Microcebus Rufus Research Articles

Arboreal mouse lemurs discovered sleeping in a burrow on the ground.

Finding sleeping sites is important for the fitness of many mammal species. Like most nonhuman primates, Madagascar's mouse lemurs (genus: Microcebus) are thought to exclusively use arboreal sleeping sites. The rufous mouse lemurs (Microcebus rufus) in Ranomafana National Park (southeastern Madagascar) have always been documented to sleep in either tree holes or leaf nests. However, in our recent field expedition, we observed, with the help of telemetry technologies, an unprecedented event of M. rufus sleeping in a burrow on the rainforest ground, curled up with a very slow heartbeat. Thus far, such behavior has not been observed in any other Microcebus species but is common in high-altitude dwarf lemurs (genus: Cheirogaleus), a closely related genus to the mouse lemurs. We believe that this discovery could illustrate an ecophysiological response strategy to habitat changes, which warrants further investigation.

Structure of Microhabitats Used by Microcebus rufus Across a Heterogeneous Landscape

Structure of Microhabitats Used by Microcebus rufus Across a Heterogeneous Landscape

Olfactory predator recognition in the brown mouse lemur (Microcebus rufus) in Ranomafana National Park, Madagascar.

Predator odors such as urine and feces are known to elicit antipredator behaviors in prey including avoidance, fear, and curiosity. We measured how wild brown mouse lemurs (Microcebus rufus) responded to odors of mammalian, avian, and snake predators as well as nonpredator controls. The first experiment took place under controlled conditions in a laboratory where we recorded the occurrence of four behavioral categories (ignore, curiosity, alert, and fear) in response to a single odor. Subjects exhibited behavioral change significantly more often in response to the predator than to control stimuli, but did not distinguish between familiar and unfamiliar predators. Mammalian predator urine and feces were most likely to elicit behavioral change. The owl was the only predator to never elicit behavioral change, possibly because owls do not provide relevant odor cues. A second experiment employing live traps in the forest found that neither predator nor control odors affected the likelihood of capture. Due to their longevity, odors do not provide accurate information of spatial and temporal risk, and while mouse lemurs may have initially hesitated to enter a trap, in the absence of additional information about risk, they may have eventually ignored the stimuli. This study found that brown mouse lemurs are able to distinguish between predator and nonpredator odors, and that risk assessment may be affected by the experience, as well as predator and sensory stimulus quality.

Brown Mouse Lemurs (Microcebus rufus) May Lack Opportunities to Learn about Predator Calls

Diurnal primates commonly modify their behaviour in response to predator vocalizations, but little is known about how nocturnal primates perceive predator calls. Mouse lemurs do not innately perceive predator calls as dangerous but may learn to do so in the wild. I hypothesized that wild brown mouse lemurs (Microcebus rufus) in Ranomafana National Park, Madagascar, differentiate between predator and non-predator vocalizations and exposed 29 adult subjects to calls of 3 sympatric predators (goshawk, fossa and owl), 3 sympatric non-predators and 4 controls. Brown mouse lemurs did not differentiate among stimuli, and there was no evidence that any recording was associated with danger, as neither activity nor spatial preferences changed in response to a stimulus, nor did subjects exhibit obvious behaviours indicative of danger such as flight or position on the branches. The failure to perceive predator calls as dangerous is common among small nocturnal solitary prey species and could be explained by a lack of learning opportunities. In social species acquired behaviours can spread quickly by social learning. Solitary species like mouse lemurs have to individually learn to associate predator calls with danger which is dependent on encounter rates. Unlike diurnal prey mouse lemurs lack the opportunity to observe diurnal raptors and associate their calls with the bird, and owls are proficient predators, which are difficult to observe in a rain forest where they hunt cryptically from perches. While sometimes nocturnal, the fossa is solitary and largely silent, and is also particularly rare at my study site. Cryptic, rare or non-vocal predators provide prey with few opportunities to associate their calls with danger. Mouse lemurs are subject to high predation rates by a wide range of predators and need to balance the costs and benefits of antipredation behaviours. Their cryptic lifestyle and preference for dense vegetation lowers the likelihood of being detected by predators, and they utilize visual and olfactory predator cues to make decisions about risk. Furthermore, their high reproductive output compensates for the high predation rates.

Effect of socio-ecological factors and parasite infection on body condition of Brown Mouse Lemur Microcebus rufus (Mammalia: Primates: Cheirogaleidae)

Various studies in ecology have shown the relationship between body condition and parasitic loads in nonhuman primates, however, little information is available regarding prosimians such as lemurs. In this study, the synergistic effect of parasite infection and socio-ecological factors on the body condition of Microcebus rufus in the family Cheirogaleidae was analyzed in Ranomafana National Park in southeastern Madagascar. This lemur species is characterized by its ability to adapt to different types of forest, and by seasonal fattening. Based on the factors considered, this species is, therefore, a good model for the study of body condition and ecology of infectious diseases in lemurs. Floatation and direct observation techniques were used for examination of parasite infection. Two indices considering body condition were analyzed: volume index (VI) and condition index (CI), the residual between the mass observed and the corrected mass. The generalized linear mixed model (GLMM) was used to model the synergistic effect of parasite infections and socio-ecological factors on variation in body condition, with the identity of individuals used as a random factor. We identified five species of helminths, one species of protist, and one species of lice which infected the 204 mouse lemurs captured. There was a sexual difference for all measures of the parasite infection. The more parasite species an individual was infected with, the smaller was its body size according to the Volume Index that reflects deposits of subcutaneous fat. Individuals with more positive Condition Index values, particularly females, excreted more parasite eggs or oocyst in their faecal matter. The results suggest that an individual’s body condition constitutes an indicator of risk of parasite infection and transmission.

Causative agent of canine heartworm (Dirofilaria immitis) detected in wild lemurs.

The lemurs of Madagascar are threatened by human activities. We present the first molecular detection of canine heartworm (Dirofilaria immitis) in a wild non-human primate, the mouse lemur (Microcebus rufus). Zoonotic D. immitis infection has been associated with clinical pathology that includes serious and often fatal cardiac and pulmonary reactions. With human encroachment and associated increases in free-roaming dog populations in Madagascar, we examined lemurs for zoonotic canid pathogens. D. immitis presents a new potential conservation threat to lemurs. We highlight the need for wide-ranging and effective interventions, particularly near protected areas, to address this growing conservation issue.

Drivers of in vivo bite performance in wild brown mouse lemurs and a comparison with the grey mouse lemur

AbstractPhysical performance is crucial for animal survival and fitness. In this context, greater bite forces can provide advantages and may allow an individual to gain access to reproductive partners and/or different food resources. Here, we explored the determinants of bite force in a wild population of the brown mouse lemur (Microcebus rufus). Our objectives were to elucidate (1) if sex, head width, heart rate (as an indicator of overall physical fitness) and body condition drive variation in bite force in this population of wild mouse lemurs; and (2) the relative importance of the ecological niche in determining bite force by comparing results from this wild population with previously published results on bite force, body mass and head width from a laboratory colony of the grey mouse lemur (Microcebus murinus). We captured 32 wild brown mouse lemurs at night in the Ranomafana National Park in Madagascar during the beginning of the rainy season from 1st to 31st October 2016. We measured bite force, heart rate, body mass and head width of all individuals, and assigned sex and body condition (estimated as the unstandardized residual of a regression of body mass against head size). Although maximum bite force was positively correlated with body mass, it was not correlated with body condition. Residual bite force was highly correlated with residual head width and heart rate. The mean bite force of wild brown mouse lemurs was much lower than that of grey mouse lemurs in captivity, but showed similar relationships to head dimension and body mass. Even when corrected by body condition, grey mouse lemurs bit significantly harder than brown mouse lemurs. The difference in bite force between species could be explained by differences in head size and niche divergence with brown mouse lemurs eating mostly soft fruits and grey mouse lemurs eating more hard insects.

Metabarcoding Gastrointestinal Nematodes in Sympatric Endemic and Nonendemic Species in Ranomafana National Park, Madagascar

Sympatric species are known to host the same parasites species. Nevertheless, surveys examining parasite assemblages in sympatric species are rare. To understand how parasite assemblages in sympatric host species differ in a given locality, we used a noninvasive identification method based on high-throughput sequencing. We collected fecal samples from sympatric species in Ranomafana National Park, Madagascar, from September to December in 2010, 2011, and 2012 and identified their parasites by metabarcoding, sequencing a region of the small ribosomal subunit (18S) gene. Our survey included 11 host species, including endemic primates, rodents, frogs, gastropods, and nonendemic rats and dogs. We collected 872 samples, of which 571 contained nematodes and 249 were successfully sequenced. We identified nine putative species of parasites, although their correspondence to actual parasite species is not clear as the resolution of the marker gene differs between nematode clades. For the host species that we successfully sampled with 10 or more positive occurrences of nematodes, i.e., mouse lemurs (Microcebus rufus), black rats (Rattus rattus), and frogs (Anura), the parasite assemblage compositions differed significantly among host species, sampling sites, and sampling years. Our metabarcoding method shows promise in interrogating parasite assemblages in sympatric host species and our results emphasize the importance of choosing marker regions for parasite identification accuracy.

Aggression and hormones are associated with heterogeneity in parasitism and parasite dynamics in the brown mouse lemur

Animal behaviours, like aggression, can directly affect host health by influencing exposure to parasites. Aggressive individuals may experience an increase in agonistic interactions and contact rates with conspecifics, which might increase their probability of acquiring parasites. However, aggression is not the only factor that shapes parasitism; proximate mechanisms like hormone-modulated immunosuppression can also have broad impacts. Here, we hypothesized that high levels of aggression, cortisol and testosterone would be positively associated with parasitism and that aggressive individuals would play a larger role spreading parasites to conspecifics than would docile individuals. We measured aggression using the level of aggressive response to human handling during capture. Our aim was to examine associations between aggression and hormones (cortisol and testosterone) on variation in endo- and ectoparasitism in a population of wild mouse lemurs ( Microcebus rufus ) over a 3-year period. By tracking the movement of lice (directly transmitted parasites) in the population, we also examined the effect of host aggression on population-wide parasite dynamics. We show that animals with high testosterone and cortisol were more likely to exhibit aggressive behaviours, and cortisol was associated with significantly higher ectoparasite infestations. Aggressive individuals were significantly more infested by lice, and also donated significantly more lice to conspecifics in the population. Taken together, our results offer insight into the individual and population health costs of aggression, and empirical support of a trade-off between aggression and ectoparasitism, which may have driven the evolution of aggression and interactions with conspecifics.

Population- and Individual-Level Dynamics of the Intestinal Microbiota of a Small Primate.

ABSTRACTLongitudinal sampling for intestinal microbiota in wild animals is difficult, leading to a lack of information on bacterial dynamics occurring in nature. We studied how the composition of microbiota communities changed temporally in free-ranging small primates, rufous mouse lemurs (Microcebus rufus). We marked and recaptured mouse lemurs during their mating season in Ranomafana National Park in southeastern mountainous rainforests of Madagascar for 2 years and determined the fecal microbiota compositions of these mouse lemurs with MiSeq sequencing. We collected 160 fecal samples from 71 animals and had two or more samples from 39 individuals. We found small, but statistically significant, effects of site and age on microbiota richness and diversity and effects of sex, year, and site on microbiota composition, while the within-year temporal trends were less clear. Within-host microbiota showed pervasive variation in intestinal bacterial community composition, especially during the second study year. We hypothesize that the biological properties of mouse lemurs, including their small body size and fast metabolism, may contribute to the temporal intraindividual-level variation, something that should be testable with more-extensive sampling regimes. IMPORTANCE While microbiome research has blossomed in recent years, there is a lack of longitudinal studies on microbiome dynamics on free-ranging hosts. To fill this gap, we followed mouse lemurs, which are small heterothermic primates, for 2 years. Most studied animals have shown microbiota to be stable over the life span of host individuals, but some previous research also found ample within-host variation in microbiota composition. Our study used a larger sample size than previous studies and a study setting well suited to track within-host variation in free-ranging mammals. Despite the overall microbiota stability at the population level, the microbiota of individual mouse lemurs can show large-scale changes in composition in time periods as short as 2 days, suggesting caution in inferring individual-level patterns from population-level data.

A preliminary investigation of sleeping site selection and sharing by the brown mouse lemurMicrocebus rufusduring the dry season

Mouse lemurs (Microcebus) are solitary, experience heavy predation, and use torpor to conserve energy, so an individual's choice of daytime sleeping site can have a major impact on its fitness. Sharing a sleeping site may strengthen social bonds or insulate against fluctuating ambient temperatures. Changing sites often may prevent tracking by predators. Choosing a site in a hole in an old, thick tree may allow for longer and deeper torpor. The aim of this paper is to provide a preliminary characterization of sleeping site selection and use in Microcebus rufus, the brown mouse lemur. We trapped mouse lemurs in disturbed forests in the peripheral zone of Ranomafana National Park, Madagascar, and tracked 7 individuals using temperature-sensitive radiocollars. Radiotracked individuals shared sleeping sites concurrently and serially and slept in same-sex and mixed-sex groups. They slept in tree holes as well as in piles of leaves on branches. Radiotracked individuals tended to use higher-diameter at breast hei...

Unpredictable environments, opportunistic responses: Reproduction and population turnover in two wild mouse lemur species (Microcebus rufus and M. griseorufus) from eastern and western Madagascar.

Small-bodied, nocturnal mouse lemurs (Microcebus) are widespread across diverse forest habitats in Madagascar. They are strict seasonal breeders and can, depending on the habitat and species, undergo daily or prolonged torpor to minimize energy expenditure during periods of food and water scarcity. Duration of reproduction, number of litters per season and timing of births vary across individuals and species. The "polyestry-seasonality" hypothesis proposes that the duration of reproduction and number of litters per year are positively correlated with rainfall but negatively correlated with longevity, whereas the "hypervariability" hypothesis suggests that the duration of reproduction is negatively correlated with the degree of predictability of food resources. We test these hypotheses in two mouse lemur species inhabiting contrasting habitats, the brown mouse lemurs, Microcebus rufus, from Ranomafana (a less seasonal and more climatically predictable habitat) and the gray-brown mouse lemurs, M. griseorufus, from Beza Mahafaly (a more seasonal and less climatically predictable environment). We use capture/mark/recapture techniques and records of female reproductive status. We found evidence of polyestry at both study sites but faster population turnover and longer duration of the reproductive season at Beza Mahafaly. The "polyestry-seasonality" hypothesis is not supported but the "hypervariability" hypothesis could not be rejected. We conclude that reproductive output cannot be tied to climatic factors in a simple manner. Paradoxically, polyestry can be expressed in contrasting habitats: less seasonal forests where females can sustain multiple reproductive events, but also highly seasonal environments where females may not fatten sufficiently to sustain prolonged torpor but instead remain active throughout the year by relying on fallback resources. Am. J. Primatol. 77:936-947, 2015. © 2015 Wiley Periodicals, Inc.

Tracking year-to-year changes in intestinal nematode communities of rufous mouse lemurs (Microcebus rufus).

While it is known that intestinal parasite communities vary in their composition over time, there is a lack of studies addressing how variation in component communities (between-hosts) manifests in infracommunities (within-host) during the host lifespan. In this study, we investigate the changes in the intestinal parasite infracommunities in wild-living rufous mouse lemurs (Microcebus rufus) from Ranomafana National Park in southeastern Madagascar from 2010 to 2012. We used high-throughput barcoding of the 18S rRNA gene to interrogate parasite community structure. Our results show that in these nematode communities, there were two frequently occurring putative species and four rarer putative species. All putative species were randomly distributed over host individuals and they did not occur in clear temporal patterns. For the individuals caught in at least two different years, there was high turnover of putative species and high variation in fecal egg counts. Our study shows that while there was remarkable variation in infracommunities over time, the component community was relatively stable. Nevertheless, the patterns of prevalence varied substantially between years in each component community.

Extension of gray-brown mouse lemur (<i>Microcebus griseorufus</i>) activity period in a disturbed forest in southwestern Madagascar

Habitat disturbances may impact behaviors of animals, including their activity patterns. In southwestern Madagascar, timing of gray - brown mouse lemur activities was investigated in adjacent forests with different levels of human disturbance. Mouse lemurs were encountered more frequently during the second part of the night in the unfenced, more disturbed forest than in the fenced, less - disturbed forest. The extension of mouse lemur activity period in the unfenced forest may be due to differences in forest composition resulting in higher travel costs or a loss of canopy cover which may limit their use of torpor.

Diversity and Prevalence of Diarrhea-Associated Viruses in the Lemur Community and Associated Human Population of Ranomafana National Park, Madagascar

Diarrhea-associated viruses are common causes of morbidity in humans in developing countries; however, they have seldom been studied in wild primates despite their pathogenic and zoonotic potential. This is of particular concern in Madagascar, one of the world’s poorest and most biodiverse countries. To improve our understanding of diarrhea-associated viruses in human and nonhuman wild primates in Madagascar, we noninvasively collected fecal samples from 7 of the 12 lemur species of Ranomafana National Park, Madagascar and concurrently sampled humans living in proximity to the lemurs. We used standard molecular techniques to screen 84 lemur and 107 human samples for viral groups commonly associated with diarrheal disease in human populations: adenovirus, enterovirus, rotavirus, and norovirus (genogroups GI and GII). Of the seven lemur taxa examined, all were positive for one or more viral group and humans were positive for all viral groups. Norovirus GII was the most common, found in 27 % of lemurs and 37 % of humans, followed by adenovirus, found in 25 % of lemurs and 35 % of humans. The high prevalence (30–50 %) of most viral groups in mouse lemurs (Microcebus rufus) highlights their potential to serve as sentinals of pathogen spillover from humans to wild lemur populations. These results improve our understanding of the diversity and prevalence of key viral taxa in lemur and associated human populations. Further characterization of these viral group sequences has the potential to reveal novel viruses in lemurs and humans in Madagascar and would be needed to determine the zoonotic potential of the viruses detected.

Séance: reference-based phylogenetic analysis for 18S rRNA studies.

BackgroundMarker gene studies often use short amplicons spanning one or more hypervariable regions from an rRNA gene to interrogate the community structure of uncultured environmental samples. Target regions are chosen for their discriminatory power, but the limited phylogenetic signal of short high-throughput sequencing reads precludes accurate phylogenetic analysis. This is particularly unfortunate in the study of microscopic eukaryotes where horizontal gene flow is limited and the rRNA gene is expected to accurately reflect the species phylogeny. A promising alternative to full phylogenetic analysis is phylogenetic placement, where a reference phylogeny is inferred using the complete marker gene and iteratively extended with the short sequences from a metagenetic sample under study.ResultsBased on the phylogenetic placement approach we built Séance, a community analysis pipeline focused on the analysis of 18S marker gene data. Séance combines the alignment extension and phylogenetic placement capabilities of the Pagan multiple sequence alignment program with a suite of tools to preprocess, cluster and visualise datasets composed of many samples. We showcase Séance by analysing 454 data from a longitudinal study of intestinal parasite communities in wild rufous mouse lemurs (Microcebus rufus) as well as in simulation. We demonstrate both improved OTU picking at higher levels of sequence similarity for 454 data and show the accuracy of phylogenetic placement to be comparable to maximum likelihood methods for lower numbers of taxa.ConclusionsSéance is an open source community analysis pipeline that provides reference-based phylogenetic analysis for rRNA marker gene studies. Whilst in this article we focus on studying nematodes using the 18S marker gene, the concepts are generic and reference data for alternative marker genes can be easily created. Séance can be downloaded from http://wasabiapp.org/software/seance/.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-014-0235-7) contains supplementary material, which is available to authorized users.

Teeth, sex, and testosterone: aging in the world's smallest primate.

Mouse lemurs (Microcebus spp.) are an exciting new primate model for understanding human aging and disease. In captivity, Microcebus murinus develops human-like ailments of old age after five years (e.g., neurodegeneration analogous to Alzheimer's disease) but can live beyond 12 years. It is believed that wild Microcebus follow a similar pattern of senescence observed in captive animals, but that predation limits their lifespan to four years, thus preventing observance of these diseases in the wild. Testing whether this assumption is true is informative about both Microcebus natural history and environmental influences on senescence, leading to interpretation of findings for models of human aging. Additionally, the study of Microcebus longevity provides an opportunity to better understand mechanisms of sex-biased longevity. Longevity is often shorter in males of species with high male-male competition, such as Microcebus, but mouse lemurs are sexually monomorphic, suggesting similar lifespans. We collected individual-based observations of wild brown mouse lemurs (Microcebus rufus) from 2003–2010 to investigate sex-differences in survival and longevity. Fecal testosterone was measured as a potential mechanism of sex-based differences in survival. We used a combination of high-resolution tooth wear techniques, mark-recapture, and hormone enzyme immunoassays. We found no dental or physical signs of senescence in M. rufus as old as eight years (N = 189, ages 1–8, mean = 2.59±1.63 SE), three years older than captive, senescent congeners (M. murinus). Unlike other polygynandrous vertebrates, we found no sex difference in age-dependent survival, nor sex or age differences in testosterone levels. While elevated male testosterone levels have been implicated in shorter lifespans in several species, this is one of the first studies to show equivalent testosterone levels accompanying equivalent lifespans. Future research on captive aged individuals can determine if senescence is partially a condition of their captive environment, and studies controlling for various environmental factors will further our understanding of senescence.

Are Tropical Small Mammals Physiologically Vulnerable to Arrhenius Effects and Climate Change?

There is some urgency in the necessity to incorporate physiological data into mechanistic, trait-based, demographic climate change models. Physiological responses at the individual level provide the mechanistic link between environmental changes and individual performances and hence population dynamics. Here we consider the causal relationship between ambient temperature (Ta) and metabolic rate (MR), namely, the Arrhenius effect, which is directly affected by global warming through increases in average global air temperatures and the increase in the frequency and intensity of extreme climate events. We measured and collated data for several small, free-ranging tropical arboreal mammals and evaluated their vulnerability to Arrhenius effects and putative heat stress associated with climate change. Skin temperatures (Tskin) were obtained from free-ranging tarsiers (Tarsius syrichta) on Bohol Island, Philippines. Core body temperature (Tb) was obtained from the greater hedgehog tenrec (Setifer setosus) and the gray brown mouse lemur (Microcebus ravelobensis) from Ankarafantsika, Madagascar. Tskin for another mouse lemur, Microcebus griseorufus, was obtained from the literature. All four species showed evidence of hyperthermia during the daytime rest phase in the form of either Tskin or Tb that was higher than the normothermic Tb during the nighttime active phase. Potentially, tropical arboreal mammals with the lowest MRs and Tb, such as tarsiers, are the most vulnerable to sustained heat stress because their Tb is already close to Ta. Climate change may involve increases in MRs due to Arrhenius effects, especially during the rest phase or during torpor and hibernation. The most likely outcome of increased Arrhenius effects with climate change will be an increase in energy expenditure at the expense of other critical functions such as reproduction or growth and will thus affect fitness. However, we propose that these hypothetical Arrhenius costs can be, and in some species probably are, offset by the use of hyperthermic daily torpor, that is, hypometabolism at high Ta.

Survey of Giardia and Cryptosporidium in lemurs from the Ranomafana National Park, Madagascar

We detected Cryptosporidium sp. by direct immunofluorescence in fecal samples from greater bamboo lemurs (Prolemur simus) and eastern rufous mouse lemurs (Microcebus rufus) inhabiting the Ranomafana National Park, Madagascar. This is the first report of an occurrence of these potentially zoonotic parasites in free-ranging lemurs in the rain forest of Madagascar.

Lumbar Vertebral Body Bone Microstructural Scaling in Small to Medium‐Sized Strepsirhines

Bone mass, architecture, and tissue mineral density contribute to bone strength. As body mass (BM) increases any one or combination of these properties could change to maintain structural integrity. To better understand the structural origins of vertebral fragility and gain insight into the mechanisms that govern bone adaptation, we conducted an integrative analysis of bone mass and microarchitecture in the last lumbar vertebral body from nine strepsirhine species, ranging in size from 42 g (Microcebus rufus) to 2,440 g (Eulemur macaco). Bone mass and architecture were assessed via µCT for the whole body and spherical volumes of interest (VOI). Allometric equations were estimated and compared with predictions for geometric scaling, assuming axial compression as the dominant loading regime. Bone mass, microarchitectural, and vertebral body geometric variables predominantly scaled isometrically. Among structural variables, the degree of anisotropy (Tb.DA) was the only parameter independent of BM and other trabecular architectural variables. Tb.DA was related to positional behavior. Orthograde primates had higher average Tb.DA (1.60) and more craniocaudally oriented trabeculae while lorisines had the lowest Tb.DA (1.25), as well as variably oriented trabeculae. Finally, lorisines had the highest ratio of trabecular bone volume to cortical shell volume (∼3x) and while there appears to be flexibility in this ratio, the total bone volume (trabecular + cortical) scales isometrically (BM(1.23) , r(2) = 0.93) and appears tightly constrained. The common pattern of isometry in our measurements leaves open the question of how vertebral bodies in strepsirhine species compensate for increased BM.