Molar crown formation times of fossil orangutan molars from Guangxi, China.

OBJECTIVES: We aimed to investigate molar enamel development in fossil orangutans from Guangxi and shed light on the evolution of Asian great apes. MATERIALS AND METHODS: We collected 32 fossil orangutan molars, most of which were from Guangxi apothecaries and the Guangxi Daxin Heidong cave, and prepared histological sections of each molar. We then characterized aspects of dental development, including long period line periodicity, number of Retzius lines and lateral enamel formation time, cuspal enamel thickness, and enamel formation time. RESULTS: The long period line periodicity in fossil orangutans ranged from 9 to 10 days (mean, 9.09 days). The molar lateral enamel formation time ranged from 1.48 to 3.17 years (540-1,152 days). Cuspal enamel thickness in fossil orangutan molars ranged from 949 to 2,535 µm, and cuspal enamel formation time ranged from 0.64 to 1.87 years. Molar enamel formation time of fossil orangutans ranged from 2.47 to 4.67 years. DISCUSSION: Long-period line periodicity of fossil orangutans from Guangxi was within the variation range of extant orangutans, and the average long period line periodicity (9.09 days) of fossil orangutans from Guangxi in this study was lower than the values for extant orangutans (9.5 days) and fossil orangutans (10.9 days) from Sumatra and Vietnam. Orangutan enamel thickness may have gradually decreased from the Middle Pleistocene to Holocene. Crown formation time of fossil orangutans was slightly longer than that of extant orangutans, and the M1 emergence age of fossil orangutans from Guangxi was about 4-6 years. These findings might indicate the regional difference or evolutionary changes in orangutans since Pleistocene. Dental development of the Guangxi fossil orangutans were more similar to that of Asian Miocene apes, suggesting the closer evolutionary relationship of orangutans to Miocene Asian fossil apes.

Dental development in living and fossil orangutans.

Numerous studies have investigated molar development in extant and fossil hominoids, yet relatively little is known about orangutans, the only great ape with an extensive fossil record. This study characterizes aspects of dental development, including cuspal enamel daily secretion rate, long-period line periodicities, cusp-specific molar crown formation times and extension rates, and initiation and completion ages in living and fossil orangutan postcanine teeth. Daily secretion rate and periodicities in living orangutans are similar to previous reports, while crown formation times often exceed published values, although direct comparisons are limited. One wild Bornean individual died at 4.5 years of age with fully erupted first molars (M1s), while a captive individual and a wild Sumatran individual likely erupted their M1s around five or six years of age. These data underscore the need for additional samples of orangutans of known sex, species, and developmental environment to explore potential sources of variation in molar emergence and their relationship to life history variables. Fossil orangutans possess larger crowns than living orangutans, show similarities in periodicities, and have faster daily secretion rate, longer crown formation times, and slower extension rates. Molar crown formation times exceed reported values for other fossil apes, including Gigantopithecus blacki. When compared to African apes, both living and fossil orangutans show greater cuspal enamel thickness values and periodicities, resulting in longer crown formation times and slower extension rates. Several of these variables are similar to modern humans, representing examples of convergent evolution. Molar crown formation does not appear to be equivalent among extant great apes or consistent within living and fossil members of Pongo or Homo.

Low testosterone correlates with delayed development in male orangutans.

Male orangutans (Pongo spp.) display an unusual characteristic for mammals in that some adult males advance quickly to full secondary sexual development while others can remain in an adolescent-like form for a decade or more past the age of sexual maturity. Remarkably little is understood about how and why differences in developmental timing occur. While fully-developed males are known to produce higher androgen levels than arrested males, the longer-term role of steroid hormones in male life history variation has not been examined. We examined variation in testosterone and cortisol production among 18 fully-developed ("flanged") male orangutans in U.S. captive facilities. Our study revealed that while testosterone levels did not vary significantly according to current age, housing condition, and species origin, males that had undergone precocious development had higher testosterone levels than males that had experienced developmental arrest. While androgen variation had previously been viewed as a state-dependent characteristic of male developmental status, our study reveals that differences in the physiology of early and late developing males are detectable long past the developmental transition and may instead be trait-level characteristics associated with a male's life history strategy. Further studies are needed to determine how early in life differences in testosterone levels emerge and what consequences this variation may have for male behavioral strategies.

Detailed comparative anatomy of the extrinsic cardiac nerve plexus and postnatal reorganization of the cardiac position and innervation in the great apes: orangutans, gorillas, and chimpanzees.

To speculate how the extrinsic cardiac nerve plexus (ECNP) evolves phyletically and ontogenetically within the primate lineage, we conducted a comparative anatomical study of the ECNP, including an imaging examination in the great apes using 20 sides from 11 bodies from three species and a range of postnatal stages from newborns to mature adults. Although the position of the middle cervical ganglion (MG) in the great apes tended to be relatively lower than that in humans, the morphology of the ECNP in adult great apes was almost consistent with that in adult humans but essentially different from that in the lesser apes or gibbons. Therefore, the well-argued anatomical question of when did the MG acquire communicating branches with the spinal cervical nerves and appear constantly in all sympathetic cardiac nerves during primate evolution is clearly considered to be after the great apes and gibbons split. Moreover, a horizontal four-chambered heart and a lifted cardiac apex with a relatively large volume in newborn great apes rapidly changed its position downward, as seen in humans during postnatal growth and was associated with a reduction in the hepatic volume by imaging diagnosis and gross anatomy. In addition, our observation using a range of postnatal stages exhibits that two sympathetic ganglia, the middle cervical and cervicothoracic ganglia, differed between the early and later postnatal stages.

Brief communication: Testing the usefulness of the basilar suture as a means to determine age in great ape skeletons.

A fused/closed basilar suture is usually treated as an indication of old age in great apes. A sample, drawn from a variety of sources, of known-aged captive great ape skeletons was analyzed to test the usefulness of using the basilar suture to categorize adult skeletons as either "adult" or "old adult". The state of closure of the basilar suture was examined in 30 chimpanzees, 19 gorillas, and 15 orangutans, all of known age. The results show that the basilar suture demonstrates a high level of uniformity in rate of closure and is closed at an early age in virtually all known-aged individuals. Thus, an old adult category most likely includes individuals who are, in fact, relatively young. This indicates that using the basilar suture as a means to classify individual skeletons as adult or old adult is very imprecise. The homogenous nature of basilar suture closure appears to prevent meaningful application of suture status for categorizing adult ape skeletons by age groups.