Propithecus Verreauxi Research Articles

Mate-Guarding as a Male Reproductive Tactic in Propithecus verreauxi

Sexual selection theory predicts that in group-living mammals, male reproductive tactics can lead to high reproductive skew in favor of dominant individuals. In sifakas (Propithecus verreauxi), a group-living primate with extremely seasonal reproduction, male reproductive success is highly skewed because dominant males sire almost all offspring despite a tendency toward an even adult group sex ratio. To understand the underlying behavioral mechanism resulting in this rank-related reproductive skew in male sifakas, we studied mate-guarding as a potential reproductive tactic. Behavioral observations of dominant males and adult females in combination with hormonal determination of timing of female receptivity in 9 groups at Kirindy Forest revealed that dominant males spent more time in proximity to females when they were receptive and were responsible for the maintenance of this proximity. Results also indicated that monopolization of receptive females was facilitated by both estrous asynchrony within groups and by the ability of dominant males to obtain olfactory cues as to the timing of female receptivity. Although dominant males engaging in mate-guarding are expected to experience various costs, there was no evidence for decreased foraging behavior and only a trend toward increased aggression between dominant and subordinate non-natal males within groups. Our results are in accordance with the hypothesis that dominant males use mate-guarding to monopolize receptive females and that it is one proximate mechanism that contributes to the high reproductive skew observed within the population of male sifakas at Kirindy.

Mating first, mating more: biological market fluctuation in a wild prosimian.

In biology, economics, and politics, distributive power is the key for understanding asymmetrical relationships and it can be obtained by force (dominance) or trading (leverage). Whenever males cannot use force, they largely depend on females for breeding opportunities and the balance of power tilts in favour of females. Thus, males are expected not only to compete within their sex-class but also to exchange services with the opposite sex. Does this mating market, described for humans and apes, apply also to prosimians, the most ancestral primate group? To answer the question, we studied a scent-oriented and gregarious lemur, Propithecus verreauxi (sifaka), showing female dominance, promiscuous mating, and seasonal breeding. We collected 57 copulations involving 8 males and 4 females in the wild (Berenty Reserve, South Madagascar), and data (all occurrences) on grooming, aggressions, and marking behaviour. We performed the analyses via exact Spearman and matrix correlations. Male mating priority rank correlated with the frequency of male countermarking over female scents but not with the proportion of fights won by males over females. Thus, males competed in an olfactory tournament more than in an arena of aggressive encounters. The copulation frequency correlated neither with the proportion of fights won by males nor with the frequency of male countermarking on female scents. Male-to-female grooming correlated with female-to-male grooming only during premating. Instead, in the mating period male-to-female grooming correlated with the copulation frequency. In short, the biological market underwent seasonal fluctuations, since males bargained grooming for sex in the mating days and grooming for itself in the premating period. Top scent-releasers gained mating priority (they mated first) and top groomers ensured a higher number of renewed copulations (they mated more). In conclusion, males maximize their reproduction probability by adopting a double tactic and by following market fluctuations.

Birth season glucocorticoids are related to the presence of infants in sifaka (Propithecus verreauxi).

The responses of plural breeding mammals to environmental stressors are little understood in free-ranging populations, but recent studies of singular breeders suggest that ecological factors and social milieu influence the variable physiological stress responses observed among individuals. Our previous studies examining faecal glucocorticoid (fGC)-behaviour interactions in plural breeding male sifaka (Propithecus verreauxi) show that fGC elevations coincide with specific dispersal events, particularly the eviction of subordinates by resident alpha males. This study examined the utility of fGC assays for assessing the stress responses of male sifaka to demographic changes in the population during the birth season. Analyses were based on 889 faecal samples collected over five birth seasons from 124 adult males residing in 55 different groups at Beza Mahafaly, Madagascar. Results show that fGC levels in males are unrelated to age, residence, group stability or rank, but are significantly and positively correlated with the presence of infants, annual elevations in weekly mean fGC concentrations in males paralleling increasing numbers of infants born annually in the population. These data are the first to show that in seasonal plural breeding species such as sifaka, elevated fGC in males reflects specific events related to reproduction rather than states or social context during the birth season.

Chest staining variation as a signal of testosterone levels in male Verreaux's Sifaka

Male Verreaux's sifaka ( Propithecus verreauxi) exhibit variation in the staining of chest hair in association with the activity of the sternal gland. Scent-marking behavior and social relationships have been shown to vary with the state of chest staining. Research on other mammals suggests that sternal gland activity is modulated by testosterone. The goal of this study was to examine the relationship among fecal testosterone, testes mass, and chest staining in sifaka. I predicted that for sifaka (1) testosterone and testes mass are positively correlated, (2) stained males have higher testosterone and larger testes than clean-chested males, (3) testes mass is greater during the mating season than the birth season, and (4) testes mass is comparable to species with multimale mating systems. Eighty fecal samples were collected from eight adult males in five social groups in the Kirindy Mitea National Park of Madagascar during the mating season and analyzed using enzyme immunoassays. Males were captured during the mating season and the subsequent birth season to measure body mass and testes size and to document chest staining. Stained males had significantly higher fT and larger testes mass than clean-chested males. Testes mass was significantly greater during the mating season than the birth season for all males. However, the stained males exhibited less testes mass reduction during birth season than clean-chested males. These results suggest that the activity of the sternal gland is regulated by testosterone and that sexual selection has shaped male morphological variation in Verreaux's sifaka.

Social Influences on Group Membership in Propithecus verreauxi verreauxi

The size and composition of a social group can influence the reproductive success of its members. I examined the hypothesis that residents actively try to manipulate group size and composition in Verreaux’s sifaka (Propithecus verreauxi verreauxi). I collected behavioral data from 21 individuals in 5 social groups from November 2000 to March 2002 in the Kirindy Forest (C.F.P.F.) of western Madagascar. I investigated the affiliative and agonistic behaviors of resident males and females toward other group members to determine whether residents attempt to manipulate the costs and benefits of group membership. I also examined 3 cases of immigration attempts and 1 case of emigration. As expected, group members facilitated the residency of subordinate members of the opposite sex. For example, females sought out the subordinate males, responded to their calls, and even groomed them more. However, intrasexual relationships were also important for establishing or maintaining residency. The data, in conjunction with the observations of immigration and emigration events, suggest that resident individuals of both sexes attempt to influence group size and composition. I suggest that single–male groups are not the norm in Propithecus verreauxi verreauxi because females encourage, and males do not discourage, multiple males to reside in the group to increase group stability and to reduce the risk of infanticide.

Berenty 2006: Census of Propithecus verreauxi and Possible Evidence of Population Stress

We provide a survey of a folivorous lemur, Propithecus verreauxi (sifaka), in the Berenty Reserve, southern Madagascar. Higher densities of folivores in small patches occur in either high-quality food areas or in less disturbed refugia. The skewness of sex ratio, which in lemurs is often male-biased, can be critically exacerbated in population stress. We predicted that sifaka would show higher densities in areas where protein-rich food is abundant (prediction 1a) and in refugium areas (prediction 1b). Owing to increased competition by brown and ring-tailed lemurs and decreased food production by tamarind trees, we expected an extremely male-biased sex ratio (prediction 2). In November–December 2006, we counted and sexed 206 adult/subadult sifaka (49 groups) during daily walks in different forest zones (Ankoba secondary forest, to the north, Malaza gallery/scrub areas, and spiny forest, to the south). Sifaka may have decreased in the gallery forest to concentrate in Ankoba (in a sort of out-of-Malaza). The area contains protein-rich food (prediction 1a confirmed). Sifaka are proportionally more concentrated in the spiny area than in the degraded scrub forest (prediction 1b confirmed). The sex ratio is extremely male biased, possibly due to either high sifaka density, in Ankoba, or food availability reduction, in Malaza (prediction 2 confirmed). The sifaka population seems to be under stress: researchers need to resume demographic studies, interrupted in Berenty in the mid-1980s, to preserve in situ a species that is difficult to protect ex situ.

Intergroup relations and home range use in Verreaux's sifaka (Propithecus verreauxi)

Relationships between neighboring groups feature prominently in socioecological theory, but few empirical studies have focused on the effects of neighbors on the behavior of primates. Interactions between neighboring groups are most conspicuous during intergroup encounters, but the likelihood of encounters with neighbors can also affect ranging and activity patterns indirectly, and, as a result, behavioral patterns in areas of exclusive use may differ from those in overlap areas of adjacent home ranges. The aim of this study was to examine intergroup relations and spatial variation in home range use in Verreaux's sifaka (Propithecus verreauxi) during the annual mating season. Based on 230 hr of focal animal data collected from ten members of five adjacent groups, we found that behavioral patterns and patterns of resource utilization were not markedly different between areas of exclusive use and overlap areas of adjacent home ranges. Group cohesion tended to be higher in overlap than in core areas, but the proportion of time spent resting and foraging did not differ between these two areas. However, dominant males exhibited a higher scent-marking rate in overlap areas. Observations during intergroup encounters revealed that chases between males occurred frequently, whereas fights involving physical contact were not observed. We also found that the intergroup dominance hierarchy depended on the relative group size or the number of males in each group, with groups of lower dominance rank exhibiting a tendency to sleep proportionally more often in their core areas. The results of this study suggest that in primate species with a moderate level of intergroup aggression, such as Verreaux's sifaka, the possibility of encountering neighbors in areas of home range overlap does not exert strong influence on their behavior and resource utilization.

Peacemaking on treetops: first evidence of reconciliation from a wild prosimian ( Propithecus verreauxi)

Reconciliation is defined as the first postconflict affinitive contact between former opponents. While reconciliation in anthropoid primates has been widely investigated, few studies have focused on postconflict mechanisms in prosimians, and only in captivity. Unlike anthropoids, Malagasy prosimians show female dominance, lack of sexual dimorphism and seasonal breeding. However, they share features with anthropoids such as cohesive societies, female philopatry and individual recognition. Comparing social prosimians with anthropoids is crucial for understanding the evolution of reconciliation dynamics. Here we present the first study on reconciliation in a wild prosimian. We focused on the Propithecus verreauxi (sifaka) of the Berenty forest (southern Madagascar). We examined postconflict behaviour in the light of theoretical expectations based on potential costs and benefits of the individuals involved. Our results indicate that P. verreauxi can evaluate possible risks and benefits of engaging in postconflict reunions. Victims were most likely to interact affinitively with the aggressor after low-intensity aggression. Moreover, only the conflicts occurring outside the feeding context were reconciled. Such results are consonant with the fact that, in P. verreauxi, social dominance is translated more into feeding priority than into a framework of despotic relationships. In agreement with the valuable relationship hypothesis, P. verreauxi were more likely to reconcile with valuable partners: reconciliation preferentially occurred between subordinates and top-ranking individuals, and between animals sharing good relationships (high levels of affinitive behaviours). Over the short term, reconciliation in P. verreauxi seems to have an important role in reducing the probability of further attacks by the aggressor.

Diurnality and nocturnality in nonhuman primates: comparative chronobiological studies in laboratory and nature

Looking for differences in circadian clock characteristics of diurnal and nocturnal nonhuman primates, this article summarizes results of chronobiological studies carried out in various nocturnal, diurnal, and cathemeral prosimian and anthropoid primate species under controlled laboratory conditions, under seminatural conditions, and in the wild. In almost all circadian parameters investigated, no differences were discernible between the two main chrono-ecotypes, either in circadian period length and the influence upon it of after-effects, of light intensity, and ambient temperature, or in the PRC, re-entrainment behavior, rhythm splitting, and internal desynchronization. Diurnal and nocturnal or cathemeral species differed only in the phase of artificial or natural LDs to which their circadian activity phase was adjusted as well as in the characteristics of masking activity upon the rhythms produced by the direct inhibiting or enhancing effects of light. Pronounced lunar periodicity—observed in the activity rhythm of nocturnal neotropical owl monkeys, genus Aotus, in seminatural and natural environments as well as in wild cathemeral Malagasy lemurs, genus Eulemur—is shown to result from masking effects of moonlight. In captive Eulemur fulvus albifrons, a change from dark-active over cathemeral to light-active behavior, without concurrently changing the circadian phase-setting of activity to D, was produced by direct masking effects of a stepwise reduction of darktime luminosity on an LD 12:12 cycle. Long-term activity recordings carried out in wild diurnal Malagasy sifakas (Propithecus verreauxi) and cathemeral redfronted lemurs (Eulemur fulvus rufus), as well as in wild nocturnal owl monkeys (Aotus a. azarai) of the North Argentinean Chaco, yielded in all species distinct bimodal long- and short-day activity patterns with pronounced peaks during dusk and dawn. Applying Pittendrigh's two-oscillator concept to these results, it is hypothesized that the differences in chrono-ecotype behavior may result from variations in internal coupling and external phase-setting of morning and evening oscillators (m, e) to dawn and dusk, interacting with direct masking effects of light.

Chemical Properties of the Diets of Two Lemur Species in Southwestern Madagascar

Seasonal dietary variations demonstrate the importance of certain plant parts during the year. A parallel analysis of their nutritional constituents provides further information on underlying patterns of consumption of the plant parts and the relative importance of key nutrients. I studied the diets of Lemur catta (ring-tailed lemurs) and Propithecus verreauxi verreauxi (sifakas), for 9 mo over a 13-mo period in the highly seasonal tropical dry forest site of Beza Mahafaly in southwestern Madagascar. I tested dietary plant parts for nutrients—protein, free amino acids, and sugars—and for 2 potential deterrents, phenolics and tannins, using plant extracts prepared in the field. I compared consumption of nutrients and secondary compounds throughout the year and between seasons. Nutrients are balanced throughout the year. The 2 lemur species do not appear nutrient-starved in either season, though actual quantities of nutrients and contributing food parts differ for each species. Lemur catta consumes high levels of sugar throughout the year, whereas Propithecus takes in higher levels of protein. The effects of phenolics and tannins are quantitative, and they appear to deter consumption of plant parts only past a certain threshold. Sifakas consume them in greater quantities than those of ring-tailed lemurs, which appear more sensitive to their effects. Sifakas may have a higher tolerance for secondary plant metabolites, which is consistent with reports for other folivores. The overall stability of nutrients throughout the year indicates no lean period that coincides with the decline in food abundance during the dry season, though actual caloric intake probably decreases.

Food Habits of an Endangered Carnivore,Cryptoprocta Ferox, in the Dry Deciduous Forests of Western Madagascar

We describe the diet of fossas (Cryptoprocta ferox) in a dry deciduous forest of western Madagascar from 376 scats collected between June 1994 and September 1996, from which 554 prey items were identified. More than 90% of these were vertebrates, and more than 50% were lemurs. No other nonprimate mammal includes such a high proportion of primate items in its diet. The principal prey comprised approximately 6 lemur species and 2 or 3 spiny tenrec species, along with snakes and small mammals. Significant differences were apparent in the composition of the scats in wet and dry seasons, with a higher proportion of Tenrec in the former, and fewer lemurs. Within the confines of a diet of vertebrates, fossas appeared to be opportunistic predators. For those prey types for which data were available, a significant relationship was found between the estimated relative number of individuals taken of any one type of prey and its abundance. Fossas were estimated to remove up to 19% of their prey populations per year. This high impact suggests that they were living close to the maximum population density possible on the available prey. Species of a wide range of body masses were included in the diet. Verreaux's sifaka (Propithecus verreauxi), weighing more than one-half of the body mass of the fossa, constituted approximately 11% of the prey biomass.

The lemur syndrome unresolved: extreme male reproductive skew in sifakas (Propithecus verreauxi), a sexually monomorphic primate with female dominance

The primates of Madagascar (Lemuriformes) are unusual among mammals in that polygynous species lack sexual dimorphism, and females dominate males socially in most species. Moreover, lemur groups are relatively small and characterized by even adult sex ratios despite the fact that one male should be able to exclude other males from the group. One hypothesis to explain this combination of behavioral, morphological, and demographic traits (the “lemur syndrome”) postulates that male–male competition is relaxed and, hence, variance in male reproductive success is low. Reproductive skew theory provides a framework for testing this and several related predictions about lemur social evolution. Specifically, low reproductive skew is also predicted if dominant males or adult females make reproductive concessions to subordinates or if the latter group successfully pursues alternative reproductive tactics. However, suitable data on paternity, demography, and behavior for a conclusive test of these predictions have not been available in the past. In this paper, we show that male reproductive success in ten groups of Verreaux’s sifakas (Propithecus verreauxi) was extremely skewed in favor of the dominant male over 9 years. Our genetic analyses also revealed that more than a third of all groups are effectively harem groups because only one male was unrelated to the resident female(s). In groups with two or more non-natal males, the dominant sired 91% of 33 infants. Together, males pursuing one of several alternative reproductive tactics, such as roaming among several groups or immigrating peacefully, sired only 11% of infants. Thus, female sifakas do not control group composition by offering reproductive opportunities to subordinate males as staying incentives, intrasexual selection is not relaxed, and dominant males prevail in a tug-of-war over subordinate males. Because male reproductive skew in sifakas is even more pronounced than in harem-living anthropoids studied to date, intrasexual selection is clearly not relaxed, and the lemur syndrome is more puzzling than ever.

Craniodental Characters in the Taxonomy of Propithecus

The indriid genus Propithecus comprises the sifakas, medium-sized lemurs endemic to the forests of Madagascar. Traditionally, scientists divided the genus into only 2 or 3 species —Propithecus diadema, P. verreauxi, and, since 1988, P. tattersalli— with 4 or 5 subspecies in each of the first 2 taxa, but recent authors have suggested that many more distinct species should be recognized. We draw from quantitative and qualitative studies of craniodental traits to evaluate further the phenetic distinctiveness and taxonomic status of each named form of Propithecus. We recognize 9–10 species in the genus. The 4 or 5 species of the Propithecus diadema group —P. diadema, P. candidus, P. perrieri, P. edwardsi, and perhaps P. holomelas, if distinct— share several derived features, including large average body size and a mandible specialized for rotational chewing, and clearly comprise a closely related complex. The 5 species of the Propithecus verreauxi group —P. verreauxi, P. coquereli, P. deckenii, P. coronatus, P. tattersalli— are each highly distinctive morphologically and likely do not comprise a monophyletic group. In particular, we point out the highly distinctive cranial features of Propithecus coronatus, which researchers have traditionally largely overlooked.

Kinematics of bipedalism in Propithecus verreauxi

AbstractBipedalism is rare in primates and has evolved in two distantly related groups: hominoids and indrids. Although copious data are available on the mechanics of bipedal locomotion in hominoids and vertical clinging and leaping (VCL) in indrids, no research has addressed the unique mode of bipedal locomotion exhibited by select indrid primates. Propithecus verreauxi is a highly specialized indrid vertical clinger and leaper that uses a peculiar form of bipedalism on the ground. The objectives of this study were to describe the bipedal gait of Propithecus, to assess the influence of VCL specializations on the kinematic patterns and propulsion mechanisms used by Propithecus during bipedalism, and to compare Propithecus bipedalism with the bipedal gaits of other primates capable of using bipedalism. Video was collected of five adult P. verreauxi moving bipedally in a seminatural setting at the Duke University Primate Center. Duty factor, footfall patterns, joint angles and center of mass movement were quantified in the sagittal plane for 73 steps. Propithecus uses a bipedal gallop, a gait unique to Propithecus. The kinematic similarities (e.g. large hip and knee angular excursions and preparatory countermovements) between bipedal galloping and VCL lead us to suggest that Propithecus takes advantage of specializations for VCL to conserve energy during bipedal galloping. Propithecus also walks bipedally at slower speeds. When Propithecus walks, it utilizes a relatively compliant gait similar to that of other primate facultative bipeds (Pan, Hylobates). During bipedal walking, energy conservation may be sacrificed for increased balance and reduced joint loads.

Influence of reproductive season and rank on fecal glucocorticoid levels in free-ranging male Verreaux's sifakas ( Propithecus verreauxi)

Studies in anthropoid primates and other mammals suggest that reproductive season, rank, reproductive skew, aggression received, and social support are the major factors influencing glucocorticoid output. In which way these are also affecting adrenal function in lemurid primates has been studied rarely. Here, we examine the influence of reproductive season and rank on glucocorticoid output in male sifakas ( Propithecus verreauxi), a species characterized by high breeding seasonality, a hierarchy among males and extreme reproductive skew towards dominant males. We established a fecal assay for non-invasively monitoring adrenal activity and collected 315 fecal samples during the reproductive and birth season from 10 male sifakas living in 5 groups in Western Madagascar. We found a significant effect of season on glucocorticoid output, with males exhibiting higher fecal glucocorticoid levels during the reproductive compared to the birth season in conjunction with an increase in overall aggression rates during the former period. Moreover, our data indicate a significant effect of rank on adrenocortical activity with dominant males exhibiting higher glucocorticoid levels than subordinate males in the reproductive season. However, dominant males did not differ significantly in rates of initiated or received aggression and rates of affiliative behavior from subordinates but showed significantly lower rates of submission. Given their highly formalized dominance relationships, we conclude that higher glucocorticoid output in dominant males during the 4-month reproductive season is likely related to higher energetic demands necessary to cope with the challenges of male reproduction rather than to physical demands of increased fighting frequency to maintain dominance status. High rank in sifakas may thus carry high costs, which, however, may be outweighed by monopolization of almost all paternities. In sum, our data generally support the findings on the relationship between environmental and social factors and glucocorticoid output found in non-lemurid primates.

Influence of Dry Season and Food Quality and Quantity on Behavior and Feeding Strategy of Propithecus verreauxi in Kirindy, Madagascar

According to optimal foraging theory, herbivores can base food choice mainly on the quality or the quantity of food, or both. Among herbivorous primates, folivorous lemurs living in the highly seasonal environment of Madagascar have to cope with the shortage of high-quality food during the dry season, at least in deciduous forests. We studied (Verreaux's sifaka) in Kirindy, western Madagascar, to understand the influence of dry season and food quality and quantity on behavioral patterns and feeding strategy (qualitative vs. quantitative dietary choice) of a folivorous lemur in a deciduous forest. We followed 7 groups (4 groups/period; 3 individuals/group/month) during 4 periods of the year (wet season: February–March; early/middle/late dry season: May–June; July–September; October–November). We collected samples of plants eaten and examined behavioral and feeding patterns, considering food quality (macronutrients, proteins/fibers ratio, and tannins) and abundance. We found 1) a significant reduction of home range, core area, and daily path length from the wet to the dry season, possibly related to dietary change and 2) a daily period of inactivity in the dry season for energy conservation. Regarding the feeding strategy, Kirindy sifakas showed 1) high variation and selection in choosing food items and 2) a dietary choice based mainly on quality: Kirindy sifakas fed on plant species/families independently from their abundance and tannins represented a feeding deterrent during the dry season. Overall, behavioral and dietary adaptations allow Kirindy sifakas to overcome the shortage of high-quality food in the lean period.

Characterization of 13 microsatellite marker loci in Verreaux's sifaka (Propithecus verreauxi)

AbstractThe Verreaux's sifaka (Propithecus verreauxi) is one of the species of Propithecus, living in the dry forest of southwest Madagascar. This species is endangered due to the loss and fragmentation of its natural habitat, a consequence of deforestation. Thirteen novel nuclear microsatellite loci were isolated and characterized in three populations of Verreaux's sifaka. The marker suite proved informative with an average of 8.9 alleles per locus and observed heterozygosity across the three populations of 0.675.

Evidence of early butchery of giant lemurs in Madagascar

We report here definitive evidence of butchery, most probably associated with hunting, of giant extinct lemurs by early human settlers in Madagascar. Specimens of Palaeopropithecus ingens and Pachylemur insignis from two sites in southwestern Madagascar, Taolambiby and Tsirave, show classic signs of butchering. We compared these to the bones (also from Taolambiby) of butchered Propithecus verreauxi, a lemur still living in the region. The characteristics of the tool-induced extinct-lemur bone alterations (sharp cuts and chop marks near joints, oblique cuts along the shafts, spiral fractures, and percussion striae) suggest skinning, disarticulation, and filleting. Conclusive evidence of megafaunal modification by humans in Madagascar was limited previously to a few hippo and elephant bird bones and one extinct aye-aye tooth. New evidence comes not from archaeological sites, but from specimens collected in the early 1900s, without stratigraphic records, at “subfossil” sites (i.e., sites renowned for their late Pleistocene or Holocene fossils, often lacking human artifacts). Whereas these are hardly the most ideal samples for analysis of this kind, careful scrutiny of the characteristics of the cut marks has allowed us to document butchery beyond any reasonable doubt. One bone with definitive cut marks has been dated to the very earliest part of the human period in Madagascar. Continued, careful research on the bones in subfossil collections is warranted.

Reach and posture hand preferences during arboreal feeding in sifakas (Propithecus sp.): A test of the postural origins theory of behavioral lateralization.

Sixteen sifakas (11 Propithecus verreauxi coquereli, 2 Propithecus verreauxi verreauxi form majori, and 3 Propithecus tattersalli) were videotaped as they fed on leaves in an arboreal context. The hand used to feed and the hand used to maintain postural stability was coded. For each subject, the lateral bias of the hand used to feed was opposite the hand used in postural support. Seven sifakas displayed no bias for feeding or posture-related hand use, 7 sifakas displayed significant feeding-related reach preferences for pulling branches to the mouth (5 left- and 2 right-hand preferences), and 9 sifakas exhibited significant hand preferences for postural support (2 left-, 7 right-hand preferent). Although these data do not strongly support the postural origins theory of behavioral lateralization, the modal preference pattern for sifakas that displayed significant hand preferences for posture and feeding involve a left bias for feeding and a right bias for postural support. s reserved).

Characterizing gastrointestinal transit time in four lemur species using barium‐impregnated polyethylene spheres (BIPS)

Differences in dietary profiles and gastrointestinal (GI) morphologies observed across lemur species suggest that there may be variation in patterns of digesta flow through the GI tract related to the method of digesta processing. Using radio-opaque barium-impregnated polyethylene spheres (BIPS), we characterized such patterns in four lemur species: Varecia variegata (VV), Eulemur fulvus (EF), Propithecus verreauxi (PV), and Hapalemur griseus (HG) (n = 2 per species). After an initial radiograph was taken under light sedation, the animals were fed the BIPS together with a small meal. A combination of 30 small (1.5 mm) and 10 large (5 mm) BIPS was administered. Radiographs were then taken on a species-dependent basis up to 48 hr post-dosage. For small BIPS, the gastric transit time (GTT; time of first exit of BIPS from stomach) was 0.25-2 hr for VV, EF, and HG, and approximately 10 hr for PV. The oro-rectal transit time (ORTT; time of first appearance in the rectum) was < 2 hr for VV and EF, and 24.0 hr for PV and HG. The intestinal transit time (ITT, measured as ORTT - GTT) was < 1.5 hr for VV and EF, and approximately 14 hr and 22 hr for PV and HG, respectively. These data suggest that the GTT of digesta as measured with BIPS was rapid for VV, EF, and HG. For VV and EF, the ORTT and ITT were also rapid, while for HG they were much slower. PV was characterized by delayed GTT, and a more rapid ITT compared to HG. Thus, patterns of flow for PV and HG, despite similar ORTT, differed in that HG emptied BIPS more rapidly and ITT was slower. The flow of BIPS did not differ for VV and EF. These data reveal new information in addition to the total tract transit time, and complement existing knowledge regarding anatomy and diet.