Orang-like manual adaptations in the fossil hominoid Hispanopithecus laietanus: first steps towards great ape suspensory behaviours

Morphological and biometrical analyses of the partial hand IPS18800 of the fossil great ape Hispanopithecus laietanus (=Dryopithecus laietanus), from the Late Miocene (about 9.5Ma) of Can Llobateres (Catalonia, Spain), reveal many similarities with extant orang-utans (Pongo). These similarities are interpreted as adaptations to below-branch suspensory behaviours, including arm-swinging and clambering/postural feeding on slender arboreal supports, due to an orang-like double-locking mechanism. This is confirmed by the long and highly curved phalanges of Hispanopithecus. The short and stout metacarpals with dorsally constricted heads, together with the dorsally extended articular facets on proximal phalanges, indicate the persistence of significant degrees of palmigrady. A powerful grasping capability is indicated by the great development of basal phalangeal tubercles, the marked insertions for the flexors on phalangeal shafts and the large pits for the collateral ligaments. The morphology of the Hispanopithecus long bones of the hand indicates a unique positional repertoire, combining orthogrady with suspensory behaviours and palmigrade quadrupedalism. The retention of powerful grasping and palmigrady suggests that the last common ancestor of hominids might have been more primitive than what can be inferred on the basis of extant taxa, suggesting that pronograde behaviours are compatible with an orthograde bodyplan suitable for climbing and suspension.     

Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999-2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.     

New reconstruction of the moroto hominoid snout and a reassessment of its affinities toAfropithecus turkanensis

Two previous reconstructions of the Moroto hominoid palate and face made during the late 1960s suffer from several errors, in particular the angle at which the facial skeleton was hafted onto the palate, and the orientation of the canines, among others. A new reconstruction is based on the discovery that there is a good contact about 1cm logn between the palatal and facial pieces, which reveals that the snout was low and elongated as inAfropithecus turkanensis. Addition of a fragment of the right orbit reveals that the interorbital region was wide as inA. turkanensis. The remaining differences between the skulls from Moroto and Kalodirr are due to five factors:−1) pathology of the Moroto right premaxilla following ante-mortem loss of the right I¹, which led to abnormal enlargement of the right half of the nasal cavity, 2) the incisive foramina of the Moroto palate are damaged, making them appear considerably larger than they were in life, 3) damage to the upper portions of the nasal aperture making it look higher than it would in life, 4) plastic deformation and crushing of the Kalodirr specimen has greatly reduced the width of the palate and has led to the cheek tooth rows converging to the rear, and 5) the slightly larger and ontogenetically older status of the Moroto individual. Resemblances between the Moroto and Kalodirr specimens far outweight the purported or real differences and it is concluded thatMorotopithecus bishopi is a junior subjective synonym ofAfropithecus turkanensis. The sites of Moroto and Kalodirr are unlikely to differ greatly in age (certainly not as much as 4 Ma).     

The relative congruence of cranial and genetic estimates of hominoid taxon relationships: implications for the reconstruction of hominin phylogeny

Previous analyses of extant catarrhine craniodental morphology have often failed to recover their molecular relationships, casting doubt on the accuracy of hominin phylogenies based on anatomical data. However, on the basis of genetic, morphometric and environmental affinity patterns, a growing body of literature has demonstrated that particular aspects of cranial morphology are remarkably reliable proxies for neutral modern human population history. Hence, it is important to test whether these intra-specific patterns can be extrapolated to a broader primate taxon level such that inference rules for understanding the morphological evolution of the extinct hominins may be devised. Here, we use a matrix of molecular distances between 15 hominoid taxa to test the genetic congruence of 14 craniomandibular regions, defined and morphometrically delineated on the basis of previous modern human analyses. This methodology allowed us to test directly whether the cranial regions found to be reliable indicators of population history were also more reliable proxies for hominoid genetic relationships. Cranial regions were defined on the basis of three criteria: developmental-functional units, individual bones, and regions differentially affected by masticatory stress. The results found that all regions tested were significantly and strongly correlated with the molecular matrix. However, the modern human predictions regarding the relative congruence of particular regions did not hold true, as the face was statistically the most reliable indicator of hominoid genetic distances, as opposed to the vault or basicranium. Moreover, when modern humans were removed from the analysis, all cranial regions improved in their genetic congruence, suggesting that it is the inclusion of morphologically-derived humans that has the largest effect on incongruence between morphological and molecular estimates of hominoid relationships. Therefore, it may be necessary to focus on smaller intra-generic taxonomic levels to more fully understand the effects of neutral and selective evolutionary processes in generating morphological diversity patterns.     

An additional specimen of a large-bodied Miocene hominoid from Chiang Muan, northern Thailand

Chiang Muan is the first Miocene fossil site in Southeast Asia, from which large-bodied Miocene hominoids have been discovered (Kunimatsu et al., Primate Res 16:299, 2000a). In this article, we describe a hominoid lower molar (CMu15-501) recovered from the Upper Lignite Member of Chiang Muan. The age of Chiang Muan is estimated to be latest Middle or earliest Late Miocene (around 11 Ma), based on the mammalian fauna.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Correlation of tooth size and body size in living hominoid primates, with a note on relative brain size in Aegyptopithecus and Proconsul.

Second molar length and body weight are used to test the correlation between tooth size and body size in living Hominoidea. These variates are highly correlated (r= 0.942, p less than 0.001), indicating that tooth size can be used in dentally unspecialized fossil hominoids as one method of predicting the average body weight of species. Based on tooth size, the average body weight of Aegyptopithecus zeuxis is estimated to have been beteen 4.5 and 7.5 kg, which is corroborated by known cranial and postcranial elements. Using Radinsky's estimates of brain size, the encephalization quotient (EQ) for Aegyptopithecus was between 0.65 and 1.04. A similar analysis for Proconsul africanus yields a body weight between 16 and 34 kg, and an EQ between 1.19 and 1.96.     

Polymorphic social organization in an ant

Identifying species exhibiting variation in social organization is an important step towards explaining the genetic and environmental factors underlying social evolution. In most studied populations of the ant Leptothorax acervorum, reproduction is shared among queens in multiple queen colonies (polygyny). By contrast, reports from other populations, but based on weaker evidence, suggest a single queen may monopolize all reproduction in multiple queen colonies (functional monogyny). Here we identify a marked polymorphism in social organization in this species, by conclusively showing that functional monogyny is exhibited in a Spanish population, showing that the social organization is stable and not purely a consequence of daughter queens overwintering, that daughter queen re-adoption is frequent and queen turnover is low. Importantly, we show that polygynous and functionally monogynous populations are not genetically distinct from one another based on mtDNA and nDNA. This suggests a recent evolutionary divergence between social phenotypes. Finally, when functionally monogynous and polygynous colonies were kept under identical laboratory conditions, social organization did not change, suggesting a genetic basis for the polymorphism. We discuss the implications of these findings to the study of reproductive skew.     

Fission-fusion social organization inAteles andPan

Recent research on the ecology and behavior of free-ranging spider monkeys (Ateles paniscus chamek) allows a more detailed comparison with the chimpanzee,Pan troglodytes, than has been possible previously. Despite their distant common ancestry, chimpanzees and spider monkeys share an unusual fission-fusion social system. In this paper, I compare subgroup size and composition, social unit structure, ranging behavior, patterns of philopatry and dispersal, and social relationships in the two taxa. It is proposed that spatial and temporal patchiness in food dispersion and abundance, resulting in a high-level of feeding competition between females within a group, has been the most important ecological selection pressure leading to the evolution of fission-fusion social organization in both species.     

The social organization of giraffes in Niger

The social organization of the last population of giraffes ( Giraffa camelopardalis ) in West Africa was studied between October 1996 and December 1997. Population size increased from 51 to 63 individuals during the study period. Groups were larger during the rainy season (mean group size 9.4) than during the dry season (mean group size 6; U = 4131; P < 0.01). Giraffes did not show strong preferential associations. Group types observed were similar to those expected on the assumption that associations are independent of sex and age. Inter-individual associations were low (mean simple ratio value = 0.1), with an individual associating with only half of its partners of the previous day. Young males interacted more frequently than other classes did. Agonistic interactions occurred mostly among males. Contact interactions (i.e. non-agonistic or sexual) occurred between both sexes and most age classes. Links between local communities, domestic animals, fauna and the environment suggest that environment and development cannot be dissociated in the sustainable exploitation of natural resources.     

Ecological polarity in primate tion

Relationships of the catarrhine primates based on morphological similarity are presented and then combined with habitat data to determine polarity of ecological change in the catarrhines. The ancestral habitat of two sister-groups is estimated by combining that of the sister groups, so that if they share one habitat type the ancestral condition is taken to be that type, or if they differ the ancestral condition is taken to be both alternatives. Analysed in this way the ancestral habitat preference of the Catarrhini is tropical lowland forest, and while the Hominoidea retain this primitive condition, the Cercopithecoidea are derived with a savanna habitat preference. Most hominoids retain the primitive forest habitat condition, and those groups that are associated with woodland-savanna habitats, notably the ramapitbecines and hominines, therefore share a derived habitat preference. There is no evidence, however, that this arose through common ancestry, and it is concluded that the functional similarities between the two groups could have arisen through parallel adaptation to the same habitat type. Similarly, many of the extant cercopithecoid groups that are now forest living may have re-entered forest independently. The derived savanna habitat preference of the cercopithecoids is linked with a number of morphological characters that also are derived with respect to catarrhine ancestry, and these include their terrestrial and/or their above-branch adaptations, and their specialized digestive, dietary and dental adaptations. In contrast, the hominoids that retain the primitive catarrhine habitat preference also retain the primitive condition in all these characters.     

社会性昆虫的组织及通讯行为

社会性昆虫在自然界有很重要的生物学意义,社会性昆虫具有3个主要特征,真社会性昆虫包括全部蚂蚁、白蚁及部分胡蜂、蜜蜂。它们具有明显的等级分化和个体分工。通过报警、集聚、化学招引、摇摆舞等方式,社会性昆虫在其内部做了很好的通讯交流从而使其社会组织得到了很好的发展。     

白蚁的社会组织和社会行为

白蚁属于等翅目昆虫,共有7科2 750种,这是昆虫纲中唯一的一个目,其所有的种类都靠消化道内共生的鞭毛类、原生动物利用和消化纤维素。介绍了白蚁社会组织和社会存为的特点,比较了白蚁与蚂蚁的异同,并指出两者相似是由于趋同进化,而两者不同是由于亲缘关系相距甚远,各来自不同祖先。     

Irrigation facilities and social organization in Taiwan

A new approach to the problem of linking social organization and the physical environment is claimed on the basis of an analysis of the relationship between the complexity of the irrigation facilities and the institutional differentiation of the service centers of a sample of townships in Taiwan. The hypothesized strong relationship holds, supporting the claim that the differentiation of institutions and technology can be interpreted by the same concept. The strategy is elaborated by showing that a wide range of data can be compressed in terms of differentiation and that the result holds when other measures of township organization are controlled and despite the blunting effect of the subregionwide irrigation association.     

Intra-specific variation in social organization of gorillas: implications for their social evolution

We analysed intra-specific variation in the social organization of gorillas and ecological and social factors influencing them, based on recent data on diet, day journey length, home range size, group size and proportion of multi-male groups in three subspecies [western lowland gorillas (WLG); eastern lowland gorillas (ELG); mountain gorillas (MG)]. Median group size was similar across subspecies and across habitats, but the extraordinarily large group including >30 gorillas was only found in habitat with dense terrestrial herbaceous vegetation. Within-group competition may determine the upper limit of group size in frugivorous WLGs and ELGs in lowland habitats with scarce undergrowth. A frugivorous diet may be a causal factor of subgrouping in multi-male groups of WLGs and ELGs, while a folivorous diet may prevent subgrouping in multi-male groups of MGs. Social factors, rather than ecological factors, may play an important role in the formation of multi-male groups and their cohesiveness in MGs. High gregariousness of female gorillas and their prolonged association with a protector male are explained by their vulnerability to both infanticide (MGs) and predators (ELGs). Comparison of long-term changes in group composition and individual movements between ELGs in Kahuzi and MGs in the Virungas suggest that the occurrence of infanticide may promote kin-male association within a group. Threat of infanticide may stimulate MG females to transfer into multi-male groups to seek reliable protection and maturing MG males to stay in their natal groups after maturity. By contrast, the absence of infanticide may facilitate ELG females to associate with infants and other females at transfer and ELG males to establish large groups in a short period by taking females from their natal groups, by luring females from neighbouring groups, or by takeover of a widow group after the death of its leading male. These conditions may prevent ELG and WLG maturing males from remaining to reproduce in their natal groups and possibly result in a rare occurrence of multi-male groups in their habitats. Similar reproductive features of MG and ELG females suggest both female strategies have been adaptive in their evolutionary history.     

The lorisiform wrist joint and the evolution of "brachiating" adaptations in the hominoidea.

In lorisines (Loris, Nycticebus, Perodicticus, Arctocebus), the tip of the ulna is reduced to the dimensions of a styloid process, a new and more proximal ulnar head is developed, and the pisiform is displaced distally away from its primitive contact with the ulna. In some Nycticebus, intra-articular tissues separate the ulna from the triquetrum. These traits are not seen in other quadrupedal primates, but they are characteristic of extant hominoids. Among hominoids, these features have been interpreted as adaptations to arm-swinging locomotion. Since hominoid-like features of the wrist joint are found in lorisines, but not in New World monkeys that practice arm-swinging locomotion, these features may have been evolved in both lorisines and large hominoids to enhance wrist mobility for cautious arboreal locomotion involving little or no leaping. Most of the other morphological traits characteristic of modern hominoids can be explained as adaptations to cautious quadrupedalism as well as to brachiation, and may have developed for different reasons in different lineages descended from an unspecialized cautious quadruped resembling Alouatta.     

Variation in social organization influences the opportunity for sexual selection in a social lizard

Social monogamy has traditionally been suggested to be maintained because of weak sexual selection on male partner acquisition. However, the ubiquitous incidence of extra-pair paternity suggests that sexual selection can be strong in monogamous systems, although studies partitioning variance in male reproductive success have come to mixed conclusions. Here, we use detailed field data to examine variance in male reproductive success and its implications for the maintenance of sociality in a population of the socially monogamous lizard Egernia whitii. We show that both within-pair and, to a lesser extent, extra-pair partner acquisition contribute to the variance in male reproductive success, resulting in considerable opportunity for sexual selection. Despite this, levels of multiple mating are lower in Egernia compared to other reptiles, suggesting that male partner acquisition is constrained. We suggest that this constraint may be a result of strong territoriality combined with sexual conflict over multiple mating generated by costs of extra-pair paternity to females as a result of facultative male care. This has the potential to limit sexual selection by reducing variance in male reproductive success and therefore contribute to the maintenance of complex social organization.     

Variation in hominoid molar enamel thickness

Enamel thickness has figured prominently in discussions of hominid origins for nearly a century, although little is known about its intra-taxon variation. It has been suggested that enamel thickness increases from first to third molars, perhaps due to varying functional demands or developmental constraints, but this has not been tested with appropriate statistical methods. We quantified enamel cap area (c), dentine area (b), and enamel-dentine junction length (e) in coronal planes of sections through the mesial and distal cusps in 57 permanent molars of Pan and 59 of Pongo, and calculated average (c/e) and relative enamel thickness (([c/e]/ radicalb) * 100). Posteriorly increasing or decreasing trends in each variable and average (AET) and relative enamel thickness (RET) were tested among molars in the same row. Differences between maxillary and mandibular analogues and between mesial and distal sections of the same tooth were also examined. In mesial sections of both genera, enamel cap area significantly increased posteriorly, except in Pan maxillary sections. In distal sections of maxillary teeth, trends of decreasing dentine area were significant in both taxa, possibly due to hypocone reduction. Significant increases in AET and RET posteriorly were found in all comparisons, except for AET in Pongo distal maxillary sections. Several significant differences were found between maxillary and mandibular analogues in both taxa. Relative to their mesial counterparts, distal sections showed increased enamel cap area and/or decreased dentine area, and thus increased AET and RET. This study indicates that when AET and RET are calculated from samples of mixed molars, variability is exaggerated due to the lumping of tooth types. To maximize taxonomic discrimination using enamel thickness, tooth type and section plane should be taken into account. Nonetheless, previous findings that African apes have relatively thinner enamel than Pongo is supported for certain molar positions.