Plesiorycteropus
Plesiorycteropus, also known as the bibymalagasy or Malagasy aardvark, is a recently extinct genus of mammals from Madagascar. Upon its description in 1895, it was classified with the aardvark, but more recent molecular evidence instead suggests that it is most closely related to the tenrecs. Two species are currently recognized, the larger P. madagascariensis and the smaller P. germainepetterae. They probably overlapped in distribution, as subfossil remains of both species have been found in the same site.
Knowledge of the skeletal anatomy is limited, as only limb, partial pelvis, and skull bones have been recovered to date. Plesiorycteropus was probably a digging animal that fed on insects such as termites and ants. It also shows adaptations for climbing and sitting. Estimates of its mass range from. When and why it became extinct remains unknown. One bone has been radiocarbon dated to 200 BCE; forest destruction by humans may have contributed to its extinction.
Taxonomy
Identification and species
French naturalist Henri Filhol first described Plesiorycteropus madagascariensis in 1895 on the basis of a partial skull found at the cave of Belo. His description was vague even by 19th-century standards, but he placed the animal close to the aardvark. The generic name combines Ancient Greek plesio- "near" with Orycteropus, the genus of the aardvark, and the specific name refers to Madagascar. Charles Lamberton, who had access to a larger sample for his 1946 review of the genus, noted substantial variation, but did not attempt to differentiate multiple species. In 1994, Ross MacPhee again reviewed Plesiorycteropus and was able to separate two species, the larger P. madagascariensis and a new, smaller species that he named Plesiorycteropus germainepetterae after scientist Germaine Petter. The two species differ in a number of morphological characters in addition to size.Remains of Plesiorycteropus have been misidentified as rodents and primates. Charles Immanuel Forsyth Major described Myoryctes rapeto in 1908 as a "giant subfossil rat" on the basis of two innominate bones. The generic name was replaced by Majoria in 1915, because Myoryctes was preoccupied by the name of a nematode worm. However, according to MacPhee, innominates of Majoria are identical to those assigned to Plesiorycteropus. Guillaume Grandidier assigned a well-preserved femur to a gigantic relative of the living votsovotsa, a large rodent, which he described as Hypogeomys boulei. Lamberton identified this femur as Plesiorycteropus and MacPhee concurred. Remains of both Majoria rapeto and Hypogeomys boulei fall at the upper end of the size range of the genus, indicating that they are referable to P. madagascariensis. Another Plesiorycteropus innominate was mistakenly assigned to Daubentonia robusta, the extinct giant aye-aye, and other material has been misidentified as of a dwarf lemur.
Relationships
Filhol had classified Plesiorycteropus as close to the aardvark on the basis of morphological similarities. In his 1946 review, Charles Lamberton was unable to provide a definitive allocation, confused by the various similarities he saw with aardvarks, pangolins, armadillos, and anteaters. He believed it was most likely a primitive, isolated member of "Edentata", a group in which he included aardvarks, pangolins, and Xenarthra. He rejected some alternatives, such as a close affinity to aardvarks or the possibility that the material assigned to Plesiorycteropus did not in fact represent a single animal. Bryan Patterson, who revised tubulidentates in the 1970s, accepted Plesiorycteropus as a member of the group, dismissing many similarities with pangolins and other animals as convergent. However, he placed it as the only member of its own subfamily Plesiorycteropodinae in view of its differences from other tubulidentates, and hypothesized that it arrived on Madagascar in the Eocene, at the same time as the lemurs. Johannes Thewissen, who critiqued some aspects of Patterson's classification in 1985, also accepted Plesiorycteropus as a tubulidentate without comment.Reviewing Patterson's and Thewissen's contributions in 1994, Ross MacPhee found little support for the classification of Plesiorycteropus as a tubulidentate in their data. MacPhee used a cladistic analysis of eutherians to ascertain the relationships of the genus, but found that while different analytic variants supported different affinities—with aardvarks, hyraxes, ungulates, and even lipotyphlans —there was no compelling evidence linking it to any other eutherian group. Therefore, he erected a separate order for Plesiorycteropus, named Bibymalagasia, arguing that it would be unacceptable to leave a Recent eutherian unassigned to any order and that discovery of more material, or further analysis, was unlikely to demonstrate close affinities of Plesiorycteropus with any other order. He considered it possible but unlikely that a few fossil taxa, such as Palaeorycteropus and Leptomanis from the Paleogene of France, would eventually be found to be bibymalagasians. Various analyses published by Robert Asher and colleagues in 2003, 2005, and 2007, based on morphology combined with DNA sequence data in some analyses, produced different estimates of the relationships of Plesiorycteropus, some placing it within Afrotheria close to aardvarks or Afrosoricida, but others supporting a relationship with the hedgehog Erinaceus. A 2004 morphological study by Inés Horovitz, focusing on extinct South American ungulates, placed Plesiorycteropus among tubulidentates and closer to the extinct aardvark relative Myorycteropus than to Orycteropus.
A 2013 study by Michael Buckley examined preserved collagen sequences in Plesiorycteropus bones. He found the animal was most closely related to the tenrecs, a family of insectivorous afrotherian mammals endemic to Madagascar. Tenrecs are believed to have diversified from a common ancestor that lived 29–37 million years ago after dispersing from Africa via a single rafting event. Buckley's analysis showed that Plesiorycteropus and the two members of subfamily Tenrecinae tested formed a monophyletic group, within a larger clade in which golden moles are the sister group; he suggested that Plesiorycteropus should be placed in the order Tenrecoidea along with tenrecs as well as African otter shrews and golden moles. He did not test members of the other two Tenrecidae subfamilies or otter shrews, leaving open the possibility that Plesiorycteropus nests within Tenrecidae.
Common names
"Madagascar aardvark" has been used as a common name for Plesiorycteropus, but MacPhee considered it inappropriate because the animal may not be related to aardvarks. Instead, he proposed "bibymalagasy" as a common name, a manufactured Malagasy word meaning "Malagasy animal".Description
Plesiorycteropus is known from a number of subfossil bones, comparable to coverage of some of the poorly known subfossil lemurs, such as Daubentonia robusta. The material includes several skulls, all of which are missing the facial bones, complete long bones such as the femur and humerus, and other bones, but some elements are still unknown, including most of the skeleton of the hand and foot. There is little reason to assume it was similar in general form to the aardvark. No teeth or jaws referable to Plesiorycteropus have been found, and it is generally assumed that the animal was toothless.Based on the area of a femur cross-section, MacPhee calculated estimates of body mass. The lowest estimate, based on comparative data from armadillos and pangolins, was for the smallest femur he had and the highest estimate, based on comparative data from caviomorph rodents, was for the largest available femur ; estimates from primates fell between those extremes. MacPhee favored the lower estimates, because those were based on armadillos, which have femora similar to those of Plesiorycteropus. On the other hand, the caviomorph model produced a better estimate of brain size in Plesiorycteropus. Any of the estimates makes it considerably larger than the largest living tenrec, Tenrec ecaudatus, at up to. The higher estimates would make it larger than any extant native Malagasy mammal. This is consistent with the trend for larger members of the late Pleistocene and Holocene faunas of Madagascar and elsewhere to have been at higher risk of extinction.
Skull
There are four known skulls, each of which is damaged. All are missing the front part, and three are broken at about the same place, suggesting that the front part of the skull was thinner and more fragile than the back part, which consists of thick bones. MacPhee estimated maximum skull length in P. madagascariensis at. The length of the frontal bone averages in P. madagascariensis and is in P. germainepetterae.The robust nasal bones, preserved in a single specimen, are widest at the front, a feature unusual among placentals that is also seen in armadillos, and are also unusually flat. The ethmoid labyrinth, in the nasal cavity, was large, suggesting that Plesiorycteropus had a good sense of smell. A much larger part of the nasal septum, which separates the left and right nasal cavities, is ossified than usual in other mammals; MacPhee could find a similar condition only in sloths, which have a very short nose. The lacrimal bone is relatively large. At it is a single lacrimal canal, which opens near the suture between the frontal and lacrimal bones, like in lipotyphlans. There is a small tubercle near this opening. The orbital cavity, which houses the eyes, is relatively short, similar to the situation in pangolins and armadillos. A distinct tubercle is present on the suture between the frontal and parietal bones in P. germainepetterae, but not P. madagascariensis. P. madagascariensis has a more expansive braincase and a less pronounced narrowing between the orbits. The foramen rotundum, an opening in the bone of the orbit, is present. The optic canal, which houses the nerves leading to the eyes, is narrow, suggesting that the eyes were small, similar to many other tenrecoids. As in pangolins and xenarthrans, little of the squamosal bone can be seen from above. The temporal lines on the braincase, which anchor muscles, are located lower in P. germainepetterae. Like in aardvarks, the parietals are relatively large. An interparietal bone is present. Unlike in anteaters and pangolins, the occiput is flat and vertical. Plesiorycteropus lacks notches above the foramen magnum, which are present in aardvarks. The nuchal crest, a projection on the occiput, is straight in P. madagascariensis, but in P. germainepetterae it is interrupted in the middle, similar to the situation in armadillos and hyraxes.
In their descriptions of Plesiorycteropus, Lamberton and Patterson posited different interpretations of the location of the mandibular fossa, where the mandible articulates with the cranium. MacPhee found problems with either interpretation and suggested that the true mandibular fossa was part of the area Lamberton identified as such, at the side of the braincase. The fossa is small and low, suggesting that the animal was not capable of powerful biting. At the back of this fossa is a pseudoglenoid proces, which is more prominent in P. germainepetterae. In P. germainepetterae but not P. madagascariensis, a small opening, perhaps the vascular foramen, is present next to the foramen ovale. The petrosal bone forms a relatively large portion of the roof of the tympanic cavity, which houses the middle ear; parts of the petrosal are more developed in P. madagascariensis. Endocasts indicate that the neopallium part of the brain was relatively small.