The evolution of the hallucial tarsometatarsal joint in the Anthropoidea

Changes in the hallucial tarsometatarsal joint, which forms the fulcrum for the grasping hallux, have played a significant role in primate evolution. Comparative studies suggest that one of the morphological novelties heralding the attainment of a monkey grade of structure was the incorporation of the prehallux within this joint. Such a joint is found in the extant Ceboidea and, paradoxically, the Hylobatidae. Hallux and prehallux then form a composite distal articular surface; the proximal surface on the medial cuneiform is completed inferomedially by a convex facet for the prehallux. Divergence of the hallux into the attitude of opposition is accompanied by conjunct rotation, screwing these joint components into a stable, close-packed position. Suppression of the prehallux is accompanied by clear osteological indicators — the absence of prehallux facets on the first metatarsal and medial cuneiform. This modification of the joint is a feature of cercopithecoid evolution, and has also occurred in the hominoid line, after divergence of the ancestral gibbons and apparently after the Dryopithecus (Proconsul) stage. The cladistic relationships indicated by these morphological changes are in striking accord with recent results on primate evolution at the molecular level.     

A survey of prepollex and prehallux variation in anuran limbs

This paper reviews skeletal variation of the prepollex and prehallux among anurans and analyses their ontogeny in neotropical species. Morphological diversity is related to the number and features of distal elements. In some groups, clear phylogenetic trends may be interpreted from prepollical and/or prehallical reduction. However, specialized patterns converge from similar habit or behaviour. Developmental data are considered for discussing homology hypotheses and also to interpret evolutionary changes of anuran prepollex and prehallux morphologies. There is an apparent invariance in the presence of these structures that suggests the prepollex and prehallux were integrated, conserved and evolved in the plan of anuran limbs.     

An integrative approach to examining a homology question: shell structures in soft‐shell turtles

Understanding the patterns of shell reduction in turtles is relevant when examining both fossils and living forms. The soft‐shelled turtles (Trionychidae) are characterized by the general reduction of the peripheral bony elements of the carapace, and some species possess structures of contested homology. By examining Remane's ‘principal criteria’, we addressed the primary homology of the prenuchal and the posterior peripheral ossicles (= PPOs) of the Asian flapshell turtles, Lissemys spp., thus evaluating their topological equivalence, their structural quality, and the presence of intermediate forms in ontogeny and phylogeny. We conducted an analysis of gross morphology, bone histology, and ontogeny of these elements in a large sample of living and fossil trionychids and their sister‐group, the carettochelyids. We conclude that the prenuchal comprises a neomorphic structure that does not fulfil any of the homology criteria examined. The assessment of the homology of PPOs is less straightforward because of the presence of partly conflicting evidence. Nevertheless, PPOs and standard peripherals share an antero‐posterior polarity of the ossification pattern, which we interpret as a significant shared underlying developmental pattern. Depending on the phylogenetic position of Lissemys in trionychid phylogeny, the hypothesis of PPOs homology with standard peripherals is a straightforward one or, alternatively, one involving homologous developmental processes at other levels of the hierarchy, resulting in similar microstructural characteristics of these bony shell features. In this respect, we consider the antero‐posterior polarity of the ossification pattern of both PPOs and standard peripherals as providing potential evidence for the homology of the genetic control regulating the expression of both these structures, and therefore we interpret these structures as homologues on the basis of a deeply homologous underlying developmental process. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 462–476.     

A Light and Scanning Electron Microscopic Study of the Closing Apparatus in Tintinnid Ciliates (Ciliophora,Spirotricha, Tintinnina): A Forgotten Synapomorphy

ABSTRACT. A membranous closing apparatus shuts the lorica opening in disturbed tintinnids of six genera belonging to four families. The homology of the apparatuses is investigated, using data from the literature and Mediterranean tintinnids studied in vivo and by scanning electron microscopy. Morphological and functional similarities indicate that the foldable closing apparatus is not only a synapomorphy of the genera Codonella (Codonellidae) and Dictyocysta (Dictyocystidae), as suggested 80 years ago, but also of Codonaria (Codonellidae) and Codonellopsis (Codonellopsidae). In Codonaria, Codonella, and Dictyocysta, the apparatuses merge posteriorly into membranous lorica sacs, which probably represent homologous structures. The diagnoses of these genera are improved according to the new findings. The close relationship of Codonella, Codonellopsis, and Dictyocysta is also inferred from small subunit rRNA phylogenies and the ultrastructure of the capsules. It contradicts the current lorica‐based classification of the tintinnids. The assumption that the diaphragm‐like apparatus in the genera Salpingacantha and Salpingella is not homologous to the foldable ones in the genera mentioned above is supported by molecular and cytological features.     

Blood group genetic variations in inhabitants of West New Guinea, with a map of the villages and linguistic groups of South West New Guinea

In non-primates the nasal fossa manifests a posterior intrasphenoidal extension, accommodating the hindermost ethmoturbinals and an accompanying series of paranasal ectoturbinal cavities within the interorbital region. Primitive primates (e.g. Tarsius, Hapale, Saimiri) show phylogenetic loss of these cavities (together with the interorbital region and posterior ethmoturbinals) and the presence of an interorbital septum: a maxillary sinus only is present and the frontal and sphenoid bones are not pneumatized. In some phylogenetically advanced primates (e.g. Lagothrix, Alouatta) trends of cerebral enlargement and resultant separation of the orbits have induced some degree of reappearance of the lost interorbital territory, and an enhanced but variable pattern of peranasal pneumatization, involving the frontal and the sphenoid. Details of this pattern are described for Tarsius, Saimiri, Hapale, Cebus, Lagothrix and Alouatta, distinction being made between homologous and merely analogous paranasal cavities.     

Levels of the genealogical hierarchy and the problem of hominoid phylogeny

Molecular data are widely used to reconstruct phylogenetic relationships among species, and these phylogenies are often used as the basis for inferences about the history of evolutionary change in other nonmolecular characters. This approach is an appropriate and powerful one in many circumstances. But when several lineages diverge over a relatively short period of time, the assumption that a molecular (gene) tree will always be a valid basis for such inferences may not hold. Empirical evidence from humans, nonhuman primates, and other mammals indicates that the relationships among molecular divergence, morphological differentiation, and the origin of reproductive isolation between diverging lineages are complex. The simple dichotomously branching trees that result from molecular systematic studies of Homo, Gorilla, and Pan may be a misleading basis for reconstructions of evolutionary change in nonmolecular characters. © 1994 Wiley-Liss, Inc.     

A comparison of proximal humeral cancellous bone of great apes and humans

The shoulder is the most mobile joint in the primate body, and is involved in both locomotor and manipulative activities. The presumed functional sensibility of trabecular bone can offer a way of decoding the activities to which the forelimbs of fossil primates were subjected. We examine the proximal humeral trabecular architecture in a relatively closely related group of similarly sized hominids (Pongo pygmaeus, Pan troglodytes, and Homo sapiens), in order to evaluate the effect of diverging habitual motion behaviors of the shoulder complex in a coherent phylogenetic group. In order to characterize and compare the humeral trabecular architectures of the three species, we imaged a large sample by high-resolution computed tomography (HrCT) and quantified their trabecular architectures by standard bone 3D morphometric parameters. Univariate statistical analysis was performed, showing significant differences among the species. However, univariate statistics could not highlight the structural particularity in the cancellous bone of each species. A principal component analysis also showed clear separation of the three taxa and enabled a structural characterization of the humeral trabecular bone of each species. We conclude that the differences in the architectural setup of the three hominids likely reflect multiple differences in their habitual activity patterns of their shoulder joint, although individual structural features are difficult to relate to specific loading conditions.     

The evolution of actinorhizal symbioses: Evidence for multiple origins of the symbiotic association

According to morphologically based classification systems, actinorhizal plants, engaged in nitrogen-fixing symbioses with Frankia bacteria, are considered to be only distantly related. However, recent phylogenetic analyses of seed plants based on chloroplast rbcL gene sequences have suggested closer relationships among actinorhizal plants. A more thorough sampling of chloroplast rbcL gene sequences from actinorhizal plants and their nonsymbiotic close relatives was conducted in an effort to better understand the phylogenetic relationships of these plants, and ultimately, to assess the homology of the different actinorhizal symbioses. Sequence data from 70 taxa were analyzed using parsimony analysis. Strict consensus trees based on 24 equally parsimonious trees revealed evolutionary divergence between groups of actinorhizal species suggesting that not all symbioses are homologous. The arrangement of actinorhizal species, interspersed with nonactinorhizal taxa, is suggestive of multiple origins of the actinorhizal symbiosis. Morphological and anatomical characteristics of nodules from different actinorhizal hosts were mapped onto the rbclL-based consensus tree to further assess homology among rbcL-based actinorhizal groups. The morphological and anatomical features provide additional support for the rbcL-based groupings, and thus, together, suggest that actinorhizal symbioses have originated more than once in evolutionary history.     

Point process models,the dimensions of biodiversity and the importance of small-scale biotic interactions

Aims Recent mechanistic explanations for community assembly focus on the debates surrounding niche-based deterministic and dispersal-based stochastic models. This body of work has emphasized the importance of both habitat filtering and dispersal limitation, and many of these works have utilized the assumption of species spatial independence to simplify the complexity of the spatial modeling in natural communities when given dispersal limitation and/or habitat filtering. One potential drawback of this simplification is that it does not consider species interactions and how they may influence the spatial distribution of species, phylogenetic and functional diversity. Here, we assess the validity of the assumption of species spatial independence using data from a subtropical forest plot in southeastern China.Methods We use the four most commonly employed spatial statistical models—the homogeneous Poisson process representing pure random effect, the heterogeneous Poisson process for the effect of habitat heterogeneity, the homogenous Thomas process for sole dispersal limitation and the heterogeneous Thomas process for joint effect of habitat heterogeneity and dispersal limitation—to investigate the contribution of different mechanisms in shaping the species, phylogenetic and functional structures of communities.Important findings Our evidence from species, phylogenetic and functional diversity demonstrates that the habitat filtering and/or dispersal-based models perform well and the assumption of species spatial independence is relatively valid at larger scales (50×50 m). Conversely, at local scales (10×10 and 20×20 m), the models often fail to predict the species, phylogenetic and functional diversity, suggesting that the assumption of species spatial independence is invalid and that biotic interactions are increasingly important at these spatial scales.     

Unique proximal tibial morphology in strepsirrhine primates

Although the morphology of the tibial plateau in primates has received very little attention in the literature, it does exhibit features of phylogenetic and functional interest. This paper describes the morphology of the tibial plateau (particularly the intercondylar region) in extant and fossil primates, and in three mammalian outgroups: the pen-tailed tree shrew (Ptilocercus), tree shrew (Tupaia), and flying lemur or dermopteran (Cynocephalus). Extant and fossil strepsirrhine primates exhibit an eminence with a single spine, which contrasts with the intercondylar morphology of haplorhine primates. Most extant platyrrhines, all catarrhine primates (including humans), and some fossil haplorhines possess an eminence with two spines (medial and lateral) connected by a ridge of bone that intersects the intercondylar groove. Tarsius and callitrichines possess an eminence with a reduced medial spine that superficially resembles that of strepsirrhine primates. Dermopterans also exhibit a morphology similar to that of strepsirrhines. In Scandentia, the intercondylar morphology of Tupaia is similar to that of rodents, whereas Ptilocercus resembles tarsiers and callitrichines. We hypothesize that proximal tibiae with either a single spine or reduced medial spine morphology facilitate a greater degree of knee rotation about the eminence relative to the double-spine condition, and are likely associated with more frequent adoption of vertical body positions. In contrast, a double-spine eminence limits knee rotation and is probably associated with greater use of horizontal supports. Although the polarity is complicated by the unknown phylogenetic status of likely sister taxa, it seems most probable that the single-spine morphology is a derived feature of strepsirrhines.     

Assessing similarity: on homology,characters and the need for a semantic approach to non‐evolutionary comparative homology

The problem of homology has been a consistent source of controversy at the heart of systematic biology, as has the step of morphological character analysis in phylogenetics. Based on a clear epistemic framework and a characterization of “characters” as diagnostic evidence units for the recognition of not directly identifiable entities, I discuss the ontological definition and empirical recognition criteria of phylogenetic, developmental and comparative homology, and how these three accounts of homology each contribute to an understanding of the overall phenomenon of homology. I argue that phylogenetic homologies are individuals or historical kinds that require comparative homology for identification. Developmental homologies are natural kinds that ultimately rest on phylogenetic homologies and also require comparative homology for identification. Comparative homologies on the other hand are anatomical structural kinds that are directly identifiable. I discuss pre‐Darwinian comparative homology concepts and their problem of invoking non‐material forces and involving the a priori assumption of a stable positional reference system. Based on Young's concept of comparative homology, I suggest a procedure for recognizing comparative homologues that lacks these problems and that utilizes a semantic framework. This formal conceptual framework provides the much needed semantic transparency and computer‐parsability for documenting, communicating and analysing similarity propositions. It provides an essential methodological framework for generalizing over individual organisms and identifying and demarcating anatomical structural kinds, and it provides the missing link to the logical chain of identifying phylogenetic homology. The approach substantially increases the analytical accessibility of comparative research and thus represents an important contribution to the theoretical and methodological foundation of morphology and comparative biology.     

Trabecular bone structure in the mandibular condyles of gouging and nongouging platyrrhine primates

The relationship between mandibular form and biomechanical function is a topic of significant interest to morphologists and paleontologists alike. Several previous studies have examined the morphology of the mandible in gouging and nongouging primates as a means of understanding the anatomical correlates of this feeding behavior. The goal of the current study was to quantify the trabecular bone structure of the mandibular condyle of gouging and nongouging primates to assess the functional morphology of the jaw in these animals. High‐resolution computed tomography scan data were collected from the mandibles of five adult common marmosets (Callithrix jacchus), saddle‐back tamarins (Saguinus fuscicollis), and squirrel monkeys (Saimiri sciureus), respectively, and various three‐dimensional morphometric parameters were measured from the condylar trabecular bone. No significant differences were found among the taxa for most trabecular bone structural features. Importantly, no mechanically significant parameters, such as bone volume fraction and degree of anisotropy, were found to vary significantly between gouging and nongouging primates. The lack of significant differences in mechanically relevant structural parameters among these three platyrrhine taxa may suggest that gouging as a habitual dietary behavior does not involve significantly higher loads on the mandibular condyle than other masticatory behaviors. Alternatively, the similarities in trabecular architecture across these three taxa may indicate that trabecular bone is relatively unimportant mechanically in the condyle of these primates and therefore is functionally uninformative. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Integration, phylogeny, and the hominid cranial base

Basicranial features were examined in catarrhine primates and early hominids in order to demonstrate how information about morphological integration can be incorporated into phylogenetic analysis. Hypotheses purporting to explain the functional and structural relationships of basicranial characters were tested using factor analysis. Characters found to be functionally or structurally related to each other were then further examined in order to determine whether there was evidence that they were phylogenetically independent. If phylogenetic independence could not be demonstrated, then the characters were presumed to be integrated and were grouped into a complex. That complex was then treated as if it were a single character for the purposes of cladistic analysis. Factor analysis revealed that five basicranial features may be structurally related to relative brain size in hominoids. Depending on how one defines phylogenetic independence, as few as two, or as many as all of those characters might be morphologically integrated. A cladistic analysis of early hominids based on basicranial features revealed that the use of integrated complexes had a substantial effect on the phylogenetic position of Australopithecus africanus, a species whose relationships are poorly resolved. Moreover, the use of complexes also had an effect on reanalyses of certain published cladistic data sets, implying that those studies might have been biased by patterns of basicranial integration. These results demonstrate that patterns of morphological integration need to be considered carefully in all morphology-based cladistic analyses, regardless of taxon or anatomical focus. However, an important caveat is that the functional and structural hypotheses tested here predicted much higher degrees of integration than were observed. This result warns strongly that hypotheses of integration must be tested before they can be adequately employed in phylogenetic analysis. The uncritical acceptance of an untested hypothesis of integration is likely to be as disruptive to a cladistic analysis as when integration is ignored.     

Morphology of locomotor appendages in Arachnida: evolutionary trends and phylogenetic implications

The morphological diversity of locomotor appendages in Arachnida is surveyed lo reconstruct phylogenetic relationships and discover evolutionary trends in form and function. The appendicular skeleton and musculature of representatives from the ten living arachnid orders ate described, and a system of homology is proposed. Character polarities are established through comparison with an outgroup. Limulus polyphemus Xiphosura). Cladistic analysis suggests that Arachnida is monophyletic and that absence of extensor muscles is a primitive condition. Extensors are primitively absent in Araneae. Amblypygi, Uropygi, Palpigradi, Ricinulei and Acari. Most similarities in the appendages of these orders are symplesiomorphic so that phylogenetic relationships among the ‘extensorless’ groups cannot be resolved solely on the basis of appendicular characters. Extensor muscles appear to have evolved once, and their presence is considered a synapomorphic feature of Opiliones, Scorpiones, Pseudoscorpiones and Solifugae. Solifugae lack extensors, but a parsimonious interpretation of other characters indicates that this is a secondary, derived condition. The phylogenetic relationships among these four orders are clarified by modifications of the patellotibial joint. Cladistic analysis indicates that Opiliones may be the sister group of the other three orders and that Scorpiones is the sister group of Pseudoscorpiones and Solifugae. Conclusions concerning phylogenetic relationships and evolutionary morphology presented here differ substantially from those of earlier studies on the locomotor appendages of Arachnida.     

Presence of punctae in the ‘plectorthoidean’ brachiopod Famatinorthis turneri (Middle Ordovician) from western Argentina: implications for early diversification of punctate orthides

Benedetto, J.L. 2013: Presence of punctae in the ‘plectorthoidean’ brachiopod Famatinorthis turneri (Middle Ordovician) from western Argentina: implications for early diversification of punctate orthides. Lethaia, Vol. 46, pp. 170–179. Famatinorthis Levy & Nullo is a distinctive orthide brachiopod of Dapingian age from the volcaniclastic rocks of the Famatina Range of western Argentina. Although it was originally classified among the plectorthoideans, a new collection from the La Escondida Formation has yielded exceptionally well‐preserved moulds of Famatinorthis turneri in which silicified infillings of punctae are clearly visible, leading to the reassignment of the genus to the dalmanellidines. In this paper, phylogenetic analyses are used to determine the evolutionary relationships of Famatinorthis, the Tremadocian linoporellid Lipanorthis, and other Ordovician Gondwanan genera. The placement of Plectorthoidea in the same major clade as linoporellids, and the separation of Dalmanellidina as an independent clade are the most important features of all shortest trees, supporting the idea that linoporellids may have originated from a plectorthoid ancestor. Cladistic analysis reveals that Lipanorthis lies close to the ancestry of the linoporellid lineage, and Famatinorthis clusters within the more derived taxa of the clade with which it shares a large septalium. It seems that the presence of endopunctae in the orthides does not necessarily indicate close phylogenetic relationships as it could have occurred at different times in different clades. If the homoplasic nature of endopunctae in the order Orthida is supported by further morphologic and phylogenetic studies, the fundamental division of orthides in non‐punctate (Orthidina) and punctate (Dalmanellidina) may need revision. □Brachiopods, Ordovician, Gondwana, Famatina, phylogeny, punctate orthides.     

Phylogenetic analysis of Cyprichromini (Perciformes: Cichlidae) endemic to Lake Tanganyika and validation of the genus <Emphasis Type="Italic">Paracyprichromis</Emphasis>

The phylogenetic relationships among two Paracyprichromis and five Cyprichromis species, included in the Tanganyikan cichlid tribe Cyprichromini, were investigated using morphological features. The previously proposed diagnostic characters of Paracyprichromis are not synapomorphies, because the nonelongated swim bladder is plesiomorphic, the numbers of dorsal and anal fin rays and scales on longitudinal line and around the caudal peduncle overlap with those of Cyprichromis, and these counts and number of vertebrae are all included within the ranges of other Tanganyikan cichlids. The monophyly of Paracyprichromis is supported by a unique condition of infraorbitals to this genus. Additionally, the monophyly of Cyprichromis was reconfirmed by one of the previously proposed diagnostic characters, the presence of an elongated swim bladder.     

Functional morphology of the palato‐maxillary apparatus in “Palatine dragging” snakes (Serpentes: Elapidae: Acanthophis,Oxyuranus)

Elapid snakes have previously been divided into two groups (palatine erectors and palatine draggers) based on the morphology and inferred movements of their palatine bone during prey transport (swallowing). We investigated the morphology and the functioning of the feeding apparatus of several palatine draggers (Acanthophis antarcticus, Oxyuranus scutellatus, Pseudechis australis) and compared them to published records of palatine erectors. We found that the palatine in draggers does not move as a straight extension of the pterygoid as originally proposed. The dragger palato‐pterygoid joint flexes laterally with maxillary rotation when the mouth opens and the jaw apparatus is protracted and slightly ventrally during mouth closing. In contrast, in palatine erectors, the palato‐pterygoid joint flexes ventrally during upper jaw protraction. In draggers, the anterior end of the palatine also projects rostrally during protraction, unlike the stability of the anterior end seen in erectors. Palatine draggers differ from palatine erectors in four structural features of the palatine and its relationships to surrounding elements. The function of the palato‐pterygoid bar in both draggers and erectors can be explained by a typical colubroid muscle contraction pattern, which acts on a set of core characters shared among all derived snakes. Although palatine dragging elapids share a fundamental design of the palato‐maxillary apparatus with all higher snakes, they provide yet another demonstration of minor structural modifications producing functional variants. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Bone histology of Silesaurus opolensisDzik, 2003 from the Late Triassic of Poland

Fostowicz‐Frelik, ?. & Sulej, T. 2009: Bone histology of Silesaurus opolensisDzik, 2003 from the Late Triassic of Poland. Lethaia, Vol. 43, pp. 137–148. The phylogenetic relationships of Silesaurus opolensis have been the subject of intense debate since its discovery. Silesaurus possesses some features characteristic of ornithischian dinosaurs, such as the presence of a beak at the front of the lower jaw, yet it lacks a number of important femoral and dental synapomorphies of Dinosauria. The microstructure of the long bones (femur, tibia and metatarsal) and ribs of this species reveals a relatively intensive rate of growth, comparable with that seen in small dinosaurs and the gracile crocodylomorph Terrestrisuchus. Cortical bone formed mainly by periosteal tissue with fibro‐lamellar matrix (in older specimens parallel fibred) shows very little secondary remodelling and only in one specimen (large tibia ZPAL Ab III/1885) few lines of arrested growth are present in the outermost cortex. The vascularization is relatively dense, mainly longitudinal and ceases towards the periphery, forming almost avascular parallel fibred bone at the bone surface. This indicates maturation and significant decrease in the growth ratio in mature specimens of S. opolensis. The delicate trabeculae exhibit cores formed by the primary cancellous tissue lined with lamellar endosteal bone. The rather intense growth of S. opolensis implies a relatively high metabolic rate. Moreover, evidence from the fibro‐lamellar tissue, predominant in the cortex, suggests that this kind of rapid bone deposition could be more typical of Archosauria than previously assumed, a prerequisite for the evolution of the very fast growth rates observed in large ornithischians, sauropods and large theropods. □Archosauria, Bone histology, Dinosauriformes, Late Triassic, Silesaurus opolensis.