Functional morphology of the hallucal metatarsal with implications for inferring grasping ability in extinct primates

Primate evolutionary morphologists have argued that selection for life in a fine branch niche resulted in grasping specializations that are reflected in the hallucal metatarsal (Mt1) morphology of extant “prosimians”, while a transition to use of relatively larger, horizontal substrates explains the apparent loss of such characters in anthropoids. Accordingly, these morphological characters—Mt1 torsion, peroneal process length and thickness, and physiological abduction angle—have been used to reconstruct grasping ability and locomotor mode in the earliest fossil primates. Although these characters are prominently featured in debates on the origin and subsequent radiation of Primates, questions remain about their functional significance. This study examines the relationship between these morphological characters of the Mt1 and a novel metric of pedal grasping ability for a large number of extant taxa in a phylogenetic framework. Results indicate greater Mt1 torsion in taxa that engage in hallucal grasping and in those that utilize relatively small substrates more frequently. This study provides evidence that Carpolestes simpsoni has a torsion value more similar to grasping primates than to any scandentian. The results also show that taxa that habitually grasp vertical substrates are distinguished from other taxa in having relatively longer peroneal processes. Furthermore, a longer peroneal process is also correlated with calcaneal elongation, a metric previously found to reflect leaping proclivity. A more refined understanding of the functional associations between Mt1 morphology and behavior in extant primates enhances the potential for using these morphological characters to comprehend primate (locomotor) evolution. Am J Phys Anthropol 156:327–348, 2015. © 2014 Wiley Periodicals, Inc.     

Electromyography of crural and pedal muscles in tufted capuchin monkeys (Sapajus apella): Implications for hallucal grasping behavior and first metatarsal morphology in euprimates

A hypertrophied peroneal process of the hallucal metatarsal, as seen in prosimians, has been linked to a powerful hallucal grasp via the contraction of the peroneus longus (PL) muscle causing adduction of the big toe. Electromyography (EMG) studies of lemurs and lorises, however, have concluded that PL is not substantially recruited during small branch locomotion when powerful hallucal grasping is needed most, and have suggested that there is no link between PL activity and peroneal process size. If this is correct, then we should also observe no change in PL activity when strong hallucal grasping is required in anthropoids because they have a relatively smaller peroneal process for PL to act on. This study addresses this hypothesis by evaluating EMG of crural and pedal muscles in capuchins (Sapajus apella) walking on substrates of different diameters. During locomotion on the narrow substrate (3.1 cm) that should elicit a strong hallucal grasp, we observed an intense increased recruitment of adductor hallucis, but only sustained, rather than markedly increased, PL activity. This indicates that PL is not involved in powerful hallucal grasping in capuchins, and confirms similar findings previously documented in prosimians. We continue to reject the hypothesis that a large peroneal process is an adaptation for powerful grasping and further argue that its morphology may not be related to PL's ability to adduct the hallux at all. In addition, the morphology of the peroneal process should not be used to assess hallucal grasping performance in fossils. Am J Phys Anthropol 156:553–564, 2015. © 2015 Wiley Periodicals, Inc.     

Hallucal grasping in Nycticebus coucang: further implications for the functional significance of a large peroneal process

Euprimate grasping feet are characterized by a suite of morphological traits, including an enlarged peroneal process on the base of the first metatarsal, which serves as the insertion site of the peroneus longus muscle. In prosimians, a large process has typically been associated with a powerful hallucal grasp via the contraction of the peroneus longus to adduct the hallux. Recent electromyography (EMG) studies have documented that peroneus longus does not contribute substantially to hallucal grasping in lemurids (Boyer et al., 2007). However, non-lemurid prosimians have a I-V opposable grasp complex that is morphologically different and phylogenetically more primitive than the I-II adductor grasp complex of the lemurids previously studied. Therefore, it is possible that peroneus longus did function during grasping in early euprimates, but lost this function in large-bodied lemurids. The present study tests the hypothesis that a large peroneal process is related to powerful grasping ability in primates displaying the more primitive I-V grasp complex. We use EMG to evaluate the recruitment of peroneus longus, other crural muscles, and adductor hallucis in static and locomotor grasping activities of the slow loris (Nycticebus coucang). Results show that peroneus longus is active during grasping behaviors that require the subject to actively resist inversion of the foot, and likely contributes to a hallucal grasp in these activities. Peroneus longus activity level does not differ between grasping and power grasping activities, nor does it differ between grasping and non-grasping locomotor modes. Conversely, the digital flexors and hallucal adductor are recruited at higher levels during power grasping and grasping locomotor modes. Consequently, we reject the hypothesis that an enlarged peroneal process represents an adaptation specifically to enhance the power of the I-V grasp, but accept that the muscle likely plays a role in adducting the hallux during grasping behaviors that require stabilization of the ankle, and suggest that further work is necessary to determine if this role is sufficient to drive selection for a large peroneal process.     

New primate first metatarsals from the Paleogene of Egypt and the origin of the anthropoid big toe

The specialized grasping feet of primates, and in particular the nature of the hallucal grasping capabilities of living strepsirrhines and tarsiers (i.e., ‘prosimians’), have played central roles in the study of primate origins. Prior comparative studies of first metatarsal (Mt1) morphology have documented specialized characters in living prosimians that are indicative of a more abducted hallux, which in turn is often inferred to be related to an increased ability for powerful grasping. These include a well-developed peroneal process and a greater angle of the proximal articular surface relative to the long axis of the diaphysis. Although known Mt1s of fossil prosimians share these characters with living non-anthropoid primates, Mt1 morphology in the earliest crown group anthropoids is not well known. Here we describe two Mt1s from the Fayum Depression of Egypt - one from the latest Eocene (from the ∼34 Ma Quarry L-41), and one from the later early Oligocene (from the ∼29-30 Ma Quarry M) - and compare them with a sample of extant and fossil primate Mt1s. Multivariate analyses of Mt1 shape variables indicate that the Fayum specimens are most similar to those of crown group anthropoids, and likely belong to the stem catarrhines Catopithecus and Aegyptopithecus specifically, based on analyses of size. Also, phylogenetic analyses with 16 newly defined Mt1 characters support the hypotheses that “prosimian”-like Mt1 features evolved along the primate stem lineage, while crown anthropoid Mt1 morphology and function is derived among primates, and likely differed from that of basal stem anthropoids. The derived loss of powerful hallucal grasping as reflected in the Mt1 morphology of crown anthropoids may reflect long-term selection for improved navigation of large-diameter, more horizontal branches at the expense of movement in smaller, more variably inclined branches in the arboreal environment.     

Telemetered electromyography of peroneus longus in Varecia variegata and Eulemur rubriventer: implications for the functional significance of a large peroneal process

A foot specialized for grasping small branches with a divergent opposable hallux (hallucal grasping) represents a key adaptive complex characterizing almost all arboreal non-human euprimates. Evolution of such grasping extremities probably allowed members of a lineage leading to the common ancestor of modern primates to access resources available in a small-branch niche, including angiosperm products and insects. A better understanding of the mechanisms by which euprimates use their feet to grasp will help clarify the functional significance of morphological differences between the euprimate grasp complex and features representing specialized grasping in other distantly related groups (e.g., marsupials and carnivorans) and in closely related fossil taxa (e.g., plesiadapiforms). In particular, among specialized graspers euprimates are uniquely characterized by a large peroneal process on the base of the first metatarsal, but the functional significance of this trait is poorly understood. We tested the hypothesis that the large size of the peroneal process corresponds to the pull of the attaching peroneus longus muscle recruited to adduct the hallux during grasping. Using telemetered electromyography on three individuals of Varecia variegata and two of Eulemur rubriventer, we found that peroneus longus does not generally exhibit activity consistent with an important function in hallucal grasping. Instead, extrinsic digital flexor muscles and, sometimes, the intrinsic adductor hallucis are active in ways that indicate a function in grasping with the hallux. Peroneus longus helps evert the foot and resists its inversion. We conclude that the large peroneal tuberosity that characterizes the hallucal metatarsal of prosimian euprimates does not correlate to "powerful" grasping with a divergent hallux in general, and cannot specifically be strongly linked to vertical clinging and climbing on small-diameter supports. Thus, the functional significance of this hallmark, euprimate feature remains to be determined.     

The skeleton of early Eocene Cantius, oldest lemuriform primate

A recently discovered partial skeleton of the adapid Cantius trigonodus from the early Eocene Willwood Formation of the Bighorn Basin, Wyoming, documents substantial new information about the anatomy of the oldest lemuriform primates. It is very similar in all features to its descendant, middle Eocene Notharctus, and both exhibit numerous resemblances to certain extant Malagasy lemurs, particularly Lepilemur, Propithecus, Lemur, and Hapalemur griseus. Like these forms, Cantius had relatively long hind limbs and short forelimbs. Forelimb traits (prominent brachialis flange of the humerus, well-developed olecranon process of the ulna, and strong shafts of the ulna and radius) suggest active use of the forelimbs in progression. Specializations in the hind limb (e.g., expanded articular surface of the femoral head, narrow and elevated patellar trochlea and prominent lateral trochlear ridge, posteriorly oriented femoral and tibial condyles, narrow and elongate talus, and hallucal metatarsal with prominent peroneal tubercle) indicate capabilities for leaping and for powerful grasping with an opposable hallux. Cantius was presumably primitive in having a relatively long ischium and much more distal inferior tibial tuberosity than most extant lemurs--traits suggesting that powerful extension of the thigh and flexion at the knee were important in its locomotion and posture. We interpret Cantius as an active arboreal quadruped with a propensity for leaping. The existence of this skeletal structure in one of the oldest primates of modern aspect suggests that it represents the primitive lemuriform morphology.     

Hallucal grasping behavior in Caluromys (Didelphimorphia: Didelphidae): Implications for primate pedal grasping

A key feature in primate evolution is a foot with a divergent opposable hallucal metatarsal bearing a large peroneal process. Extant primates are characterized by a powerful hallucal grasp—an either “euprimate” or “plesiadapoid-euprimate” ancestor acquisition—that facilitates the exploitation of fine branches, an ability that increased the fitness of ancestral euprimates. In this context, the didelphid marsupial Caluromys has been used as the extant analog to this primate morphotype stage due to some morphological, ecological, and behavioral features. However, the extent to which and the positional and support contexts in which Caluromys uses powerful hallucal grasping are not known. This renders analogies to any mode of “euprimate” or “stem primate” grasping poorly substantiated. The present paper quantifies locomotor and postural behavior, support use, and associated frequencies of hallucal grasping in captive Caluromys philander via analysis of video recordings. During locomotion, Caluromys primarily used diagonal sequence walk, clamber, and climb, whereas stand, foot-hang, and bipedal stand were the dominant postures. Small, fine, horizontal, and moderately inclined branches were frequently used. Overall rates of “apparently powerful hallucal grasps” were high, but were exceptionally high during clamber, climb, foot-hang, and bipedal stand. Additionally, an “apparently powerful hallucal grasp” was very common upon fine, small, steep, and vertical branches. The extensive use of such powerful hallucal grasping provided stability and safety that enabled Caluromys to proficiently utilize fine branches of various orientations. The ability to negotiate such unstable supports, further reflected in foot anatomy, provides evidence that the morphobehavioral complex of Caluromys can serve as an extant analog to the plesiadapoid-euprimate ancestor, represented as a terminal branch feeder with effective hallucal grasping.     

The morphological basis of hallucal orientation in extant birds

The perching foot of living birds is commonly characterized by a reversed or opposable digit I (hallux). Primitively, the hallux of nonavian theropod dinosaurs was unreversed and lay parallel to digits II-IV. Among basal birds, a unique digital innovation evolved in which the hallux opposes digits II-IV. This digital configuration is critical for grasping and perching. I studied skeletons of modern birds with a range of hallucal designs, from unreversed (anteromedially directed) to fully reversed (posteriorly directed). Two primary correlates of hallucal orientation were revealed. First, the fossa into which metatarsal I articulates is oriented slightly more posteriorly on the tarsometatarsus, rotating the digit as a unit. Second, metatarsal I exhibits a distinctive torsion of its distal shaft relative to its proximal articulation with the tarsometatarsus, reorienting the distal condyles and phalanges of digit I. Herein, I present a method that facilitates the re-evaluation of hallucal orientation in fossil avians based on morphology alone. This method also avoids potential misinterpretations of hallucal orientation in fossil birds that could result from preserved appearance alone.     

Evolution and Allometry of Calcaneal Elongation in Living and Extinct Primates

Specialized acrobatic leaping has been recognized as a key adaptive trait tied to the origin and subsequent radiation of euprimates based on its observed frequency in extant primates and inferred frequency in extinct early euprimates. Hypothesized skeletal correlates include elongated tarsal elements, which would be expected to aid leaping by allowing for increased rates and durations of propulsive acceleration at takeoff. Alternatively, authors of a recent study argued that pronounced distal calcaneal elongation of euprimates (compared to other mammalian taxa) was related primarily to specialized pedal grasping. Testing for correlations between calcaneal elongation and leaping versus grasping is complicated by body size differences and associated allometric affects. We re-assess allometric constraints on, and the functional significance of, calcaneal elongation using phylogenetic comparative methods, and present an evolutionary hypothesis for the evolution of calcaneal elongation in primates using a Bayesian approach to ancestral state reconstruction (ASR). Results show that among all primates, logged ratios of distal calcaneal length to total calcaneal length are inversely correlated with logged body mass proxies derived from the area of the calcaneal facet for the cuboid. Results from phylogenetic ANOVA on residuals from this allometric line suggest that deviations are explained by degree of leaping specialization in prosimians, but not anthropoids. Results from ASR suggest that non-allometric increases in calcaneal elongation began in the primate stem lineage and continued independently in haplorhines and strepsirrhines. Anthropoid and lorisid lineages show stasis and decreasing elongation, respectively. Initial increases in calcaneal elongation in primate evolution may be related to either development of hallucal-grasping or a combination of grasping and more specialized leaping behaviors. As has been previously suggested, subsequent increases in calcaneal elongation are likely adaptations for more effective acrobatic leaping, highlighting the importance of this behavior in early euprimate evolution.     

Anatomy of the bony pelvis in parapithecid primates

Four partial innominate bones, attributed to the parapithecid primates Parapithecus grangeri and Apidium phiomense, have recently been recovered from Oligocene deposits in the Fayum of Egypt. These fossils provide the first documentation of pelvic morphology for early anthropoids. In pelvic anatomy, parapithecids show definite similarities to higher primates rather than to prosimians, but cannot be clearly allied with any one extant group. Functionally, the fossils indicate quadrupedal or leaping habits rather than suspensory or bipedal behaviors.     

Articular to diaphyseal proportions of human and great ape metatarsals

This study proposes a new way to use metatarsals to identify locomotor behavior of fossil hominins. Metatarsal head articular dimensions and diaphyseal strength in a sample of chimpanzees, gorillas, orangutans, and humans (n = 76) are used to explore the relationships of these parameters with different locomotor modes. Results show that ratios between metatarsal head articular proportions and diaphyseal strength of the hallucal and fifth metatarsal discriminate among extant great apes and humans based on their different locomotor modes. In particular, the hallucal and fifth metatarsal characteristics of humans are functionally related to the different ranges of motion and load patterns during stance phase in the forefoot of humans in bipedal locomotion. This method may be applicable to isolated fossil hominin metatarsals to provide new information relevant to debates regarding the evolution of human bipedal locomotion. The second to fourth metatarsals are not useful in distinguishing among hominoids. Further studies should concentrate on measuring other important qualitative and quantitative differences in the shape of the metatarsal head of hominoids that are not reflected in simple geometric reconstructions of the articulation, and gathering more forefoot kinematic data on great apes to better understand differences in range of motion and loading patterns of the metatarsals. Am J Phys Anthropol 143:198–207, 2010. © 2010 Wiley‐Liss, Inc.     

Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela

I observed leaping behavior in the white-faced saki (Pithecia pithecia) and the black-bearded saki (Chiropotes satanas satanas) for 15 and 10 months, respectively, as part of a larger study of positional behavior in the tribe Pitheciini. I used focal animal instantaneous sampling to observe the two species on separate islands in their natural habitat at Guri Lake, Venezuela. Leaping behavior correlates with patterns of forest use and body size, and differences between the species relate more to habitat preferences than to habitat differences per se. Pithecia usually chose vertical or highly angled supports of lower tree portions for take-off and landing, and took off from a stationary posture. Chiropotes took off from the main crown or terminal branches, gaining momentum from locomotor movement before performing a leaping take-off. Pithecia's vertical body orientation and longer leap distance allowed it to assume a mid-flight tuck to prepare for a hindlimb-first landing onto a solid support, and to absorb landing forces with its relatively longer hindlimbs. Chiropotes remained more pronograde throughout its leaps, and minimized landing forces by landing on all four limbs onto numerous flexible supports in the terminal branches. The smaller-bodied P. pithecia is specialized for vertical clinging and leaping, and exhibits behavioral and morphological parallels with other vertical clingers and leapers. The larger C. satanas is a generalized leaper that lacks morphological specializations for leaping. Pithecia's use of solid supports in the lower tree portions allows it to move quietly through the forest-one of a suite of behaviors related to predator avoidance. This example of variation within one behavioral category has implications for devising locomotor classifications and interpreting fossil remains.     

Cortical Structure of Hallucal Metatarsals and Locomotor Adaptations in Hominoids

Diaphyseal morphology of long bones, in part, reflects in vivo loads experienced during the lifetime of an individual. The first metatarsal, as a cornerstone structure of the foot, presumably expresses diaphyseal morphology that reflects loading history of the foot during stance phase of gait. Human feet differ substantially from those of other apes in terms of loading histories when comparing the path of the center of pressure during stance phase, which reflects different weight transfer mechanisms. Here we use a novel approach for quantifying continuous thickness and cross-sectional geometric properties of long bones in order to test explicit hypotheses about loading histories and diaphyseal structure of adult chimpanzee, gorilla, and human first metatarsals. For each hallucal metatarsal, 17 cross sections were extracted at regularly-spaced intervals (2.5% length) between 25% and 65% length. Cortical thickness in cross sections was measured in one degree radially-arranged increments, while second moments of area were measured about neutral axes also in one degree radially-arranged increments. Standardized thicknesses and second moments of area were visualized using false color maps, while penalized discriminant analyses were used to evaluate quantitative species differences. Humans systematically exhibit the thinnest diaphyseal cortices, yet the greatest diaphyseal rigidities, particularly in dorsoplantar regions. Shifts in orientation of maximum second moments of area along the diaphysis also distinguish human hallucal metatarsals from those of chimpanzees and gorillas. Diaphyseal structure reflects different loading regimes, often in predictable ways, with human versus non-human differences probably resulting both from the use of arboreal substrates by non-human apes and by differing spatial relationships between hallux position and orientation of the substrate reaction resultant during stance. The novel morphological approach employed in this study offers the potential for transformative insights into form-function relationships in additional long bones, including those of extinct organisms (e.g., fossils).     

Submaximal leaping in the grey mouse lemur

In arboreal animals such as the grey mouse lemur (Microcebus murinus Miller, 1777), leaping is the most frequent strategy for predator avoidance. The aim of this study was to characterise the locomotor adaptation in response to the structural constraint of the habitat (i.e., position of the landing substrate). Thus, we characterised the push-off phase by inducing the lemurs to leap up to a range of heights from horizontal to their own individual highest performance. Using uniplanar high-frequency cineradiographs collected in a sagittal plane, the relative contributions of the centre of mass (CoM) velocity vector magnitude and orientation to leaping performance were evaluated. The kinematics of the push-off phase showed that for low landing heights, leaping performance was essentially due to hip and knee extensions. Higher leaps seemed to be related to an increase in ankle contribution. At all leaping heights, the proximal-to-distal sequence of the hind limb joints controlled the orientation and magnitude of the M. murinus CoM velocity vector while pushing off. Finally, the analysis of the velocity vector at the onset of take-off suggested that the optimal solution for predator avoidance was to leap for horizontal distance and not for vertical distance.     

Morphometry and allometry of the primate humerus

The primate distal humerus has been used both in phylogenetic reconstruction and in assessing locomotor and postural adaptations. This study uses an allometric approach to predict locomotor patterns of extant primates regardless of phylogenetic position. By showing the relationship between form and function in living primate taxa it will be possible to use this data set to predict locomotor behavior of extinct primates. Several linear measurements were taken from the distal humerus of 71 extant primate species (anthropoids and prosimians). Allometric regressions of each measurement were performed with mandibular M₂ area as a surrogate for body size. These measurements were used to determine if significant differences in distal humerus morphology exist among locomotor groups. The results were then used to test several hypotheses about the relationship between humeral form and function. For example, the hypothesis that suspensory primates have a large medial epicondyle is confirmed; the hypothesis that terrestrial quadrupeds have a deep olecranon fossa could not be confirmed with quantitative data. In addition to this hypothesis testing, the residuals from the allometric regressions of the humeral measurements were used in a discriminant functions analysis to estimate locomotor behavior from distal humerus morphology. The discriminant functions analysis correctly reclassified 64/71 (90%) species.     

Diets of fossil primates from the Fayum Depression of Egypt: a quantitative analysis of molar shearing

Over the last 90 years, Eocene and Oligocene aged sediments in the Fayum Depression of Egypt have yielded at least 17 genera of fossil primates. However, of this diverse sample the diets of only four early Oligocene anthropoid genera have been previously studied using quantitative methods. Here we present dietary assessments for 11 additional Fayum primate genera based on the analysis of body mass and molar shearing crest development. These studies reveal that all late Eocene Fayum anthropoids were probably frugivorous despite marked subfamilial differences in dental morphology. By contrast, late Eocene Fayum prosimians demonstrated remarkable dietary diversity, including specialized insectivory (Anchomomys), generalized frugivory (Plesiopithecus), frugivory+insectivory (Wadilemur), and strict folivory (Aframonius). This evidence that sympatric prosimians and early anthropoids jointly occupied frugivorous niches during the late Eocene reinforces the hypothesis that changes in diet did not form the primary ecological impetus for the origin of the Anthropoidea. Early Oligocene Fayum localities differ from late Eocene Fayum localities in lacking large-bodied frugivorous and folivorous prosimians, and may document the first appearance of primate communities with trophic structures like those of extant primate communities in continental Africa. A similar change in primate community structure during the Eocene-Oligocene transition is not evident in the Asian fossil record. Putative large anthropoids from the Eocene of Asia, such as Amphipithecus mogaungensis, Pondaungia cotteri, and Siamopithecus eocaenus, share with early Oligocene Fayum anthropoids derived features of molar anatomy related to an emphasis on crushing and grinding during mastication. However, these dental specializations are not seen in late Eocene Fayum anthropoids that are broadly ancestral to the later-occurring anthropoids of the Fayum's upper sequence. This lack of resemblance to undisputed Eocene African anthropoids suggests that the "progressive" anthropoid-like dental features of some large-bodied Eocene Asian primates may be the result of dietary convergence rather than close phyletic affinity with the Anthropoidea.     

In vivo baseline measurements of hip joint range of motion in suspensory and nonsuspensory anthropoids

Hominoids and atelines are known to use suspensory behaviors and are assumed to possess greater hip joint mobility than nonsuspensory monkeys, particularly for range of abduction. This assumption has greatly influenced how extant and fossil primate hip joint morphology has been interpreted, despite the fact that there are no data available on hip mobility in hominoids or Ateles. This study uses in vivo measurements to test the hypothesis that suspensory anthropoids have significantly greater ranges of hip joint mobility than nonsuspensory anthropoids. Passive hip joint mobility was measured on a large sample of anesthetized captive anthropoids (nonhuman hominids = 43, hylobatids = 6, cercopithecids = 43, Ateles = 6, and Cebus = 6). Angular and linear data were collected using goniometers and tape measures. Range of motion (ROM) data were analyzed for significant differences by locomotor group using ANOVA and phylogenetic regression. The data demonstrate that suspensory anthropoids are capable of significantly greater hip abduction and external rotation. Degree of flexion and internal rotation were not larger in the suspensory primates, indicating that suspension is not associated with a global increase in hip mobility. Future work should consider the role of external rotation in abduction ability, how the physical position of the distal limb segments are influenced by differences in ROM proximally, as well as focus on bony and soft tissue differences that enable or restrict abduction and external rotation at the anthropoid hip joint. Am J Phys Anthropol 153:417–434, 2014. © 2013 Wiley Periodicals, Inc.     

Sexual dimorphism in large-bodied primates: The case of the subfossil lemurs

Large body size has evolved repeatedly in the order Primates, not merely among anthropoids but also among prosimians. Whereas high degrees of sexual size dimorphism characterize many of the large-bodied anthropoids, this is not the case for extinct large-bodied lemurs. This paper uses finite mixture analysis and other techniques to ascertain just how much skull length dimorphism might be embedded in the generally unimodal distributions of skull lengths of giant extinct lemurs from single localities, and then compares these results with known skull length dimorphisms in extant lemurs and large-bodied catarrhines. We show that low levels of skull length sexual dimorphism (or none at all) characterize subfossil lemurs, and we explore several possible explanations for this phenomenon. Traditional explanations of sexual size dimorphism generally focus on body size or mating systems. These are not sufficient to explain the variation in sexual dimorphism that can be observed in the order Primates. © 1993 Wiley-Liss, Inc.     

Functional anatomy of the limbs of erethizontidae (Rodentia, Caviomorpha): Indicators of locomotor behavior in Miocene porcupines

Functional analysis of the limb bones of the erethizontid Steiromys duplicatus, one of the most abundant Miocene porcupines from Patagonia, provides evidence to infer their locomotor behavior. Remains of the giant Neosteiromys pattoni (Late Miocene of Northeast Argentina) are also analyzed. Osteological and myological features of extant porcupines were evaluated and used as a model to interpret the functional significance of Miocene species' limbs. Several features in erethizontids are compatible with the ability to climb: the low humeral tuberosities indicate a mobile gleno-humeral joint; the prominent and distally extended deltopectoral crest indicates a powerful pectoral muscle, which is particularly active when climbing; the humero-ulnar and humero-radial joints are indicative of pronation-supination movements; the well-developed lateral epicondylar ridge and the medially protruding entepicondyle are in agreement with an important development of the brachioradialis, supinator, flexor digitorum profundus, and pronator teres muscles, acting in climbing and grasping functions; the mechanical advantage of the biceps brachii would be emphasized because of its distal attachment on the bicipital tuberosity. As with extant porcupines, in Miocene species, the large femoral head would have permitted a broad range of abduction of the femur, and the medially protruding lesser trochanter would have emphasized the abduction and outward rotation of the femur by the action of the ilio-psoas complex. In S. duplicatus, the shape of the hip, knee, and cruro-astragalar, calcaneo-astragalar, and astragalo-navicular joints would have allowed lateral and rotational movements, although probably to a lesser degree than in extant porcupines. Foot features of S. duplicatus (e.g., great medial sesamoid bone, medial astragalar head, complete hallux) indicate that this species would have had grasping ability, but would not have achieved the high degree of specialization of Coendou. Steiromys duplicatus would have been a semiarboreal dweller, resembling Erethizon dorsatum.     

A volumetric comparison of the vestibular nuclei in primates

The volumes of each of the four vestibular nuclei, superior, lateral, medial and descending, were measured in 80 brains from 2 species of Scandentia, 18 species of prosimians, and 26 species of anthropoids. Size indices were calculated by comparing species-specific points to the nucleus volume-body weight allometry in prosimians, where the average prosimian was set at 1.00. The indices range from 1.78 in Saimiri to 0.48 in Gorilla, and the distributions by families overlap partially or completely. The observed trend in size indices is independent of changes in the neocortex and the ventral pons; average indices are 1.35 in New World monkeys, 1.20 in Old World monkeys, 0.74 in apes, 0.82 in man. Among prosimians, Galago, Galagoides and Tarsius (leaping locomotion) show significantly higher indices than Nycticebus, Loris and Perodicticus (slow movement without leaping). The lateral vestibular nuclear indices in Pongidae and man are extremely low, about half of those of the average prosimians. Correlation coefficients of size indices between the vestibular nuclei and other motor nuclei, such as the cerebellar nuclei, ventral pons and striatum, are analysed. The ratio of the vestibular nuclear volumes to the total brain volumes and the distribution of percentages of each vestibular nuclear volume to the total complex are also obtained.