No brain expansion in Australopithecus boisei

The endocranial volumes of robust australopithecine fossils appear to have increased in size over time. Most evidence with temporal resolution is concentrated in East African Australopithecus boisei. Including the KNM-WT 17000 cranium, this sample comprises 11 endocranial volume estimates ranging in date from 2.5 million to 1.4 million years ago. But the sample presents several difficulties to a test of trend, including substantial estimation error for some specimens and an unusually low variance. This study reevaluates the evidence, using randomization methods and a related test using an explicit model of variability. None of these tests applied to the A. boisei endocranial volume sample produces significant evidence for a trend in that species, whether or not the early KNM-WT 17000 specimen is included.     

Hominid cranium from Omo: Description and taxonomy of Omo-323-1976-896

Omo-323-1976-896, a partial hominid cranium dated to ca. 2.1 from the Member G, Unit G-8 of the Shungura Formation, lower Omo Basin of Ethiopia, is described. It is suggested that the specimen is an adult male based on the well-developed and completely fused sagittal crest; heavily worn teeth; relatively large canine; and size of the articular eminence. Omo-323 consists of fragments of the frontal, both temporals, occipital, parietals, and the right maxilla, and is attributed to Australopithecus boisei, making it the oldest known cranium of this species. The specimen shares features with Australopithecus aethiopicus (KNM-WT 17000), thus supporting the existence of an evolving East African robust lineage between ca. 2.6-1.2 Ma. The morphology of Omo-323 increases our knowledge of the intraspecific variability of A.boisei.     

The functional significance of the squamosal suture in Australopithecus boisei.

A juvenile Australopithecus boisei specimen from the Omo basin, southern Ethiopia, is found to exhibit and extraordinarily large overlap of the temporal squama on the parietal, a phenomenon shared with at least two adult specimens of A. boisei. An attempt is made to interpret the overlap as a structural (bony/ligamentous) adaptation necessitated by the unique combination of certain components of the masticatory system of A. boisei. These are: (1) the massiveness and strength of the temporalis muscle, (2) its relatively anterior location, and (3) the lateral position of the masseter muscle due to the flaring of the zygomatic arches. The effect of the temporalis muscle is to create excessive pressure on the portion of the squamosal suture along the parietal, while the lateral placement of the masseter and the resultant increase of pressure on the temporal squama via the zygomatic arch tend to "loosen" the contact between the temporal and parietal bones.     

Cranial variability in East African ‘Robust’ hominis

Among Plio-Pleistocene hominins the East African ‘robust’ group [Australopithecus (Paranthropus) boisei sensu lato] has the largest sample. This makes it an important test case for examining within-group variability and its implications for early hominin systematics. When using the CV to test for mixed-species samples, sexual dimorphism and diachronic variation are important additional (confounding) sources of variability. After examining nineteen variables, five craniometric variables in the East African ‘robust’ group are identified that have low sexual dimorphism. All are characterised by CVs from a combined KNM-WT 17000 andA. (P.) boisei sample less than the CV of the bonobo. Diachronic variation is found to be an important source of variability for cranial capacity. This form of variation cannot be detected in other variables studied here, but its possible presence cannot be ignored (owing mostly to small sample sizes). It is concluded that there is insufficient evidence to refute the hypothesis that all East African ‘robust’ fossils belong toA. (P.) boisei (Walker and Leakey, 1988).     

Temporal squama shape in fossil hominins: relationships to cranial shape and a determination of character polarity

In 1943, Weidenreich described the squamosal suture of Homo erectus as long, low, and simian in character and suggested that this morphology was dependent upon the correlation between the size of the calvarium and the face. Many researchers now consider this character to be diagnostic of H. erectus. The relationship between cranial size and shape and temporal squama morphology, however, is unclear, and several authors have called for detailed measurements of squamosal variation to be collected before any conclusions are drawn regarding the nature of the morphology observed in H. erectus. Thirteen fossil and extant taxa were examined to address two questions: 1) Are size and shape of the temporal squama correlated with cranial vault morphology? and 2) Is the H. erectus condition plesiomorphic? To answer these questions, measurements were collected and indices were calculated for squamosal suture height, length, and area in relation to metric variables describing cranial size and shape. A two‐dimensional morphometric study was also completed using High Resolution‐Polynomial Curve Fitting (HR‐PCF) to investigate correlations between curvature of the squamosal suture and curvature of the cranial vault. Results of both analyses indicate that squamosal suture form is related to cranial size and shape. Furthermore, the plesiomorphic condition of the squamosal suture for hominins was identified as high and moderately arched; this condition is retained in H. erectus and is distinct from the great ape condition. It is suggested that this similarity is the result of increased cranial length without a corresponding increase in cranial height. Am J Phys Anthropol, 2008. © 2008 Wiley‐Liss, Inc.     

Relationship of squamosal suture to asterion on external skull surfaces versus endocasts of pongids: Implications for Hadar early hominid AL 162-28

The relationship between the squamosal suture and asterion was quantified in 15 hemispheres of eight chimpanzee endocasts that were aligned in the conventional lateral view (i.e., with frontal pole [FP]–occipital pole [OP] horizontal). Using a three-dimensional digitizer, x, y, and z coordinates were collected for the highest and lowest points of the squamosal suture, and the most rostral point of the suture approximate to the coronal suture. Our results were compared to a similar study of the squamosal suture on the external surfaces of chimpanzee skulls that were oriented in the Frankfurt horizontal (Holloway and Shapiro, 1992). The relationship between the squamosal suture and asterion differs markedly between the outsides of skulls and endocasts. Whereas the squamosal suture is very rarely below asterion on the external skull, we found that most of the squamosal suture is located inferior to asterion on endocasts. We also found that the squamosal suture courses approximately 2.0 mm lower on the right side than the left. (An asymmetry of the same magnitude was reported for the external skull but, curiously, in the opposite direction.) It may be that a lowered right squamosal endosuture on chimpanzee endocasts is associated with earlier closure on that side. The discrepancy in results for the external skull versus endocast is partially attributable to orienting chimpanzee skulls in the Frankfurt horizontal, which usually results in the endocasts being tilted so that FP is above OP, i.e., FP-OP is not parallel with the Frankfurt horizontal. Falk's (1985) orientation of the early hominid endocast from Hadar (AL 162-28) is consistent with data determined from endocasts of chimpanzees. © 1994 Wiley-Liss, Inc.     

Functional implications of squamosal suture size in paranthropus boisei

It has been hypothesized that the extensively overlapping temporal and parietal bones of the squamosal sutures in Paranthropus boisei are adaptations for withstanding loads associated with feeding. Finite element analysis (FEA) was used to investigate the biomechanical effects of suture size (i.e., the area of overlap between the temporal and parietal bones) on stress, strain energy, and strain ratio in the squamosal sutures of Pan troglodytes and P. boisei (specimen OH 5) during biting. Finite element models (FEMs) of OH 5 and a P. troglodytes cranium were constructed from CT scans. These models contain sutures that approximate the actual suture sizes preserved in both crania. The FEM of Pan was then modified to create two additional FEMs with squamosal sutures that are 50% smaller and 25% larger than those in the original model. Comparisons among the models test the effect of suture size on the structural integrity of the squamosal suture as the temporal squama and parietal bone move relative to each other during simulated premolar biting. Results indicate that with increasing suture size there is a decreased risk of suture failure, and that maximum stress values in the OH 5 suture were favorable compared to values in the Pan model with the normal suture size. Strain ratios suggest that shear is an important strain regime in the squamosal suture. This study is consistent with the hypothesis that larger sutures help reduce the likelihood of suture failure under high biting loads. Am J Phys Anthropol 153:260–268, 2014. © 2013 Wiley Periodicals, Inc.     

Relationship of squamosal suture to asterion in pongids (Pan): relevance to early hominid brain evolution.

Based on 244 measurements of the relationship of the squamosal suture to the landmark asterion in 49 chimpanzee skulls, it is shown that in the normal lateral view the squamosal suture is very rarely inferior to asterion. In hominid crania, the squamosal suture is always well superior to asterion. Even in Pan, that part of the squamosal suture most homologous with the remnant found on the Hadar AL 162-28 Australopithecus afarensis hominid cranial fragment is very rarely inferior to asterion. Such variability suggests that Falk's (Nature 313:45-47, 1985) orientation of the Hadar specimen is incorrect; she places asterion superior to the position of the squamosal suture if projected endocranially. The implication for the brain endocast is that, however the fragment is oriented, the posterior aspect of the intraparietal (IP) sulcus is in a very posterior position relative to any chimpanzee brain. The distance from the posterior aspect of IP to occipital pole is twice as great in chimpanzee brain casts than on the Hadar AL 162-28 endocast, even though the chimpanzee brain casts are smaller in overall size. This suggests that brain reorganization, at least as exemplified as a reduction in primary visual striate cortex (area 17 of Brodmann), occurred early in hominid evolution, prior to any major brain expansion.     

Stature-at-death of KNM-WT 15000

The specimen KNM-WT 15000 is an exceptionally complete 1.53 Myr juvenile skeleton of Homo erectus from West Turkana, Kenya. It therefore provides a unique opportunity to examine stature estimates of fossil hominids based strictly on long bone lengths. Using recovered axial and appendicular elements of KNM-WT 15000 that contributed to stature during life, we conclude that KNM-WT 15000 was much shorter at time-of-death than previous estimates that used only appendicular elements. We conservatively estimate stature-at-death at about 147 cm, although this individual could have been as short as 141 cm. Because long bone based estimates of stature also imply the axial skeletal proportion, our new stature estimate stems from the recognition of axial/appendicular disproportion in the individual KNM-WT 15000. It is possible that the peripubescent age-at-death of this specimen, and any resulting differential maturity between the appendicular and axial skeleton, may have contributed to previous overestimates of stature-at-death. However, the possibility that this individual was abnormal, as implied by axial/appendicular disproportion, remains to be fully tested. Regardless, these results suggest that some interpretations of the biology of early African Homo erectus, largely based upon KNM-WT 15000, should be viewed with caution. 5 Primate Evolution and Morphology Group, Department of Human     

Endocranial suture closure in rhesus macaques (Macaca mulatta)

Endocasts from skulls of 330 rhesus monkeys (Macaca mulatta) of known age are scored for closure of nine bilateral and three unilateral sutures or segments of sutures. A variety of tests reveals a strong relationship between age and stages of suture closure, although increasingly broad confidence intervals prevent sutures from being very useful for precisely aging older macaques. The order in which endosutures begin to close, as well as that in which closure is finally achieved, is determined for macaques, and these sequences compared to those for endosutures of humans (Todd and Lyon, 1924). The basilar suture is the earliest to close, while the masto-occipital and rostral and caudal squamosal sutures achieve closure quite late in both species. On the other hand, humans and macaques differ in their schedules for the sphenofrontal suture and in the initiation of closure for the rostral portion of the squamosal suture. Two sutures close significantly sooner on the right than on the left side (the rostral squamosal and masto-occipital) and asymmetry favoring closure of the right lateral lambdoid suture also approaches significance at the 0.05 level. No sutures close significantly sooner on the left side. It is suggested that macaque sutures may close from the inside out, that endosutures are more sensitive than ectosutures for detecting sequences in which cranial sutures begin to close, and that directional asymmetries in suture closure of macaques may be related to minor asymmetries in brain/skull shape (petalias).     

Enlargement of the temporalis muscle and alterations in the lateral cranial vault

The purpose of this study was to test the hypothesis that increasedmasticatory muscle accompanied morphologic changes in the temporalbone and squamosal suture. Ten mice deficient for the proteinmyostatin (Mstn –/–) had significantly increasedskeletal muscle mass and were compared with nine controls (Mstn+/+). Variables measured include linear and areal metrics describingtemporal size and temporal bone shape as well as the extentof the area of the squamosal suture that overlaps, or bevels,with parietal bones. Mstn–/– mice showed significantlylarger temporalis muscles. Their temporal bones showed significantlydecreased size as well as decreased beveling of the squamosalsuture. These decreases were absolute as well as relative andwere not restricted to either vertical or horizontal axes. Theincreased masticatory musculature of Myostatin-null mice hada shrinking effect on the temporal aspect of the cranium. Theseresults are inconsistent with the interpretation that increasedtemporalis mass induces morphologic changes in temporal bonethat compensate for putative increases in compressive forcestransduced at this region. Rather than increase in the areaof overlap between two calvarial bones, potential increase inbiomechanical loading along the temporal squama led to a smallerbevel which would presumably weaken this joint. It is unclearwhy this is so. Either compressive forces are not anabolic tosuture beveling or they do upregulate growth of the suture bevel,with compression not being the primary loading regime at thissuture.     

Skeletal age, dental age, and the maturation of KNM-WT 15000

The skeleton of the Homo erectus boy from West Lake Turkana, Kenya (KNM-WT 15000), is remarkably complete, and this individual has thus provided a case study for several researchers examining Homo erectus growth. Using data from a longitudinal study of Montreal French-Canadian children, it is shown that while dental and skeletal ages match reasonably well at the level of a sample of children, individuals can display differences between skeletal and dental ages of 2 years or more. Furthermore, the relationship between these two markers may change over time in individual children. It is also possible to find children with patterns of dental maturation similar to KNM-WT 15000's pattern in the Montreal sample. Therefore, neither the discrepancy between skeletal age and dental age alone nor the pattern of dental maturation as assessed by dental stages precludes a human-like pattern of growth, including an adolescent growth spurt, for this individual. Some indicators (e.g., estimated body size for predicted age, and enamel formation) do suggest possible growth-patterning differences from modern humans, and therefore earlier maturation is a reasonable hypothesis, but caution is warranted, given the large degree of modern human variation in developmental markers and the inherent uncertainty in precise estimation of KNM-WT 15000's maturational parameters.     

First occurrence of early Homo in the Nachukui Formation (West Turkana, Kenya) at 2.3-2.4 Myr

Cognitive abilities and techno-economic behaviours of hominids in the time period between 2.6-2.3 Myr have become increasingly well-documented. This time period corresponds to the oldest evidence for stone tools at Gona (Kada Gona, West Gona, EG 10-12, OGS 6-7), Hadar (AL 666), lower Omo valley (Ftji1, 2 & 5, Omo 57, Omo 123) in Ethiopia, and West Turkana (Lokalalei sites -LA1 & LA2C-) in Kenya. In 2002 a new palaeoanthropological site (LA1alpha), 100 meters south of the LA1 archaeological site, produced a first right lower molar of a juvenile hominid (KNM-WT 42718). The relative small size of the crown, its marked MD elongation and BL reduction, the relative position of the cusps, the lack of a C6 and the mild expression of a protostylid, reinforced by metrical analyses, demonstrate the distinctiveness of this tooth compared with Australopithecus afarensis, A. anamensis, A. africanus and Paranthropus boisei, and its similarity to early Homo. The LA1alpha site lies 2.2 m above the Ekalalei Tuff which is slightly younger than Tuff F dated to 2.34+/-0.04 Myr. This juvenile specimen represents the oldest occurrence of the genus Homo in West Turkana.     

Functional morphology of the asterionic region in extant hominoids and fossil hominids

Asterionic sutural patterns in Plio-Pleistocene hominid crania have never been examined in detail. We present an analysis of this anatomical region in Australopithecus and Homo and relate different sutural patterns to functional changes in the masticatory apparatus. The great apes and A. afarensis share the common adult higher primate sutural pattern referred to as the "asterionic notch," which develops in response to the hypertrophy of posterior temporalis muscle fibers and the consequent formation of compound temporal/nuchal crests. This sutural configuration also appears to be present on the early Homo cranium KNM-ER 1805. In contrast, adult male A. boisei crania exhibit a unique pattern where the temporal squama overlaps the parietal which, in turn, overlaps the par mastoidea and the upper scale of the occipital bone. We relate this arrangement to the need to reinforce the rear of a thin-walled braincase against the net tensile forces exerted by the temporalis and nuchal muscles. The common juvenile hominoid edge-to-edge asterionic articulation is maintained in adult A. africanus, A. robustus, female A. boisei, and most Homo crania. We discuss the latter pattern in regard to anterior temporalis hypertrophy in A. africanus, A. robustus, and A. boisei and to craniofacial paedomorphosis in Homo.     

四川自贡晚侏罗世西蜀鳄一新种

记述了四川省自贡市汇东新区自贡市乳品厂发现的西蜀鳄一新种——周氏西蜀鳄(Hsisosuchus chowi sp.nov.)。新种区别于西蜀鳄已知种的特征是:鼻骨后部沿缝合线有一浅的纵凹,额骨的眶缘向上凸起成嵴,沿额骨缝合线也隆起成一微弱的纵嵴,上颞窝的内侧缘向上凸起呈明显的嵴,顶骨具一前中突,侧视颧骨腹缘呈明显的波曲状,眶后骨前侧角约90°,鳞骨后侧突特别拉长,向侧下后方伸展,使鳞骨侧缘明显向内侧弓曲,左右外枕骨的枕髁部分不相接,翼骨的腹中嵴源于翼骨主体部分,内鼻孔位置比较靠前。此外,齿骨外面和夹板骨腹面具有发达的沟和嵴状雕饰,夹板骨参与下颌联合的部分比较长,肩胛片异常扩展,乌喙骨远端宽于近端,肱骨头增厚并强烈向内侧扩展,三角肌嵴发达,桡侧腕骨具发达的尺骨突,尺侧腕骨远端宽于近端,6列荐前部腹部骨板和3列尾部腹部骨板,也可能是周氏西蜀鳄的衍生特征,但这些性状在大山铺西蜀鳄中情况不明,有待更多的材料来证实。杨钟健、周明镇(1953)在建立西蜀鳄属之初就已注意到西蜀鳄是一种特化的鳄类,认为西蜀鳄不仅将原始特征和进步特征混存于一身,而且还具有一些一般鳄类所没有的独特性质。以此为基础,他们建立了西蜀鳄科。目前西蜀鳄类动物发现并不多,仅有1属2种,即重庆西蜀鳄和大山铺西蜀鳄,而且材料不完整,特别是头后骨骼保存不理想。周氏西蜀鳄的发现不仅扩大了西蜀鳄类的分布范围,而且还增加了我们对这一特化鳄类的认识。     

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 new skeleton of Theropithecus brumpti (Primates: Cercopithecidae) from Lomekwi,West Turkana,Kenya

A relatively complete skeleton of the fossil papionin, Theropithecus brumpti, from the site of Lomekwi, west of Lake Turkana, Kenya, is here described. The specimen, KNM-WT 39368, was recovered at the site of LO 5 (3 degrees 51'N and 35 degrees 45'E), from sediments dated to approximately 3.3Ma. The skeleton is that of an old adult male and preserves a number of articulated elements, including most of the forelimbs and tail. The cranial morphology is that of a large, early T. brumpti, exhibiting a deep mandible with a deeply excavated mandibular corpus fossa, and mandibular alveoli and cheek teeth arrayed in a reversed Curve of Spee. The forelimb skeleton exhibits a unique mixture of characteristics generally associated with a terrestrial locomotor habitus, such as a narrow scapula and a highly stable elbow joint, combined with those more representative of habitual arborealists, such as muscle attachments reflecting a large rotator cuff musculature and a flexible shoulder joint. The forelimb of KNM-WT 39368 also presents several features, unique toTheropithecus, which represent adaptations for manual grasping and fine manipulation. These features include a large, retroflexed medial humeral epicondyle (to which large pronator, and carpal and digital flexor muscles attached) and proportions of the digital rays that denote capabilities for precise opposition between the thumb and index finger. Taken together, these features indicate that one of the earliest recognized representatives of Theropithecus exhibited the food harvesting and processing anatomy that distinguished the genus through time and that contributed to its success throughout the later Pliocene and Pleistocene. Based on the anatomy of KNM-WT 39368 and the known habitat preference of T. brumpti, the species is reconstructed as being a generally terrestrial but highly dexterous, very large-bodied, sexually dimorphic, and possibly folivorous papionin. T. brumpti was adapted for propulsive quadrupedal locomotion over generally even ground, and yet was highly adept at manual foraging. The estimate of 43.8kg body mass for KNM-WT 39368 renders unlikely the possibility that the species, or at least adult males of the species, were highly arboreal. T. brumpti, as represented by KNM-WT 39368, is seen as a large, colorfully decorated, and basically terrestrial papionin that was restricted to riverine forest habitats in the Lake Turkana Basin from the middle to latest Pliocene.     

Basicranial anatomy of Plio-Pleistocene hominids from East and South Africa

The results of a metrical analysis of the basicranium of 19 Plio-Pleistocene fossil hominid crania are presented. The sample includes crania attributed to Australopithecus africanus, Australopithecus boisei, and robustus, and Homo erectus as well as crania whose attribution is still under discussion. These results confirm significant differences between the cranial base patterns of the "gracile" and "robust" australopithecines and the three crania attributed to Homo erectus have a pattern which resembles that of modern humans. None of the crania examined from East Africa sites have base patterns which resemble that of the "gracile" australopithecines. The crania KNM-ER 407 and 732 have patterns which are compatible with them being smaller-bodied females of Australopithecus boisei; KNM-ER 1470 and 1813 have base patterns which most closely resemble that of Homo erectus. The cranial base pattern of KNM-ER 1805 is compatible with its inclusion in either Australopithecus boisei or Homo. When account is taken of the immaturity of Taung, the evidence of its cranial base pattern suggests that if it had reached adulthood it would have resembled the "gracile" australopithecine crania from Sterkfontein and Makapansgat.     

Dental microwear and diet of the Plio-Pleistocene hominin Paranthropus boisei

The Plio-Pleistocene hominin Paranthropus boisei had enormous, flat, thickly enameled cheek teeth, a robust cranium and mandible, and inferred massive, powerful chewing muscles. This specialized morphology, which earned P. boisei the nickname "Nutcracker Man", suggests that this hominin could have consumed very mechanically challenging foods. It has been recently argued, however, that specialized hominin morphology may indicate adaptations for the consumption of occasional fallback foods rather than preferred resources. Dental microwear offers a potential means by which to test this hypothesis in that it reflects actual use rather than genetic adaptation. High microwear surface texture complexity and anisotropy in extant primates can be associated with the consumption of exceptionally hard and tough foods respectively. Here we present the first quantitative analysis of dental microwear for P. boisei. Seven specimens examined preserved unobscured antemortem molar microwear. These all show relatively low complexity and anisotropy values. This suggests that none of the individuals consumed especially hard or tough foods in the days before they died. The apparent discrepancy between microwear and functional anatomy is consistent with the idea that P. boisei presents a hominin example of Liem's Paradox, wherein a highly derived morphology need not reflect a specialized diet.