Deciphering the mandibular shape variation in a group of Malagasy primates using Fourier outline analysis.

OBJECTIVES: Among living Malagasy primates, the family Lemuridae has previously been recognized as presenting a higher mandibular morphological variation than other families. We conducted a quantitative analysis of mandibular size and shape within the five genera (Lemur, Eulemur, Hapalemur, Prolemur, and Varecia) associated with a set of covariables that could explain this variation. MATERIALS AND METHODS: We used Fourier outline analysis on the left hemimandible of 182 specimens covering the Lemuridae family. The influence of the phylogeny but also seven covariables (genus, diet, sex, sexual behavior, mating system, ecoregion, and forest type) on mandibular variation was examined using multivariate statistics and model selection. RESULTS: Our results indicate that the high level of morphological variation within the family, associated with a phylogenetic effect and differences in diet, is due to a strong distinction between the genera Prolemur and Hapalemur and the other genera of the family. A second analysis, correcting this strong effect, indicates that mandibular shape variation is influenced not only by the phylogeny and the diet but by a combination of all the covariables. DISCUSSION: The analysis of morphological variation is a powerful tool with major applications, both for the estimation of biological diversity and for the understanding of the fundamental parameters of species' ecology. Our work indicates that, if mandibular shape variation is mainly driven by dietary adaptation, other variables describing ecology and habitat should be considered and taken into account for an integrative understanding of species resources and the establishment of conservation measures.

Myological variation in the forearm anatomy of Callitrichidae and Lemuridae.

The anatomy of the primate forearm is frequently investigated in terms of locomotor mode, substrate use, and manual dexterity. Such studies typically rely upon broad, interspecific samples for which one or two representative taxa are used to characterize the anatomy of their genus or family. To interpret variation between distantly related taxa, however, it is necessary to contextualize these differences by quantifying variation at lower hierarchical levels, that is, more fine-grained representation within specific genera or families. In this study, we present a focused evaluation of the variation in muscle organization, integration, and architecture within two speciose primate families: the Callitrichidae and Lemuridae. We demonstrate that, within each lineage, several muscle functional groups exhibit substantial variation in muscle organization. Most notably, the digital extensors appear highly variable (particularly among callitrichids), with many unique configurations represented. In terms of architectural variables, both families are more conservative, with the exception of the genus Callimico-for which an increase is observed in forearm muscle mass and strength. We suggest this reflects the increased use of vertical climbing and trunk-to-trunk leaping within this genus relative to the more typically fine-branch substrate use of the other callitrichids. Overall, these data emphasize the underappreciated variation in forearm myology and suggest that overly generalized typification of a taxon's anatomy may conceal significant intraspecific and intrageneric variation therein. Thus, considerations of adaptation within the forearm musculature should endeavor to consider the full range of anatomical variation when making comparisons between multiple taxa within an evolutionary context.

Vertical Clinging and Leaping Ahead: How Bamboo Has Shaped the Anatomy and Physiology of Hapalemur.

Hapalemur sps. and Prolemur simus (bamboo lemurs, collectively) stand out from the relatively homogeneous lemurids because they are bamboo feeders and vertical clingers and leapers. This unique diet presents equally unique challenges, like its verticality, toughness, and toxicity. The bamboo lemurs share the generalized anatomy of the other lemurids, but also display some well-documented skeletal adaptations, perhaps to overcome the problems presented by their specialization. Soft-tissue adaptations, however, remain largely unexplored. Explored here are possible soft-tissue adaptations in Hapalemur griseus. We compare H. griseus with other lemurids, Propithecus, Galago, Tarsier, and a tree shrew. Based on the available anatomical and physiological data, we hypothesize that Hapalemur and Prolemur species will have differences in hindlimb morphology when compared with other lemurids. We predict that H. griseus will have more hindlimb muscle mass and will amplify muscle mass differences with increased type II muscle fibers. Relative hindlimb muscle mass in H. griseus is less than other prosimians sampled, yet relative sural muscle mass is significantly heavier (P < 0.01) in H. griseus. Results show that the soleus muscle of H. griseus has a higher amount of type II (fast) fibers in plantarflexors. These findings indicate although H. griseus shares some generalized lemurid morphology, its diet of bamboo may have pushed this generalized lemurid to an anatomical extreme. We suspect additional bamboo-specific adaptations in their anatomy and physiology will be uncovered with further examination into the anatomy of the bamboo lemurs. Anat Rec, 2019. © 2019 Wiley Periodicals, Inc. Anat Rec, 303:295-307, 2020. © 2019 American Association for Anatomy.

Use of space and home range characteristics of Lepilemur mittermeieri, an endangered sportive lemur endemic to the Ampasindava peninsula, north-west Madagascar.

Lepilemur mittermeieri, a little-studied sportive lemur of north-west Madagascar, endemic to the Ampasindava Peninsula, faces habitat loss through forest degradation and rapid fragmentation. Understanding its habitat requirement is the first step toward preservation of this threatened forest-dependent species. In this study, we gathered data on the use of space and home range characteristics of L. mittermeieri. We studied individuals from early March to the end of June 2015 and 2016, in three sites of the Ampasindava peninsula. We radio-tracked 15 individuals to obtain detailed information on the size and location of home ranges (around 450 hr of tracking). Direct observation and morphometric measurements provided additional data sets. Both kernel density estimation (KDE) and minimum convex polygon (MCP) methods yielded similar home range sizes (an average of 2.01 ha with KDE method and 1.96 ha with MCP method). We did not find differences in home range size between males and females, with respect to forest type or proximity to the forest edge. Home ranges overlapped and individuals showed low levels of territoriality. We highlighted a sexually-dimorphic trait: males have longer upper canine than females. Our results constitute the first set of ecological information on Lepilemur mittermeieri and could be the basis for a conservation strategy for this endangered species with a very small distribution area.

Pitch control and speed limitation during overground deceleration in lemurid primates.

An animal's fitness is influenced by the ability to move safely through its environment. Recent models have shown that aspects of body geometry, for example, limb length and center of mass (COM) position, appear to set limits for pitch control in cursorial quadrupeds. Models of pitch control predict that the body shape of these and certain other primates, with short forelimbs and posteriorly positioned COM, should allow them to decelerate rapidly while minimizing the risk of pitching forward. We chose to test these models in two non-cursorial lemurs: Lemur catta, the highly terrestrial ring-tailed lemur, and Eulemur fulvus, the highly arboreal brown lemur. We modeled the effects of changes in limb length and COM position on maximum decelerative potential for both species, as well as collecting data on maximal decelerations across whole strides. In both species, maximum measured decelerations fell below the range of pitch-limited deceleration values predicted by the geometric model, with the ring-tailed lemur approaching its pitch limit more closely. Both lemurs showed decelerative potential equivalent to or higher than horses, the only comparative model currently available. These data reinforce the hypothesis that a relatively simple model of body geometry can predict aspects of maximum performance in animals. In this case, it appears that the body geometry of primates is skewed toward avoiding forward pitch in maximal decelerations.

Anatomy, histology, and ultrasonography of the normal adrenal gland in brown lemur: Eulemur fulvus.

BACKGROUND: The medical care currently to brown lemurs (Eulemur fulvus) is limited by a lack of knowledge of their anatomy. The aim of this study was to describe the anatomy and histology and obtain ultrasonographic measurements of normal adrenal glands in these animals. METHODS: The adrenal glands of four lemurs cadavers were used for the anatomical and histological studies, and those of 15 anesthetized lemurs were examined by ultrasonography. RESULTS: Anatomically, the adrenal glands of brown lemurs are comparable to those of other species. The histological findings showed that the cortex is organized into three distinct layers, whereas most domestic mammals have an additional zone. The surface area of the adrenal glands increased with body weight, and the area of the right adrenal was slightly larger than the left. CONCLUSIONS: We suggest using ultrasonography to aid the etiological diagnosis of behavioral abnormalities that might be due to dysfunctions of the adrenal gland.

Djebelemur, a tiny pre-tooth-combed primate from the Eocene of Tunisia: a glimpse into the origin of crown strepsirhines.

BACKGROUND: Molecular clock estimates of crown strepsirhine origins generally advocate an ancient antiquity for Malagasy lemuriforms and Afro-Asian lorisiforms, near the onset of the Tertiary but most often extending back to the Late Cretaceous. Despite their inferred early origin, the subsequent evolutionary histories of both groups (except for the Malagasy aye-aye lineage) exhibit a vacuum of lineage diversification during most part of the Eocene, followed by a relative acceleration in diversification from the late Middle Eocene. This early evolutionary stasis was tentatively explained by the possibility of unrecorded lineage extinctions during the early Tertiary. However, this prevailing molecular view regarding the ancient origin and early diversification of crown strepsirhines must be viewed with skepticism due to the new but still scarce paleontological evidence gathered in recent years. METHODOLOGICAL/PRINCIPAL FINDINGS: Here, we describe new fossils attributable to Djebelemur martinezi, a≈50 Ma primate from Tunisia (Djebel Chambi). This taxon was originally interpreted as a cercamoniine adapiform based on limited information from its lower dentition. The new fossils provide anatomical evidence demonstrating that Djebelemur was not an adapiform but clearly a distant relative of lemurs, lorises and galagos. Cranial, dental and postcranial remains indicate that this diminutive primate was likely nocturnal, predatory (primarily insectivorous), and engaged in a form of generalized arboreal quadrupedalism with frequent horizontal leaping. Djebelemur did not have an anterior lower dentition as specialized as that characterizing most crown strepsirhines (i.e., tooth-comb), but it clearly exhibited a transformed antemolar pattern representing an early stage of a crown strepsirhine-like adaptation ("pre-tooth-comb"). CONCLUSIONS/SIGNIFICANCE: These new fossil data suggest that the differentiation of the tooth-comb must postdate the djebelemurid divergence, a view which hence constrains the timing of crown strepsirhine origins to the Middle Eocene, and then precludes the existence of unrecorded lineage extinctions of tooth-combed primates during the earliest Tertiary.

Clinal variation in a brown lemur (Eulemur spp.) hybrid zone: combining morphological, genetic and climatic data to examine stability.

Studies of hybrid zones can inform our understanding of reproductive isolation and speciation. Two species of brown lemur (Eulemur rufifrons and E. cinereiceps) form an apparently stable hybrid zone in the Andringitra region of southeastern Madagascar. The aim of this study was to identify factors that contribute to this stability. We sampled animals at 11 sites along a 90-km transect through the hybrid zone and examined variation in 26 microsatellites, the D-loop region of mitochondrial DNA, six pelage and nine morphological traits; we also included samples collected in more distant allopatric sites. Clines in these traits were noncoincident, and there was no increase in either inbreeding coefficients or linkage disequilibrium at the centre of the zone. These results could suggest that the hybrid zone is maintained by weak selection against hybrids, conforming to either the tension zone or geographical selection-gradient model. However, a closer examination of clines in pelage and microsatellites indicates that these clines are not sigmoid or stepped in shape but instead plateau at their centre. Sites within the hybrid zone also occur in a distinct habitat, characterized by greater seasonality in precipitation and lower seasonality in temperature. Together, these findings suggest that the hybrid zone may follow the bounded superiority model, with exogenous selection favouring hybrids within the transitional zone. These findings are noteworthy, as examples supporting the bounded superiority model are rare and may indicate a process of ecologically driven speciation without geographical isolation.

Shouldering the burdens of locomotion and posture: glenohumeral joint structure in prosimians.

Despite its importance in movement of the upper limb, the soft-tissue morphology of the shoulder joint complex (the acromioclavicular, coracoclavicular, and glenohumeral joints) across primates is poorly understood. This study compares soft-tissue morphology of these three shoulder joint components among broad phylogenetic, locomotor, and postural behavior ranges in prosimian primates. Two adult specimens of Galago moholi (a vertical clinger and leaper) were dissected for study, along with one adult each of Cheirogaleus medius (an arboreal quadruped), Eulemur macaco (an arboreal quadruped that also frequently engages in suspensory behavior), and Tarsius syrichta (a vertical clinger and leaper). Because of their role in glenohumeral joint movement and stabilization, the rotator cuff muscles were also dissected and weighed among the species. Results showed that muscle mass of individual components of the rotator cuff musculature may be adaptive to locomotor and postural behaviors of the taxa in this study. Two soft-tissue components of the glenohumeral joint, but not the acromioclavicular and coracoclavicular joints, were also considered adaptive. The quadrupedal species, C. medius and E. macaco, both had glenohumeral ligaments and E. macaco had a relatively deeper glenoid articular surface for the humerus because of the shape of the glenoid labrum. Additionally, this study noted a lack of a teres minor muscle in G. moholi, C. medius, and E. macaco despite previous studies describing them. A relatively robust teres minor muscle was found in T. syrichta. Even with the limited sample dissected here, these results suggest that soft-tissue joint morphology itself may be as adaptive to locomotory and postural styles as osseous morphology.

Gastrointestinal morphology of the crowned lemur (Eulemur coronatus).

I provide measurements of the gastrointestinal tract of a captive female Eulemur coronatus, obtained at necropsy, and describe its morphology. The small intestine, caecum and colon were short when compared with those of other lemur species. The ratio of intestine length to body length was low. Distinct sacculations were present in the caecum, but not in the colon. The results suggest that E. coronatus is able to digest a certain amount of structural polysaccharides as contained in plant cell wall. The main fermentation chamber seems to be the caecum. Its digestive morphology implies that E. coronatus relies on a diet higher in energy-rich cell contents than those of other lemur species of similar body size.

Lemuriform origins as viewed from the fossil record.

Fossils relevant to lemuriform origins are reviewed. Omanodon seems very close to the other early tooth-combed lemuriforms Karanisia, Wadilemur and Saharagalago, whereas Bugtilemur is rejected from the Lemuriformes. The Djebelemurinae, including Djebelemur and 'Anchomomys' milleri, are considered as stem lemuriforms preceding tooth comb differentiation; they are shown to be very distinct from European adapiforms. With tooth-combed lemuriforms present in Africa around 40 million years ago, and stem lemuriforms without tooth combs present on the same continent around 50-48 million years ago, a reasonable scenario can be proposed: tooth comb differentiation and lemuriform dispersal to Madagascar between 52-40 million years ago. The possible significance of Plesiopithecus for daubentoniid origins is raised. A critique of molecular dates is presented in the light of the fossil record. Azibiids are possibly early African prosimians. The timing of the dispersal of primates to Africa and the problem of strepsirhine origins are briefly examined.

Feeding patterns in a small-bodied nocturnal folivore (Avahi laniger) and the influence of leaf chemistry: a preliminary study.

We present data on feeding patterns in Avahi laniger and compare these data to temporal variations in leaf chemistry. Because A. laniger is one of the smallest folivorous primates and has a monogastric stomach, we hypothesized that this lemur would display behavioural adaptations to a leaf-based diet by scheduling feeding times when leaves were of highest quality. Data were collected from May to August 2004 at the Vatoharanana site in Ranomafana National Park, Madagascar. A. laniger fed during different time periods despite leaf carbohydrate and protein concentrations exhibiting little variation throughout the night. Although tannin concentrations exhibited temporal fluctuations, they did not covary with A. laniger feeding times. We suggest that A. laniger feeding times cannot be explained entirely by variations in leaf chemistry.

Shape of the lateral mandibular outline in Lemuridae: a quantitative analysis of variability using elliptical Fourier analysis.

While several morphometric analyses in lemurids have focused on the craniofacial complex, the characterization of their mandibular morphology has received less attention. The mandibular outline, in lateral perspective, was quantified using elliptical Fourier analysis, in an osteological sample encompassing 189 lemurid mandibles (66 Eulemur, 51 Hapalemur, 22 Lemur and 50 Varecia), and compared using multivariate statistical techniques. The taxonomic value of this outline in Lemuridae was demonstrated by the existence of significant separations between the four genera studied. In particular, the mandibular morphology of Hapalemur was markedly different from that in the group Eulemur-Lemur-Varecia. Excluding Hapalemur from analysis, the distinctions between Eulemur, Lemur and Varecia were enhanced suggesting the existence of more subtle intergeneric differences in mandibular morphology. Variation in mandibular form was greatest in Hapalemur and smallest in Eulemur and Varecia (as demonstrated by the mean values of interindividual distances); variation was higher in Lemur than in Eulemur and Varecia, but not higher than in Hapalemur. This morphological diversity may be related to functional adaptation in response to particular dietary habits. The patterns of intergeneric and intrageneric shape variations of the mandible in Lemuridae presented here provide a valuable resource for the analysis of variation among living and fossil lemurids.

The hands and feet of Archaeolemur: metrical affinities and their functional significance.

Recent expeditions to Madagascar have recovered abundant skeletal remains of Archaeolemur, one of the so-called "monkey lemurs" known from Holocene deposits scattered across the island. These new skeletons are sufficiently complete to permit reassembly of entire hands and feet--postcranial elements crucial to drawing inferences about substrate preferences and positional behavior. Univariate and multivariate analysis of intrinsic hand and foot proportions, phalangeal indices, relative pollex and hallux lengths, phalangeal curvature, and distal phalangeal shape reveal a highly derived and unique morphology for an extinct strepsirrhine that diverges dramatically from that of living lemurs and converges in some respects on that of Old World monkeys (e.g., mandrills, but not baboons or geladas). The hands and feet of Archaeolemur are relatively short (extremely so relative to body size); the carpus and tarsus are both "long" relative to total hand and foot lengths, respectively; phalangeal indices of both the hands and feet are low; both pollex and hallux are reduced; the apical tufts of the distal phalanges are very broad; and the proximal phalanges are slightly curved (but more so than in baboons). Overall grasping capabilities may have been compromised to some extent, and dexterous handling of small objects seems improbable. Deliberate and noncursorial quadrupedalism was most likely practiced on both the ground and in the trees. A flexible locomotor repertoire in conjunction with a eurytopic trophic adaptation allowed Archaeolemur to inhabit much of Madagascar and may explain why it was one of the latest surviving subfossil lemurs.

Phylogeny and character behavior in the family Lemuridae.

A phylogenetic analysis of the family Lemuridae was accomplished using multiple gene partitions and morphological characters. The results of the study suggest that several nodes in the lemurid phylogeny can be robustly resolved; however, the relationships of the species within the genus Eulemur are problematically nonrobust. The genus Varecia is strongly supported as the basal genus in the family. Hapalemur and Lemur catta are strongly supported as sister taxa and together are the sister group to the genus Eulemur. E. mongoz is the most basal species in the genus Eulemur. E. fulvus subspecies form a monophyletic group with three distinct lineages. E. coronatus is strongly supported as the sister taxon to E. macaco. The relationships of E. rubriventer, E. fulvus, and the E. macaco-E. coronatus pair are unresolved. Our combined molecular and morphological analysis demonstrates the lack of influence that morphology has on the simultaneous analysis tree when these two kinds of data are given equal weight. The effects of several extreme weighting schemes (removal of transitions and of third positions in protein-coding regions) and maximum-likelihood analysis were also explored. We suggest that these other forms of inference add little to resolving the problematic relationships of the species in the genus Eulemur.

Variation in quantitative measures of enamel prisms from different species as assessed using confocal microscopy.

This study presents a statistical analysis of variability in six measures of enamel prism and ameloblast size and spacing gathered using confocal microscopy, and applies the results to a consideration of appropriate sampling strategies for taxonomic analyses. Variability within individuals was examined within depth series. Individual variability was also assessed within a nested analysis of variation for prism measurements between micrographs, specimens and species. While sample depth was not often significantly associated with differences in prism and ameloblast measures, there was significant variation between micrographs taken from the same region of a tooth. The highest levels of variation were found between species, while variation between conspecific individuals was relatively small. These results demonstrate that data gathered from several micrographs are likely to be representative of a specimen, but that several micrographs of a single specimen will rarely illustrate the range of variation contained within a species. It is essential for systematic and taxonomic analyses that several micrographs be used to characterize an individual. It is also recommended that samples from several individuals be used to characterize species. While data from isolated specimens is often of great interest, taxonomic or systematic conclusions based on isolated individuals should be approached cautiously.

Enamel microstructure in Lemuridae (Mammalia, Primates): assessment of variability.

This study describes the molar enamel microstructure of seven lemurid primates: Hapalemur griseus, Varecia variegata, Lemur catta, Lemur macaco, Lemur fulvus rufus, Lemur fulvus fulvus, and Lemur fulvus albifrons. Contrary to earlier accounts, which reported little or no prism decussation in lemurid enamel, both Lemur and Varecia molars contain a prominent inner layer of decussating prisms (Hunter-Schreger bands), in addition to an outer radial prism layer, and a thin, nonprismatic enamel surface layer. In contrast, Hapalemur enamel consists entirely of radial and, near the surface, nonprismatic enamel. In addition, for all species, prism packing patterns differ according to depth from the tooth surface, and for all species but Varecia (which also has the thinnest enamel of any lemurid), average prism area increases from the enamel-dentine junction to the surface; this may be a developmental solution to the problem of accommodating a larger outer surface area with enamel deposited from a fixed number of cells. Finally, contradicting some previous reports, Pattern 1 prisms predominate only in the most superficial prismatic enamel. In the deeper enamel, prism cross-sections include both closed (Pattern 1) and arc-shaped (Pattern 2 or, most commonly, Pattern 3). This sequence of depth-related pattern change is repeated in all taxa. It should also be emphasized that all taxa can exhibit all three prism patterns in their mature enamel. The high degree of quantitative and qualitative variation in prism size, shape, and packing suggests that these features should be used cautiously in phylogenetic studies. Hapalemur is distinguished from the other lemurids by unique, medially constricted or rectangular prism cross-sections at an intermediate depth and the absence of prism decussation, but, without further assessment of character polarity, these differences do not clarify lemurid phylogenetic relations. Some characters of enamel microstructure may represent synapomorphies of Lemuridae, or of clades within Lemuridae, but homoplasty is likely to be common. Homoplasy of enamel characters may reflect functional constraints.

[Presence of neuritic plaques and neurofibrillary changes in the cerebral cortex of an aged lemuride primate]. / Présence de plaques neuritiques et de modifications neurofibrillaires dans le cortex cérébral d'un primate lémurien âgé.

Some aged Microcebus murinus brains (8 to 11-years old animals) displayed a dramatic atrophy particularly in the neocortex, hippocampus, basal ganglia, hypothalamus, brainstem and cerebellum, associated with a conspicuous increase in the size of the cerebral ventricles. These morphological changes were accompanied by certain histological profiles indicative of pathology. In the cortex, these histological changes consisted of (1) a large number of neuritic plaques formed by degenerated neurites sometimes surrounding an amyloid plaque, (2) dense bundles of argyrophilic filaments in many pyramidal neurons, and (3) amyloid deposits in and around the vascular walls. So far these preliminary studies cannot prone that this degeneration is similar to the human senile dementia of Alzheimer's type, but they may indicate that Microcebus murinus may well be a good model to study cerebral aging, providing comparisons with humans. Furthermore, the size, life span and cost of this species provide further advantages over other nonhuman primates.

[Comparative anatomic studies of the tongue of Pan troglodytes (Blumenbach, 1799) and other primates. I. The chimpanzee tongue]. / Vergleichend-anatomische Untersuchungen an den Zungen von Pan troglodytes (Blumenbach, 1799) und weiteren Primates. Teil I: Befunde an der Schimpansenzunge.

This study was carried out on a comprehensive collection of tongues of Pan troglodytes. Material of tongues of Tupaia glis, Microcebus murinus, Chirogaleus major, Lemur catta, Galago crassicaudatus, Cebus apella, and Macaca mulatta were used for comparison of the different stages of the development of the papilla foliata. The tongue of Pan exhibits longitudinal growth correlated with the growth of length of the jaws and individual age. A typical foramen caecum as it occurs in man was gross anatomically not recognizable. Seldom a small circular depression appeared in the midline close to the papilla circumvallata centralis which could be the remnant of the foramen caecum. There is a remarkable accumulation of papillae filiformes and fungiformes at the apex linguae; the same seems to be true for other genera of primates. It is self-explanatory that the accumulation of papillae makes the tip of the tongue a particularly sensitive area; this has been found to be true even in man. One specimen of the prenatal age just before term was studied. It exhibited remarkably elongated, thread-like papillae, their function is unknown as yet. They are no longer present in a newborn specimen of 4 d of age. In the aboral part of the dorsum linguae, a special kind of papillae occur; they are flat lobes which carry moderately elongated, tiny processes at their free margin. The function of these papillae is unknown; taste buds are absent.

Size, space, and adaptation in some subfossil lemurs from Madagascar.

We examine several explanations for the geographic pattern of body size variation exhibited by the subfossil lemur Archaeolemur. Part and partial correlation analysis and multiple regression analysis are applied in a stepwise, hierarchical fashion to help to determine variable interdependencies. Variance in site means for body size is best explained by the richness of the plant community and by several correlated climatic variables (bioclimatic zone and mean annual rainfall). Body size differentiation in Archaeolemur roughly mirrors patterns observed among many other Malagasy lemur species and subspecies groups. This consistency alone suggests that common ecological factors have strongly affected size differentiation in lemurs, most probably (as suggested by our correlation analyses) by uniformly influencing the productivity of their niches. Smaller individuals tend to inhabit arid regions, and larger individuals tend to inhabit wetter regions. The interplay between selective differentiation and allopatric speciation appears to have yielded the concordant pattern of size variation observed in Malagasy lemurs.