Longevity in some Malagasy rainforest passerines

Data on longevity is a prerequisite to understand the life-history strategies of a species. For Malagasy songbirds no information has been previously published. Therefore, we studied their longevity in a capture–recapture effort in a rainforest in eastern Madagascar (Maromizaha, Andasibe) between 2003 and 2016. We present first data on the longevity of 23 species of Malagasy songbirds. A female Dark Newtonia Newtonia amphichroa (Vangidae) and a Grey-crowned Greenbul Bernieria cinereiceps (Bernieridae) attained an age of at least 12 years, followed by two male Madagascar Brush Warblers Nesillas typica (Acrocephalidae), female Madagascar Bulbul Hypsipetes madagascariensis (Pycnonotidae) and Madagascar Drongo Dicrurus forficatus (Dicruridae) that attained at least 10 years. There was much variation within some taxonomic groups, longevity did not increase with the mass of a bird species and most insectivorous birds lived longer than granivorous ones.     

Global climate cycles and cyclones: consequences for rainfall patterns and lemur reproduction in southeastern Madagascar

Most studies that examine the influence of climatic change on flora and fauna have focused on northern latitudes; however, there is increasing recognition that tropical regions are also being affected. Despite this, regions such as Madagascar, which are rich in endemic biodiversity but may have low adaptive capacity to climatic change, are poorly represented in studies examining the effects of climate variability on biota. We investigated how El Niño Southern Oscillations (ENSO) influence precipitation patterns in the rainforest region of southeastern Madagascar (1962–2006) and then constructed models to assess the potential contribution of climatic variables on the reproductive parameters of the Milne Edward's sifaka, a threatened lemur species (Propithecus edwardsi), over a 20‐year period. The Southern Oscillation Index of sea surface temperature (SST) anomalies in the tropical Pacific was associated with precipitation patterns including wetter wet seasons during warmer phases and drier dry seasons following cooler phases. The best‐supported models of lemur fecundity (female offspring per female that survive to 1 year of age per year) included cyclone presence during gestation and ENSO phase before conception and during the first 6 months of life. Models also suggested that heavy rains during gestation may limit birth rates and that prolonged drought during female lactation may limit first year offspring survival; although these variables were given little importance for predicting overall fecundity relative to ENSO phases and cyclone presence. Our results linking lemur reproduction with climatic variability suggest that climatic changes may be an additional threat to Madagascar's unique and already endangered flora and fauna. The association between precipitation in southeastern Madagascar and SST anomalies in the tropical Pacific suggests that dynamics of wildlife populations even in tropical areas such as Madagascar can be affected by global climate cycles making them potentially vulnerable to global climate change.     

Effects of spatial proximity and alliances on dominance relations among female ring-tailed lemurs (Lemur catta) at Berenty Reserve, Madagascar

In the present study, we describe a change in the dominance rank of the top-ranking female in a wild troop of ring-tailed lemurs (Lemur catta) at Berenty Reserve, Madagascar. After the top-ranking female fell to the bottom-ranking position, she was able to outrank a low-ranking female with the support of her adult daughter or an unrelated high-ranking female. These results indicate that, as in cercopithecine monkeys such as macaques and baboons, close proximity and alliances influence dominance relations among adult females in a wild troop of ring-tailed lemurs.     

Matching dimorphic sexes and immature stages with adults: resolving the systematics of the Bekilya group of Malagasy assassin bugs (Hemiptera: Reduviidae: Peiratinae)

The Madagascar endemic assassin bugs Bekilya Villiers and Hovacoris Villiers were described from macropterous male specimens with striking colour patterns, and are currently monotypic. Mutillocoris Villiers, with two species from Madagascar, was based on brachypterous female specimens that resemble female mutillid wasps. To investigate the validity of the three genera, recently collected specimens from Madagascar were studied with both morphological and molecular techniques. Morphology alone appeared to be of limited value for associating males with females, and immature stages with adults, because of drastic differences between the sexes and the life stages. However, the use of morphology in conjunction with molecular data resolved these associations and showed that species of Mutillocoris represent females of Bekilya and Hovacoris, which we transfer accordingly to these two genera and refer to them as the Bekilya group. The type species of Mutillocoris belongs in Bekilya, resulting in the synonymy of these two genera (Mutillocoris syn.n. ). The Bekilya group is diagnosed and several new species are described: Bekilya mahafalya sp.n. , Bekilya tenebra sp.n. , Bekilya tuleara sp.n. , Hovacoris bicolornotum sp.n. , Hovacoris melanoceps sp.n. and Hovacoris rufiventris sp.n. A total of ten species are recognized within the Bekilya group. The monophyly of Bekilya, Hovacoris and the Bekilya group is confirmed by morphological and molecular phylogenetic analyses.     

TRAPPING AND RINGING OF EGYPTIAN GEESE AND AFRICAN SHELDUCK AT VOGELVLEI,CAPE

Many small plovers Charadrius spp. have sexually monomorphic plumage and cryptic sexual size dimorphism. The objective of our study was to assess the variation in body sizes between male and female plovers breeding in Madagascar. We collected blood samples and data on adult body sizes of four small plovers (Madagascar Plover Charadrius thoracicus, Kittlitz's Plover C. pecuarius, White-fronted Plover C. marginatus and Three-banded Plover C. tricollaris), and used molecular genetic markers to sex the adults. We found significant differences in body size among the four species, and between sexes. Furthermore, individuals from the southern ecoregion tended to be larger than in the western ecoregion. The Madagascar Plover's body size was significantly more dimorphic than the Kittlitz's and White-fronted Plovers. Breeding Malagasy plovers' show significant sexual size dimorphism (SSD): Madagascar Plover females were heavier and had longer wings than males, whereas the males had longer tarsi; in White-fronted Plover only wing length was different between the sexes. Taken together, our work reports SSD in small African plovers that exhibit monomorphic plumage, and we propose that SSD may be more common than currently acknowledged; we term this 'cryptic sexual size dimorphism'. Our results also suggest sexual selection and/or natural selection exert different pressures on body size in different Malagasy plover species.     

Phylogeography of Ptychadena mascareniensis suggests transoceanic dispersal in a widespread African-Malagasy frog lineage

Aim The Mascarene ridged frog, Ptychadena mascareniensis, is the only African amphibian species thought to occur on Madagascar and on the Seychelles and also Mascarene islands. We explored its phylogenetic relationships and intraspecific genetic differentiation to contribute to the understanding of transoceanic dispersal in amphibians. Methods Fragments of the mitochondrial 16S rRNA gene were sequenced from specimens collected over most of the distribution area of P. mascareniensis, including populations from Madagascar, Mascarenes and Seychelles. Results We identified five deeply divergent clades having pairwise divergences >5%, which probably all represent cryptic species in a P. mascareniensis complex. One of these seems to be restricted to Madagascar, the Mascarenes and the Seychelles. Sequences obtained from topotypic material (Réunion) were identical to the most widespread haplotype from Madagascar. The single Mauritian/Seychellean haplotype differed by only one mutation from a Malagasy haplotype. Main conclusions It is likely that the Mascarene and Seychellean populations were introduced from Madagascar by humans. In contrast, the absence of the Malagasy haplotypes from Africa and the distinct divergences among Malagasy populations (16 mutations in one divergent hapolotype from northern Madagascar) suggest that Madagascar was populated by Ptychadena before the arrival of humans c. 2000 years ago. Because Madagascar has been separated from Africa since the Jurassic, this colonization must have taken place by overseas rafting, which may be a more widespread dispersal mode in amphibians than commonly thought.     

Madagascar Fish Eagle productivity in the Tsimembo-Manambolomaty Protected Area and surrounding habitat of western Madagascar

We monitored the productivity of the critically endangered Madagascar Fish Eagle Haliaeetus vociferoides inside and outside of the Tsimembo-Manambolomaty Protected Area (T-M PA), western Madagascar from 2010 to 2015. We recorded 14 breeding pairs inside and 13 outside T-M PA. The T-M PA and surrounding habitat hosted respectively 10 and six breeding polyandrous pairs, composed of one adult female and two adult males. During the six-year study period, 101 eggs were laid in nests in T-M PA of which 60 hatched and 58 young fledged. We recorded 62 eggs laid in nests outside the T-M PA of which 39 hatched and 36 young fledged. Productivity was similar at both sites, inside and outside T-M PA, with 0.84 (58/69) and 0.76 (36/47) fledgling per nesting attempt and 0.69 (58/84) and 0.5 (36/72) fledglings per territorial pair, respectively. Polyandrous pairs have higher productivity compared with normal pairs. Threats to Madagascar Fish Eagles and their habitat were low due to the existence of a community-based resource management system called the Local Management Secured System (GELOSE) inside and outside the T-M PA. This system is based on strengthening local traditional customs and rules, and involving local people in managing their natural resources sustainably along with biodiversity conservation.     

The importance of littoral forest remnants for indigenous bird conservation in southeastern Madagascar

The littoral forests of Madagascar are relatively unexplored ecosystems that are considered seriously threatened by deforestation and habitat fragmentation. We set out to describe the bird communities inhabiting the littoral forest remnants in three different sub-regions of southeastern Madagascar to determine the national importance of these forests for bird conservation. In total, 77 bird species were found inhabiting 14 littoral forest remnants. Of these species, 40 are endemic to Madagascar and a further 21 are endemic to the Indian Ocean sub-region, consisting of Madagascar, the Comoros and the Mascarenes. The matrix habitats (Melaleuca forests, marécage swamp forest, Eucalyptus plantations and Erica grassland) that immediately surround the littoral forests were depauperate of bird species and contained few species that were found within the littoral forests. The geographic location of littoral forest remnants had an important role in determining what bird species occurred within them, with the northern remnants having similar bird communities to nearby humid forest whilst the most southern remnant had a bird community that resembled those of nearby spiny forest habitats. Eleven bird species that have been previously described as being habitat-restricted endemics to either spiny forests or humid forests, were found in littoral forest remnants. These results suggest that these littoral forests may play an important transitional role between the two other major natural habitats (spiny forest and humid forest) of southeastern Madagascar. On this basis we advocate that the littoral forest remnants of southeastern Madagascar should be afforded continuing conservation priority.     

Ecology,conservation, and phylogenetic position of the Madagascar Jacana Actophilornis albinucha

The Madagascar Jacana Actophilornis albinucha (Jacanidae) is an endemic shorebird found in the threatened wetlands of western Madagascar. This species is presumed to exhibit classical polyandry; however, few data are available to support that assumption. More generally, a lack of basic understanding of this species hinders conservation efforts. We conducted the most extensive study of the Madagascar Jacana to date, and report on its: 1) distribution, population size and density; 2) degree of sexual size dimorphism; and 3) phylogenetic position. The surveys were conducted at 54 lakes, between January and October in 2016. Madagascar Jacana were found at 22 lakes, and within these were distributed at a mean density of 3.5 ± 0.74 [SE] individuals per hectare of surveyed habitat. We estimate the global population size to be between 975 and 2 064 individuals, and habitat destruction appears to be the main threat to the species. Females were significantly larger than males, consistent with reports for other Jacanidae species. Using a mitochondrial DNA fragment, we expanded the Jacanidae genetic phylogeny, and confirmed that Madagascar Jacana is the sister species to the African Jacana Actophilornis africanus. Further studies are urgently needed to thoroughly re-assess the threat status and population trend of the Madagascar Jacana.     

Phylogeography of the introduced species Rattus rattus in the western Indian Ocean,with special emphasis on the colonization history of Madagascar

Aim To describe the phylogeographic patterns of the black rat, Rattus rattus, from islands in the western Indian Ocean where the species has been introduced (Madagascar and the neighbouring islands of Réunion, Mayotte and Grande Comore), in comparison with the postulated source area (India). Location Western Indian Ocean: India, Arabian Peninsula, East Africa and the islands of Madagascar, Réunion, Grande Comore and Mayotte. Methods Mitochondrial DNA (cytochrome b, tRNA and D‐loop, 1762 bp) was sequenced for 71 individuals from 11 countries in the western Indian Ocean. A partial D‐loop (419 bp) was also sequenced for eight populations from Madagascar (97 individuals), which were analysed in addition to six previously published populations from southern Madagascar. Results Haplotypes from India and the Arabian Peninsula occupied a basal position in the phylogenetic tree, whereas those from islands were distributed in different monophyletic clusters: Madagascar grouped with Mayotte, while Réunion and Grand Comore were present in two other separate groups. The only exception was one individual from Madagascar (out of 190) carrying a haplotype that clustered with those from Réunion and South Africa. ‘Isolation with migration’ simulations favoured a model with no recurrent migration between Oman and Madagascar. Mismatch distribution analyses dated the expansion of Malagasy populations on a time‐scale compatible with human colonization history. Higher haplotype diversity and older expansion times were found on the east coast of Madagascar compared with the central highlands. Main conclusions Phylogeographic patterns supported the hypothesis of human‐mediated colonization of R. rattus from source populations in either the native area (India) or anciently colonized regions (the Arabian Peninsula) to islands of the western Indian Ocean. Despite their proximity, each island has a distinct colonization history. Independent colonization events may have occurred simultaneously in Madagascar and Grande Comore, whereas Mayotte would have been colonized from Madagascar. Réunion was colonized independently, presumably from Europe. Malagasy populations may have originated from a single successful colonization event, followed by rapid expansion, first in coastal zones and then in the central highlands. The congruence of the observed phylogeographic pattern with human colonization events and pathways supports the potential relevance of the black rat in tracing human history.     

Genetically and geographically isolated lineages of a tropical bat (Chiroptera: Molossidae) show demographic stability over the late Pleistocene

The newly described molossid bat, Chaerephon atsinanana Goodman et al., 2010, endemic to eastern Madagascar, shows notably high levels of phylogeographic and genetic structure compared with allopatric Chaerephon leucogaster Grandidier, 1869 from western Madagascar. Such highly significant structuring of haplotypes among altitudinally and latitudinally stratified population groups is contrary to the expected panmixia in strong flying bats. The null model of concordance in historical demographic patterns across these two Chaerephon species was not supported. Mismatch and Bayesian skyline analyses indicated ancient stable C. atsinanana populations of constant size during the last two major Pleistocene glacial periods, making retreat into and expansion from glacial refugia an unlikely explanation for such high levels of structure, in accordance with expectations for tropical bats. Analyses were consistent with post‐refugial population expansion in the less diverse and structured C. leucogaster during the end of the last Pleistocene glacial period. We hypothesise that the pronounced genetic structuring in C. atsinanana may result from female philopatry. Furthermore, differing demographic histories of the two species may have been shaped by differing climate or habitat preferences, consistent with evidence from MaxEnt ecological niche modelling, which shows differences in variables influencing the current predicted distributions. Fossil Quaternary pollen deposits further indicate greater stability in past climatic patterns in eastern versus western Madagascar. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 18–40.     

First nest descriptions,nesting biology and food habits for Bernier's Vanga,Oriolia bernieri,in Madagascar

Four nests of the rare and endemic Bernier's Vanga, Oriolia bernieri, were discovered; one in 1997, one in 1998, and two in 1999 on the Masoala Peninsula, northeastern Madagascar. At the 1998 nest, the female made 189 visits with 186 deliveries of nesting material during 34.6 h of observation. The female spent 9.2% (194.2 min) of the observation time building the nest while an immature male delivered nest material six times and spent 3.2 min at the nest placing the material. Nesting material included: 67.2% (125) decomposed root material, 24.7% (46) palm fibres, 6.5% (12) dry leaves, 1.1% (2) moss, and 0.5% (1) white plant dawn. In 41.0 h of observation during the incubation period the female incubated for 53% (21.7 h) of the time, the adult male for 32.3% (13.2 h), the immature male for 4.3% (1.8 h), and the nest was unattended for 10.4% (4.3 h). This breeding attempt foiled on day 13 of incubation when a Madagascar Harrier-Hawk, Polyboroides radiatus, ate the egg(s). At one of the 1999 nests, the incubation and nestling periods were 17 days each. Three young fledged during the middle of November. Of the 82 identified prey items recorded during the nestling period, 91% were invertebrates and 9% vertebrates. Spiders, crickets, cockroaches, and geckos represented the most numerous prey taken, totaling 77% of the identified prey.     

Coalescent analyses support multiple mainland-to-island dispersals in the evolution of Malagasy Triaenops bats (Chiroptera: Hipposideridae)

Aim We investigate the directionality of mainland‐to‐island dispersals, focusing on a case study of an African‐Malagasy bat genus, Triaenops (Hipposideridae). Taxa include T. persicus from east Africa and three Triaenops species from Madagascar (T. auritus, T. furculus, and T. rufus). The evolution of this bat family considerably post‐dated the tectonic division of Madagascar from Africa, excluding vicariance as a viable hypothesis. Therefore, we consider three biogeographical scenarios to explain these species' current ranges: (A) a single dispersal from Africa to Madagascar with subsequent speciation of the Malagasy species; (B) multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy lineages; or (C) early dispersal of a proto‐species from Africa to Madagascar, with later back‐dispersal of a descendant Malagasy taxon to Africa. Location East Africa, Madagascar, and the Mozambique Channel. Methods We compare the utility of phylogenetic and coalescent methodologies to address the question of directionality in a mainland‐to‐island dispersal event for recently diverged taxa. We also emphasize the application of biologically explicit demographic systems, such as the non‐equilibrium isolation‐with‐migration model. Here, these methods are applied to a four‐species haploid genetic data set, with simulation analyses being applied to validate this approach. Results Coalescent simulations favour scenario B: multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy bat lineages. From coalescent dating, we estimate that the genus Triaenops was still a single taxon approximately 2.25 Ma. The most recent Africa to Madagascar dispersal occurred much more recently (c. 660 ka), and led to the formation of the extant Malagasy species, T. rufus. Main conclusions Haploid genetic data from four species of Triaenops are statistically most consistent with multiple, unidirectional dispersals from mainland Africa to Madagascar during the late Pleistocene.     

Characterization of 11 microsatellite marker loci in the Malagasy big‐headed turtle (Erymnochelys madagascariensis)

The Malagasy big‐headed turtle (Erymnochelys madagascariensis) is the only Erymnochelys species living in lakes, rivers and watersheds of western Madagascar. This species is endangered due to over harvesting of natural populations for human consumption. Eleven nuclear microsatellite loci were isolated from a genomic DNA library derived from a free‐ranging Malagasy big‐headed turtle from the Beroboka Classified Forest, Madagascar. Population genetic parameters were estimated on 10 individuals sampled from Ampijoroa and Andranohobaka River, Madagascar, to determine marker utility and as preliminary baseline values to study future populations in these locations.     

Guloptiloides: an Extraordinary New Carnivorous Genus of Baetidae (Ephemeroptera)

Guloptiloides (Ephemeroptera, Baetidae), gen.n., is described to accomodate the species G. gargantua (larvae) and G. spA (female imagoes) based on specimens from the Eastern coast of Madagascar. Larvae of Guloptiloides are remarkable by the adaptation of the mouthparts to a predatory diet. It is also distinct from other genera in that adults have hindwings with 3 well-marked longitudinal veins and a single extremely developed spur on the costal margin. Affinities and ecology are discussed.     

Linking seed dispersal and genetic structure of trees: a biogeographical approach

Aim Natural and human‐induced differences in frugivore assemblages can influence the seed dispersal distances of trees. An important issue in seed dispersal systems is to understand whether differences in seed dispersal distances also affect the genetic structure of mature trees. One possible approach to test for a relationship between seed dispersal and the genetic structure of mature trees is to compare the genetic structure of two closely related tree species between two biogeographical regions that differ in frugivore assemblages and seed dispersal distances. Previous studies on two Commiphora species revealed that Commiphora guillauminii in Madagascar has a much lower seed dispersal distance than Commiphora harveyi in South Africa. We tested whether the lower seed dispersal distance might have caused decreased gene flow, resulting in a stronger genetic structure in Madagascar than in South Africa. Location Madagascar and South Africa. Methods Using amplified fragment length polymorphism markers we investigated the genetic structure of 134 trees in Madagascar and 158 trees in South Africa at a local and a regional spatial scale. Results In concordance with our hypothesis, kinship analysis suggests that gene flow was restricted mostly to 3 km in Madagascar and to 30 km in South Africa. At the local spatial scale, the genetic differentiation among groups of trees within sample sites was marginally significantly higher in Madagascar (F_ST = 0.069) than in South Africa (F_ST = 0.021). However, at a regional spatial scale genetic differentiation was lower in Madagascar (F_ST = 0.053) than in South Africa (F_ST = 0.163). Main conclusions Our results show that lower seed dispersal distances of trees were linked to higher genetic differentiation of trees only at a local spatial scale. This suggests that seed dispersal affects the genetic population structure of trees at a local, but not at a regional, spatial scale.     

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.     

Characterization of 20 microsatellite marker loci in the Malagasy tree boa (Sanzinia madagascariensis madagascariensis)

The Madagascar tree boa, Sanzinia madagascariensis madagascariensis, is one of four snakes in the family Boidae living in Madagascar. This species is considered ‘vulnerable’ due to habitat loss and utilization as a food source by locals. Twenty species-specific microsatellite loci were isolated and characterized from Sanzinia m. madagascariensis to assess population genetic parameters. Individuals were collected from two populations in the east and the northeastern coast of Madagascar: Torotofotsy wetlands and Mananara-Nord Biosphere Reserve, respectively. This marker suite will provide a useful tool for future research to determine and validate the taxonomic status of the tree boa with samples collected extensively throughout the island.     

Male Mating Success in the Presence of a Conspecific Opponent in a Madagascar Hissing Cockroach,Gromphadorhina portentosa (Dictyoptera: Blaberidae)

The influence of a conspecific competitor on male mating behavior was examined in a Madagascar hissing cockroach, Gromphadorhina portentosa. Previous studies have suggested that both male-male competition and female discrimination during courtship interactions may influence male mating success. Familiar pairs of males with a known social association were placed in an arena with a single virgin female and observed. As expected, subordinate males mated significantly less often than their dominant opponents. In pairs in which one male mated, dominant individuals limited the access of subordinates to females. Dominant males displayed an increased frequency and duration of interaction with the female. However, in pairs where both males remained unmated, the mating behavior of dominant and subordinate males did not differ significantly. As interactions progressed, as in the case of males that remained unmated, subordinate males gained increased access to the female. Mated males tended to be larger than their opponent although within a rank, males that mated were no larger than those that remained unmated. These results are discussed in light of the possible roles of male-male competition and female discrimination during courtship interactions.