Jonathan I. Bloch

New Paleocene skeletons and the relationship of plesiadapiforms to crown-clade primates

Plesiadapiforms are central to studies of the origin and evolution of primates and other euarchontan mammals (tree shrews and flying lemurs). We report results from a comprehensive cladistic analysis using cranial, postcranial, and dental evidence including data from recently discovered Paleocene plesiadapiform skeletons (Ignacius clarkforkensis sp. nov.; Dryomomys szalayi, gen. et sp. nov.), and the most plesiomorphic extant tree shrew, Ptilocercus lowii. Our results, based on the fossil record, unambiguously place plesiadapiforms with Euprimates and indicate that the divergence of Primates (sensu lato) from other euarchontans likely occurred before or just after the Cretaceous/Tertiary boundary (65 Mya), notably later than logistical model and molecular estimates. Anatomical features associated with specialized pedal grasping (including a nail on the hallux) and a petrosal bulla likely evolved in the common ancestor of Plesiadapoidea and Euprimates (Euprimateformes) by 62 Mya in either Asia or North America. Our results are consistent with those from recent molecular analyses that group Dermoptera with Scandentia. We find no evidence to support the hypothesis that any plesiadapiforms were mitten-gliders or closely related to Dermoptera.

Giant boid snake from the Palaeocene neotropics reveals hotter past equatorial temperatures

The largest extant snakes live in the tropics of South America and southeast Asia where high temperatures facilitate the evolution of large body sizes among air-breathing animals whose body temperatures are dependant on ambient environmental temperatures (poikilothermy). Very little is known about ancient tropical terrestrial ecosystems, limiting our understanding of the evolution of giant snakes and their relationship to climate in the past. Here we describe a boid snake from the oldest known neotropical rainforest fauna from the Cerrejón Formation (58–60 Myr ago) in northeastern Colombia. We estimate a body length of 13 m and a mass of 1,135 kg, making it the largest known snake. The maximum size of poikilothermic animals at a given temperature is limited by metabolic rate, and a snake of this size would require a minimum mean annual temperature of 30–34 °C to survive. This estimate is consistent with hypotheses of hot Palaeocene neotropics with high concentrations of atmospheric CO2 based on climate models. Comparison of palaeotemperature estimates from the equator to those from South American mid-latitudes indicates a relatively steep temperature gradient during the early Palaeogene greenhouse, similar to that of today. Depositional environments and faunal composition of the Cerrejón Formation indicate an anaconda-like ecology for the giant snake, and an earliest Cenozoic origin of neotropical vertebrate faunas.

Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum

Warming and Shrinking In most mammals, individual body sizes tend to be smaller in warmer regions and larger in cooler regions. Secord et al. (p. 959; see the Perspective by Smith) examined a high-resolution 175,000-year record of equid fossils deposited over a past climate shift—the Paleocene-Eocene Thermal Maximum—for changes in body size. Using oxygen isotopes collected from the teeth of co-occurring mammal species to track prevailing environmental temperature, a clear decrease in equid body size was seen during 130,000 years of warming, followed by a distinct increase as the climate cooled at the end of the period. These results indicate that temperature directly influenced body size in the past and may continue to have an influence as our current climate changes. Oxygen isotope measurements of fossil teeth show that the body size of the horse Sifrhippus decreased as temperature increased. Body size plays a critical role in mammalian ecology and physiology. Previous research has shown that many mammals became smaller during the Paleocene-Eocene Thermal Maximum (PETM), but the timing and magnitude of that change relative to climate change have been unclear. A high-resolution record of continental climate and equid body size change shows a directional size decrease of ~30% over the first ~130,000 years of the PETM, followed by a ~76% increase in the recovery phase of the PETM. These size changes are negatively correlated with temperature inferred from oxygen isotopes in mammal teeth and were probably driven by shifts in temperature and possibly high atmospheric CO2 concentrations. These findings could be important for understanding mammalian evolutionary responses to future global warming.

Intrinsic hand proportions of euarchontans and other mammals: implications for the locomotor behavior of plesiadapiforms.

Arboreal primates have distinctive intrinsic hand proportions compared with many other mammals. Within Euarchonta, platyrrhines and strepsirrhines have longer manual proximal phalanges relative to metacarpal length than colugos and terrestrial tree shrews. This trait is part of a complex of features allowing primates to grasp small-diameter arboreal substrates. In addition to many living and Eocene primates, relative elongation of proximal manual phalanges is also present in most plesiadapiforms. In order to evaluate the functional and evolutionary implications of manual similarities between crown primates and plesiadapiforms, we measured the lengths of the metacarpal, proximal phalanx, and intermediate phalanx of manual ray III for 132 extant mammal species (n=702 individuals). These data were compared with measurements of hands in six plesiadapiform species using ternary diagrams and phalangeal indices. Our analyses reveal that many arboreal mammals (including some tree shrews, rodents, marsupials, and carnivorans) have manual ray III proportions similar to those of various arboreal primates. By contrast, terrestrial tree shrews have hand proportions most similar to those of other terrestrial mammals, and colugos are highly derived in having relatively long intermediate phalanges. Phalangeal indices of arboreal species are significantly greater than those of the terrestrial species in our sample, reflecting the utility of having relatively long digits in an arboreal context. Although mammals known to be capable of prehensile grips demonstrate long digits relative to palm length, this feature is not uniquely associated with manual prehension and should be interpreted with caution in fossil taxa. Among plesiadapiforms, Carpolestes, Nannodectes, Ignacius, and Dryomomys have manual ray III proportions that are unlike those of most terrestrial species and most similar to those of various arboreal species of primates, tree shrews, and rodents. Within Euarchonta, Ignacius and Carpolestes have intrinsic hand proportions most comparable to those of living arboreal primates, while Nannodectes is very similar to the arboreal tree shrew Tupaia minor. These results provide additional evidence that plesiadapiforms were arboreal and support the hypothesis that Euarchonta originated in an arboreal milieu.

Affinities of ‘hyopsodontids’ to elephant shrews and a Holarctic origin of Afrotheria

Macroscelideans (elephant shrews or sengis) are small-bodied (25–540 g), cursorial (running) and saltatorial (jumping), insectivorous and omnivorous placental mammals represented by at least 15 extant African species classified in four genera. Macroscelidea is one of several morphologically diverse but predominantly African placental orders classified in the superorder Afrotheria by molecular phylogeneticists. The distribution of modern afrotheres, in combination with a basal position for Afrotheria within Placentalia and molecular divergence-time estimates, has been used to link placental diversification with the mid-Cretaceous separation of South America and Africa. Morphological phylogenetic analyses do not support Afrotheria and the fossil record favours a northern origin of Placentalia. Here we describe fossil postcrania that provide evidence for a close relationship between North American Palaeocene–Eocene apheliscine ‘hyopsodontid’ ‘condylarths’ (early ungulates or hoofed mammals) and extant Macroscelidea. Apheliscine postcranial morphology is consistent with a relationship to other ungulate-like afrotheres (Hyracoidea, Proboscidea) but does not provide support for a monophyletic Afrotheria. As the oldest record of an afrothere clade, identification of macroscelidean relatives in the North American Palaeocene argues against an African origin for Afrotheria, weakening support for linking placental diversification to the break-up of Gondwana.

Cranial Anatomy of the Earliest Marsupials and the Origin of Opossums

Background The early evolution of living marsupials is poorly understood in part because the early offshoots of this group are known almost exclusively from jaws and teeth. Filling this gap is essential for a better understanding of the phylogenetic relationships among living marsupials, the biogeographic pathways that led to their current distribution as well as the successive evolutionary steps that led to their current diversity, habits and various specializations that distinguish them from placental mammals. Methodology/Principal Findings Here we report the first skull of a 55 million year old peradectid marsupial from the early Eocene of North America and exceptionally preserved skeletons of an Oligocene herpetotheriid, both representing critical groups to understand early marsupial evolution. A comprehensive phylogenetic cladistic analysis of Marsupialia including the new findings and close relatives of marsupials show that peradectids are the sister group of living opossums and herpetotheriids are the sister group of all living marsupials. Conclusions/Significance The results imply that North America played an important role in early Cenozoic marsupial evolutionary history and may have even been the center of origin of living marsupials and opossums. New data from the herpetotheriid postcranium support the view that the ancestral morphotype of Marsupialia was more terrestrial than opossums are. The resolution of the phylogenetic position of peradectids reveals an older calibration point for molecular estimates of divergence times among living marsupials than those currently used.

Virtual endocast of Ignacius graybullianus (Paromomyidae, Primates) and brain evolution in early primates

Extant primates are distinctive among mammals in having relatively large brains. As stem primates, Paleogene plesiadapiforms provide direct information relevant to the earliest stages in the evolution of this characteristic. Here we describe a virtual endocast reconstructed from ultra high resolution X-ray computed tomography data for the paromomyid plesiadapiform Ignacius graybullianus (USNM 421608) from the early Eocene of Wyoming. This represents the most complete endocast known for a stem primate, allowing for an unprecedented study of both size and fine details of anatomy. Relative to fossil and extant euprimates, I. graybullianus had large olfactory lobes, but less caudal development of the cerebrum and a poorly demarcated temporal lobe, suggesting more emphasis on olfaction and a less well developed visual system. Although its brain was small compared to those of extant primates, the encephalization quotient of I. graybullianus is higher than that calculated for Paleocene Plesiadapis cookei and overlaps the lower portion of the range documented for fossil euprimates. Comparison to the basal gliroid Rhombomylus suggests that early primates exhibited some expansion of the cerebrum compared to their ancestors. The relatively small brain size of I. graybullianus, an arboreal frugivore, implies that neither arboreality nor frugivory was primarily responsible for the expanded brains of modern primates. However, the contrasts in features related to the visual system between I. graybullianus and fossil and extant euprimates suggest that improvements to these portions of the brain contributed to increases in brain size within Euprimates.

New Skeletons of Paleocene-Eocene Plesiadapiformes: A Diversity of Arboreal Positional Behaviors in Early Primates

Knowledge of plesiadapiform skeletal morphology and inferred ecological roles are critical for establishing the evolutionary context that led to the appearance and diversification of Euprimates (see Silcox, this volume). Plesiadapiform dentitions are morphologically diverse, representing over 120 species usually classified in 11 families from the Paleocene and Eocene of North America, Europe, and Asia (Hooker et al., 1999; Silcox, 2001; Silcox and Gunnell, in press). Despite this documented diversity in dentitions,

Endocasts of Microsyops (Microsyopidae, Primates) and the evolution of the brain in primitive primates

We describe a virtual endocast produced from ultra high resolution X-ray computed tomography (CT) data for the microsyopid, Microsyops annectens (middle Eocene, Wyoming). It is the most complete and least distorted endocast known for a plesiadapiform primate and because of the relatively basal position of Microsyopidae, has particular importance to reconstructing primitive characteristics for Primates. Cranial capacity is estimated at 5.9 cm(3), yielding encephalization quotients (EQ) of 0.26-0.39 (Jerisons equation) and 0.32-0.52 (Eisenbergs equation), depending on the body mass estimate. Even the lowest EQ estimate for M. annectens is higher than that for Plesiadapis cookei, while the range of estimates overlaps with that of Ignacius graybullianus and with the lower end of the range of estimates for fossil euprimates. As in other plesiadapiforms, the olfactory bulbs of M. annectens are large. The cerebrum does not extend onto the cerebellum or form a ventrally protruding temporal lobe with a clear temporal pole, suggesting less development of the visual sense and a greater emphasis on olfaction than in euprimates. Contrasts between the virtual endocast of M. annectens, and both a natural endocast of the same species and a partial endocast from the earlier-occurring Microsyops sp., cf. Microsyops elegans, suggest that the coverage of the caudal colliculi by the cerebrum evolved within the Microsyops lineage. This implies that microsyopids expanded their cerebra and perhaps evolved an improved visual sense independent of euprimates. With a growing body of data on the morphology of the brain in primitive primates, it is becoming clear that many of the characteristics of the brain common to euprimates evolved after the divergence of stem primates from other euarchontans and likely in parallel in different lineages. These new data suggest a different model for the ancestors of euprimates than has been assumed based on the anatomy of the brain in visually specialized diurnal tree shrews.

Systematics and biogeography of crocodylians from the Miocene of Panama

ABSTRACT Despite the fact that fossil crocodylians have been recovered from the Panama Canal Zone starting with initial excavations in 1912, detailed studies have been lacking. Recent excavations of the canal have resulted in new discoveries of many vertebrate fossils, including the first known Miocene crocodylian skulls from Central America. These fossil skulls from the early-middle Miocene represent two new taxa with distinct morphology that is shared with extinct and extant caimans (Caimaninae). A cladistic analysis of 32 alligatorid and three outgroup taxa, scored for 75 characters, resulted in 1210 equally most parsimonious cladograms, all of which suggest that Culebrasuchus mesoamericanus, gen. et sp. nov., is the sister taxon to all previously known Caimaninae. Additionally, the analysis suggests that Centenariosuchus gilmorei, gen. et sp. nov., is the sister taxon to a caimanine clade that includes Purussaurus from the Miocene of South America. In fact, teeth very similar to those of Purussaurus have also been recovered from the Panama Canal. Given these South American affinities, we suggest that these early caimanines dispersed across saltwater. This is a potentially surprising result, because all extant alligatorids lack the salt glands that would have been necessary for the marine dispersal required to reach Central America during the Miocene. Unlike Miocene mammals that all have North American affinities, the Miocene crocodylians of Panama represent a ‘melting pot’ with taxa of disparate origins living together at the southern extreme of Central America.