) seasonal exudate feeding and occasional trunk foraging (many Saguinus spp.); (2) exploitation of bark surface insects and the use of trunks as a platform to locate terrestrial prey (Saguinus fuscicollis, S. nigricollis, and Callimico); (3) manipulative foraging and bark stripping to locate concealed insects and small vertebrates (Leontopithecus); and (4) tree gouging and year-round exudate feeding (many Callithrix). Large-branch feeding and the use of vertical clinging postures appear to be a primary adaptation among virtually all callitrichines, distinguishing them ecologically from other platyrrhine taxa. Given the anatomy and behavior of extant callitrichines, Saguinus appears to be the most ecologically generalized member of this subfamily, and species of this genus may provide useful models for reconstructing the feeding and foraging adaptations of early callitrichines.

A recently proposed scenario for the evolution of a unified complex of exudativory, vertical clinging, and claw-like nails in callitrichids is examined. Consideration of behaviors and morphologies exhibited by Callimico goeldii leads to a decoupling of this "complex," with the early occurrence of claws and vertical clinging in the callitrichid lineage and only later, in some descendents, the development of exudativory and possible related behaviors.