Diets of Two Lemur Species in Different Microhabitats in Beza Mahafaly Special Reserve,Madagascar

Studies of primate diets usually focus on differences that distinguish species or populations. However, variation in diet can occur at a more local level of groups within a population, especially in a non-homogeneous habitat. I compared dietary variation in food composition and toughness across groups of 2 lemur species in Beza Mahafaly special reserve, Madagascar. Beza Mahafaly contains an 80-ha reserve (Parcel 1) that, while small, hosts a dense population of Lemur catta (ring-tailed lemurs) and Propithecus verreauxi verreauxi (sifakas). Microhabitats in the eastern vs. western sides of the parcel are structurally and floristically distinct. Sifakas in this parcel have small, discrete home ranges and are morphological folivores. For these reasons, I expected that the 6 groups studied would eat a different menu of food plants but with similar toughness values. Ring-tailed lemurs have comparatively large, overlapping home ranges, and I expected that the 5 study groups would eat similar foods. Despite living in different microhabitats across the parcel, sifakas exhibit high dietary uniformity both in dietary plant species composition and the toughness of the foods. Food selection in sifakas operates on two distinct levels. Sifaka groups share many key food species that appear independent of local abundances, but the ranking of the foods within each group appears related to availability. Ring-tailed lemur groups are more heterogeneous in the composition of their diets relative to sifakas, though the time spent feeding on individual foods reveals a marked preference for the fruits of Tamarindus indica by all groups. Food toughness is consistent across the parcel with the exception of the most western group. Ring-tailed lemurs are highly specific feeders, but indiscriminate nibblers. Sifakas are targeted, balanced feeders. There does not appear to be a consistent microhabitat effect operating across species. Differences within sifaka and ring-tailed lemur populations in food composition and toughness, however, correspond to an east-west microhabitat gradient. Measures of dietary flexibility must take into account not only the plant species consumed and the different parts eaten but also their associated food properties and proportion of time spent feeding on them.     

Food choice from endemic North Carolina tree species by captive prosimians (Lemur fulvus)

Seven captive-born lemurs (Lemur fulvus) at the Duke University Primate Center were presented with leaves of different maturity from five species of North Carolina trees. The animals demonstrated three distinct behaviors toward the novel plant material. They sniffed, tasted, and/or ingested it. New leaves were sniffed, but little tasting and ingestion was observed. Intermediate pine needles were sniffed and ingested but little tasted. Mature leaves were sniffed equally, but the mature leaves of tulip trees and honeysuckle were tasted significantly more than pine, sweetgum, and red maple. Pine, sweetgum, and red maple were ingested significantly more than tulip trees and honeysuckle. Male lemurs ate significantly more mature pine needles and new sweetgum leaves than did the females. Chemical analysis of these plant materials indicated that the new and mature leaves of tulip tree and honeysuckle contained alkaloids. Captive-born lemurs apparently use their sense of smell and taste in choosing what to eat and seem just as capable as free-ranging animals in finding food when faced with the chemical defenses that protect trees from insect predation.     

Habitat Characteristics and Lemur Species Richness in Madagascar1

In the extant lemur communities of Madagascar the number of lemur species increases with increasing number of tree species. In forests with few tree species lemurs use patches with higher number of tree species than average. However, in forest plots with high number of tree species, lemurs stay in places with lower number of tree species than average. At low tree species diversity a minimum number of different tree species seems to be required within the animals' home range to assure year-round food availability. At high tree species diversity tree species essential for survival might be diluted by resources which are of no use for lemurs, thus increasing energetic expenses for traveling between suitable patches. According to the present analyses, structural diversity is of subordinate importance to the number of tree species as a correlate of lemur species richness. Within limits of disturbance intensity and on a small geographic scale, disturbances increase forest productivity. Lemurs reach higher species numbers and population densities in slightly disturbed areas compared with undisturbed sites. This peaked curve of the number of lemur species over disturbance, however, may not only be a consequence of “resource dilution” in undisturbed sites and higher food abundance in slightly disturbed areas, but also a consequence of selective extinction of lemur species which were unable to cope with the disturbance regime exaggerated by human interference over the last few hundred or thousand years.     

Feeding Ecology of Sympatric Mouse Lemur Species in Northwestern Madagascar

We performed a comparative pilot study on vertical space use and feeding ecology of 2 closely related sympatric mouse lemur species in northwestern Madagascar. We recorded feeding behavior and the use of vertical forest strata for 6 gray and 4 golden-brown mouse lemur females that we followed for 120 h. We carried out feeding experiments and analyzed fecal samples for food remains. Both species showed no significant difference in use of forest strata. They had a similar overall diet, with insect secretions and gum as main plant food resources; arthropod remains occurred in about half of all fecal samples. Both lemurs used in common >50% of the plant species that each consumed, which accounts for ≥70% of all individual plants used. However, both species used >40% of their feeding plant species exclusively and seemed to differ in their degree of specialization on certain plant species.     

A 24-Year Record of Female Reproductive Dynamics in Two Sympatric Mouse Lemur Species in Northwestern Madagascar

International Journal of Primatology - Seasonal reproduction is widespread among primates but the degree of reproductive synchrony and plasticity can vary, even between closely related species....     

Estimating Encounter Rates and Densities of Three Lemur Species in Northeastern Madagascar

Primate populations, including Madagascar’s lemurs, are threatened worldwide and conservationists need accurate population estimates to develop targeted conservation plans. We sought to fill knowledge gaps for three lemur taxa —white-fronted brown lemur (Eulemur albifrons); eastern woolly lemur (Avahi laniger); and Allocebus/Microcebus, a category combining observations of hairy-eared dwarf lemurs (Allocebus trichotis) and mouse lemurs (Microcebus spp.)— in northeastern Madagascar by estimating their density, examining how their encounter rates and densities vary across three different forest types, and monitoring trends in encounter rates and densities at resurveyed sites, using data from surveys at six forest sites over a 4-year period (2010–2013). Landscape density for white-fronted brown lemur, eastern woolly lemur, and Allocebus/Microcebus was 21.5 (SE 3.67), 57.7 (SE 9.17), and 39.1 (SE 9.55) individuals/km², respectively. There was no difference in density estimates at intact and intermediately degraded forest sites; however, we encountered white-fronted brown lemurs more often in intact forest (1.64 ± SE 0.40 individuals/km) than in intermediately degraded and degraded forest (0.15 ± SE 0.06 and 0.16 ± SE 0.06 individuals/km). In addition, we encountered white-fronted brown lemurs at lower rates in 2013 (0.15 ± SE 0.06 individuals/km) compared to 2010 (0.82 ± SE 0.12 individuals/km) at a resurveyed site. Our findings emphasize that primate researchers must account for variation in how lemur encounter rates and densities differ between intact and degraded forests, and although we observed a decline in white-fronted brown lemur encounter rate at our resurveyed site, we caution that changes in lemur encounter rates may simply reflect lower detection rates rather than lower density. Future research should focus on using conventional distance sampling techniques, which are infrequently used in primate studies, to provide more robust density estimates as a way to accurately assess trends and the effects of anthropogenic pressures on lemur populations.     

Habitat Utilization of Three Sympatric Cheirogaleid Lemur Species in a Littoral Rain Forest of Southeastern Madagascar

I compared the habitat utilization in 3 sympatric species of Cheirogaleidae (Microcebus murinus [81 g], Cheirogaleus medius [183 g] and Cheirogaleus major [362 g]) in a littoral rain forest in southeastern Madagascar during 3 rainy seasons. Females of promiscuous Microcebus murinus had small home ranges and the males had large overlapping home ranges. Home ranges of family groups of monogamous Cheirogaleus medius and C. major overlapped extensively. Home ranges of all 3 species overlapped completely in the study area but home range sizes differed among species and correlate positively with body masses. Male Microcebus murinus slept in open vegetation (79%) and alone (71%), whereas female M. murinus and family group members of Cheirogaleus spp. preferred communal sleeping in tree holes. There are significant interspecific differences in the choice of sleeping sites: smaller lemurs chose smaller trees and used more sleeping sites than larger lemurs did. Species also differed significantly in the vertical dimension of forest utilization: Cheirogaleus major used the upper part of the trees, C. medius used the middle parts, and Microcebus murinus used the understory during nocturnal activities. The 3 species differed mainly in vertical habitat utilization and showed vertical stratification.     

Diet, Nutritional Ecology, and Birth Season of Eulemur macaco in an Anthropogenic Forest in Madagascar

We investigated the feeding ecology of Eulemur macaco macaco in an old coastal secondary forest of northwestern Madagascar. We analyzed whether the local combination of introduced and native plant species could provide viable anthropic conditions for sustaining the black lemurs. Fruits (79 spp.) dominated the annual diet (>104 species from 50 families via observations ad libitum and use of a feeding frequency methods). Records from the early dry (mating) and late dry (birth) seasons show that a few major fruit species are staples in conjunction with a variety of other plant items in much lower proportions. We further estimated daily food intake and analyzed nutrient/antinutrient content in the diet during the birth season to evaluate the possibility that black lemurs undergo nutritional stress. They exhibited a high-energy input/low energy output foraging strategy then and had limited use of alternative resources such as leaves throughout the study period. We conclude that the potential for feeding flexibility is low because specialization on fruit results in protein requirements being achieved probably by a narrow margin. We hypothesize that patchy distribution of preferred cash-crop plants and indigenous species currently has a major limiting effect on population size through feeding competition.     

栎黄枯叶蛾生长发育与几种寄主内含物的关系

【目的】探讨饲喂不同寄主植物对栎黄枯叶蛾Trabala vishnou gigantina Yang生长发育及繁殖的影响,分析寄主植物内含物与其生长发育及繁殖的关系,为研究不同寄主植物对栎黄枯叶蛾种群动态的影响提供理论依据。【方法】在室内条件下,利用沙棘Hippophae rhamnoides、山杏Armeniaca sibirica、山杨Populus davidiana、旱柳Salix matsudana饲养该虫,观察幼虫发育历期、蛹历期、蛹重及产卵量等指标,同时测定寄主叶片中可溶性糖、可溶性蛋白质、水分、单宁及黄酮的含量,并进行线性回归分析。【结果】不同寄主植物饲喂栎黄枯叶蛾幼虫的发育历期、蛹历期、蛹重及产卵量均存在显著差异。取食沙棘的幼虫历期最短,仅为(74.50?1.76)d,而取食旱柳历期最长,达(106.00?1.51)d。取食山杨的蛹历期最长,为(44.00?1.41)d,最短的是取食旱柳的,仅为(32.70?3.13)d。饲喂沙棘和山杏的雌、雄蛹重和成虫产卵量显著高于饲喂山杨和旱柳的。对不同寄主的营养物质、次生物质和水分含量测定结果表明:不同寄主间营养物质、次生物质与水分的含量差异显著。回归相关分析表明:幼虫发育历期与寄主植物可溶性蛋白质含量呈显著负相关;雌蛹重和产卵量与寄主植物可溶性糖含量呈显著负相关;雌、雄蛹重与寄主植物含水量呈显著正相关;蛹历期与单宁含量呈显著正相关,但黄酮含量的高低与栎黄枯叶蛾生长发育等指标无相关性。【结论】沙棘是栎黄枯叶蛾生长发育及繁殖的最适寄主植物;并且寄主植物中可溶性糖、单宁的含量低、含水量和可溶性蛋白质含量高有利于栎黄枯叶蛾的生长发育和繁殖。     

The Use of an Invasive Species Habitat by a Small Folivorous Primate: Implications for Lemur Conservation in Madagascar

The lemurs of Madagascar are among the most threatened mammalian taxa in the world, with habitat loss due to shifting cultivation and timber harvest heavily contributing to their precarious state. Deforestation often leads to fragmentation, resulting in mixed-habitat matrices throughout a landscape where disturbed areas are prone to invasion by exotic plants. Our study site, the Mandena littoral forest (southeast Madagascar), is a matrix of littoral forest, littoral swamp, and Melaleuca swamp habitats. Here, Melaleuca quinquenervia has invaded the wetland ecosystem, creating a mono-dominant habitat that currently provides the only potential habitat corridor between forest fragments. We sought to understand the role of this invasive Melaleuca swamp on the behavioral ecology of a threatened, small-bodied folivore, the southern bamboo lemur (Hapalemur meridionalis). We collected botanical and behavioral data on four groups of H. meridionalis between January and December 2013. Our results confirm Melaleuca swamp as an important part of their home range: while lemurs seasonally limited activities to certain habitats, all groups were capable of utilizing this invasive habitat for feeding and resting. Furthermore, the fact that Hapalemur use an invasive plant species as a dispersal corridor increases our knowledge of their ecological flexibility, and may be useful in the conservation management of remaining threatened populations.     

Social Relationships Among Ring-Tailed Lemurs (Lemur catta) in Two Free-Ranging Troops at Berenty Reserve, Madagascar

We observed two free-ranging troops of ring-tailed lemurs at the Berenty Reserve, Madagascar. Kinship affinities in these troops are known only for mothers and their offspring 4 years of age. We attempted to quantify social relationships. Almost all agonistic interactions were dyadic, and triadic agonistic interactions, such as alliances, were very rare. Dominance hierarchies in both sexes in the two troops were not linear. As in cercopithecine monkeys, mothers were dominant over their adult daughters. However, the daughters were not ranked immediately below their mothers. Close proximity and social grooming occurred more frequently between closely related females, such as mother–daughter and sister–sister dyads, than between unrelated females. Frequent-proximity relations also occurred between adult males that had emigrated from another troop and entered the present troop together, even though they did not rank closely to one another. Subordinates were likely to groom and to greet dominants more frequently than vice versa. During group encounters, particular females were involved in agonistic interactions with animals of other troops, regardless of dominance rank. Adult males, regardless of their dominance rank, but not adult females, constantly tried to drive solitary males away.     

Biosynthesis of plant phenolic compounds in elevated atmospheric CO2

Experiments were carried out to determine the effects of elevated atmospheric carbon dioxide (CO₂) on phenolic biosynthesis in four plant species growing over three generations for nine months in a model plant community. Results were compared to those obtained when the same species were grown individually in pots in the same soils and controlled environment. In the model herbaceous plant community, only two of the four species showed any increase in biomass under elevated CO₂, but this occurred only in the first generation for Spergula arvensis and in the second generation for Poa annua. Thus, the effects of CO₂ on plant biomass and carbon and nitrogen content were species‐ and generation‐specific. The activity of the principle phenolic biosynthetic enzyme, phenylalanine ammonia lyase (PAL), increased under elevated CO₂ in Senecio vulgaris only in Generation 1, but increased in three of the four plant species in Generation 2. There were no changes in the total phenolic content of the plants, except for P. annua in Generation 1. Lignin content decreased under elevated CO₂ in Cardamine hirsuta in Generation 1, but increased in Generation 2, whilst the lignin content of P. annua showed no change, decreased, then increased in response to elevated CO₂ over the three generations. When the species were grown alone in pots, elevated CO₂ increased PAL activity in plants grown in soil taken from the Ecotron community after nine months of plant growth, but not in plants grown in the soil used at the start of the experiment (‘initial' soil). In P. annua, phenolic biosynthesis decreased under elevated CO₂ in initial soil, and in both P. annua and S. vulgaris there was a significant interaction between effects of soil type and CO₂ level on PAL activity. In this study, plant chemical composition altered more in response to environmental factors such as soil type than in response to carbon supply. Results were species‐specific and changed markedly between generations.     

MHC Variability of a Small Lemur in the Littoral Forest Fragments of Southeastern Madagascar

Habitat fragmentation inhibits gene flow between populations often resulting in a loss of genetic diversity with possible negative effects on fitness parameters. In vertebrates, growing evidence suggests that such genetic diversity is particularly important at the level of the major histocompatibility complex (MHC) because its gene products play an important role in immune functions. Diversity in the MHC is assumed to improve population viability. Here, we investigated the impact of forest fragmentation on the genetic variability of one of the functionally important parts of the MHC, DRB exon 2, of the endemic mouse lemur Microcebus murinus by comparing populations inhabiting two littoral forest fragments of different size in southeastern Madagascar. Twelve different alleles of DRB exon 2 were found in 145 individuals of M. murinus with high levels of sequence divergence between alleles. In both subpopulations, levels of genetic diversity were high, and the genetic analyses revealed only limited effects of fragmentation. Significantly more non-synonymous than synonymous substitutions were found in the functionally important antigen recognition and binding sites indicating selection processes maintaining MHC polymorphism. This is the first study on MHC variation in a free-ranging Malagasy lemur population.     

The relationship between diet and the size of the midgut caeca in grasshoppers (Insecta: Orthoptera: Acridoidea)

A survey of the size and form of the midgut caeca in relation to diet has been carried out on 173 species from 21 families and subfamilies of Acridoidea (grasshoppers). Although differences exist in the size of the anterior caecal arms relative to body length, these differences are not related to the type of food eaten. Assuming that the anterior arms have a key role in digestive and absorptive processes, this suggests that different foods make similar demands on these processes. The posterior caecal arms are smaller in graminivorous species than in species eating other types of plants as a whole or part of their diet. This is true across all the taxa, including those families and subfamilies that are predominantly forb-feeding. It is suggested that the posterior caecal arms have a special role in the detoxification of plant secondary compounds and that the requirement for this is reduced in graminivorous species because of the lower levels of toxic secondary compounds in grasses. A specialized pocket region is present in the posterior caecal arms of some forb-feeding species. Its occurrence across the taxa is spasmodic. It may be concerned with the removal of phenolic compounds.     

Effects of Transect Selection and Seasonality on Lemur Density Estimates in Southeastern Madagascar

I investigated how transect type (trails vs. cut transects) and seasonality influenced density estimates for 5 lemur taxa (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus) in the Vohibola III Classified Forest in SE Madagascar. I surveyed tree height and diameter and lemur populations from June 1 to December 28, 2004 along 2 1250-m trails local people used and 2 1250-m transects cut parallel to the trails in primary rain forest. Despite dendrometric variations within and between trails and transects, only density estimates of Hapalemur griseus griseus differed significantly by transect type. The spatial variation may be a result of removal by local people of giant bamboo, which is the main food for Hapalemur griseus griseus, along trails. Conversely, seasonality influenced density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus. The temporal variations may be related to seasonal torpor for Cheirogaleus major and increased detection probabilities during periods of fruit exploitation for Eulemur rubriventer, Microcebus rufus, and Hapalemur griseus griseus. Transect type and seasonality did not affect density estimates for Avahi laniger, which may be related to the highly folivorous and low-energy diet of the nocturnal lemur. Researchers surveying lemurs along line transects should be aware that transect selection may influence density estimates for Hapalemur griseus griseus and that seasonality may influence density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus.     

Methyl jasmonate-induced nicotine production in Nicotiana attenuata: inducing defenses in the field without wounding

The functional significance of herbivore-induced plant traits known to directly or indirectly influence herbivore performance remains largely untested under field conditions due to the difficulty of uncoupling the response to herbivory from the act of herbivory. The signals that activate many of the induced responses in plants are endogenously produced in response to wounding, unlike many of the predator-induced responses found in aquatic invertebrates (which are activated by exogenous cues derived from predators). Jasmonates, endogenously-produced damage signals, activate diverse wound-induced responses in plants including induced nicotine production in Nicotiana sylvestris. The results presented here are from two experiments which illustrate the use of jasmonates to uncouple induced nicotine production in Nicotiana attenuata (Torrey ex. Watson) from wounding. The exogenous addition of methyl jasmonate (MJ) in small quantities (11 g for a 1.4 g dry mass plant) to roots of hydroponically-grown plants induces de novo nicotine synthesis and increases whole-plant nicotine concentrations just as wounding does. The MJ-induced changes were proportional to the quantity of MJ given. Moreover, the effects of MJ were additive to the effects of damage. Applications of MJ to shoots were less effective. Root treatments also worked with plants growing in a field plot. The application of MJ represents a promising tool for examining the functional significance of induced nicotine responses in plants growing in their native environments.     

Demography and social organization of free-ranging Lemur catta in the Beza Mahafaly Reserve,Madagascar

In 1987, a long-term study of the demography of Lemur catta was begun in southern Madagascar. Eighty-five ringtailed lemurs were captured, marked, and released. Adult age classes were estimated using patterns of dental attrition. Including young, 155 individuals from nine groups were identified and monitored over 18 months. The study population of the reserve remained stable, with a growth rate of 0.98. Group sizes ranged from nine to 22 individuals (mean 14). Home ranges were larger (32 ha) and population densities lower (135/km²) than those for previously studied populations, and there was a relationship between habitat quality (e.g., no. of large trees) and these factors. At the beginning of the study, there were more adult males than females, but the sex ratio reached 1.00 by the last census. Females first gave birth at 3 years of age, and 80% or more of the females gave birth in 2 consecutive years. Fifty-two percent of the infants died in the first year and, given preliminary findings, only 40% of those born reach adulthood. Age-specific fertility patterns were similar to those reported for anthropoid primates. Forty-seven percent of the adult males migrated or were missing within a year. This included 78% of the 3–4 year olds and 38% of older age classes. No females were observed to migrate. One group split during the study. Demographic patterns are discussed and related to patterns in other populations of ringtailed lemurs as well as in anthropoids.     

Preliminary observations on the role of Acacia gum chemistry in Acacia utilization by Galago senegalensis in Kenya

A census and preliminary observations on the ranging and feeding behavior of Galago senegalensis were made in a savannah-woodland site in Kenya. Population densities in three slightly different habitats were all approximately 1.5 animals/ha. Ranging patterns suggested the existence of territorial behavior. The diet of the galagos was exclusively insects and gums of two species of Acacia. The gum of A. drepanolobium appeared to be preferred to that of A. xanthophloea. The chemistry of the gums suggests that this preference is not due simply to the total level of phenolics or to avoidance of condensed tannins but may relate to the presence of compounds (e. g., flavonoids) having nutritional or hormonelike action.     

Diet of the Brown Mouse Lemur (Microcebus rufus) in Ranomafana National Park, Madagascar

The conventional notion is that small-bodied primates should be highly insectivorous in order to obtain protein and other nutrients from a food source that is more easily digestible than plant matter. I studied feeding behavior of Microcebus rufus for 16 months in the east coast rainforest of Ranomafana National Park. I determined the diet primarily through analysis of 334 fecal samples from live-trapped individuals. They consumed a mixed diet basically of fruit and insects year-round. I identified 24 fruits, while 40–52 remain unidentified. Bakerella, a high-lipid epiphytic semiparasitic plant, was in 58% of fecal samples that contained fruit seeds, and was consumed year-round irrespective of general resource availability. It served both as a staple and keystone resource. Fruit was less frequently totally absent from fecal samples of individual mouse lemurs than insect matter was. For Microcebus rufus, fruit may be a primary source of energy, not just complementary to insects. Fruit consumption increased in quantity and diversity during the latter part of the rainy season and the very early part of the dry season, when fruit production was relatively high. This pattern in fruit feeding is similar to that for mouse lemurs in the west coast dry forests and is related to specific nutritional needs dictated by the highly seasonal character of the life cycle. Coleoptera were present in 67% of samples examined and were consumed year-round by the subjects, but insect consumption did not increase during the rainy season when insect abundance was highest.