Frank L’Engle Williams

RECONSTRUCTING THE PALEOECOLOGY OF TAUNG, SOUTH AFRICA FROM LOW MAGNIFICATION OF DENTAL MICROWEAR FEATURES IN FOSSIL PRIMATES

Abstract Taung, South Africa yielded the first Pliocene Hominini fossil, Australopithecus africanus, recovered from a lime quarry in 1924. To identify whether the habitat of the site differed from present-day conditions, dietary preferences of fossil papionins from Taung, including Parapapio antiquus (n  =  8), Papio izodi (n  =  12), and indeterminate specimens (n  =  10) were examined under low magnification to discern patterns of dental microwear. The comparative fossil sample from Sterkfontein Member 4 includes Parapapio broomi (n  =  10) and Parapapio jonesi (n  =  5). Extant Papio ursinus (n  =  20), a savanna-dwelling baboon from South Africa, provides a modern analogue. Six dental use-wear scars on the paracone of the second molar (M2) were recorded and the data analyzed using ANOVA with Tukeys test to detect whether group differences were present for each feature; linear regression identified significant covariation of microwear features. Principal components analysis and discriminant function analysis were utilized to identify species-specific dietary signals. Extant Papio ursinus is separated from the extinct taxa solely by a relatively greater number of fine scratches with respect to the other microwear features. Papio izodi overlaps primarily with extant Papio and secondarily with Parapapio, which forms a more discrete grouping that includes Parapapio antiquus from Taung. A wetter, more closed environment is suggested for Taung and Sterkfontein Member 4 compared to the habitat of present-day central South Africa.

EVIDENCE OF TERRESTRIAL DIETS IN PLIOCENE EURASIAN PAPIONINS (MAMMALIA: PRIMATES) INFERRED FROM LOW-MAGNIFICATION STEREOMICROSCOPY OF MOLAR ENAMEL USE-WEAR SCARS

ABSTRACT Pliocene papionin monkeys of Eurasia, such as Paradolichopithecus arvernensis, Procynocephalus subhimalayanus, and Macaca sp. indet. from Yushe, are large-bodied and interpreted as at least partly terrestrial. Terrestriality in primates has implications for the dietary items consumed. Dietary proclivities from the evaluation of pit and scratch counts under low-magnification (35×) were inferred to address whether the diets of these Eurasian papionins differ and whether they correspond to extant primates (Papio ursinus, n  =  24, Pan troglodytes, n  =  9, and Gorilla gorilla, n  =  10), three Plio-Pleistocene southern African fossil papionins with known isotopic values (Parapapio broomi, n  =  14; Papio robinsoni, n  =  10; Theropithecus oswaldi danieli, n  =  8), or a Pliocene bipedal hominin (Australopithecus africanus, n  =  8). Macaca sp. indet. from Yushe most closely resembles Australopithecus africanus from South Africa, which is characterized by evidence of some hard-object feeding. Paradolichopithecus from Romania and to a lesser extent Procynocephalus from northern India also exhibit use-wear scars associated with the ingestion of grit possibly associated with the exploitation of corms and bulbs of CAM or C4 plants. The dental microwear signal of grassland foraging typical of such C4 grazers as Theropithecus oswaldi danieli does not characterize other Plio-Pleistocene papionins. Pliocene papionin diets converge with those of Australopithecus, suggesting that global climate cooling intensifying during the Pliocene may have led both early baboon-like monkeys and australopiths to increase consumption of nutrient-rich edible plant storage organs in these fragmented and partially open habitats of the temperate Old World.

Ontogeny and Phylogeny of the Pelvis in Gorilla, Pongo, Pan, Australopithecus and Homo

To examine the evolutionary differences between hominoid locomotor systems, a number of observations concerning the growth of the pelvis among the great apes as compared to modern and fossil hominids are reported. We are interested in the size and shape of the coxal bones at different developmental stages across species that may elucidate the relationship between ontogeny and phylogeny (i.e., heterochrony) in the hominoid pelvis. Our hypotheses are: (1) do rates of absolute growth differ?, (2) do rates of relative growth differ?, and (3) does heterochrony explain these differences? Bivariate and multivariate analyses of pelvic dimensions demonstrate both the diversity of species-specific ontogenetic patterns among hominoids, and an unequivocal separation of hominids and the great apes. Heterochrony alone fails to account for the ontogenetic differences between hominids and the great apes. Compared to recent Homo,Australopithecus can be described as ‘hyper-human’ from the relative size of the ischium, and short but broad ilium. Australopithecus afarensis differs from Australopithecus africanus by its relatively long pubis. In multivariate analyses of ilium shape, the most complete coxal bone attributed to Homo erectus, KNM-ER 3228, falls within the range of juvenile and adult Australopithecus, whereas Broken Hill falls within the range of modern Homo, suggesting that the modern human ilium shape arose rather recently. Among the great apes, patterns of pelvic ontogeny do not exclusively separate the African apes from Pongo.

RECONSTRUCTING THE DIET AND PALEOECOLOGY OF PLIO-PLEISTOCENE CERCOPITHECOIDES WILLIAMSI FROM STERKFONTEIN, SOUTH AFRICA

ABSTRACT The Plio-Pleistocene cave site of Sterkfontein has yielded the remains of Cercopithecoides williamsi, a colobine monkey reconstructed as a terrestrial folivore given its dental morphology, heavily worn teeth and robustly built postcranial skeleton. To address the validity of these dietary and habitat inferences, patterns of dental microwear obtained from low-magnification stereomicroscopy for C. williamsi specimens from Sterkfontein Member 4 (n  =  13) are compared to a broad comparative sample of extant primates, including Cebus apella (n  =  10), Colobus polykomos, (n  =  6), Papio anubis (n  =  13), and Papio ursinus (n  =  39) as well as extinct Parapapio broomi (n  =  12) from Sterkfontein Member 4 and Theropithecus oswaldi danieli (n  =  6) from Swartkrans Member 1. Cercopithecoides williamsi exhibits numerous large pits and puncture pits, and few fine scratches. When diet is considered, C. williamsi is closely associated with C. polykomos and C. apella, suggesting folivorous resources and some hard-objects were consumed, possibly from seeds or grit on fallen fruit, but more likely from grit adhering to the underground storage organs of C4 plants given its mixed C3/C4 isotopic signal. This fossil taxon groups with terrestrial primates more than arboreal ones when habitat is considered. Individuals attributed to C. williamsi largely cluster as a group and a comparison of multiple use-wear scars differentiates open or partially open habitats from forested ones, suggesting dental microwear provides a strong habitat signal.

Quantifying Mental Foramen Position in Extant Hominoids and Australopithecus: Implications for its Use in Studies of Human Evolution

The location of the mental foramen on the mandibular corpus has figured prominently in debates concerning the taxonomy of fossil hominins and Gorilla gorilla. In this study we quantify the antero/posterior (A/P) position of the mental foramen across great apes, modern humans and Australopithecus. Contrary to most qualitative assessments, we find significant differences between some extant hominoid species in mental foramen A/P position supporting its potential usefulness as a character for taxonomic and phylogenetic analyses of fossil hominoids. Gorilla gorilla, particularly the eastern subspecies, with a comparatively longer dental arcade and fossil and extant hominins with reduced canines and incisors tend to exhibit more anteriorly positioned mental foramina. Conversely, Pan troglodytes exhibits more posteriorly positioned mental foramina. Variation in this character among Gorilla gorilla subspecies supports recent taxonomic assessments that separate eastern and western populations. In all taxa other than Pan troglodytes the A/P position of the mental foramen is positively allometric with respect to dental arcade length. Thus, within each of these species, specimens with longer dental arcades tend to have more posteriorly positioned mental foramina. Those species with greater sexual dimorphism in canine size and dental arcade length (i.e., Gorilla gorilla and Pongo pygmaeus) exhibit more extreme differences between smaller and larger individuals. Moreover, among great apes those individuals with greater anterior convergence of the dental arcade tend to exhibit more posteriorly positioned mental foramina. Dental arcade length, canine crown area and anterior convergence are all significantly associated with mental foramen A/P position, suggesting that these traits may influence taxonomic variation in the A/P position of the mental foramen. Anat Rec 293:1337–1349, 2010.

Dental microwear texture analysis of late Pliocene Procynocephalus subhimalayanus (Primates: Cercopithecidae) of the Upper Siwaliks, India

Late Pliocene Procynocephalus subhimalayanus from the Upper Siwaliks, India is known from only three specimens. The dietary proclivities of this taxon have implications for reconstructing the paleoecology of the Upper Siwaliks. The dental microwear texture properties of Procynocephalus are compared to those from extant tropical forest primates including Alouatta palliata (n = 11), Cebus apella (n = 13), Gorilla gorilla (n = 9), Lophocebus albigena (n = 15) and Trachypithecus cristatus (n = 12). Dental microwear textures are generated by scanning the surface enamel of Facet 9 using white-light confocal microscopy at 100x. Four variables were extracted from scale-sensitive fractal analysis, and the data were ranked before ANOVA with post-hoc tests of significance and multivariate analyses were performed. Procynocephalus clusters closest to Lophocebus, Cebus and some Gorilla specimens suggesting hard-object feeding characterized a portion of its diet. The dental microwear texture of Procynocephalus supports interpretations of widespread grasslands of the Late Pliocene Kansal Formation (Pinjor zone). The extreme enamel complexity characterizing Procynocephalus may derive from consumption of underground storage organs, or other foods with high grit loads. Foods consumed near ground level carry a heavy load of abrasive minerals possibly contributing to greater enamel surface complexity and textural fill volume.

The mid-face of lower Pleistocene hominins and its bearing on the attribution of SK 847 and StW 53

SK 847 and StW 53 have often been cited as evidence for early Homo in South Africa. To examine whether midfacial morphology is in agreement with these attributions, we analyze Euclidean distances calculated from 3-D coordinates on the maxillae of SK 847 and StW 53, as well as Australopithecus africanus (Sts 5, Sts 71), Paranthropus robustus (SK 46, SK 48, SK 52, SK 83), early Homo (KNM-ER 1813, KNM-ER 1805, KNM-ER 3733, KNM-WT 15000), P. boisei (KNM-ER 406, KNM-WT 17000, KNM-WT 17400), Gorilla gorilla (n=116), Homo sapiens (n=342), Pan paniscus (n=21) and P. troglodytes (n=65). Multivariate analyses separate extant hominoids suggesting we have captured taxonomic affinity. With the exception of SK 847 and SK 52, South African fossils tend to cluster together. P. robustus differs substantially from East African robust megadonts. SK 847 and StW 53 resemble the East African Homo specimens that are the most australopith-like, such as KNM-WT 15000 and KNM-ER 1813. The resemblance between StW 53 and Homo is driven partly by similarities in maxillary size. When distances are scaled, StW 53 aligns with A. africanus, while SK 847 clusters primarily with early Homo.

A secondary mandibular condylar articulation and collateral effects on a Late Neolithic mandible from Bois Madame rockshelter in Arbre, Belgium

A Neolithic Belgian mandible from Bois Madame rockshelter in Arbre presents an asymmetrical morphology resulting from a secondary, or false, articulation of the right mandibular condyle. The pathological articulation produced enlarged masseter, medial pterygoid and mylohyoid musculature on the right side as well as a flattening of the right incisal alveolus curvature. The secondary condylar articulation did not lead to pronounced asymmetry of attrition on the antimeres of the dental arcade. This is the most complete mandible from this Late Neolithic collective burial dating to the beginning of the Bronze Age circa 4000 years BP. It is possible that a fall or blow to the mental symphysis during early adolescence could have resulted in the partial intrusion of the mandibular condyle into the articulation disc of the temporomandibular joint capsule. When the affected condyle healed, a secondary, but serviceable articulation developed, producing unique stresses on the involved muscular tissue and ultimately resulted in an asymmetry of mandibular form.

Dietary reconstruction of Pliocene Parapapio whitei from Makapansgat, South Africa, using dental microwear texture analysis.

Pliocene Parapapio whitei fossils from Makapansgat have yielded stable isotope values suggesting some foraging on C4 plants and possibly underground storage organs (USOs). Dental microwear texture analysis on Pp.whitei (M 3147, MP 62, MP 223 and MP 239) from Makapansgat Members 3-4 is performed to examine whether tooth surface damage from mastication agrees with prior dietary inferences from isotopes. The enamel surface texture of Pp. whitei is relatively complex, resembling Lophocebus albigena and Cebusapella, and lacks the anisotropy of Trachypithecuscristatus and Alouattapalliata. The textural fill volume of Pp. whitei is distinct from extant forest primates suggesting extremes in hard-object consumption. Grit adhering to USOs is offered as an explanation for these enamel textural properties, corroborating the inference that Pp. whitei supplemented its diet with terrestrial resources.

Expression of Nonmetric Dental Traits in Western European Neanderthals

Specific dental abnormalities are considered pathognomonic of congenital syphilis (CS); however, European physicians recognized their variation during the late 19th to mid 20th centuries. Observations of syphilis-related dental abnormalities in American individuals from similar time periods are made to determine types of variation among the American population. From a survey of the Smithsonian Institution’s National Museum of Natural History anatomical human skeletal collection, five individuals demonstrated dental characteristics consistent with CS (P00011R, P219398, P000707, P000679, and P000161). Hutchinson’s three categories of dental anomalies were used to describe variations among syphilitic individuals. Previously identified pathological dental characteristics related to CS were present in the analyzed individuals. P00011R, 24-year-old Black female, has a maxillary right Moon’s molar. P219398, approximately 20-year-old Black female, has Hutchinson’s incisors and Fournier’s molars. P000707, 26-year-old Black male, displays severe hypoplasia on all incisors, canines and maxillary first molars. P000679, 33-year-old Black female has “screwdriver” shaped maxillary central incisors, altered occlusal morphology of first maxillary molars and hypoplasia. P000161, 45-year-old Black female, demonstrates severe hypoplasia on incisors and canines (molars lost). “Classic” dental characteristics of CS are not ubiquitous to all identified cases. This study exemplifies that dental anomalies associated with CS do not all have to be present for diagnosis. Although other causes for some of these anomalies are possible, observations in these five cases are most consistent with CS. Five Cases of Dental Anomalies Attributable to Congenital Syphilis from Early 20 Century American Anatomical Collections Stella Ioannou*, David Hunt, and Maciej Henneberg 1 Biological Anthropology and Comparative Anatomy Research Unit, Adelaide Medical School, The University of Adelaide, Adelaide, South Australia, 5005 2 Department of Anthropology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013-7012 Prior to the introduction of penicillin in the 1940s, syphilis was a public health problem in the United States (Lancet, 1930; Lancet, 1937a). The prevalence of syphilis in the United States at that time is difficult to determine, as data collection for syphilis by state health departments did not begin until the early 20th Century, and the Venereal Disease Division of the U.S. Public Health Service was not created until 1918 (Nakashima et al., 1996). To control venereal disease, various states implemented programs (free treatment, and clinics that offered free, pay, and part pay clinics) (Lancet, 1937a), and legislation (marital examination law and prenatal law) (Lancet, 1917; Prebble, 1938; Lancet, 1940; DePorte, 1941). In cases of medical intervention, mercury was used to treat congenital syphilis in the 19th and early 20th centuries throughout the United States (Conrad and McCann, 1922; Cole et al., 1929; Scheer and Fraser, 1930; Cole et al., 1933; Chargin and Saunders, 1939). Treatments of syphilis also included chemotherapies of arsenic and bismuth compounds (Lee, 1878; Cole et al., 1929; Eller and Maloney, 1929). The chemotherapies most favored in the treatment of congenital syphilis included mercury, arsphenamine and potassium iodide (United States. Public Health Service. Division of Venereal Diseases, 1930). The effectiveness of mercury as a treatment for syphilis has been questioned (Miller, 1858: Weatherill, 1833); although, the treatment remained popular with some physicians. In some cases, syphilitic lesions completely healed and patients became seronegative (Wakerlin, 1934). In syphilitic women treated with mercury during their pregnancy, 91.5% were efficient in completing their pregnancies successfully by live birth, while 47.6% non-treated women experienced fetal death (United States. Public Health Service. Division of Venereal Diseases, 1930). Mercury and its compounds were seen to *Correspondence to: Stella Ioannou The University of Adelaide Adelaide 5005, South Australia, Australia email: stelzy_25@hotmail.com