Systematic and locomotor diversification of the Adapis group (Primates, Adapiformes) in the late Eocene of the Quercy (Southwest France), revealed by humeral remains.

Twenty humeral specimens from the old and new Quercy collections attributed to the fossil primates Adapis and Palaeolemur are described and analysed together. We provide a qualitative and quantitative analysis of the different humeri, revealing that high variability is present within the "Adapis group" sample. Six different morphotypes are identified, confirming that what has often been called "Adapis parisiensis" is a mix of different species that present different locomotor adaptations. Such a relatively high level of locomotor diversity is unique in the Paleogene primate fossil record. The humeral proportions of Adapis overlap with different groups of extant strepsirrhines and platyrrhines depending on the specimen, so the popular view of Adapis as a loris-like slow climbing primate does not apply to the whole sample presented here. Moreover, different humeral features traditionally associated with "Adapis parisiensis", such as the absence of a zona conoidea and a reduced brachioradialis flange, are variable depending on the sample studied. In addition, results of our analyses show that adapine and omomyid humeral morphology overlap extensively, leading us to question the accuracy of taxonomic attributions based on morphology of isolated humeri at localities where omomyids and adapines of similar size coexist. Finally, assuming our different morphotypes represent different species within two genera, we propose a phylogenetic hypothesis relating these morphotypes, which inhabited a small geographic area.

Nasolacrimal anatomy and haplorhine origins.

Computed tomography X-ray imaging of the internal face in well-preserved primate fossil crania permits reconstruction of the nature of their nasal anatomy, including some soft-tissue features. These features are diagnostic of the primate suborder Haplorhini, and allow reevaluation of the phylogenetic status of several purported early members of the group. Here we examine the nasolacrimal morphology of a broad sample of extant primates, as well as a number of Paleogene fossils. The extant sample confirms the distinctiveness of the two suborders. Of the fossils studied, only Shoshonius cooperi from the late-early Eocene exhibits evidence of a haplorhine nose. This suggests that the haplorhine oronasal complex may have evolved before the postorbital septum, and strengthens the claim that Shoshonius is a close relative of tarsiers and anthropoids. These results indicate that Omomyiformes is not a monophyletic group, and that few of its members possessed the derived oronasal morphology that characterizes crown haplorhines.

Postcrania of the most primitive euprimate and implications for primate origins.

The fossil record of early primates is largely comprised of dentitions. While teeth can indicate phylogenetic relationships and dietary preferences, they say little about hypotheses pertaining to the positional behavior or substrate preference of the ancestral crown primate. Here we report the discovery of a talus bone of the dentally primitive fossil euprimate Donrussellia provincialis. Our comparisons and analyses indicate that this talus is more primitive than that of other euprimates. It lacks features exclusive to strepsirrhines, like a large medial tibial facet and a sloping fibular facet. It also lacks the medially positioned flexor-fibularis groove of extant haplorhines. In these respects, the talus of D. provincialis comes surprisingly close to that of the pen-tailed treeshrew, Ptilocercus lowii, and extinct plesiadapiforms for which tali are known. However, it differs from P. lowii and is more like other early euprimates in exhibiting an expanded posterior trochlear shelf and deep talar body. In overall form, the bone approximates more leaping reliant euprimates. The phylogenetically basal signal from the new fossil is confirmed with cladistic analyses of two different character matrices, which place D. provincialis as the most basal strepsirrhine when the new tarsal data are included. Interpreting our results in the context of other recent discoveries, we conclude that the lineage leading to the ancestral euprimate had already become somewhat leaping specialized, while certain specializations for the small branch niche came after crown primates began to radiate.

Scale insect larvae preserved in vertebrate coprolites (Le Quesnoy, France, Lower Eocene): paleoecological insights.

Coprolites of terrestrial vertebrates from the Sparnacian Le Quesnoy locality (Ypresian, Eocene, MP7, 53 Ma; Oise, France) were examined for possible parasitic helminth eggs. The extraction of the coprolite components was performed by a weak acetolyse and a slide mounting in glycerin. This long examination did not reveal paleoparasite remains, which may be explained through several arguments. However, some pollen grains, some enigmatic components, and two well-preserved first-instar cochineal nymphs (Hemiptera: Sternorrhyncha: Coccoidea) were evidenced in coprolites. Identified as Coccidae, these larvae are the earliest stage of the scale insect development ever reported as fossil, revealing the specific environment of preservation that fossilized scats may provide. These observations, combined to the coprolites morphotype, enable to ascribe the fossil scats producer to a small herbivorous mammal present in the deposit (early perissodactyls or Plesiadapidae). Regarding the ecology of extant representatives of Coccidae, this mammal was a likely foliage consumer, and the abundant Juglandaceae and/or Tiliaceae from Le Quesnoy might have lived parasitized by scale insects. These Early Eocene parasites had an already well-established dissemination strategy, with prevalent minute first-instar larvae. The herein performed extraction technique appears well-suited for the study of carbonate coprolites and could certainly be useful for evidencing other kind of microorganisms (including internal parasites).

Testing for Depéret's Rule (Body Size Increase) in Mammals using Combined Extinct and Extant Data.

Whether or not evolutionary lineages in general show a tendency to increase in body size has often been discussed. This tendency has been dubbed "Cope's rule" but because Cope never hypothesized it, we suggest renaming it after Depéret, who formulated it clearly in 1907. Depéret's rule has traditionally been studied using fossil data, but more recently a number of studies have used present-day species. While several paleontological studies of Cenozoic placental mammals have found support for increasing body size, most studies of extant placentals have failed to detect such a trend. Here, we present a method to combine information from present-day species with fossil data in a Bayesian phylogenetic framework. We apply the method to body mass estimates of a large number of extant and extinct mammal species, and find strong support for Depéret's rule. The tendency for size increase appears to be driven not by evolution toward larger size in established species, but by processes related to the emergence of new species. Our analysis shows that complementary data from extant and extinct species can greatly improve inference of macroevolutionary processes.

Hands of early primates.

Questions surrounding the origin and early evolution of primates continue to be the subject of debate. Though anatomy of the skull and inferred dietary shifts are often the focus, detailed studies of postcrania and inferred locomotor capabilities can also provide crucial data that advance understanding of transitions in early primate evolution. In particular, the hand skeleton includes characteristics thought to reflect foraging, locomotion, and posture. Here we review what is known about the early evolution of primate hands from a comparative perspective that incorporates data from the fossil record. Additionally, we provide new comparative data and documentation of skeletal morphology for Paleogene plesiadapiforms, notharctines, cercamoniines, adapines, and omomyiforms. Finally, we discuss implications of these data for understanding locomotor transitions during the origin and early evolutionary history of primates. Known plesiadapiform species cannot be differentiated from extant primates based on either intrinsic hand proportions or hand-to-body size proportions. Nonetheless, the presence of claws and a different metacarpophalangeal [corrected] joint form in plesiadapiforms indicate different grasping mechanics. Notharctines and cercamoniines have intrinsic hand proportions with extremely elongated proximal phalanges and digit rays relative to metacarpals, resembling tarsiers and galagos. But their hand-to-body size proportions are typical of many extant primates (unlike those of tarsiers, and possibly Teilhardina, which have extremely large hands). Non-adapine adapiforms and omomyids exhibit additional carpal features suggesting more limited dorsiflexion, greater ulnar deviation, and a more habitually divergent pollex than observed plesiadapiforms. Together, features differentiating adapiforms and omomyiforms from plesiadapiforms indicate increased reliance on vertical prehensile-clinging and grasp-leaping, possibly in combination with predatory behaviors in ancestral euprimates.

New postcrania of Deccanolestes from the Late Cretaceous of India and their bearing on the evolutionary and biogeographic history of euarchontan mammals.

Extant species of the supraordinal mammal clade Euarchonta belong to the orders Primates, Scandentia, or Dermoptera. The fossil record of euarchontans suggests that they underwent their initial radiation during the Paleocene (65-55 million years ago) in North America, Eurasia, and Africa. The time and place of origin is poorly resolved due to lack of definitive fossils of euarchontan stem taxa. We describe a fragmentary humerus and two fragmentary ulnae from the latest Cretaceous of India that bear significantly on this issue. The fossils are tentatively referred to Deccanolestes cf. hislopi due to their small size and the fact that Deccanolestes is the only eutherian dental taxon to have been recovered from the same locality. The new fossils are used to evaluate the existing behavioral hypothesis that Deccanolestes was arboreal, and the competing phylogenetic hypotheses that Deccanolestes is a stem eutherian versus a stem euarchontan. The humerus resembles those of euarchontans in possessing a laterally keeled ulnar trochlea, a distinct zona conoidea, and a spherical capitulum. These features also suggest an arboreal lifestyle. The ulnar morphology is consistent with that of the humerus in reflecting an arboreal/scansorial animal. Detailed quantitative comparisons indicate that, despite morphological correlates to euarchontan-like arboreality, the humerus of Deccanolestes is morphologically intermediate between those of Cretaceous "condylarthran" mammals and definitive Cenozoic euarchontans. Additionally, humeri attributed to adapisoriculids are morphologically intermediate between those of Deccanolestes and definitive euarchontans. If adapisoriculids are euarchontans, as recently proposed, our results suggest that Deccanolestes is more basal. The tentative identification of Deccanolestes as a basal stem euarchontan suggests that (1) Placentalia began to diversify and Euarchonta originated before the Cretaceous-Tertiary boundary and (2) the Indian subcontinent, Eurasia, and Africa are more likely places of origin for Euarchonta than is North America.

Semicircular canal system in early primates.

Mammals with more rapid and agile locomotion have larger semicircular canals relative to body mass than species that move more slowly. Measurements of semicircular canals in extant mammals with known locomotor behaviours can provide a basis for testing hypotheses about locomotion in fossil primates that is independent of postcranial remains, and a means of reconstructing locomotor behaviour in species known only from cranial material. Semicircular canal radii were measured using ultra high resolution X-ray CT data for 9 stem primates ("plesiadapiforms"; n=11), 7 adapoids (n=12), 4 omomyoids (n=5), and the possible omomyoid Rooneyia viejaensis (n=1). These were compared with a modern sample (210 species including 91 primates) with known locomotor behaviours. The predicted locomotor agilities for extinct primates generally follow expectations based on known postcrania for those taxa. "Plesiadapiforms" and adapids have relatively small semicircular canals, suggesting they practiced less agile locomotion than other fossil primates in the sample, which is consistent with reconstructions of them as less specialized for leaping. The derived notharctid adapoids (excluding Cantius) and all omomyoids sampled have relatively larger semicircular canals, suggesting that they were more agile, with Microchoerus in particular being reconstructed as having had very jerky locomotion with relatively high magnitude accelerations of the head. Rooneyia viejaensis is reconstructed as having been similarly agile to omomyids and derived notharctid adapoids, which suggests that when postcranial material is found for this species it will exhibit features for some leaping behaviour, or for a locomotor mode requiring a similar degree of agility.

Lemuriform origins as viewed from the fossil record.

Fossils relevant to lemuriform origins are reviewed. Omanodon seems very close to the other early tooth-combed lemuriforms Karanisia, Wadilemur and Saharagalago, whereas Bugtilemur is rejected from the Lemuriformes. The Djebelemurinae, including Djebelemur and 'Anchomomys' milleri, are considered as stem lemuriforms preceding tooth comb differentiation; they are shown to be very distinct from European adapiforms. With tooth-combed lemuriforms present in Africa around 40 million years ago, and stem lemuriforms without tooth combs present on the same continent around 50-48 million years ago, a reasonable scenario can be proposed: tooth comb differentiation and lemuriform dispersal to Madagascar between 52-40 million years ago. The possible significance of Plesiopithecus for daubentoniid origins is raised. A critique of molecular dates is presented in the light of the fossil record. Azibiids are possibly early African prosimians. The timing of the dispersal of primates to Africa and the problem of strepsirhine origins are briefly examined.