New basicrania of Paleocene-Eocene Ignacius: re-evaluation of the Plesiadapiform-Dermopteran link

Plesiadapiformes has long been considered to be an archaic group of Primates. Discovery of a paromomyid plesiadapiform skull and independent analysis of referred postcrania have led investigators to conclude that Plesiadapiformes shares a closer relationship to extant flying lemurs (Dermoptera) than to Primates (= Euprimates of Hoffstetter [1977] Bull Mem Soc Anthropol Paris Ser 13 4:327-346). Despite challenges to this interpretation, the plesiadapiform-dermopteran relationship has gained currency in recent years. Here we show that newly discovered crania of Ignacius graybullianus, preserving previously undocumented portions of the ear, are more similar to primates than to dermopterans. New specimens confirm that paromomyids lacked the petrosal bulla of primates. However, these new specimens also demonstrate that paromomyids likely had: 1) a small promontorial branch of the internal carotid artery; 2) a lateral route for the internal carotid nerves crossing the promontorium; and 3) a ring-like ectotympanic with an annular bridge. This pattern is similar to primitive primates and fundamentally different from dermopterans, which have: 1) no internal carotid artery; 2) internal carotid nerves that take a more medial route; and 3) no annular bridge. Recognition of some primate-like traits, documented here by new evidence, indicates that Paromomyidae is likely to be more closely related to other Paleogene Plesiadapiformes and Eocene Primates than to extant Dermoptera. In view of these findings, a link between paromomyids and extant dermopterans ("Eudermoptera") is not convincingly supported by a single characteristic of the basicranium.     

Cranial anatomy of the Paleocene plesiadapiform Carpolestes simpsoni (Mammalia, Primates) using ultra high-resolution X-ray computed tomography, and the relationships of plesiadapiforms to Euprimates

Central to issues surrounding the origin of euprimates, affinities of Paleocene Carpolestidae have been controversial. Carpolestids have been classified as plesiadapoid primates, tarsiiform euprimates, dermopterans, or the sister taxon of euprimates to the exclusion of other plesiadapiforms, based exclusively on dental or postcranial data. Newly discovered crania of Carpolestes simpsoni from the latest Paleocene of the Clarks Fork Basin, Wyoming, are the first described for the family Carpolestidae. The two best preserved skulls were studied using ultra high-resolution X-ray computed tomography. Comparison of these specimens to those of other stem primates (Plesiadapiformes) demonstrates that the diversity of cranial morphology in this group is greater than previously thought. Carpolestes differs from euprimates and is similar to other plesiadapiforms (Ignacius and Plesiadapis) in lacking a postorbital bar and having a relatively long rostrum. Carpolestes is similar to fossil euprimates and Plesiadapis in having a bullar morphology consistent with a petrosal origin, and differs from Ignacius, in which the bulla is composed of the entotympanic. Carpolestes differs from primitive euprimates and all other known plesiadapiforms in possessing a two-chambered auditory bulla, similar to that of modern Tarsius. However, Carpolestes had an internal carotid artery (ICA) that took a transpromontorial route from a posteromedially positioned posterior carotid foramen (pcf), unlike Tarsius, in which this artery takes a perbullar route from an anterolaterally positioned pcf. Carpolestes has clear grooves on the promontorium for both the promontorial and stapedial arteries, indicating that it had an unreduced internal carotid circulation, similar to that of early euprimates. Carpolestes differs from primitive euprimates and some specimens of Ignacius in not having bony tubes surrounding the branches of the ICA. Cladistic analysis of cranial data fails to support a close relationship of Carpolestidae to either tarsiiform euprimates or extant Dermoptera, but suggests a close relationship between Carpolestidae, Plesiadapidae, and Euprimates.     

A pragmatic approach to the species problem from a paleontological perspective

The ideal scenario for paleontologists would be for the species they designate to be equivalent to the species recognized for modern animals, in the sense that they were formed as a result of the same evolutionary processes. This would mean, for example, that we could be confident that in combining extant and extinct taxa in phylogenetic analyses we would be dealing with equivalent operational taxonomic units. Notwithstanding the many thousands of pages that have been spent arguing over species concepts, the only concept that has won widespread acceptance for the designation of modern species is Mayr's Biological Species Concept (BSC).¹ In fact, whenever we complete a cladistic analysis, we assume reproductive isolation of our terminal taxa because otherwise their similarities could be the product of interbreeding rather than common ancestry. Fundamentally, we all behave as though the BSC is true.     

New discoveries on the middle ear anatomy of Ignacius graybullianus (Paromomyidae,Primates) from ultra high resolution X-ray computed tomography

A skull of Ignacius graybullianus (USNM 421608) was studied using ultra high resolution X-ray computed tomography (uhrCT). The anatomy of the middle ear in this specimen was previously studied through partial removal of the auditory bulla on one side. The data now available allow for examination of the others unprepared ear, which is more completely preserved, as well as adding to the information available about the previously studied ear. Analysis of the relationships between the bones making up the auditory bulla confirms previous assertions that it is formed from the entotympanic, and not from the petrosal, basioccipital, or basisphenoid. Contrary to previous reconstructions of the middle ear anatomy in all known plesiadapiforms, this specimen exhibits a bony canal for the promontorial artery and/or internal carotid nerves running across the lateral extreme of the promontorium. The identification of this structure is confirmed by the clear presence of a lumen, and its origination at a posterior carotid foramen (pcf) in a position that corresponds to that identified in previous studies of the paromomyid basicranium (Am. J. Phys. Anthropol., 36 (1972) 59, Am. J. Phys. Anthropol., 89 (1992) 477). Remnants of this canal are present bilaterally in USNM 421608, which additionally supports its identification. The presence of bony canals for branches of the internal carotid artery and the internal carotid nerves is a feature seen in scandentians and euprimates that is missing in dermopterans. The unusual lateral route followed by the internal carotid nerves is a primitive euprimate feature missing in all other archontans. As such, this evidence is consistent with a close euprimate-paromomyid relationship, and the inclusion of the latter in the order Primates. The discovery of this feature in paromomyids after almost 30 years of study of the ear region of this family acts as a cautionary note to the interpretation of the middle ear in damaged specimens.     

孟氏苏崩猴(哺乳动物纲:近兔猴超科)的齿列以及亚洲窃果猴科系统发育关系的再研究（英文）

格沙头期的窃果猴类(carpolestid)孟氏苏崩猴(Subengius mengi)属于亚洲已知最早的近兔猴形类(plesiadapiform)。苏崩猴的新标本澄清了该类群的牙齿解剖结构。孟氏苏崩猴的齿列比以前认为的要原始得多,下齿列的齿式2.1.3.3,p4低冠,具有3个主尖,但不似Elphidotarsius的相应主尖那样完全并生,P3的舌侧缘更窄,结构更简单,m1不具有高度扩展的下前尖和下后尖。苏崩猴P3的独特结构以及对Elphidotarsius sp.,cf.E.florencae的P3解剖特征的重新研究表明,过去对窃果猴科P3的某些齿尖的同源性的解释是错误的。在详细的特征分析基础上,重新建立了窃果猴类及其近亲的系统发育关系。崩班期(Bumbanian)的同时猴(Chronolestes simul)被重新认定是窃果猴科最基部的成员。孟氏苏崩猴以及崩班期的另一个种旭日多脊食果猴(Carpocristes oriens)也是窃果猴类相对靠基部的成员,但这些亚洲窃果猴类之间似乎都没有特殊的相互关系。虽然北美和亚洲的窃果猴类均延续到古新世-始新世界线附近,但它们在两个大陆之间的扩散似乎仅限于古新世的较早期。     

山东五图早始新世更猴科(Plesiadapidae,Mammalia)化石

记述了山东五图盆地早始新世五图组的一种更猴类 (plesiadapid)化石 :杨氏亚洲更猴(Asioplesiadapisyoungigen .etsp .nov .)。在它增大的下门齿上具有更猴科特有的下缘尖和下缘脊 ,但其下颊齿性状相当原始 ,p4跟座呈盆状 ,m3下次小尖呈锥状更类似白垩纪晚期出现的更猴形动物Purgatorius。Asioplesiadapis的下臼齿形态可以说明亚洲更猴类虽然有北美和欧洲更猴的某些典型特征 ,但并不在同一进化支系。因此 ,另立一新亚科———亚洲更猴亚科(Asioplesiadapinaesubf.nov.)。     

Discovery of a highly-specialized plesiadapiform primate in the early-middle Eocene of northwestern Africa

In this paper we report the first occurrence of an endemic African plesiadapiform primate from the early-middle Eocene locality of Glib Zegdou (Hammada du Dra, Algeria). Dralestes (new genus) is a very specialized taxon, and its closest known relative is the enigmatic and controversial genus Azibius from Gour Lazib (Hammada du Dra). We group both together as the Azibiidae (new rank). Dralestes provides the first evidence of the upper dentition in this group. Some critical dental characters, such as a postprotocingulum on upper teeth, consistently reveal a primate status for the azibiids. Dralestes exhibits, however, a very unusual configuration of the upper molars by the enlarged parastyle, the lack of a metaconule, and the ectoloph structure (preparacrista, centrocrista and postmetacrista are aligned in a high blade-like structure). The apparent dental specializations of both lower premolars and molars of azibiids (exaenodonty, large P(4) bearing sharp apical cusps, and M(1) having a highly elongated trigonid) point to potential relationships with Chronolestes and carpolestid plesiadapiforms. A phylogenetic analysis, performed on 55 dental characters scored for 19 primate genera, clarifies the euprimate status of Altiatlasius, and thus indicates that azibiids are the only known plesiadapiforms from Africa. Azibiids are the sister group of the clade carpolestids/Chronolestes in the superfamily Plesiadapoidea. However, the azibiids differ fundamentally from carpolestids by the combined lack of a centroconule and multiple buccal cusps on P(4). The exact position of both Chronolestes and azibiids in the plesiadapoids appears difficult to resolve. A basal position of Chronolestes in this superfamily cannot be ruled out because it exhibits a simple morphology of I(1) and no conule on P(3). Considering this ad hoc hypothesis, azibiids are found to lie outside a clade including carpolestids/plesiadapids/saxonellids, and they are the sister group to Chronolestes. The clade including the carpolestid, saxonellid, and plesiadapid families is characterized by the occurrence of a centroconule on P(3-4). The lack of this trait in Dralestes and Chronolestes could mean that azibiids are basal plesiadapoids that diverged before the evolution of the common ancestor of the three derived plesiadapoid families, i.e. at least around the Paleocene-Eocene boundary or more probably during the Paleocene. The report of the first offshoot in Africa of plesiadapoids enhances the role of Africa in the early primate radiation.     

Hallucal grasping behavior in Caluromys (Didelphimorphia: Didelphidae): Implications for primate pedal grasping

A key feature in primate evolution is a foot with a divergent opposable hallucal metatarsal bearing a large peroneal process. Extant primates are characterized by a powerful hallucal grasp—an either “euprimate” or “plesiadapoid-euprimate” ancestor acquisition—that facilitates the exploitation of fine branches, an ability that increased the fitness of ancestral euprimates. In this context, the didelphid marsupial Caluromys has been used as the extant analog to this primate morphotype stage due to some morphological, ecological, and behavioral features. However, the extent to which and the positional and support contexts in which Caluromys uses powerful hallucal grasping are not known. This renders analogies to any mode of “euprimate” or “stem primate” grasping poorly substantiated. The present paper quantifies locomotor and postural behavior, support use, and associated frequencies of hallucal grasping in captive Caluromys philander via analysis of video recordings. During locomotion, Caluromys primarily used diagonal sequence walk, clamber, and climb, whereas stand, foot-hang, and bipedal stand were the dominant postures. Small, fine, horizontal, and moderately inclined branches were frequently used. Overall rates of “apparently powerful hallucal grasps” were high, but were exceptionally high during clamber, climb, foot-hang, and bipedal stand. Additionally, an “apparently powerful hallucal grasp” was very common upon fine, small, steep, and vertical branches. The extensive use of such powerful hallucal grasping provided stability and safety that enabled Caluromys to proficiently utilize fine branches of various orientations. The ability to negotiate such unstable supports, further reflected in foot anatomy, provides evidence that the morphobehavioral complex of Caluromys can serve as an extant analog to the plesiadapoid-euprimate ancestor, represented as a terminal branch feeder with effective hallucal grasping.     

The phylogenetic significance of strepsirhinism in Paleogene primates

Two lines of evidence reviewed here argue against the recent proposal that strepsirhinism is an apomorphous feature in primates, shared only by adapiforms and lemuriforms. These are (1) the presence of strepsirhinism in several extant taxa of nonprimate mammals, including Tupaiidae, Tenrecidae, Erinaceus,and Didelphis,and (2) the inferred presence of strepsirhinism not only in adapiforms, but also in all plesiadapiforms and omomyids for which the relevant anatomical regions are known. Therefore, strepsirhinism cannot be invoked as an adaptive innovation underlying the initial strep-sirhine/haplorhine dichotomy. Likewise, the apparent retention of strepsirhinism in omomyids suggests either that the haplorhine oronasal configuration of extant tarsiids and anthropoids was acquired independently or that tarsiids and anthropoids form a clade to the exclusion of omomyids.