Dental eruption sequence of a fossil 'baboon' (Paradolichopithecus arvernensis)

The fossil remains of this large-sized 'baboon' found at Vatera, Lesvos, Greece (Plio-Pleistocene boundary, or Late Pleistocene now that the boundary has been moved up again) includes a mandible of a young male with unerupted wisdom molar. We X-rayed the specimen to reveal the unerupted elements and the mineralization of their crowns. The chronological, or individual age of our Paradolichopithecus at the time of its untimely death is estimated at between 5.0 and 5.3 years of age, based on eruption patterns and root formation times of similar-sized living papionins. Mandrills, yellow baboons and Japanese monkeys are in their puberty at this age and stage of dental development, from which we conclude that our male was in his puberty as well. During this period, young males disperse from their natal groups and live at the periphery of their troupes. Risks of predation, disease, and injury are higher than before; a quarter of male mandrills dies before reaching adulthood. Among known Paradolichopithecus specimens, nearly half have died before the third molars erupted. This may be explained either by some taphonomic factor or due to higher mortality levels during puberty.
We further found that the eruption sequence of the permanent mandibular dentition of this male is {m1 i1–2 m2} p4, p3, c, m3. The order of the already fully erupted elements (between curly brackets) is based on data from the living baboons, mandrills, macaques and geladas. The p4 p3 sequence as seen in our Paradolichopithecus occurs at high frequency is other papionins as well: Macaca nemestrina, M. mulatta, Mandrillus sphinx, Papio cynocephalus, but not in Papio anubis, Macaca fuscata and M. fascicularis. Theearlier root formation of p4 seems thus not to be related with body size or phylogeny. There is a considerable delay in the canine development relative to the premolars, as in other papioninmales; the m3 is delayed in formation relative to the premolars and the canine. In total, the dental eruption sequence of Paradolichopithecus is very similar to that of the living papionins.

Read more in Van der Geer, A.A.E., Dermitzakis, M. 2008. Dental eruption sequence in the Pliocene Papionini Paradolichopithecus arvernensis (Mammalia: Primates) from Greece. Journal of Vertebrate Paleontology 28 (4): 1238-1244. (Ask a pfd, geeraae@geol.uoa.gr)

Locomotor behavior of Paradolichopithecus arvernensis as inferred from the functional morphology of its ankle and elbow

Taking all ankle and elbow elements of Paradolichopithecus into account, the picture emerges of a highly terrestrial monkey. This is not surprising as many fossil cercopithecines are found in open country habitats and show terrestrial adaptations, such as Dinopithecus (Late Pliocene, Africa), Procynocephalus (Late Pliocene, China and India), Paradolichopithecus (Pliocene, Spain and Asia), Theropithecus (Middle Pleistocene - Holocene, Africa) and, among the colobines, Paracolobus (Pliocene, East Africa) and Dolichopithecus (Pliocene, Europa) (Szalay & Delson, 1979). In addition, the larger species tend to be terrestrial, possibly as a response to predator pressure. This, too, makes a terrestrial adaptation of our large Paradolichopithecus very probable.
Body weight was carried more posterior, as the architecture of the olecranon and the trochlear notch are less apted for sustaining heavy load than is the case in the extant baboons. The morphology of the arm indicates an increased mobility in the elbow joint, with a departure from the sagittal plane during flexion. Paradolichopithecus could very well have used his strong arms for carrying food while walking or standing. Another option is the use of the arms in fights and defense.
The massive medial malleolus of the tibia also shows that a larger (part of the) body weight was carried on the hindlimbs. The suspensory facet for the fibular malleolus indicates an increased importance of the lateral malleolus in transferring body weight, and an increased fixation of the talus in the malleolar fork, formed by both the malleoli together.
As to the ankle joint, a remarkable parallel is seen with Australopithecus. Unique features that distinguish Paradolichopithecus, and probably also Procynocephalus, from the other papionins are seen also in Australopithecus, though the overall architecture of the Paradolichopithecus talus is typically cercopithecoid (pronounced lateral trochlear ridge, hardly developed groove for large toe flexor), whereas it is typically hominoid for Australopithecus (symmetrical trochlea, pronounced large toe flexor).The terrestrial traits in the postcranial elements show that this large monkey was clearly adapted to the habitat: an open savanna/bushland environment with seasonal availability of food, and large distances between the food sources.

Read more in SONDAAR P.Y., VAN DER GEER A.A.E., DERMITZAKIS M. (2006). The unique postcranial of the Old World monkey Paradolichopithecus: more similar to Australopithecus than to baboons. Hellenic Journal of Geosciences 41, 1: 19-28. Special volume in the memory of P.Y. Sondaar

and in VAN DER GEER A.A.E., SONDAAR P.Y. (2002). The postcranial elements of Paradolichopithecus arvernensis (Primates, Cercopithecidae, Papionini) from Lesvos, Greece. Annales Géologiques des Pays Helléniques 1e Série 39, A: 71-86. Free pdf at http://users.uoa.gr/~geeraae/publications/2002-agph-Paradolichopithecus.pdf .

and in SONDAAR P.Y., VAN DER GEER A.A.E. (2002). Arboreal and terrestrial traits as revealed by the primate ankle joint. Annales Géologiques des Pays Helléniques 1e Série 39, A: 87-98. Free pdf at http://users.uoa.gr/~geeraae/publications/2002-agph-terrestriality.pdf .

The unique postcranial of the extinct Old World monkey Paradolichopithecus

The talus (astragalus), distal tibia and the humerus of Paradolichopithecus arvernensis show some unique features, not seen in other monkeys.
The humerus has an increased articulation area on the head compared to Papio, a wide and deep groove for the biceps tendon, a gradually descending capitulum, and an oblique axis for flexion-extension through the elbow joint. During flexion, the ulna deviates from the parasagittal plane, and ends in a position medially to the humerus instead of parallel above it, due to the trochlear shape and axis. This unique feature yields a significant increased mobility.
The distal tibia bears a more massive, square and blunt malleolus that lacks the typically pronounced ball-shaped area, a wider groove (sulcus malleolaris) for the tendon of the M. tibialis posterior, a more square cross-section, clear scars for the fibula, and a double tendon groove on the dorsal surface (either for a bifurcated tendon for the M. flexorum tibialis posterior or a pronounced groove for the long toe flexor), which follows the parasagittal plane. None of these features is unique, and they make Paradolichopithecus resemble Australopithecus, a trained Japanese macaque and to a lesser extent some other macaques. The combination indicates a maintainance of the close-packed situation from dorsiflexion to plantar flexion, an increased importance of the fibula in weight transfer, a stronger plantar flexion, and possibly a slightly abducted foot. The flat tibial malleolus in Paradolichopithecus and Australopithecus, compared to baboons (Papio) and chimps (Pan) respectively, in combination with the corresponding facet on the talus acts as a blocking mechanism, preventing further dorsiflexion rotation during maximal dorsiflexion. This makes this ankle unsuitable for climbing.
The talus has an almost parallel trochlea, a large flap-like, protruding fibular suspensory facet, and a slightly deeper facet for the spring ligament on the talar head. These features are suggestive for a baboon-like ankle joint with the body weight more evenly distributed over the talar trochlea, a greater proportion of the weight transfer through the lateral (fibular) side, and with approximate the same stability in maximal dorsiflexion as in maximal plantar flexion. In these aspects Paradolichopithecus resembles Australopithecus.
Considering the unique features of the ankle and elbow of Paradolichopithecus, it may be expected that its locomotion differed from that of baboons. Main differences are the increased fibular component, the increased stability in plantar flexion, a more evenly distribution of stability during locomotion, and an equal medio-lateral stability in maximal plantiflexion and in maximal dorsiflexion. In our view, such a type of locomotion finds a parallel in Australopithecus and in trained Japanese macaques. The latter appear to develop significant modifications during training, especially in the hind limb, to satisfy the functional requirements for increased habitual bipedalism. Amongst others, the malleolus of the tibia has been remodeled under the influence of the greater stress and became less cusp-shaped, and the talar malleolar facet correspondingly more planar. The varus knee in the trained macaque further requires an increased fibular compound. This may have its parallel in Paradolichopithecus and Australopithecus, in whom we also find an increased fibular component. It should be stressed, however, that the kind of bipedalism of the trained macaque differs essentially from the striding gait bipedalism with erect trunk and straight knees of the genus Homo. The macaque bipedalism is characterised by high energy cost and bent knees. Considering the similar biomechanical features in Paradolichopithecus, Australopithecus and the trained macaque, it is tempting to conclude that also the two former genera had an all-round, energetically expensive bipedal mode with bent knees. This development then was not restricted to the hominoid clade, but appeared also in the papionins, as evidenced by the difference between Australopithecus and Pan on one hand and Paradolichopithecus and Papio on the other hand. The pattern shared indicates similar mechanical stresses, and reflects a shared increased frequency of bipedalism in the daily locomotor behavior, possibly but not necessarily, accompagnied by an increased mobility of the arm.

Read more in SONDAAR P.Y., VAN DER GEER A.A.E., DERMITZAKIS M.D. (2006). The unique postcranial of the Old World monkey Paradolichopithecus: more similar to Australopithecus than to baboons. Hellenic Journal of Geosciences 41, 1: 19-28. Special volume in the memory of Paul Yves Sondaar. Free pdf [868 kb] at http://users.uoa.gr/~geeraae/publications/2006-HJG-Paradolichopithecus.