Evidence for a grooming claw in a North American adapiform primate: implications for anthropoid origins

Among fossil primates, the Eocene adapiforms have been suggested as the closest relatives of living anthropoids (monkeys, apes, and humans). Central to this argument is the form of the second pedal digit. Extant strepsirrhines and tarsiers possess a grooming claw on this digit, while most anthropoids have a nail. While controversial, the possible presence of a nail in certain European adapiforms has been considered evidence for anthropoid affinities. Skeletons preserved well enough to test this idea have been lacking for North American adapiforms. Here, we document and quantitatively analyze, for the first time, a dentally associated skeleton of Notharctus tenebrosus from the early Eocene of Wyoming that preserves the complete bones of digit II in semi-articulation. Utilizing twelve shape variables, we compare the distal phalanges of Notharctus tenebrosus to those of extant primates that bear nails (n = 21), tegulae (n = 4), and grooming claws (n = 10), and those of non-primates that bear claws (n = 7). Quantitative analyses demonstrate that Notharctus tenebrosus possessed a grooming claw with a surprisingly well-developed apical tuft on its second pedal digit. The presence of a wide apical tuft on the pedal digit II of Notharctus tenebrosus may reflect intermediate morphology between a typical grooming claw and a nail, which is consistent with the recent hypothesis that loss of a grooming claw occurred in a clade containing adapiforms (e.g. Darwinius masillae) and anthropoids. However, a cladistic analysis including newly documented morphologies and thorough representation of characters acknowledged to have states constituting strepsirrhine, haplorhine, and anthropoid synapomorphies groups Notharctus tenebrosus and Darwinius masillae with extant strepsirrhines rather than haplorhines suggesting that the form of pedal digit II reflects substantial homoplasy during the course of early primate evolution.     

An articulated titanosaur from Patagonia (Argentina): New evidence of neosauropod pedal evolution

Most titanosaur dinosaurs are represented by incomplete skeletal elements lacking articulated pes. An exceptionally preserved specimen from the Late Campanian–Early Maastrichtian strata of Patagonia (Argentina) provides new data on pedal morphology and the evolutionary trends of these huge dinosaurs. This finding is one of the few articulated titanosaur pes known in the world, and shows a phalangeal formula of 2-2-2-2-0. The first three digits possess sickle-shaped claws and the articular facets of ungual phalanges, suggesting mobility in horizontal and vertical planes. A comparative analysis of available record suggests that titanosaurs had a progressive reduction of size and number of pedal phalanges in digits III and IV during the Late Cretaceous.     

Post-autotomy claw regrowth and functional recovery in the snapping shrimp Alpheus angulosus

The ability to regenerate lost tissues, organs or whole body parts is widespread across animal taxa; in some animals, regeneration includes transforming a remaining structure to replace the one that was lost. The transformation of one limb into another involves considerable plasticity in morphology, physiology and behavior, and snapping shrimp offer excellent opportunities for studying this process. We examined the changes required for the transformation of the small pincer to a mature snapping claw in Alpheus angulosus. First molt claws differ from mature claws in overall shape as well as in morphology related to snapping function; nonetheless, shrimp with first molt claws do produce snaps. While most shape variables of second molt claws do not differ significantly from mature claws, the plunger (structure required for snap production) does not reach mature size until the third molt for females, or later for males. Thus, the pincer claw can be transformed into a functional snapping claw in one molt, although both the underlying morphology and superficial shape are not fully regenerated at this stage. The rapid production of a functional snapping claw that we observe in this study suggests that this particular function is of significant importance to snapping shrimp behavior and survival.     

Claw retraction and protraction in the carnivora: Skeletal microvariation in the phalanges of the Felidae

All carnivorans retract and protract their claws. In felids and some viverrids the claws of digits II through V of both the manus and pes have a larger arc of rotation than those of other carnivorans; the claws retract to the lateral side of the middle phalanx rather than onto its dorsal surface as in most other carnivorans. This condition should be termed hyper-retraction. Morphological features of the middle and distal (ungual) phalanges that have been purported to be necessary for hyper-retraction in felids vary considerably among digits within the manus and pes. These features include the lateral projection of the distal head and the asymmetry of the shaft of the middle phalanx, and the oblique orientation of the articular surface on the distal phalanx. None of these features is necessary in every instance for hyperretraction, and some of the variation in these features is associated instead with protraction. Differences among digits in the orientation of the articular surface on the distal phalanx are associated with differences in the degree to which the claws must move laterally to rotate from the protracted to the retracted position. Differences in the orientation of the distal head on the middle phalanx are associated with the spreading of the claws during protraction. The manual claws are hook-shaped, whereas the pedal claws are more blade-like; this morphological difference is associated with differences in function between the manus and pes. In the manus the medial claws have a larger radius of curvature and a smaller angle of arc as compared to the more lateral claws; in the pes, the claws on digits III and IV have larger radii of curvature and smaller angles of arc. Digit I of the manus lacks the hyper-retraction mechanism; nonetheless, this digit shares many of the attributes that are associated with this mechanism. © 1996 Wiley-Liss, Inc.     

Morphological and functional diversity in therizinosaur claws and the implications for theropod claw evolution

Therizinosaurs are a group of herbivorous theropod dinosaurs from the Cretaceous of North America and Asia, best known for their iconically large and elongate manual claws. However, among Therizinosauria, ungual morphology is highly variable, reflecting a general trend found in derived theropod dinosaurs (Maniraptoriformes). A combined approach of shape analysis to characterize changes in manual ungual morphology across theropods and finite-element analysis to assess the biomechanical properties of different ungual shapes in therizinosaurs reveals a functional diversity related to ungual morphology. While some therizinosaur taxa used their claws in a generalist fashion, other taxa were functionally adapted to use the claws as grasping hooks during foraging. Results further indicate that maniraptoriform dinosaurs deviated from the plesiomorphic theropod ungual morphology resulting in increased functional diversity. This trend parallels modifications of the cranial skeleton in derived theropods in response to dietary adaptation, suggesting that dietary diversification was a major driver for morphological and functional disparity in theropod evolution.     

Evolutionary and developmental aspects of phalangeal formula variation in pig-nose and soft-shelled turtles (Carettochelyidae and Trionychidae)

In order to examine the evolution of the phalangeal formula in a diverse clade of turtles, including hyperphalangy as a rare condition in this group, we studied 210 specimens representing all extant genera of Trionychidae and their sister taxon, Carettochelyidae. Both groups consist of highly aquatic species with elongated autopods that are either paddle-like (Trionychidae) or transformed to flippers (Carettochelyidae). Phalangeal formulae were obtained mostly by radiographs of alcohol-preserved or dry specimens, as well as by direct counts from skeletons. All trionychids and Carettochelys are pentadactylous, but their phalangeal formulae differ. Carettochelys exhibits the turtle-plesiomorphic state (manus and pes: 2-3-3-3-3), with no variation in adults. Trionychids exhibit intraspecific variation, ranging from 2-3-3-3-2 to 2-3-3-6-5 for the manus, and from 2-3-3-3-2 to 2-3-3-5-3 for the pes. The extant Carettochelys as well as the Middle Eocene Allaeochelys crassesculpta are characterized by an elongation of phalanges, whereas trionychids consistently have shorter phalanges. All trionychid genera exhibit some degree of hyperphalangy in digits IV and V, in both the manus and pes. Phalanges of the clawed digits I–III are very robust compared to phalanges of the non-clawed digits IV and V. The latter contribute significantly to the enlargement of the paddle by their additional phalanges. We hypothesize that this phalangeal pattern is coupled with prolongation of growth processes in the non-clawed digits. The differences in autopod morphology between carettochelyids and trionychids reflect different locomotor patterns related to different natural histories (elongated flippers for high-speed escape in the mainly herbivorous Carettochelys; broad paddles for rapid turns during hunting in the mainly carnivorous trionychids). The autopod of Pelodiscus sinensis is proposed as an experimental model to examine the developmental basis of adult autopod variation.     

Ungual asymmetry in the context of pedal symmetry in Ailuronyx (Reptilia Gekkonidae): modification for an opposable grip

The claws of the gekkonine gecko genus Ailuronyx are unique in being asymmetrically disposed. Those of digits I-III are curved towards the body, whereas those of digits IV and V curve away from the body. This ungual asymmetry is imposed upon the essentially symmetrical pedal morphology that typifies pad-bearing geckos. The resultant functional organization of manus and pes parallels that seen in chameleons. The pedal configuration of Ailuronyx seychellensis appears to provide a mechanical advantage for locomotion on at least some of the animal's natural substrata.     

THE DESIGN OF A BEAUTIFUL WEAPON: COMPENSATION FOR OPPOSING SEXUAL SELECTION ON A TRAIT WITH TWO FUNCTIONS

Male fiddler crabs, genus Uca, have one greatly enlarged claw with which they court females and threaten and fight other males. Longer claws are more effective signals but are thought to be less effective weapons because the relative closing force at the tip of the claw decreases with claw length. We studied claw morphology and fighting in Uca terpsichores and Uca beebei and found a mechanism that may resolve opposing selection for signaling and fighting ability. When males fought they delivered gripping forces not at the tips but at the tubercles on the inner margins of their claws’ fingers. As claws grow, these tubercles remain relatively close to the apex of the gape. Consequently, the mechanical advantage that governs the forces that can be delivered at these tubercles decreases only slightly with increasing claw length allowing the claw to be an effective signal and a powerful weapon. Animal weapons are exceptionally diverse in form and detail of armature and the causes of this diversity are poorly understood. We suggest that the designs of weapons may often reflect compensatory patterns of growth and placement of armature that enhances the weapon's overall utility for multiple uses in competition for mates.     

Metatarsophalangeal joint function and positional behavior in Australopithecus afarensis

Recent discussions of the pedal morphology of Australopithecus afarensis have led to conflicting interpretations of australopithecine locomotor behavior. We report the results of a study using computer aided design (CAD) software that provides a quantitative assessment of the functional morphology of australopithecine metatarsophalangeal joints. The sample includes A. afarensis, Homo sapiens, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus. Angular measurements of the articular surfaces relative to the long axes of the metatarsals and phalanges were taken to determine whether the articular surfaces are plantarly or dorsally oriented. Humans have the most dorsally oriented articular surfaces of the proximal pedal phalanges. This trait appears to be functionally associated with dorsiflexion during bipedal stride. Pongo has the most plantarly oriented articular surfaces of the proximal pedal phalanges, probably reflecting an emphasis on plantarflexion in arboreal positional behaviors, while the African hominoids are intermediate between Pongo and Homo for this characteristic. A. afarensis falls midway between the African apes and humans. Results from an analysis of metatarsal heads are inconclusive with regard to the functional morphology of A. afarensis. Overall, the results are consistent with other evidence indicating that A. afarensis was a capable climber. © 1994 Wiley-Liss, Inc.     

First report of dinosaurian claws from the Late Triassic of India

The Late Triassic Tiki Formation has yielded five isolated nearly complete claws or ungual phalanges from a fossil locality, which are described in detail and compared with other Late Triassic tetrapods. Of these, four ungual phalanges are slender, asymmetric, ventrally recurved, transversely compressed, and contain deep collateral grooves on either side, a low median keel on the proximal articular surface and a prominent proximoventral flexor tubercle showing their high similarity to the theropod dinosaurs. The remaining claw is unlike that of any theropods in terms of high robusticity and near symmetry. However, as in dinosaurs it is ventrally recurved and contains deep lateral grooves, a small flexor tubercle, lateromedially extended proximal articular surface with a distinct median keel and is considered as belonging to an indeterminate dinosaur. Although it is not possible to ascertain whether the unguals belong to a single taxon or multiple taxa, this new find points towards the presence of small dinosaurs in the Late Triassic Tiki fauna.     

Claw morphometrics in monitor lizards: Variable substrate and habitat use correlate to shape diversity within a predator guild

Numerous studies investigate morphology in the context of habitat, and lizards have received particular attention. Substrate usage is often reflected in the morphology of characters associated with locomotion, and, as a result, claws have become well‐studied ecomorphological traits linking the two. The Kimberley predator guild of Western Australia consists of 10 sympatric varanid species. The purpose of this study was to quantify claw size and shape in the guild using geometric morphometrics, and determine whether these features correlated with substrate use and habitat. Each species was assigned a Habitat/substrate group based on the substrate their claws interact with in their respective habitat. Claw morphometrics were derived for both wild caught and preserved specimens from museum collections, using a 2D semilandmark analysis. Claw shape significantly separated based on Habitat/substrate group. Varanus gouldii and Varanus panoptes claws were associated with sprinting and extensive digging. Varanus mertensi claws were for shallow excavation. The remaining species’ claws reflected specialization for some form of climbing, and differed based on substrate compliance. Varanus glauerti was best adapted for climbing rough sandstone, whereas Varanus scalaris and Varanus tristis had claws ideal for puncturing wood. Phylogenetic signal also significantly influenced claw shape, with Habitat/substrate group limited to certain clades. Positive size allometry allowed for claws to cope with mass increases, and shape allometry reflected a potential size limit on climbing. Claw morphology may facilitate niche separation within this trophic guild, especially when considered with body size. As these varanids are generalist predators, morphological traits associated with locomotion may be more reliable candidates for detecting niche partitioning than those associated directly with diet.     

A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu,Inner Mongolia

Abstract: A new oviraptorid is described on the basis of a partial forelimb collected from the Upper Cretaceous redbeds of Bayan Mandahu, Inner Mongolia. Machairasaurus leptonychus, gen. et sp. nov. is diagnosed by slender, weakly curved manual unguals, reduced flexor tubercles, penultimate phalanges that are subequal in length to the preceding phalanges, and short, robust manual digits. Machairasaurus is found to be a member of the Ingeniinae, along with Ingenia yanshini, Heyuannia huangi, Conchoraptor gracilis, and Nemegtomaia barsboldi. Machairasaurus exhibits unusual proportions of the manus, suggesting that the manus was not primarily used to grasp prey. Instead, Machairasaurus and other oviraptorids are likely to have fed largely on plant material. The recognition of a previously unknown oviraptorid at Bayan Mandahu provides further evidence that the Bayan Mandahu dinosaur assemblage is distinct from that found at the Djadokhta Formation exposures at Bayn Dzak, Tugriken Shireh, and Ukhaa Tolgod. Given that these localities are separated by just a few hundred kilometres and represent similar palaeoenvironments, marked differences in the fauna suggest that the Bayan Mandahu Formation of Inner Mongolia is not coeval with the known Djadokhta localities in Mongolia.     

An unusual pattern of limb morphology in the Tithonian marine turtle Neusticemys neuquina from the Vaca Muerta Formation,Neuquén Basin,Argentina

de la Fuente M.S. & Fernández, M.S. 2010: An unusual pattern of limb morphology in the Tithonian marine turtle Neusticemys neuquina from the Vaca Muerta Formation, Neuquén Basin, Argentina. Lethaia, Vol. 44, pp. 15–25. Here, we report an unusual pattern in the manus and pes morphology of the Tithonian marine turtle Neusticemys neuquina. We analyse the forelimbs of two previously known specimens and describe the hind limbs of two previously undescribed specimens. Neusticemys neuquina is characterized by a relative elongation of both the forelimb and hind limb, compared with stem Chelonoidea, as well as an elongation of the pedal digit V, achieved through the elongation of the bones, as well as a moderate hyperphalangy. The elongation of pedal digit V is the most striking feature of N. neuquina, a feature unknown in other turtles. □Limb morphology, Neusticemys neuquina, Tithonian turtle.     

The origin and early evolution of birds

Birds evolved from and are phylogenetically recognized as members of the theropod dinosaurs; their first known member is the Late Jurassic Archaeopteryx, now represented by seven skeletons and a feather, and their closest known non-avian relatives are the dromaeosaurid theropods such as Deinonychus. Bird flight is widely thought to have evolved from the trees down, but Archaeopteryx and its outgroups show no obvious arboreal or tree-climbing characters, and its wing planform and wing loading do not resemble those of gliders. The ancestors of birds were bipedal, terrestrial, agile, cursorial and carnivorous or omnivorous. Apart from a perching foot and some skeletal fusions, a great many characters that are usually considered ‘avian’ (e.g. the furcula, the elongated forearm, the laterally flexing wrist and apparently feathers) evolved in non-avian theropods for reasons unrelated to birds or to flight. Soon after Archaeopteryx, avian features such as the pygostyle, fusion of the carpometacarpus, and elongated curved pedal claws with a reversed, fully descended and opposable hallux, indicate improved flying ability and arboreal habits. In the further evolution of birds, characters related to the flight apparatus phylogenetically preceded those related to the rest of the skeleton and skull. Mesozoic birds are more diverse and numerous than thought previously and the most diverse known group of Cretaceous birds, the Enantiornithes, was not even recognized until 1981. The vast majority of Mesozoic bird groups have no Tertiary records: Enantiornithes, Hesperornithiformes, Ichthyornithiformes and several other lineages disappeared by the end of the Cretaceous. By that time, a few Linnean ‘Orders’ of extant birds had appeared, but none of these taxa belongs to extant ‘families’, and it is not until the Paleocene or (in most cases) the Eocene that the majority of extant bird ‘Orders’ are known in the fossil record. There is no evidence for a major or mass extinction of birds at the end of the Cretaceous, nor for a sudden ‘bottleneck’ in diversity that fostered the early Tertiary origination of living bird ‘Orders’.     

Pincer-like claws in centipedes (Chilopoda): multiple evolutionary origin of similar form and serial pattern

In most Chilopoda, the walking legs end in a single-tip claw usually accompanied by short accessory spines. Instead, in all species of three small and only distantly related geophilomorph taxa (Diphyonyx, Neogeophilidae, Eucratonyx), the claws of an anterior set of leg pairs are unusually pincer-like. By integrating different microscopic techniques, including confocal laser scanning microscopy, we found that these modified claws are very similar in form, internal structure, and pattern of variation in shape along the trunk in all three taxa: the claws are distinctly swollen and bent, provided with peculiar bulges, and flanked by a conspicuous additional branch, either cylindrical or flattened, which overreaches the tip of the claw; instead, the internal cuticular features are not modified with respect to the condition in the other centipedes, claiming against the possibility of controlled abduction/adduction between claw and branch. Irrespective of the total number of leg pairs (63–129), the claws change gradually from pincer-like to usual shape invariantly in the range spanning between the 34 and the 45% of the total number of leg pairs. Despite these similarities, pincer-like claws originated independently in the three taxa, and by way of fundamentally different changes, either by the dramatic modification of the already existent anterior accessory spine (Diphyonyx, Neogeophilidae) or by the production of a novel cuticular projection (Eucratonyx). Moreover, their shared pattern of variation along the body was most probably constrained by already operating developmental processes controlling the longitudinal patterning of the trunk.     

Large Carnivore Footprints from the Late Pleistocene of Argentina

The aim of the present contribution is to describe large felipedid footprints from a new ichnological site from the Late Pleistocene of Buenos Aires Province, Argentina. The prints are referred as the new ichnospecies Felipeda miramarensis nov. ichnosp. Based on size and morphology, this new ichnotaxon may have been produced by the large machairodontine felid Smilodon populator. Track analysis indicates that the producer of the tracks had fully retractile claws, a plantigrade feet, and lacked strong supination capabilities on pes and manus. The size and depth differences between manus and pes prints indicate that the producer had notably robust anterior limbs. If correctly assigned, the new ichnospecies reinforces the idea that Smilodon was an ambush predatory mammal.     

Postcranial morphology,functional adaptations and palaeobiology of Callistoe vincei,a predaceous metatherian from the Eocene of Salta,north‐western Argentina

Abstract: Callistoe vincei Babot et al., 2002 is a Paleogene borhyaenoid known from exceptionally complete postcranial elements, which provides rare information about the anatomy and evolutionary history of metatherian predators during the South American Cenozoic. The axial skeleton of Callistoe is characterized by the peculiar transverse processes of the cervical vertebrae emphasizing lateral instead of sagittal traction. There is no clavicle and eighteen thoracolumbar vertebrae, of which only five are lumbars. The shoulder and elbow joints suggest movements restricted to parasagittal flexion/extension that are consistent with primarily terrestrial locomotion, as is also emphasized in Borhyaena tuberata and Lycopsis longirostrus. On the manus, the pollex is not reduced and the ungual phalanges indicate very long claws, similar to those observed in some extant digging taxa. This feature is unique to C. vincei among borhyaenoids. The knee joint is characterized by the presence of ossified patellae but shallow femoral trochleae. This joint suggests that the leg was nearly parasagittal, a position also inferred for Borhyaena. The astragalus shape is consistent with parasagittal flexion/extension, as in all Miocene–Pliocene borhyaenoids. The hind foot is characterized by reduced claws in comparison with the manus as well as the slenderness of the first and fifth digits, another peculiarity of C. vincei. The habitat of Callistoe was a temperate humid forest and according to the known fossil record, Callistoe was the largest mammalian predator of its time, sharing the predator ecological niche with crocodiles.     

Flow visualisation and high speed video analysis of water jets in the snapping shrimp (Alpheus heterochaelis)

Snapping shrimp (Alpheus heterochaelis) produce a fast, well-focused water jet by rapid closure of their specialised snapper claw. As shown previously, water jets may injure the opponent in interspecific encounters (e.g. with small crabs) although no damage was observed in intraspecific encounters. For conspecific receivers the jet represents a potential hydrodynamic signal and can be analysed with the help of mechanosensory hairs. To gain more insight in the biophysical characteristics of the water jet we visualised and analysed jets of tethered snapping shrimp using standard and high speed video recordings. Water jet width increases with increasing distance from the snapper claw tip, and both width and distance increase with increasing snapper claw size. Water jet distances do not increase with increasing claw cocking duration (building up muscle tension) but medium cocking durations of about 550 ms result in longest distances. Mean water jet velocity is 6.5 m s⁻¹ shortly after claw closure but rapidly decreases subsequently. At the mean distance between snapping conspecifics (9 mm) water jet velocities produced by snapping shrimp with larger snapper claws are significantly higher than those of animals with smaller claws. Interestingly, males with equal snapper claw size as females produce significantly faster water jets. Accepted: 31 March 1999