Computational Approaches and Observer Variation in the 3D Musculoskeletal Modeling of the Heads of Anolis.

High-resolution imaging, 3D modeling, and quantitative analyses are equipping evolutionary biologists with new approaches to understanding the variation and evolution of the musculoskeletal system. However, challenges with interpreting DiceCT data and higher order use of modeled muscles have not yet been fully explored, and the error in and accuracy of some digital methods remain unclear. West Indian Anolis lizards are a model clade for exploring patterns in functional adaptation, ecomorphology, and sexual size dimorphism in vertebrates. These lizards possess numerous jaw muscles with potentially different anatomies that sculpt the adductor chamber of the skull. Here we test approaches to quantifying the musculoskeletal shape of the heads of two species of Anolis: A. pulchellus and A. sagrei. We employ comparative approaches such as DiceCT segmentation of jaw muscles, 3D surface attachment mapping, and 3D landmarking with the aim of exploring muscle volumes, 3D muscle fiber architecture, and sexual dimorphism of the skull. We then compare sources of measurement error in these 3D analyses while also presenting new 3D musculoskeletal data from the Anolis feeding apparatus. These findings demonstrate the accessibility and repeatability of these emerging techniques as well as provide details regarding the musculoskeletal anatomy of the heads of A. pulchellus and A. sagrei which show potential for further research of comparative biomechanics and evolution in the clade.

New fossils of Australopithecus anamensis from Kanapoi, West Turkana, Kenya (2012-2015).

Kanapoi, Kenya, has yielded the earliest evidence of the genus Australopithecus, Australopithecus anamensis. Renewed fieldwork from 2012 through 2015 yielded 18 new fossils attributable to this species. The new specimens include the second maxillary fragment known from a Kanapoi hominin and the first from a relatively young adult. The new maxilla has the distinctive rounded nasal aperture margin characteristic of A. anamensis. A second partial proximal tibia from the site is the first postcranial element from a small A. anamensis individual. A new partial mandible and complete mandibular dentition display distinctive Kanapoi hominin morphology, but the mandible displays a larger trigonid on its fourth premolar than any known so far. Two new complete sets of mandibular incisors are also notably large, especially the lateral ones, a distinctive feature of A. anamensis compared with Australopithecus afarensis. The new fossils also highlight the distinctive morphology of Kanapoi A. anamensis compared to later hominins.

New fossils of Australopithecus anamensis from Kanapoi, West Turkana, Kenya (2003-2008).

Renewed fieldwork from 2003 through 2008 at the Australopithecus anamensis type-site of Kanapoi, Kenya, yielded nine new fossils attributable to this species. These fossils all date to between 4.195 and 4.108 million years ago. Most were recovered from the lower fluvial sequence at the site, with one from the lacustrine sequence deltaic sands that overlie the lower fluvial deposits but are still below the Kanapoi Tuff. The new specimens include a partial edentulous mandible, partial maxillary dentition, two partial mandibular dentitions, and five isolated teeth. The new Kanapoi hominin fossils increase the sample known from the earliest Australopithecus, and provide new insights into morphology within this taxon. They support the distinctiveness of the early A. anamensis fossils relative to earlier hominins and to the later Australopithecus afarensis. The new fossils do not appreciably extend the range of observed variation in A. anamensis from Kanapoi, with the exception of some slightly larger molars, and a canine tooth root that is the largest in the hominin fossil record. All of the Kanapoi hominins share a distinctive morphology of the canine-premolar complex, typical early hominin low canine crowns but with mesiodistally longer honing teeth than seen in A. afarensis, and large, probably dimorphic, canine tooth roots. The new Kanapoi specimens support the observation that canine crown height, morphology, root size and dimorphism were not altered from a primitive ape-like condition as part of a single event in human evolution, and that there may have been an adaptive difference in canine function between A. anamensis and A. afarensis.

Associated cranial and forelimb remains attributed to Australopithecus afarensis from Hadar, Ethiopia.

A partial skeleton from Hadar, Ethiopia (A.L. 438-1) attributed to Australopithecus afarensis is comprised of part of the mandible, a frontal bone fragment, a complete left ulna, two second metacarpals, one third metacarpal, plus parts of the clavicle, humerus, radius, and right ulna. It is one of only a few early hominin specimens to preserve both cranial and postcranial elements. It also includes the first complete ulna from a large A. afarensis individual, and the first associated metacarpal and forelimb remains. This specimen, dated to approximately 3Ma, is among the geologically youngest A. afarensis fossils and is also one of the largest individuals known. Its ulnar to mandibular proportions are similar to those of the geologically older and much smaller A.L. 288-1, suggesting that body size increased without disproportional enlargement of the mandible. Overall, however, analysis of this large specimen and of the diminutive A.L. 288-1 demonstrates that the functional morphology of the A. afarensis upper limb was similar at all body sizes; there is no evidence to support the hypothesis that more than one hominin species is present at Hadar. Morphologically, all apparent apomorphic traits of the elbow, forearm, wrist, and hand of A.L. 438-1 are shared uniquely with humans. Compared to humans, A.L. 438-1 does have a more curved ulna, although A.L. 288-1 does not, and it appears to have had slightly less well-developed manipulatory capabilities of its hands, although still more derived than in apes. We conclude that selection for effective arboreality in the upper limb of Australopithecus afarensis was weaker than in non-hominins, and that manipulative ability was of greater selective advantage than in extant great apes.

Morphology of Australopithecus anamensis from Kanapoi and Allia Bay, Kenya.

The hominid species Australopithecus anamensis was originally described in 1995, with new specimens and more secure dates given in 1998. This paper lists all fossils attributed to A. anamensis, and provides anatomical descriptions of those not yet described in detail with photographs of all but undiagnostic fragments. We also provide comparative analysis of these specimens. The A. anamensis holotype mandible was found at Kanapoi, as were most of the paratypes. The Allia Bay sample is less well represented, and does not preserve many anatomical elements diagnostic of this species. Still, the Allia Bay sample most closely resembles that from Kanapoi, and we suggest that for the time being it be retained as A. anamensis. A. anamensis most closely resembles A. afarensis, but can be distinguished from it in many features. Most of these features are inferred to be primitive for the genus. Based on the limited postcranial evidence available, A. anamensis appears to have been habitually bipedal, although it retained some primitive features of its upper limbs. A. anamensis differs from A. afarensis in having narrower, more parallel jaws with a very slightly more ape-like canine/premolar complex than is found in A. afarensis, although not as ape-like as in Ardipithecus ramidus. It had slightly larger lower lateral incisors, a unique upper canine morphology, and a different structure of the lateral nasal aperture than A. afarensis. A. anamensis had at least as great a range of body size, and perhaps slightly greater canine dimorphism, although this is difficult to determine. At present, there appears to be no autapomorphies precluding A. anamensis from ancestry of A. afarensis.

South Turkwel: a new pliocene hominid site in Kenya.

New fossils discovered south of the Turkwel River in northern Kenya include an associated metacarpal, capitate, hamate, lunate, pedal phalanx, mandibular fragment, and teeth. These fossils probably date to around 3.5 m.y.a. Faunal information suggests that the environment at South Turkwel was predominantly bushland. The mandibular and dental remains are fragmentary, but the postcranial fossils are informative. Comparisons with Australopithecus, modern human, chimpanzee and gorilla hand bones suggest that the Turkwel hominid was most like Australopithecus afarensis and A. africanus. Carpometacarpal articulations are intermediate between those of modern humans and African apes, suggesting enhanced gripping capabilities compared with extant apes. The hamulus was strikingly large, similar in proportion only to Neandertals and some gorillas, suggesting the presence of powerful forearms and hands. There are no indicators of adaptations to knuckle-walking or suspensory locomotion in the hand, and the pedal phalanx suggests that this hominid was habitually bipedal.

Torso morphology and locomotion in Proconsul nyanzae.

A new partial skeleton of Proconsul nyanzae from Mfangano Island, Kenya (KNM-MW 13142) includes five lumbar vertebrae, a partial sacrum, and nearly complete hipbone. Until recently, little was known of the axial and pelvic skeletons of Proconsul, or any early Miocene hominoid. Because torso morphology directly reflects locomotor behavior, these elements provide important new information about posture and locomotion of P. nyanzae. Two basic patterns of locomotor behavior and corresponding torso morphology exist among extant anthropoids. Monkeys have long, flexible spines with powerful epaxial muscles and mediolaterally narrow thoraces. This morphology represents a response to the mechanical requirements of habitual pronograde quadrupedalism. In contrast, hominoids, particularly great apes, have shorter, stiffer spines and broader torsos than do monkeys. The hominoid pattern reflects an emphasis on forelimb abduction-adduction, and on stability and control of the lower back during forelimb-dominated arboreal locomotion. KNM-MW 13142 had craniocaudally elongate vertebral bodies, and probably six lumbar vertebrae, reflecting a long, flexible vertebral column. The narrow, laterally facing iliac blades of KNM-MW 13142 imply the presence of a mediolaterally narrow, dorsoventrally deep thorax. Its wide iliac tuberosities, ventrally located lumbar transverse processes, and distinct accessory processes suggest that Proconsul nyanzae also had strap-like epaxial muscles similar to those characterizing most monkeys. Axial and pelvic morphology of KNM-MW 13142 indicate that P. nyanzae had not undergone an ape-like rearrangement of its torso to adapt to forelimb-dominated arboreality. Instead, P. nyanzae probably retained a more primitive pattern of pronograde quadrupedalism. This morphology indicates that the extant hominoid pattern of torso anatomy arose subsequent to the establishment of the hominid clade.

New Australopithecus boisei calvaria from East Lake Turkana, Kenya.

The calvaria of an adult Australopithecus boisei from Area 104, Koobi Fora, Lake Turkana, is described. The specimen, KNM-ER 23000, comes from sediments dated to about 1.9 Ma. It consists of the frontal, both parietals, both temporals, most of the occipital as well as two small pieces of sphenoid, and a mandibular tooth root. The specimen is presumed to be an adult male, based on its size and the great development of features associated with the masticatory apparatus. KNM-ER 23000 is close in general size and shape to KNM-ER 406, KNM-ER 13750, and Olduvai Hominid 5 and it has a mixture of features seen in these three roughly contemporaneous crania. The frontal, especially the tori, resembles that of OH 5; the parietals are most like those of KNM-ER 13750; the occipital is like those of the two other Turkana specimens, and the temporals have a mixture of features from all of these, This specimen adds to our knowledge of variability in A. boisei.

Partial skeleton of Proconsul nyanzae from Mfangano Island, Kenya.

A partial skeleton attributed to Proconsul nyanzae (KNM-MW 13142) is described. The fossils were found at a site on Mfangano Island, Kenya, which dates to 17.9 +/- .1 million years ago. KNM-MW 13142 consists of six partial vertebrae (T12-S1), a nearly complete hipbone, most of the right femur and left femoral shaft, a fragmentary tibia and fibula, and a nearly complete talus and calcaneus. This skeleton provides the first pelvic fossil known for any East African Miocene hominoid. The new Proconsul specimen is compared to a large sample of extant anthropoids to determine its functional and phylogenetic affinities. In most aspects of its anatomy, KNM-MW 13142 closely resembles nonhominoid anthropoids. This individual had a long, flexible spine, narrow torso, and habitually pronograde posture, features characteristic of most extant monkeys. Evidence of spinal musculature suggests a generalized condition intermediate between that of cercopithecoids and hylobatids. The hindlimb of KNM-MW 13142 exhibits relatively mobile hip and ankle joints, with structural properties of the femur like those of hominoids. This mix of features implies a pattern of posture and locomotion that is unlike that of any extant primate. Many aspects of the Proconsul nyanzae locomotor skeleton may represent the primitive catarrhine condition.