Jean-Baptiste Ramanamanjato

Extinction vulnerability of tropical montane endemism from warming and upslope displacement: a preliminary appraisal for the highest massif in Madagascar

One of the predicted biological responses to climate warming is the upslope displacement of species distributions. In the tropics, because montane assemblages frequently include local endemics that are distributed close to summits, these species may be especially vulnerable to experiencing complete habitat loss from warming. However, there is currently a dearth of information available for tropical regions. Here, we present a preliminary appraisal of this extinction threat using the herpetological assemblage of the Tsaratanana Massif in northern Madagascar (the islands highest massif), which is rich with montane endemism. We present meteorological evidence (individual and combined regional weather station data and reanalysis forecast data) for recent warming in Madagascar, and show that this trend is consistent with recent climate model simulations. Using standard moist adiabatic lapse rates, these observed meteorological warming trends in northern Madagascar predict upslope species displacement of 17–74 m per decade between 1993 and 2003. Over this same period, we also report preliminary data supporting a trend for upslope distribution movements, based on two surveys we completed at Tsaratanana. For 30 species, representing five families of reptiles and amphibians, we found overall mean shifts in elevational midpoint of 19–51 m upslope (mean lower elevation limit 29–114 m; mean upper elevation limit −8 to 53 m). We also found upslope trends in mean and median elevational observations in seven and six of nine species analysed. Phenological differences between these surveys do not appear to be substantial, but these upslope shifts are consistent with the predictions based on meteorological warming. An elevational range displacement analysis projects complete habitat loss for three species below the 2 °C ‘dangerous’ warming threshold. One of these species is not contracting its distribution, but the other two were not resampled in 2003. A preliminary review of the other massifs in Madagascar indicates potential similar vulnerability to habitat loss and upslope extinction. Consequently, we urgently recommend additional elevational surveys for these and other tropical montane assemblages, which should also include, when possible, the monitoring of local meteorological conditions and habitat change.

Edge effects and extinction proneness in a herpetofauna from Madagascar

Edge effects are thought to play a key role in fragmented habitats. It is often assumed that edge-avoiding species are more prone to local extinction than non-edge-avoiding species, but there are few data to support this assumption. Also, few data are available on the effects of edges on some groups, and there is little understanding of seasonal changes in edge effect intensity. To better understand the role that edge effects play in fragmented tropical forests, we assessed the distribution of reptiles and amphibians in six littoral rainforest fragments in southeastern Madagascar in 1999 (dry season) and 2000 (wet season). Using randomization tests, we found all three types of edge responses: edge-avoiders, interior-avoiders, and omnipresent species. However, edge responses varied considerably among seasons. For example, some species (e.g., Mantidactylus bicalcaratus, Phelsuma quadriocellata) were strong edge-avoiders in the dry season, but showed no preference for edge or interior habitats during the wet season. Also, edge-avoiding species tended to be more extinction-prone than non-edge-avoiding species. Abiotic data documented significantly higher temperatures and dew point temperatures near edges. Wind speed also tended to be higher and relative humidity tended to be lower near edges. Our results indicate that many amphibians and reptiles respond to altered microclimates near edges and these responses have a strong seasonal component. For many species, edge sensitivity does seem to be correlated with extinction vulnerability and therefore should be of primary consideration in plans to conserve biodiversity in fragmented tropical forests.

Effects of forest fragmentation, introduced Rattus rattus and the role of exotic tree plantations and secondary vegetation for the conservation of an endemic rodent and a small lemur in littoral forests of southeastern Madagascar

We sought to assess the effects of forest fragmentation, introduced Rattus rattus , exotic tree plantations and secondary vegetation on the endemic rodent Eliurus webbi (Nesomyinae) and the lemur Microcebus murinus in the littoral forests of southern Madagascar. For E. webbi the number of individuals caught, the body mass of males and the percentage of females in the population were positively correlated with the size of the forest fragments. Capture rates and population characteristics of the other two species were uncorrelated with fragment size. None of the endemic species was caught outside the native forest while R. rattus inhabited all vegetation formations except for a newly planted corridor of tree saplings. Capture rates of both endemic species were uncorrelated with the number of R. rattus caught at the same site and thus did not indicate replacement of native species by R. rattus . The study demonstrated negative effects of fragmentation on capture rates of E. webbi and changes in their population characteristics. Exotic tree plantations or secondary vegetation seem to represent unsuitable or marginal habitats for the endemic species.

Sensory Basis of Food Detection in Wild Microcebus murinus

Very little is known about how nocturnal primates find their food. Here we studied the sensory basis of food perception in wild-caught gray mouse lemurs (Microcebus murinus) in Madagascar. Mouse lemurs feed primarily on fruit and arthropods. We established a set of behavioral experiments to assess food detection in wild-born, field-experienced mouse lemurs in short-term captivity. Specifically, we investigated whether they use visual, auditory, and motion cues to find and to localize prey arthropods and further whether olfactory cues are sufficient for finding fruit. Visual cues from motionless arthropod dummies were not sufficient to allow reliable detection of prey in choice experiments, nor did they trigger prey capture behavior when presented on the feeding platform. In contrast, visual motion cues from moving prey dummies attracted their attention. Behavioral observations and experiments with live and recorded insect rustling sounds indicated that the lemurs make use of prey-generated acoustic cues for foraging. Both visual motion cues and acoustic prey stimuli on their own were sufficient to trigger approach and capture behavior in the mouse lemurs. For the detection of fruit, choice experiments showed that olfactory information was sufficient for mouse lemurs to find a piece of banana. Our study provides the first experimental data on the sensory ecology of food detection in mouse lemurs. Further research is necessary to address the role of sensory ecology for food selection and possibly for niche differentiation between sympatric Microcebus species.

Reptile, amphibian, and lemur diversity of the Malahelo Forest, a biogeographical transition zone in southeastern Madagascar

Ambatorongorongo Mountain lies at the historical intersection betweenhumid, spiny, and littoral forests in southeastern Madagascar. We report theresults of surveys of the herpetofauna and lemurs occurring in Malahelo Forest,a small (<25 ha) forest fragment lying on the western slope ofAmbatorongorongo Mountain. There are at least 41 reptile, 11 amphibian, and 7lemur species in this forest, including several that are endemic to southeasternMadagascar and are at severe risk of extinction. The species richness of theMalahelo fauna is comparable to that of even the largest forest reserves in theregion. We also evaluate the similarity of the Malahelo herpetofauna to that ofnearby humid, spiny, and littoral forests to assess the biogeographic affinitiesof its amphibians and reptile assemblages. Both groups contain speciescharacteristic of each of the three surrounding forest types, but thebiogeographic patterns appear to differ for amphibians and reptiles. Overall,the herpetofauna and lemurs of the Malahelo Forest indicate that it is a remnantof a biogeographic transition zone between the major forest types ofsoutheastern Madagascar. The combination of high species richness, regionalendemics, and unique herpetofaunal and lemur assemblages should make MalaheloForest a high conservation priority, and we give recommendations for protectingwhat remains of this important transitional forest.

Better few than hungry: flexible feeding ecology of collared lemurs Eulemur collaris in littoral forest fragments.

Background Frugivorous primates are known to encounter many problems to cope with habitat degradation, due to the fluctuating spatial and temporal distribution of their food resources. Since lemur communities evolved strategies to deal with periods of food scarcity, these primates are expected to be naturally adapted to fluctuating ecological conditions and to tolerate a certain degree of habitat changes. However, behavioral and ecological strategies adopted by frugivorous lemurs to survive in secondary habitats have been little investigated. Here, we compared the behavioral ecology of collared lemurs (Eulemur collaris) in a degraded fragment of littoral forest of south-east Madagascar, Mandena, with that of their conspecifics in a more intact habitat, Sainte Luce. Methodology/Principal Findings Lemur groups in Mandena and in Sainte Luce were censused in 2004/2007 and in 2000, respectively. Data were collected via instantaneous sampling on five lemur groups totaling 1,698 observation hours. The Shannon index was used to determine dietary diversity and nutritional analyses were conducted to assess food quality. All feeding trees were identified and measured, and ranging areas determined via the minimum convex polygon. In the degraded area lemurs were able to modify several aspects of their feeding strategies by decreasing group size and by increasing feeding time, ranging areas, and number of feeding trees. The above strategies were apparently able to counteract a clear reduction in both food quality and size of feeding trees. Conclusions/Significance Our findings indicate that collared lemurs in littoral forest fragments modified their behavior to cope with the pressures of fluctuating resource availability. The observed flexibility is likely to be an adaptation to Malagasy rainforests, which are known to undergo periods of fruit scarcity and low productivity. These results should be carefully considered when relocating lemurs or when selecting suitable areas for their conservation.

Biogeography of Dwarf Lemurs: Genetic Evidence for Unexpected Patterns in Southeastern Madagascar

Seven species of dwarf lemurs (Cheirogaleus spp.) are currently recognized after a recent revision of the genus. During a field study in southeastern Madagascar, we observed 3 distinct morphotypes of Cheirogaleus resembling Cheirogaleus medius, C. major and C. crossleyi. In particular, for Cheirogaleus crossleyi southeastern Madagascar was far away from the known distribution range of the species when referring to the recent revision of the genus. In order to clarify the taxonomic status of the 3 morphotypes, we compared field samples from southeastern Madagascar to museum specimens from several locations in Madagascar and to field samples from Kirindy/CFPF, western Madagascar by genetic analyses using the mitochondrial cytochrome b gene. Genetic data yield strong support for 2 of the 3 morphotypes to represent Cheirogaleus medius and C. major. The third morphotype is genetically closely related to Cheirogaleus crossleyi, albeit more distantly than the other 2 morphotypes to the respective reference samples. The genetic distance is related to a considerable geographic distance between the location of our field study and the origin of the respective reference specimen, because Cheirogaleus crossleyi had been reported only from northern Madagascar. Our field observations indicate an isolated population in southeastern Madagascar. We propose to identify them as Cheirogaleus crossleyi and discuss the population genetic and biogeographic considerations supporting this conclusion.

Extinction Risks and the Conservation of Madagascar's Reptiles

Background An understanding of the conservation status of Madagascars endemic reptile species is needed to underpin conservation planning and priority setting in this global biodiversity hotspot, and to complement existing information on the islands mammals, birds and amphibians. We report here on the first systematic assessment of the extinction risk of endemic and native non-marine Malagasy snakes, lizards, turtles and tortoises. Methodology/Principal Findings Species range maps from The IUCN Red List of Threatened Species were analysed to determine patterns in the distribution of threatened reptile species. These data, in addition to information on threats, were used to identify priority areas and actions for conservation. Thirty-nine percent of the data-sufficient Malagasy reptiles in our analyses are threatened with extinction. Areas in the north, west and south-east were identified as having more threatened species than expected and are therefore conservation priorities. Habitat degradation caused by wood harvesting and non-timber crops was the most pervasive threat. The direct removal of reptiles for international trade and human consumption threatened relatively few species, but were the primary threats for tortoises. Nine threatened reptile species are endemic to recently created protected areas. Conclusions/Significance With a few alarming exceptions, the threatened endemic reptiles of Madagascar occur within the national network of protected areas, including some taxa that are only found in new protected areas. Threats to these species, however, operate inside and outside protected area boundaries. This analysis has identified priority sites for reptile conservation and completes the conservation assessment of terrestrial vertebrates in Madagascar which will facilitate conservation planning, monitoring and wise-decision making. In sharp contrast with the amphibians, there is significant reptile diversity and regional endemism in the southern and western regions of Madagascar and this study highlights the importance of these arid regions to conserving the islands biodiversity.

Effects of rainforest fragmentation and correlates of local extinction in a herpetofauna from Madagascar

Habitat destruction in tropical forests often produces fragmented landscapes where relatively small forest remnants are present in a human-dominated matrix. To estimate rates of species loss after fragmentation and identify the traits of species which are most vulnerable to extinction, we assessed the distribution of amphibians and reptiles in an old (>50 yr) fragmented landscape in southeastern Madagascar. In the six rainforest fragments (10-457 ha) surveyed between 1999 and 2002, we detected forty-five species (28 reptiles, 17 amphibians). These species were not distributed randomly among the fragments but rather were very highly nested. The herpetofauna of small fragments consisted of smaller subsets of the species pool found in larger fragments. Species loss rates were high, with up to 64% of reptile and 88% of amphibian species lost when compared to the largest fragment. Species that were particularly prone to local extinction were generally rare, habitat specialists, and/or intolerant of the matrix habitat. Edge-avoidance was a good predictor of extinction vulnerability for frogs, but not for reptiles. Body size was generally not a good predictor of extinction proneness. In contrast with some other studies of amphibians and reptiles in fragmented landscapes, the results of this study indicate that species loss is a likely consequence of persistent isolation.

An Intersite Comparison of Fruit Characteristics in Madagascar: Evidence for Selection Pressure Through Abiotic Constraints Rather Than Through Co-Evolution

Fruit and seed characteristics are compared between a dry deciduous forest in the west and a humid littoral forest in the south-east of Madagascar to discriminate between the role of abiotic factors (humidity, climate, soil characteristics) and frugivorous vertebrates for the evolution of morphological and biochemical fruit characteristics. The sites differed in abiotic conditions but contain very similar communities of frugivorous vertebrates. Fruit selection by two lemur species (Eulemur fulvus and Cheirogaleus medius) that are important for seed dispersal and that are present at both study sites, was compared between sites to examine fixed selection criteria that could give rise to possible co-evolution between frugivores and their fruit species on the one hand or to dietary flexibility of the frugivores on the other hand. Our results show that most fruit characteristics differ significantly between study sites. Food selection by both lemur genera in relation to morphological and biochemical fruit characteristics co-varies closely with their representation at a given site. These results indicate that morphological and biochemical characteristics are more likely the result of abiotic conditions rather than of interactions between frugivorous lemurs and their food.