Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length

Previous studies of daily energy expenditure (DEE) in hominin fossils have estimated locomotor costs using a formula that was based on six species, all 18 kg or less in mass, including no primates, and that has a number of other problems when applied in an ecological context. It is well established that the energetic cost of human walking is lower than that of representative mammals, particularly for individuals with long lower limbs. The current study reevaluates the daily energy expenditures of a variety of hominin species using more appropriate approaches to estimating locomotor costs. To estimate DEE for primates, I relied on published data on body mass, day range, and the percentage of time spent in various activities. Based on those data, I calculated a value for nonlocomotor DEE. I then used a variant of a method that I have suggested elsewhere to calculate the daily cost due to locomotion (DEEL) and summed the two to calculate total DEE. The more up-to-date methods for calculating the cost of travel result in lower estimates of this aspect of the energy budget than seen in previous studies. Values obtained here for DEE in various representatives of Australopithecus are lower than reported previously by around 200 kcal/day. Taking into account the greater economy of human walking, particularly the effect of the longer lower limbs found in many later Homo species, also results in lowered estimates of DEE. Elongation of the lower limbs in H. erectus reduced relative travel costs nearly 50% in comparison to A.L. 288-1 (A. afarensis). The present method for calculating DEE indicates that female H. erectus DEE was 84% greater than that of female Australopithecus; this disparity is even larger than that suggested by previous workers.     

Tooth components of mandibular deciduous molars of Homo sapiens sapiens and Homo sapiens neanderthalensis: a radiographic study.

Tooth components of deciduous molars were measured from standardized radiographs of Homo sapiens sapiens and Homo sapiens neanderthalensis. Enamel height and width were greater in deciduous teeth of Homo sapiens sapiens than in Homo sapiens neanderthalensis and the differences were statistically significant (p less than 0.01). Dentin height showed no significant differences between the two groups, but enamel to floor of pulp chamber and pulp height and width dimensions were significantly greater in Homo sapiens neanderthalensis. Discriminant analysis carried out between groups, using deciduous tooth components, showed an accuracy of 98-100% for identification of Homo sapiens sapiens and 83-92% for identification of Homo sapiens neanderthalensis. The results obtained in this study on dental dimensions support the hypothesis of a distinct evolutionary line for Neanderthals.     

Cranial integration in Homo: singular warps analysis of the midsagittal plane in ontogeny and evolution

This study addresses some enduring issues of ontogenetic and evolutionary integration in the form of the hominid cranium. Our sample consists of 38 crania: 20 modern adult Homo sapiens, 14 sub-adult H. sapiens, and four archaic Homo. All specimens were CT-scanned except for two infant H. sapiens, who were imaged by MR instead. For each specimen 84 landmarks and semi-landmarks were located on the midsagittal plane and converted to Procrustes shape coordinates. Integration was quantified by the method of singular warps, a new geometric-statistical approach to visualizing correlations among regions. The two classic patterns of integration, evolutionary and ontogenetic, were jointly explored by comparing analyses of overlapping subsamples that span ranges of different hypothetical factors. Evolutionary integration is expressed in the subsample of 24 adult Homo, and ontogenetic integration in the subsample of 34 H. sapiens. In this data set, vault, cranial base, and face show striking and localized patterns of covariation over ontogeny, similar but not identical to the patterns seen over evolution. The principal differences between ontogeny and phylogeny pertain to the cranial base. There is also a component of cranial length to height ratio not reducible to either process. Our methodology allows a separation of these independent processes (and their impact on cranial shape) that conventional methods have not found.     

The utility of cranial ontogeny for phylogenetic inference: a case study in crocodylians using geometric morphometrics

The degree to which the ontogeny of organisms could facilitate our understanding of phylogenetic relationships has long been a subject of contention in evolutionary biology. The famed notion that ‘ontogeny recapitulates phylogeny’ has been largely discredited, but there remains an expectation that closely related organisms undergo similar morphological transformations throughout ontogeny. To test this assumption, we used three‐dimensional geometric morphometric methods to characterize the cranial morphology of 10 extant crocodylian species and construct allometric trajectories that model the post‐natal ontogenetic shape changes. Using time‐calibrated molecular and morphological trees, we employed a suite of comparative phylogenetic methods to assess the extent of phylogenetic signal in these trajectories. All analyses largely demonstrated a lack of significant phylogenetic signal, indicating that ontogenetic shape changes contain little phylogenetic information. Notably, some Mantel tests yielded marginally significant results when analysed with the morphological tree, which suggest that the underlying signal in these trajectories is correlated with similarities in the adult cranial morphology. However, despite these instances, all other analyses, including more powerful tests for phylogenetic signal, recovered statistical and visual evidence against the assumption that similarities in ontogenetic shape changes are commensurate with phylogenetic relatedness and thus bring into question the efficacy of using allometric trajectories for phylogenetic inference.     

Relevance of aquatic environments for hominins: a case study from Trinil (Java, Indonesia)

Knowledge about dietary niche is key to understanding hominin evolution, since diet influences body proportions, brain size, cognition, and habitat preference. In this study we provide ecological context for the current debate on modernity (or not) of aquatic resource exploitation by hominins. We use the Homo erectus site of Trinil as a case study to investigate how research questions on possible dietary relevance of aquatic environments can be addressed. Faunal and geochemical analysis of aquatic fossils from Trinil Hauptknochenschicht (HK) fauna demonstrate that Trinil at ∼1.5 Ma contained near-coastal rivers, lakes, swamp forests, lagoons, and marshes with minor marine influence, laterally grading into grasslands. Trinil HK environments yielded at least eleven edible mollusc species and four edible fish species that could be procured with no or minimal technology. We demonstrate that, from an ecological point of view, the default assumption should be that omnivorous hominins in coastal habitats with catchable aquatic fauna could have consumed aquatic resources. The hypothesis of aquatic exploitation can be tested with taphonomic analysis of aquatic fossils associated with hominin fossils. We show that midden-like characteristics of large bivalve shell assemblages containing Pseudodon and Elongaria from Trinil HK indicate deliberate collection by a selective agent, possibly hominin.     

Morphological study on early ontogeny of theGinkgo leaf

The early ontogeny of theGinkgo leaf was studied by scanning electron microscope. The lamina of the leaf is derived from the protuberance of the adaxial side of a leaf buttress. The protuberance shows successive bifurcations. The second bifurcation occurs approximately at a right angle to the first. Thus, the abaxial surface of the leaf is derived from the outer surface of the protuberance and the adaxial is derived from the depressed inner surface of the protuberance formed by the first and second bifurcations. It is supposed that thick veins along both edges of the lamina are differentiated by an uneven dichotomous branching system of procambia caused by positional relationships: both edges of the lamina are derived from the parts of the protuberance just above a pair of procambia which come from the stem. On the shoot apex, there are a central cell group, which consists of only a few cells, and radially-arranged cell files. The leaf primordium is initiated from a sector area which consists of several files among radially-arranged cell files.     

Cosmos pollen ontogeny: A scanning electron microscope study

Summary Ontogenetic data concerning pollen not only clarifies the mode of deposition of the elaborate walls but has considerable functional and taxonomic relevance. Hitherto such studies have used optical or transmission electron microscopy but here a recently devised preparative technique has enabled pollen development inCosmos bipinnatus to be studied using the scanning electron microscope. The technique involves freeze-fracturing of osmium fixed, cryoprotected anthers, maceration in dilute osmium tetroxide, critical point drying, sputter coating and examination. The processes of pollen wall development can then be observed in three dimensions, an important aid to understanding the spatial relationships involved in the determination of ornamentation and apertures. Details of the pollen and tapetum are described at various stages between meiosis and anthesis. A close conformity is demonstrated between the results obtained and those of earlier transmission electron microscopic studies of the same and related species although very different interpretations are made.     

Patterns of intraspecific variation through ontogeny: a case study of the Cretaceous nautilid Eutrephoceras dekayi and modern Nautilus pompilius

The magnitude and ontogenetic patterns of intraspecific variation can provide important insights into the evolution and development of organisms. Understanding the intraspecific variation of organisms is also a key to correctly pursuing studies in major fields of palaeontology. However, intraspecific variation has been largely overlooked in ectocochleate cephalopods, particularly nautilids. Furthermore, little is known regarding the evolutionary pattern. Here, we present morphological data for the Cretaceous nautilid Eutrephoceras dekayi (Morton) and the modern nautilid Nautilus pompilius Linnaeus through ontogeny. The data are used to describe conch morphology and to elucidate the evolutionary patterns of intraspecific variation. We discovered a similar overall pattern of growth trajectories and the presence of morphological changes at hatching and maturity in both taxa. We also found that intraspecific variation is higher in earlier ontogeny than in later ontogeny in both taxa. The high variation in earlier ontogeny may imply increased flexibility in changing the timing of developmental events, which probably played an important role in nautilid evolution. We assume that the decrease in variation in later ontogeny reflects developmental constraints. Lastly, we compared the similarity/dissimilarity of ontogenetic patterns of variation between taxa. Results reveal that the similarity/dissimilarity of the ontogenetic pattern differs between E. dekayi and N. pompilius. We conclude that this shift in the ontogenetic pattern of variation may be rooted in changes in the developmental programme of nautilids through time. We propose that studying ontogenetic patterns of intraspecific variation can provide new insights into the evolution and development of organisms.     

现代人起源问题的新争论

现代人在分类学上称为晚期智人。他们是怎样起源的?长期来有两种理论,一种叫直接演化说(Direct evolution hypothesis)或系统说,也叫多地区起源说。这种理论认为现代人是由当地的早期智人以至猿人演化而来的,各人种在很久以前即已分歧,各自平行发展,演化成现代人,但长时期来互相有基因的交流。例如欧洲的白种人是由当地的尼安德特人逐渐演变来的;亚洲的黄种人是由当地的早期智人和猿人演变来的。另一种理论叫入侵论(Invasion hypothesis),也叫迁徙说或代替说,这种理论认为欧洲是典型尼人的家乡, 而解剖结构上的现代人是在欧洲以外地区演化出来, 然后侵人欧洲的尼人区域, 消灭了土著的尼人而形成的;侵人亚洲的也一样, 形成现代的黄种人。这种单一地区起源说过去一般认为亚洲西部是现代人最早形成的地区, 近年来有人提出非洲南部才是现代人最早形成的地区, 然后迁徙到世界各地。     

南京1号直立人头骨与肯尼亚KNM-ER 3733人类头骨化石的形态比较

KNM-ER 3733人类头骨化石的年代为距今1.78百万年,1975年发现于肯尼亚。Walker和Leakey注意到这具头骨与周口店直立人的在脑颅形态上很相近,但二者在年代上相差大约1百万年,故认为直立人形态在这1百万年期间是稳定的。长期来此观点缺乏更多的人类化石证据来支持。1993年在中国发现了南京1号人类头骨化石。该头骨与KNM-ER 3733头骨一样兼具脑颅和面颅,且都属于成年女性个体,但南京1号人类头骨化石的年代比KNM-ER 3733人类头骨化石的要晚大约1百万年。因此,南京1号人类头骨是目前所知的可用来验证直立人头骨形态是否在1百万年期间保持稳定的唯一合适的人类头骨化石材料。形态比较表明,这两个人类头骨化石的脑颅虽然在眶上圆枕上沟的发育程度、眶后收缩的程度、额骨横向隆起的程度、角圆枕和乳后突的发育与否、顶骨形状以及骨壁厚度的表现上有所差异,但有更多的形态性状显示出相近。这些相近表现在脑颅的长、宽、高值上;颅容量上;脑颅的低矮性上;脑颅最大宽之位置上;额骨、顶骨、枕骨之矢弧值的比例上;眶上圆枕的纤细上;顶骨的大小和矢向扁平性上;颞线位置和颞鳞顶缘的形状上;枕鳞的低宽形状上;上枕鳞与下枕鳞之间的转折形状和比例上;枕骨圆枕和枕骨圆枕上沟的发育程度上等。这两具头骨的面颅虽然有同属突颌型的面角、皆发育有鼻骨间嵴、两鼻骨组成的上部宽度与下部宽度皆差别很大,但有更多的形态性状显示出差别。这些差别表现在面型上、颜面上部扁平度上、眶形和眶型上、上颌额突外侧面的朝向上、鼻骨横向隆起程度上、鼻梁外突程度上、鼻型上、颧骨下缘外展程度上、颊高上、颧上颌下缘的形状上、上颌颧突基部的位置上以及颧结节的位置上等。因此,南京1号头骨与KNM-ER 3733头骨之间在脑颅上显示出较多的相近性状,在面颅上则显示出较多的相异性状。脑颅方面的相近性状大多具有分类上的鉴别价值。这两个头骨脑颅形态的相近支持把KNM-ER 3733头骨鉴定为"直立人"的观点;也提示了南京1号头骨的脑颅似乎保持着1百多万年前的"祖先"形态。如果直立人的某些成员在至少1百万年期间保持着形态稳定的话,则这种形态上的稳定主要是表现在脑颅形态上。这两具头骨的面颅形态上较大差异的意义,目前尚不清楚。     

Brief Communication: Lumbar lordosis in extinct hominins: Implications of the pelvic incidence

Recently, interest has peaked regarding the posture of extinct hominins. Here, we present a new method of reconstructing lordosis angles of extinct hominin specimens based on pelvic morphology, more specifically the orientation of the sacrum in relation to the acetabulum (pelvic incidence). Two regression models based on the correlation between pelvic incidence and lordosis angle in living hominoids have been developed. The mean values of the calculated lordosis angles based on these models are 36°?45° for australopithecines, 45°?47° for Homo erectus, 27°?34° for the Neandertals and the Sima de los Huesos hominins, and 49°?51° for fossil H. sapiens. The newly calculated lordosis values are consistent with previously published values of extinct hominins (Been et al.: Am J Phys Anthropol 147 (2012) 64–77). If the mean values of the present nonhuman hominoids are representative of the pelvic and lumbar morphology of the last common ancestor between humans and nonhuman hominoids, then both pelvic incidence and lordosis angle dramatically increased during hominin evolution from 27° ± 5 to 22° ± 3 (respectively) in nonhuman hominoids to 54° ± 10 and 51° ± 11 in modern humans. This change to a more human‐like configuration appeared early in the hominin evolution as the pelvis and spines of both australopithecines and H. erectus show a higher pelvic incidence and lordosis angle than nonhuman hominoids. The Sima de los Huesos hominins and Neandertals show a derived configuration with a low pelvic incidence and lordosis angle. Am J Phys Anthropol 154:307–314, 2014. © 2014 Wiley Periodicals, Inc.     

Embryonic development of the bullseye puffer Sphoeroides annulatus (Tetraodontidae): A morphofunctional approach to ontogenetic steps

The embryonic development of the bullseye puffer, Sphoeroides annulatus, was characterized on the basis of the theory of saltatory ontogeny. This theory predicts a correlative relationship between the ontogeny-type in an altricial-precocial spectrum and the habitat that a species occupies within an unstable-stable environmental spectrum. Because S. annulatus inhabits a variety of unstable environments along a wide latitudinal range, the hypothesis that this species presents one of the most altricial embryonic developments among tetraodontids was tested. Based on major developmental events that marked the ontogenetic thresholds nine embryonic steps were identified. Developmental features such as small adhesives eggs, lack of vitelline circulation, small free embryos swimming up at hatching guided by positive phototaxis, and small first-feeding larvae actively swam in the water column, suggest that S. annulatus belongs to the reproductive guild of the nonguarders-lithopelagophils. Moreover, a comparative analysis of the developmental sequences, egg size, and first-feeding larvae size between tetraodontids confirms the hypothesis of this study and supports the evolutionary principle of the altricial-precocial spectrum postulated in the theory of saltatory ontogeny.     

Acclimation responses to short‐term temperature treatments during early life stages causes long lasting changes in spontaneous activity of adult Drosophila melanogaster

Ecotherms adjust their physiology to environmental temperatures. Long‐term exposures to heat or cold typically induce acclimation responses that generate directional, but reversible shifts in thermal tolerance and performance. However, less is known about how short exposure in different life stages will affect the adult phenotype. In the present study, we compared the effects of long‐term temperature exposure to 15, 19 and 31 °C with that of brief (16 h) exposure periods at the same temperatures in Drosophila melanogaster eggs, larvae, pupae, or adults, respectively. The acclimation responses are evaluated using activity measurements at 11, 15, 19, 27, 31 and 33 °C and by measuring upper and lower thermal limits (CT_max and CT_min) in 5‐day‐old adult males. As expected, long‐term cold exposure reduces relative CT_min, whereas long‐term heat exposure increases relative CT_max. By contrast, we find little effect on thermal limits when using short‐term exposures at different life stages. Long‐term exposures to 31 and 15 °C both suppressed activity relative to the 19 °C control, suggesting that development at high and low temperatures may lead to reduced activity later in life. Short‐term cold exposure early in development reduces activity in the adult stage, whereas the effects of short‐term heat exposure on behaviour are dependent on life stage and test temperature. Together, our results highlight how the thermal sensitivity of the trait measured determines the ability to detect acclimation responses.     

印度纳尔玛达地区发现的粗壮智人化石及其与亚洲直立人的连续性

1999年,在印度西部的纳尔玛达(Narmada)河的一条支流——奥尔桑(Orsang)河谷发现了一具孤立的智人头骨,这对于了解印度人类进化历史和南亚直立人与最古老的亚洲智人(或AMH,解剖学上现代的人)之间的联系有着重要的意义。化石是在古老的河流沉积物中发现的。对主体沉积和颅内沉积物的红外光释光(IRSL)测年结果显示,其年代为3—5万年。然而,对化石的直接测年(放射性碳加速器质谱测年AMS)结果表明,其年代最小为4981—5579年前。头骨被归类为圆颅型智人。头骨上最引人注意的特征是颅外有亚洲直立人头后部很发达的角圆枕。头骨最宽处在下部(颞骨)如直立人,这样的情况从来不见于智人。这可能是由于乳突的气窦化而形成的。乳突上脊发育,从破损的眉间区(20mm)可以看到很宽的额窦。所有这些粗壮的特征都表明Orsang头骨和晚期亚洲直立人之间具有遗传连续性。     

Dynamic body acceleration increases by 20% during flight ontogeny of greylag geese Anser anser

Despite our knowledge of the biophysical and behavioural changes during flight ontogeny in juvenile birds, little is known about the changes in the mechanical aspects of energy expenditure during early flight development, particularly in migratory species. Here, we investigate in a unique experimental setup how energy expended during flights changes over time beginning with early ontogeny. We calculate overall dynamic body acceleration (ODBA) as a proxy for energy expenditure in a group of hand raised greylag geese Anser anser trained to fly behind a microlight aircraft. We propose two potential hypotheses; energy expenditure either increases with increasing physiological suitability (the ‘physical development hypothesis’), or decreases as a result of behavioural improvements mitigating flight costs (the ‘behavioural development hypothesis’). There was a significant temporal increase of flight duration and ODBA over time, supporting the ‘physical development hypothesis’. This suggests that early on in flight ontogeny behavioural development leading to flight efficiency plays a weaker role in shaping ODBA changes than the increased physical ability to expend energy in flight. We discuss these findings and the implications of flight development on the life history of migratory species.     

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.