The role of developmental rate, body size, and positional behavior in the evolution of covariation and evolvability in the cranium of strepsirrhines and catarrhines.

Recent studies on hominin craniofacial evolution have focused on phenotypic integration or covariation among traits. Covariation is thought to significantly affect evolutionary trajectories, shaping the ways in which hominins and other primates could have evolved. However, the ways in which covariation itself evolves are not well understood. This study aims to investigate the role of phylogeny, development, body size, and positional behavior in shaping the strength of covariation in strepsirrhine and catarrhine primate crania (n = 1009, representing 11 genera). These factors may have been catalysts for change in the magnitude of covariation, and they have changed significantly during primate evolution and particularly hominin evolution. Modern humans in particular have slow developmental trajectories, large bodies, and a unique form of locomotion in the form of orthograde bipedalism. Variance of eigenvalues, mean integration, mean evolvability, and mean conditional evolvability was estimated and their relationship to the various factors described earlier was assessed using phylogenetic and nonphylogenetic analyses. Results indicate that some phylogenetic signal is present, but it is not equivalent across integration statistics or cranial regions. In particular, these results suggest that closely related species are more similar than more distantly related species in evolvability of the cranial base and integration of the face. Two divergent patterns were also identified, in which covariation and evolvability of the cranial base are linked to developmental rate, but those of the face are linked to body size. Neither locomotion nor posture appears related to covariation or evolvability of the primate cranium. These results suggest that overall low covariation observed in the hominin cranium may be a result of separate trends in different cranial regions.

First navicular remains of a European adapiform (Anchomomys frontanyensis) from the Middle Eocene of the Eastern Pyrenees (Catalonia, Spain): implications for early primate locomotor behavior and navicular evolution.

We describe the first known navicular bones for an Eocene euprimate from Europe and assess their implications for early patterns of locomotor evolution in primates. Recovered from the fossil site of Sant Jaume de Frontanyà-3C (Barcelona, Spain), the naviculars are attributed to Anchomomys frontanyensis. The small size of A. frontanyensis allows us to consider behavioral implications of comparisons with omomyiforms, regardless of allometric sources of navicular variation. Researchers usually consider omomyiforms to be more prone to leaping than contemporaneous adapiforms partly because of the more pronounced elongation of omomyiform tarsal elements. However, A. frontanyensis differs from other adapiforms and is similar to some omomyiforms in its more elongated navicular proportions. Although this might raise questions about attribution of these naviculars to A. frontanyensis, the elements exhibit clear strepsirrhine affinities leaving little doubt about the attribution: the bones' mesocuneiform facets contact their cuboid facets. We further propose that this strepsirrhine-specific feature in A. frontanyensis and other adapiforms reflects use of more inverted foot postures and potentially smaller substrates than sympatric omomyiforms that lack it. Thus substrate differences may have influenced niche partitioning in Eocene euprimate communities along with differences in locomotor agility. As previous studies on the astragalus and the calcaneus have suggested, this study on the navicular is consistent with the hypothesis that the locomotor mode of A. frontanyensis was similar to that of extant cheirogaleids, especially species of Microcebus and Mirza.

A primate with a Panda's thumb: The anatomy of the pseudothumb of Daubentonia madagascariensis.

OBJECTIVES: Accessory digits have evolved independently within several mammalian lineages. Most notable among these is the pseudothumb of the giant panda, which has long been considered one of the most extraordinary examples of contingent evolution. To date, no primate has been documented to possess such an adaptation. Here, we investigate the presence of this structure within the aye-aye (Daubentonia madagascariensis), a species renowned for several other specialized morphological adaptations in the hand, including a morphologically unique third digit. MATERIALS AND METHODS: We combine physical dissection techniques with digital imaging processes across a sample of seven individuals (six adults and one immature individual) to describe and visualize the anatomy of the wrist and hand within the aye-aye. RESULTS: A distinct pseudothumb, which consists of both a bony component (an expanded radial sesamoid) and a dense cartilaginous extension (the "prepollex") was observed in all specimens. We demonstrate that this pseudodigit receives muscular attachments from three muscles, which collectively have the potential to enable abduction, adduction, and opposition. Finally, we demonstrate that the pseudothumb possesses its own distinct pad within the palm, complete with independent dermatoglyphs. DISCUSSION: Pseudothumbs have been suggested to improve palmar dexterity in taxa with overly -generalized first digits (e.g., pandas) and to widen the hand for digging (e.g., some fossorial moles), but the aye-aye's pseudothumb represents what we believe is a heretofore unrecognized third functional role: its accessory digit compensates for overspecialization of its fingers for non-gripping functions (in this case, the aye-aye's unique "tap foraging" practices).

The evolution of palate shape in the Lepilemur-Cheirogaleidae clade (Primates: Strepsirrhini).

OBJECTIVES: Phylogenies consistently group the folivorous Lepilemur species with the small-bodied insectivorous-frugivorous cheirogaleids. Juvenile lepilemurs and adult cheirogaleids share allometries in most aspects of skull morphology, except the palate. We investigated potential influences on palate shape in these taxa and several outgroups using geometric morphometrics. MATERIALS AND METHODS: Our sample included representatives of four extant strepsirrhine families, Cheirogaleidae (including Lepilemurinae), Lemuridae, Indriidae, and Galagidae, and one subfossil Megaladapis. Our dataset comprised 32 landmarks collected from 397 specimens representing 15 genera and 28 species, and was analyzed using generalized procrustes analyses and between group principal component analysis. We explored the influence of size, phylogeny, diet, and the propagation of loud vocalizations on palate shape. RESULTS: While congeneric species clustered within the morphospace, the phylomorphospace did not mirror molecular phylogenetic hypotheses of higher-order relationships. Four palate forms were distinguished within the Cheirogaleidae. Diet, strongly linked to body size, had the single greatest influence on palate shape. The production of long-distance advertisement calls was most often associated with positive scores on the PC1 axis. DISCUSSION: Our results suggest that the extensive variation in palate shape among Cheirogaleidae is related to dietary shifts that accompanied changes in body size during the clade's radiation. Molecular phylogenies indicate that cheirogaleid diversification involved repeated dwarfing events, which in turn drove dietary shifts from ancestral folivory-frugivory to frugivory, gummivory, and faunivory in the descendant species. The elongated Lepilemur palate is probably related to accelerated eruption of the cheek teeth to render juveniles competent to shear leaves upon weaning.

A comparison of relief estimates used in three-dimensional dental topography.

OBJECTIVES: Topographic estimates of dental relief are now commonly used to make dietary inferences from the teeth of extant and extinct primates. We thoroughly compared commonly used relief estimates in an effort to help researchers decide which variable best suits their objectives. MATERIALS AND METHODS: We combined a total of three datasets: five theoretical models built to compare the effect of tooth complexity and basin depth on relief estimates, a dataset of 110 lower molars of prosimians, and a dataset of 25 upper molars of apes. We investigated intra-mesh variation and tooth average relief, estimated from slope and three different relief indices, according to four criteria: (1) the ability to map relief on topographic maps, (2) the correlation with other relief estimates, (3) the ability to separate high-relief molars of folivores from deep-relief molars of insectivores in prosimians, and (4) the influence of surface complexity on relief estimates in apes. RESULTS: We found that polygon slope and relief index are linked by a mathematical relation. Tooth average slope and all relief indices are strongly correlated. In contrast, relief estimates are moderately correlated to cusp elevation. One relief index of four relief estimates had an excellent ability to separate high-relief from deep-relief molars in prosimians, whereas slope could not separate them. No significant effect of tooth complexity on dental relief could be detected in apes. CONCLUSIONS: Because slope and relief indices are highly correlated, it is strongly recommended not to combine them in multivariate analysis. Still, slope and relief indices show interesting differences in scaling, graphical representation, computation method, and ability to separate high-relief and deep-relief molars. Our results also suggest that slope and relief indices can vary independently of tooth complexity and are moderately affected by mean cusp elevation in apes.

Middle ear pneumatization in nonhuman primates: A comparative analysis.

OBJECTIVES: We test the effects of body mass and phylogeny on middle ear cavity pneumatization, and the role of pneumatization in hearing function, spanning the anatomical, ecological, and behavioral diversity of nonhuman primates. MATERIALS AND METHODS: All cavities were segmented in middle ear scans of 96 specimens, from 12 strepsirrhine and 15 haplorhine extant species. We measured the tympanic cavity (TC) separately, and all other middle ear spaces together (MES), calculating the degree of pneumatization with the surface area-to-volume ratio. We tested body mass effect with linear regression; we evaluated the phylogenetic signal and selection patterns, using a Kappa statistic test, and Ornstein-Uhlenbeck models (OU). We investigated the link between pneumatization and hearing sensitivity using phylogenetic regression. RESULTS: Testing body mass reveals an allometric pattern for both TC and MES dimensions. Degree of pneumatization in MES is dependent on body mass in haplorhines: larger animals have more pneumatized MES. Differences at various taxonomic ranks were observed for MES, while no phylogenetic influence was observed for TC. Infraorder selection patterns are different. Auditory performance is significantly related to degree of pneumatization, indicating that a pneumatized middle ear is associated with better perception of low frequencies. DISCUSSION: Pneumatization in MES is under differential selective pressure, indicating several optima for this trait. Pneumatization in MES probably modifies hearing sensitivity through pressure regulation mechanisms, auditory bulla size reduction, and frequency modulation. This could explain strepsirrhine adaptation to high-frequency perception, while haplorhine auditory perception is adapted to a broader sound range, including high and low frequencies.

Convergent evolution in the Euarchontoglires.

Convergence-the independent evolution of similar phenotypes in distantly related clades-is a widespread and much-studied phenomenon. An often-cited, but hitherto untested, case of morphological convergence is that between the aye-aye and squirrels. The aye-aye (Daubentonia madagascariensis) is a highly unusual lemuriform primate that has evolved a dentition similar to that of rodents: it possesses large, ever-growing incisors which it uses to strip the bark from trees in order to feed on wood-boring beetle larvae. Indeed, such is the similarity that some of the earliest classifications of the aye-aye placed it in the squirrel genus Sciurus Here, we aimed to test the degree of convergence between the skulls and lower jaws of squirrels and the aye-aye. Three-dimensional landmarks were recorded from the crania and mandibles of 46 taxa representing the majority of families in the Euarchontoglires. Results were plotted as phylomorphospaces and convergence measures were calculated. The convergence between squirrels and the aye-aye was shown to be statistically significant for both the cranium and mandible, although the mandibles seem to converge more closely in shape. The convergence may indicate strong functional drivers of morphology in these taxa, i.e. the use of the incisors to produce high bite forces during feeding. Overall, we have shown that this classic case of convergence stands up to quantitative analysis.

Do forelimb shape and peak forces co-vary in strepsirrhines?

OBJECTIVES: In this study, we explore whether ground reaction forces recorded during horizontal walking co-vary with the shape of the long bones of the forelimb in strepsirrhines. To do so, we quantify (1) the shape of the shaft and articular surfaces of each long bone of the forelimb, (2) the peak vertical, mediolateral, and horizontal ground reaction forces applied by the forelimb during arboreal locomotion, and (3) the relationship between the shape of the forelimb and peak forces. MATERIALS AND METHODS: Geometric morphometric approaches were used to quantify the shape of the bones. Kinetic data were collected during horizontal arboreal walking in eight species of strepsirrhines that show variation in habitual substrate use and morphology of the forelimb. These data were then used to explore the links between locomotor behavior, morphology, and mechanics using co-variation analyses in a phylogenetic framework. RESULTS: Our results show significant differences between slow quadrupedal climbers (lorises), vertical clinger and leapers (sifaka), and active arboreal quadrupeds (ring-tailed lemur, ruffed lemur) in both ground reaction forces and the shape of the long bones of the forelimb, with the propulsive and medially directed peak forces having the highest impact on the shape of the humerus. Co-variation between long bone shape and ground reaction forces was detected in both the humerus and ulna even when accounting for differences in body mass. DISCUSSION: These results demonstrate the importance of considering limb-loading beyond just peak vertical force, or substrate reaction force. A re-evaluation of osseous morphology and functional interpretations is necessary in light of these findings.

Subcortical barrelette-like and barreloid-like structures in the prosimian galago (Otolemur garnetti).

Galagos are prosimian primates that resemble ancestral primates more than most other extant primates. As in many other mammals, the facial vibrissae of galagos are distributed across the upper and lower jaws and above the eye. In rats and mice, the mystacial macrovibrissae are represented throughout the ascending trigeminal pathways as arrays of cytoarchitecturally distinct modules, with each module having a nearly one-to-one relationship with a specific facial whisker. The macrovibrissal representations are termed barrelettes in the trigeminal somatosensory brainstem, barreloids in the ventroposterior medial subnucleus of the thalamus, and barrels in primary somatosensory cortex. Despite the presence of facial whiskers in all nonhuman primates, barrel-like structures have not been reported in primates. By staining for cytochrome oxidase, Nissl, and vesicular glutamate transporter proteins, we show a distinct array of barrelette-like and barreloid-like modules in the principal sensory nucleus, the spinal trigeminal nucleus, and the ventroposterior medial subnucleus of the galago, Otolemur garnetti. Labeled terminals of primary sensory neurons in the brainstem and cell bodies of thalamocortically projecting neurons demonstrate that barrelette-like and barreloid-like modules are located in areas of these somatosensory nuclei that are topographically consistent with their role in facial touch. Serendipitously, the plane of section that best displays the barreloid-like modules reveals a remarkably distinct homunculus-like patterning which, we believe, is one of the clearest somatotopic maps of an entire body surface yet found.

Another look at the foramen magnum in bipedal mammals.

A more anteriorly positioned foramen magnum evolved in concert with bipedalism at least four times within Mammalia: once in macropodid marsupials, once in heteromyid rodents, once in dipodid rodents, and once in hominoid primates. Here, we expand upon previous research on the factors influencing mammalian foramen magnum position (FMP) and angle with four new analyses. First, we quantify FMP using a metric (basioccipital ratio) not previously examined in a broad comparative sample of mammals. Second, we evaluate the potential influence of relative brain size on both FMP and foramen magnum angle (FMA). Third, we assess FMP in an additional rodent clade (Anomaluroidea) containing bipedal springhares (Pedetes spp.) and gliding/quadrupedal anomalures (Anomalurus spp.). Fourth, we determine the relationship between measures of FMP and FMA in extant hominoids and an expanded mammalian sample. Our results indicate that bipedal/orthograde mammals have shorter basioccipitals than their quadrupedal/non-orthograde relatives. Brain size alone has no discernible effect on FMP or FMA. Brain size relative to palate size has a weak influence on FMP in some clades, but effects are not evident in all metrics of FMP and are inconsistent among clades. Among anomaluroids, bipedal Pedetes exhibits a more anterior FMP than gliding/quadrupedal Anomalurus. The relationship between FMA and FMP in hominoids depends on the metric chosen for quantifying FMP, and if modern humans are included in the sample. However, the relationship between FMA and FMP is nonexistent or weak across rodents, marsupials, and, to a lesser extent, strepsirrhine primates. These results provide further evidence that bipedal mammals tend to have more anteriorly positioned foramina magna than their quadrupedal close relatives. Our findings also suggest that the evolution of FMP and FMA in hominins may not be closely coupled.

Reversible Deactivation of Motor Cortex Reveals Functional Connectivity with Posterior Parietal Cortex in the Prosimian Galago (Otolemur garnettii).

We examined the functional macrocircuitry of frontoparietal networks in the neocortex of prosimian primates (Otolemur garnettii) using a microfluidic thermal regulator to reversibly deactivate selected regions of motor cortex (M1). During deactivation of either forelimb or mouth/face movement domains within M1, we used long-train intracortical microstimulation techniques to evoke movements from the rostral division of posterior parietal cortex (PPCr). We found that deactivation of M1 movement domains in most instances abolished movements evoked in PPCr. The most common effect of deactivating M1 was to abolish evoked movements in a homotopic domain in PPCr. For example, deactivating M1 forelimb lift domains resulted in loss of evoked movement in forelimb domains in PPCr. However, at some sites, we also observed heterotopic effects; deactivating a specific domain in M1 (e.g., forelimb lift) resulted in loss of evoked movement in a different movement domain in PPCr (e.g., hand-to-mouth or eye-blink). At most sites examined in PPCr, rewarming M1 resulted in a reestablishment of the baseline movement at the same amplitude as that observed before cooling. However, at some sites, reactivation did not result in a return to baseline movement or to the full amplitude of the baseline movement. We discuss our findings in the context of frontoparietal circuits and how they may subserve a repertoire of ecologically relevant behaviors. SIGNIFICANCE STATEMENT: The posterior parietal cortex (PPC) of primates integrates sensory information used to guide movements. Different modules within PPC and motor cortex (M1) appear to control various motor behaviors (e.g., reaching, defense, and feeding). How these modules work together may vary across species and may explain differences in dexterity and even the capacity for tool use. We investigated the functional connectivity of these modules in galagos, a prosimian primate with relatively simple frontoparietal circuitry. By deactivating a reaching module in M1, we interfered with the function of similar PPC modules and occasionally unrelated PPC modules as well (e.g., eye blink). This circuitry in galagos, therefore, is more complex than in nonprimates, indicating that it has been altered with the expansion of primate PPC.

New euprimate postcrania from the early Eocene of Gujarat, India, and the strepsirrhine-haplorhine divergence.

The oldest primates of modern aspect (euprimates) appear abruptly on the Holarctic continents during a brief episode of global warming known as the Paleocene-Eocene Thermal Maximum, at the beginning of the Eocene (∼56 Ma). When they first appear in the fossil record, they are already divided into two distinct clades, Adapoidea (basal members of Strepsirrhini, which includes extant lemurs, lorises, and bushbabies) and Omomyidae (basal Haplorhini, which comprises living tarsiers, monkeys, and apes). Both groups have recently been discovered in the early Eocene Cambay Shale Formation of Vastan lignite mine, Gujarat, India, where they are known mainly from teeth and jaws. The Vastan fossils are dated at ∼54.5 Myr based on associated dinoflagellates and isotope stratigraphy. Here, we describe new, exquisitely preserved limb bones of these Indian primates that reveal more primitive postcranial characteristics than have been previously documented for either clade, and differences between them are so minor that in many cases we cannot be certain to which group they belong. Nevertheless, the small distinctions observed in some elements foreshadow postcranial traits that distinguish the groups by the middle Eocene, suggesting that the Vastan primates-though slightly younger than the oldest known euprimates-may represent the most primitive known remnants of the divergence between the two great primate clades.

Locomotor pattern fails to predict foramen magnum angle in rodents, strepsirrhine primates, and marsupials.

Foramen magnum position has traditionally been used as an indicator of bipedality because it has been thought to favor a more "balanced" skull position. Here, we analyzed foramen magnum angle (FMA) in relation to locomotion in three mammalian orders that include bipedal or orthograde species in addition to quadrupedal or pronograde species. In marsupials and strepsirrhine primates, we found that there is no relationship between locomotor pattern and FMA. In rodents, we found that there is a significant difference in FMA between bipedal and quadrupedal rodents. However, when these species are analyzed in the context of enlarged auditory bullae, this relationship is no longer significant. Additionally, we find a significant relationship between relative brain size and FMA in strepsirrhine primates. Taken together, these data indicate that several developmental modules of the cranium influence FMA, but that locomotion does not. We caution that basicranial evolution is a complex phenomenon that must be explored in the context of each taxon's unique evolutionary and developmental history.

Lower molar shape and size in prosimian and platyrrhine primates.

The goal of this research is to evaluate the relative strength of the influences of diet, size, and phylogenetic signal on dental geometric shape. Accurate comprehension of these factors and their interaction is important for reconstructing diet and deriving characters for a cladistic analysis in fossil primates. Geometric morphometric analysis is used to identify axes of shape variation in the lower second molars of (a) prosimian primates and (b) platyrrhines. Landmarks were placed on µCT-generated surface renderings. Landmark configurations were aligned using generalized Procrustes analysis. Principal components analysis and phylogenetic principal components analysis (pPCA) were performed on species average landmark co-ordinates. pPCs were examined with phylogenetic generalized least squares analysis for association with size and with diet. PCs from both phylogenetic and non-phylogenetic analyses were sufficient to separate species by broad dietary categories, including insectivores and folivores. In neither analysis was pPC1 correlated with tooth size, but some other pPCs were significantly correlated with size. The pattern of association between pPCs and size altered when centroid size and dietary variables were combined in the model; effects of diet factors typically exceeded effects of size. These results indicate a dominant phylogenetic and dietary signal in molar shape but also show some shape change correlated with size in the absence of obvious dietary associations. Geometric morphometric analysis appears to be useful for tracking functional traits in molars, particularly in tracking differences between folivorous and insectivorous species.

Craniomandibular signals of diet in adapids.

OBJECTIVES: The craniomandibular morphology of the adapid primates of Europe, especially Adapis and Leptadapis (sensu lato), suggests that they possessed enormous jaw adductor muscles. The goal of this study is to estimate jaw adductor muscle mass, physiological cross-sectional area (PCSA), and fiber length in adapid primates from the Eocene of Europe. We also estimated muscle leverage, bite force, and gape parameters. MATERIALS AND METHODS: We use bony morphology and osteological correlates of soft tissues in a sample of extant strepsirrhines to estimate these soft-tissue and performance variables in Adapis and Leptadapis. RESULTS: Our results suggest that, compared to a broad sample of extant strepsirrhines, Adapis and Leptadapis had relatively great jaw adductor muscle mass, PCSA, and bite force. They had moderately great jaw adductor leverage but no sign of adaptation for wide gapes. There is no support for the hypothesis that either adapid was a gouger. DISCUSSION: Our estimates support the inference that Adapis and Leptadapis were primarily folivorous, perhaps also consuming small to medium-sized tough fruits, nuts, and seeds. Explanations for the likely extreme development of the jaw adductor muscles in adapids remain speculative. These include (1) foods that were generally tougher and/or of higher yield strength than those eaten by strepsirrhines today, (2) using the muscles "in shifts" to avoid muscle fatigue in the context of an obdurate diet, and (3) potential constraints on reshaping of the skull for more efficient food processing.

Dental topography of platyrrhines and prosimians: convergence and contrasts.

Dental topographic analysis is the quantitative assessment of shape of three-dimensional models of tooth crowns and component features. Molar topographic curvature, relief, and complexity correlate with aspects of feeding behavior in certain living primates, and have been employed to investigate dietary ecology in extant and extinct primate species. This study investigates whether dental topography correlates with diet among a diverse sample of living platyrrhines, and compares platyrrhine topography with that of prosimians. We sampled 111 lower second molars of 11 platyrrhine genera and 121 of 20 prosimian genera. For each tooth we calculated Dirichlet normal energy (DNE), relief index (RFI), and orientation patch count (OPCR), quantifying surface curvature, relief, and complexity respectively. Shearing ratios and quotients were also measured. Statistical analyses partitioned effects of diet and taxon on topography in platyrrhines alone and relative to prosimians. Discriminant function analyses assessed predictive diet models. Results indicate that platyrrhine dental topography correlates to dietary preference, and platyrrhine-only predictive models yield high rates of accuracy. The same is true for prosimians. Topographic variance is broadly similar among platyrrhines and prosimians. One exception is that platyrrhines display higher average relief and lower relief variance, possibly related to lower relative molar size and functional links between relief and tooth longevity distinct from curvature or complexity. Explicitly incorporating phylogenetic distance matrices into statistical analyses of the combined platyrrhine-prosimian sample results in loss of significance of dietary effects for OPCR and SQ, while greatly increasing dietary significance of RFI.

New insights into the ear region anatomy and cranial blood supply of advanced stem Strepsirhini: evidence from three primate petrosals from the Eocene of Chambi, Tunisia.

We report the discovery of three isolated primate petrosal fragments from the fossiliferous locality of Chambi (Tunisia), a primate-bearing locality dating from the late early to the early middle Eocene. These fossils display a suite of anatomical characteristics otherwise found only in strepsirhines, and as such might be attributed either to Djebelemur or/and cf. Algeripithecus, the two diminutive stem strepsirhine primates recorded from this locality. Although damaged, the petrosals provide substantial information regarding the ear anatomy of these advanced stem strepsirhines (or pre-tooth-combed primates), notably the patterns of the pathway of the arterial blood supply. Using µCT-scanning techniques and digital segmentation of the structures, we show that the transpromontorial and stapedial branches of the internal carotid artery (ICA) were present (presence of bony tubes), but seemingly too small to supply enough blood to the cranium alone. This suggests that the ICA was not the main cranial blood supply in stem strepsirhines, but that the pharyngeal or vertebral artery primitively ensured a great part of this role instead, an arterial pattern that is reminiscent of modern cheirogaleid, lepilemurid lemuriforms and lorisiforms. This could explain parallel loss of the ICA functionality among these families. Specific measurements made on the cochlea indicate that the small strepsirhine primate(s) from Chambi was (were) highly sensitive to high frequencies and poorly sensitive to low frequencies. Finally, variance from orthogonality of the plane of the semicircular canals (SCs) calculated on one petrosal (CBI-1-569) suggests that Djebelemur or cf. Algeripithecus likely moved (at least its head) in a way similar to that of modern mouse lemurs.

Brief communication: Effect of size biases in the coefficient of variation on assessing intraspecific variability in the prosimian skeleton.

This study examines the effect of a measurement size bias in coefficients of variation on the evaluation of intraspecific skeletal variability in a sample of eight prosimian species (Eulemur fulvus, Hapalemur griseus, Lemur catta, Varecia variegata, Galago senegalensis, Otolemur crassicaudatus, Nycticebus coucang, and Tarsius syrichta). Measurements with smaller means were expected to have higher coefficients of variation (CVs) due to the impact of instrumental precision on the ability to assess variability. This was evaluated by testing for a negative correlation between CVs and means in the total sample, within each species, and within each measurement, and by testing for the leveraging impact of small measurements on the significance of comparisons of variability between regions of the prosimian skeleton. Three comparisons were made: cranial versus postcranial variability, epiphysis versus diaphysis variability, and forelimb versus hindlimb variability. CVs were significantly negatively correlated with means within the total sample (r(2) = 0.208, P < 0.0001) and within each species. CVs and means were significantly correlated within only three of the measurements, which may reflect the relatively low body size range of the species studied. As predicted by the higher variability of smaller measurements, removing the smallest measurements from comparisons of variable classes containing measurements of different mean magnitudes pushed the comparisons below significance. These results indicate caution should be exercised when using CVs to assess variability across sets of measurements with different means.

Females are the ecological sex: sex-specific body mass ecogeography in wild sifaka populations (Propithecus spp.).

Previous work in primates has shown that body size often covaries with ecological parameters related to resource or energy availability in the environment. This relationship may differ for males and females as access to resources has greater importance for reproductive success in females. We test the hypotheses that (1) female body mass may be more tightly constrained than male body mass by ecological variables, and (2) female body mass may respond more strongly than male body mass to changes in ecological variables (i.e., population-specific female mass may vary more across an ecological gradient than male mass). Specifically, we investigate the relationship between climatic variables and sex-specific body mass in Propithecus, a genus in which species-specific body mass has already been demonstrated to covary significantly with climatic variables. Data from 733 wild sifakas are used to identify sex-specific body mass for 27 populations representing all nine described sifaka species, and climatic data for each population are derived from the WorldClim database. We use phylogenetic generalized least squares models to demonstrate that body mass in both sexes is significantly correlated with annual rainfall and number of dry months. Furthermore, coefficients of determination are always higher for female models, and coefficients for each climatic variable are higher for females in all significant models. These results support the two hypotheses tested, indicating that ecological forces can have a greater impact on female mass than on male mass in primates.

Seeing the wood through the trees: the current state of higher systematics in the Strepsirhini.

Strepsirhines comprise 10 living or recently extinct families, ≥50% of extant primate families. Their phylogenetic relationships have been intensively studied, but common topologies have only recently emerged; e.g. all recent reconstructions link the Lepilemuridae and Cheirogaleidae. The position of the indriids, however, remains uncertain, and molecular studies have placed them as the sister to every clade except Daubentonia, the preferred sister group of morphologists. The node subtending Afro-Asian lorisids has been similarly elusive. We probed these phylogenetic inconsistencies using a test data set including 20 strepsirhine taxa and 2 outgroups represented by 3,543 mtDNA base pairs, and 43 selected morphological characters, subjecting the data to maximum parsimony, maximum likelihood and Bayesian inference analyses, and reconstructing topology and node ages jointly from the molecular data using relaxed molecular clock analyses. Our permutations yielded compatible but not identical evolutionary histories, and currently popular techniques seem unable to deal adequately with morphological data. We investigated the influence of morphological characters on tree topologies, and examined the effect of taxon sampling in two experiments: (1) we removed the molecular data only for 5 endangered Malagasy taxa to simulate 'extinction leaving a fossil record'; (2) we removed both the sequence and morphological data for these taxa. Topologies were affected more by the inclusion of morphological data only, indicating that palaeontological studies that involve inserting a partial morphological data set into a combined data matrix of extant species should be interpreted with caution. The gap of approximately 10 million years between the daubentoniid divergence and those of the other Malagasy families deserves more study. The apparently contemporaneous divergence of African and non-daubentoniid Malagasy families 40-30 million years ago may be related to regional plume-induced uplift followed by a global period of cooling and drying.