An endemic radiation of Malagasy songbirds is revealed by mitochondrial DNA sequence data

The bird fauna of Madagascar includes a high proportion of endemic species, particularly among passerine birds (Aves: Passeriformes). The endemic genera of Malagasy songbirds are not allied obviously with any African or Asiatic taxa, and their affinities have been debated since the birds first were described. We used mitochondrial sequence data to estimate the relationships of 13 species of endemic Malagasy songbirds, 17 additional songbird species, and one species of suboscine passerine. In our optimal trees, nine of the 13 Malagasy species form a clade. although these birds currently are classified in three different families. In all optimal trees, the sister to this endemic clade is a group of Old World warblers including both African and Malagasy birds. The endemic Malagasy songbird clade rivals other island radiations, including the vangas of Madagascar and the finches of the Galapagos, in ecological diversity.     

Integrative taxonomy and phylogeny of leafless Vanilla orchids from the South-West Indian Ocean region reveal two new Malagasy species

The leafless Vanilla species complex from the South-West Indian Ocean (SWIO) region has long been a taxonomic challenge, due to limited patterns of morphological differentiation and an absence of variation within chloroplast sequences. This complex includes seven known morphospecies: V. madagascariensis, V. bosseri, V. decaryana, and V. perrieri endemic to Madagascar, V. humblotii presumed as endemic to the Comoros Archipelago, but also present in Madagascar, V. roscheri from the East African coast, and V. phalaenopsis endemic to Seychelles. A previous population genetic study using microsatellite markers allowed us to distinguish, in addition to the five recognized Malagasy taxa, two other genetic clusters present in the East of the island. An integrative taxonomy approach was therefore conducted by combining microsatellite and morphological data used in the previous study with new data sets, and by adding ITS sequencing data, to validate the taxonomic level of these Malagasy genetic clusters and unravel phylogenetic relationships between SWIO species. As a result, based on phylogenetic, genotypic and morphological evidence, nine species were discriminated in the SWIO region, including seven in Madagascar, with two new eastern species. The leafless Vanilla group originated and diversified in Madagascar, from an ancestor of African descent, with three subsequent independent colonization events from Madagascar to the other territories of SWIO within the two main lineages (white versus yellow flower species). The new Malagasy species, V. allorgeae Andriamihaja & Pailler sp. nov., and V. atsinananensis Andriamihaja & Pailler sp. nov., are described and a new identification key is proposed.     

New species of the Ornithomya biloba-group and records of other Hippoboscidae (Diptera) from Africa

Two new species of the Ornithomya biloba-group are described, one from Kenya and the second from Madagascar, and a revised key to the African species of this group is given. Records of other African Hippoboscidae from birds are included.     

Diversification and the adaptive radiation of the vangas of Madagascar

The vangas of Madagascar exhibit extreme diversity in morphology and ecology. Recent studies have shown that several other Malagasy species also are part of this endemic radiation, even as the monophyly of the clade remains in question. Using DNA sequences from 13 genes and representatives of all 15 vanga genera, we find strong support for the monophyly of the Malagasy vangids and their inclusion in a family along with six aberrant genera of shrike-like corvoids distributed in Asia and Africa. Biogeographic reconstructions of these lineages include both Asia and Africa as possible dispersal routes to Madagascar. To study patterns of speciation through time, we introduce a method that can accommodate phylogenetically non-random patterns of incomplete taxon sampling in diversification studies. We demonstrate that speciation rates in vangas decreased dramatically through time following the colonization of Madagascar. Foraging strategies of these birds show remarkable congruence with phylogenetic relationships, indicating that adaptations to feeding specializations played a role in the diversification of these birds. Vangas fit the model of an 'adaptive radiation' in that they show an explosive burst of speciation soon after colonization, increased diversification into novel niches and extraordinary ecomorphological diversity.     

Taxonomy of the hyper-diverse ant genus Tetramorium Mayr in the Malagasy region (Hymenoptera,Formicidae, Myrmicinae) – first record of the T. setigerum species group and additions to the Malagasy species groups with an updated illustrated identification key

In this study we provide an update to the taxonomy of the ant genus Tetramorium Mayr in Madagascar. We report the first record of the Tetramorium setigerum species group in Madagascar and describe the only Malagasy representative as Tetramorium cavernicola sp. n., which is known only from a cave in Ankarana. In addition, we provide an overview of the 19 proposed Malagasy species groups, and discuss their zoogeography and relationships to other groups and larger lineages within the hyper-diverse genus Tetramorium. At present, we recognise a highly unique Malagasy Tetramorium fauna with 113 species endemic to the island of Madagascar out of a total of 125 translating into an endemism rate of 93%. We hypothesise that this fauna is based on one or a few colonisation events from the Afrotropical region, with subsequent adaptive radiation in Madagascar. Furthermore, we present an updated and illustrated identification key to the Tetramorium species groups in the Malagasy region.     

Rapid assessments and local knowledge reveal high bird diversity in mangroves of north-west Madagascar

Although the importance of regulating and provisioning services provided by mangroves is widely recognised, our understanding of their role in the maintenance of terrestrial biodiversity is patchy globally and largely lacking for many regions, including conservation priorities such as Madagascar. We carried out the first multi-site bird inventory of mangroves in Madagascar and complemented our data with assessments of local knowledge, in order to broaden our knowledge of which species use this habitat. We directly observed 73 species across three sites in Ambanja and Ambaro Bays, while local respondents indicated the presence of 18 additional species: four observed species are globally threatened, while 37 are endemic to Madagascar or the Malagasy region. Over half the species observed are typically terrestrial, of which 22 have not previously been recorded in mangrove habitats in Madagascar. Local knowledge provided a useful complement to our observed data but we are likely to have underestimated total richness; nevertheless, our findings greatly increased our knowledge of mangrove use by Madagascar’s birds. However, further research is required to investigate the functional role of mangroves in the ecology of the observed species and provide insights into the factors influencing mangrove use.     

A new species of Plasmodium from Malagasy vangas

During a recent examination of blood smears from Malagasy birds, a species of avian Plasmodium unlike those currently known was observed. All infected birds were members of the Vangidae, which is endemic to Madagascar and the Comoro Islands. Plasmodium parvulum n. sp. is described, and classified as a member of the subgenus Haemamoeba because of gametocyte and schizont shape, displacement of the host cell nucleus, as well as distortion of the host cell. Round, rosettelike schizonts with 6-8 merozoites, clumped refractile granules, and little cytoplasm were observed. Both schizonts and mature, round gametocytes rotated and displaced the erythrocyte nucleus. A brief comparison to P. relictum is included.     

Molecular phylogeny and biogeography of the Malagasy and South Asian cichlids (Teleostei: Perciformes: Cichlidae)

Phylogenetic relationships of the Malagasy and South Asian cichlids are investigated using nucleotide characters from two mitochondrial genes, a 544 bp region of the large ribosomal subunit (16S) and a 649 bp region of cytochrome c oxidase subunit I (COI). This is the first molecular analysis to include a thorough taxonomic sampling of all Malagasy-South Asian genera, and a near complete taxonomic inventory of all valid species. Parsimony analysis of the combined data set (1193 aligned nucleotide positions) results in a single, completely resolved phylogenetic hypothesis. Results of this analysis, and that based on more comprehensive taxonomic sampling across Cichlidae for 16S alone (554 bp for 73 taxa), indicate that the Malagasy cichlids are paraphyletic, whereas the Malagasy and South Asian cichlids comprise a monophyletic group. In both analyses, the African and Neotropical assemblages are monophyletic. The Malagasy-South Asian cichlids are not recovered as plesiomorphic members of the family in either analysis. Two major clades are recovered within the Malagasy-South Asian assemblage and given subfamilial rank, Etroplinae, comprising Paretroplus (Madagascar) and Etroplus (southern India and Sri Lanka), and Ptychochrominae, comprising Ptychochromis, Ptychochromoides, and Oxylapia, all endemic Malagasy genera. Placement of the endemic Malagasy genus Paratilapia is equivocal depending on the gene fragment(s) analyzed. Inter- and intrageneric relationships within Ptychochrominae and Etroplinae are presented and discussed. The hypothesis of relationships for Cichlidae based on nucleotide characters from 16S alone, arguably the most comprehensive and broadly sampled data set across all major geographic assemblages to date, is congruent with prevailing hypotheses regarding the sequence of Gondwanan fragmentation and a vicariance scenario to explain the current distribution of cichlid fishes.     

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.     

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.     

Springs and wire plants: anachronistic defences against Madagascar's extinct elephant birds

The extinction of large vertebrates in the last few millennia has left a legacy of evolutionary anachronisms. Among these are plant structural defences that persist long after the extinction of the browsers. A peculiar, and controversial, example is a suite of traits common in divaricate (wide-angled branching) plants from New Zealand. Divaricate architecture has been interpreted as an adaptive response to cold climates or an anachronistic defence against the extinct moas. Madagascar, a larger tropical island, also had a fauna of large flightless birds, the elephant birds. If these extinct ratites selected for similar plant defences, we expected to find convergent features between New Zealand and Malagasy plants, despite their very different climates. We searched the southern thickets of Madagascar for plants with putative anti-ratite defences and scored candidate species for a number of traits common to many New Zealand divaricates. We found many Malagasy species in 25 families and 36 genera shared the same suite of traits, the 'wire plant' syndrome, as divaricates in New Zealand that resist ratite browsing. Neither ecologically, nor phylogenetically, matched species from South Africa shared these traits. Malagasy wire plants differ from many New Zealand divaricates in lacking the distinctive concentration of leaves in the interior of shrubs. We suggest that New Zealand divaricates have a unique amalgam of traits that acted as defences and also confer tolerance to cold. We conclude that many woody species in the thickets of southern Madagascar share, with New Zealand, anachronistic structural defences against large extinct bird browsers.     

The Kilopilopitsofy, Kidoky, and Bokyboky: Accounts of Strange Animals from Belo-sur-mer, Madagascar, and the Megafaunal "Extinction Window"

In July and August 1995, the authors interviewed elderly Malagasy with knowledge of the traditions and natural history of their home areas, centered on the villages of Belo-sur-mer, Antsira, and Ambararata, on the southwest coast of Madagascar.
Several individuals related personal experiences in which they claim to have seen and heard animals that do not match any known extant animal of Madagascar. Two of the mystery animals, known locally as the kilopilopitsofy and kidoky, were described in terms similar to creatures detailed in historical accounts and folklore recorded in Madagascar between the mid-1600s and the end of the nineteenth century. The former of these has been compared by some authors to the dwarf hippopotamus and the latter to a giant lemur, animals generally inferred from radiocarbon dating of last known occurrences to have gone extinct early in the present millennium or perhaps a few centuries later.
Stories by these same informants concerning extant animals demonstrate their accurate knowledge of the fauna. Magical powers are ascribed to some animals, such as the bokyboky, a viverrid (Mungotictis) that occurs in the region. Radiocarbon dates obtained recently on some of the extinct megafauna, showing that some extirpated taxa may have survived until recent centuries, confirm that ethnographic sources of information on these species should not automatically be dismissed as irrelevant or unreliable. [Madagascar, extinctions, megafauna, hippopotamus, lemur]     

Shifting continents, not behaviours: independent colonization of solitary and subsocial Anelosimus spider lineages on Madagascar (Araneae, Theridiidae)

Agnarsson, I., Kuntner, M., Coddington, J. A. & Blackledge, T. A. (2009). Shifting continents, not behaviours: independent colonization of solitary and subsocial Anelosimus spider lineages on Madagascar (Araneae, Theridiidae). —Zoologica Scripta, 39, 75–87. Madagascar is a biodiversity hotspot, thought to be colonized mostly via Cenozoic dispersal from Africa, followed by endemic radiation of multiple lineages. Anelosimus spiders are diverse in Madagascar, and, like their congeners in the Americas, are most diverse in wet montane forests. Most Anelosimus species are social in that they cooperate in web building and prey capture either during a part of their life cycles (subsocial), including hitherto studied Malagasy species, or permanently (quasisocial). One Central American coastal species, Anelosimus pacificus, has secondarily switched to solitary living, and available evidence suggests that its closest relatives from S. America and Europe are likely also solitary. Here, we show that the only known coastal Anelosimus species in Madagascar and Comoros –Anelosimus decaryi and Anelosimus amelie sp. n. – are also solitary. Using a phylogenetic approach, we test two competing hypotheses: (i) that Malagasy Anelosimus are monophyletic and thus represent a second example of reversal to solitary living in a littoral habitat or (ii) that solitary and subsocial lineages independently colonized Madagascar. We find that solitary Malagasy Anelosimus are closely related to their solitary counterparts from Europe and the Americas, while subsocial Malagasy species nest sister to Anelosimus nelsoni from S. Africa. This finding suggests that (i) the two Anelosimus lineages colonized Madagascar independently and (ii) a reversal to solitary behaviour has occurred only once in Anelosimus. Thus, solitary littoral Malagasy species did not descend from Malagasy mountains, but arrived from much further afar. African and possibly American origin of the two lineages is implied by our findings. To restore natural classification of Anelosimus, Seycellocesa Koçak & Kemal is synonymized with it.     

Altitudinal and latitudinal distribution of east Malagasy forest bird communities

Aim To describe variation in forest bird communities with altitude and latitude. Location Eastern Madagascar. Methods Extraction of data from forest bird inventories conducted in eastern Madagascar. Results There is a strong decline in species richness with altitude, above about 1300 m. Below this altitude, species richness is about constant or declines slightly. Seventy-eight percent of species occurring regularly in forest are absent from at least one of low, mid-altitude or high altitude forest. Of eighty-seven species occurring regularly in forest, only four or possibly five have latitudinally limited distributions, over a latitudinal range of over 1200 km. Three or possibly four are limited to the northern two-thirds, and one appears to be at least much more common in the southern half. Main conclusions Eastern Malagasy rain forest birds show previously unanalysed variation in altitudinal distribution. There is much less latitudinal variation. Species currently considered threatened are concentrated in the lowland and high-altitude zones. This may be at least partly due to lack of survey effort giving the impression that these species are rare, but lowland forests at least are under great human pressure. Bird conservation initiatives would probably have most effect if targeted at lowland east Malagasy rain forest.     

A report on a collection of pontoniine shrimps from Madagacar and adjacent seas

A collection of 41 species of pontoniine shrimp form the Madagascar region is reported upon. Most of the material has been collected from the north-west of Madagascar in the vicinity of Nosy Bé. Six new species, belonging to four different genera, are described: Palaemonella crosnieri, Periclimenes lepidus, P. granulimanus, P. perturbans, Periclimenaeus lobiferus and Onycocaris trullata . A key to the species of Onycocaris is also given. Nineteen species are recorded from Madagascar for the first time, and four species, Periclimenes perlucidus, Periclimenaeus djiboutensis, Pontonia sibogae and Coralliocaris viridis are reported for the second time only in the Indo-West Pacific region. Pontonides unciger Calman is considered to be distinct from P. unciger Holthuis and also is recorded for the second time only. The records of pontoniine shrimp previously reported from Madagascar are briefly summarized. A total of 61 species are now known to be represented in the fauna of Madagascar, indicating a diversity almost as rich as the Indonesian region, with 69 species. Une collection de quarante et une espèces de crevettes pontoniine de la région de Madagascar est analysée. La plus grande partie du matériel provient du nord-ouest de Madagascar, dans le voisinage de Nosy Bé. Six espèces nouvelles appartenant à quatre genres différents sont décrites: Palaemonella crosnieri, Periclimenes lepidus, P. granulimanus, P. perturbans, Periclimenaeus lobiferus et Onycocaris trullata. Une clé des espèces du genre Onycocaris est aussi donnée. Dix-neuf espèces sont notées à Madagascar pour la première fois, et quatre espèces ne sont notées que pour la seconde fois dans la region Indo-Pacifique occidentale. Pontonides unciger Calman est consideréêtre distinct de P. unciger Holthuis, 1952, et n'est aussi noté que pour la seconde fois. Les rapports notant l'occurrence de crevettes pontoniines à Madagascar sont rapidement resumés. Un total de soixante-et-une espèces sont maintenant connues dans la faune de Madagascar, indiquant une diversité presque aussi riche que celle de la region Indonesienne, qui possède soixante-neuf espèces.     

Beta-globin haplotype analysis suggests that a major source of Malagasy ancestry is derived from Bantu-speaking Negroids.

The origins of the inhabitants of Madagascar have not been fully resolved. Anthropological studies and preliminary genetic data point to two main sources of ancestry of the Malagasy, namely, Indonesian and African, with additional contributions from India and Arabia. The sickle-cell (beta s) mutation is found in populations of African and Indian origin. The frequency of the beta s-globin gene, derived from 1,425 Malagasy individuals, varies from 0 in some highland populations to .25 in some coastal populations. The beta s mutation is thought to have arisen at least five times, on the basis of the presence of five distinct beta s-associated haplotypes, each found in a separate geographic area. Twenty-five of the 35 Malagasy beta s haplotypes were of the typical "Bantu" type, 1 "Senegal" haplotype was found, and 2 rare or atypical haplotypes were observed; the remaining 7 haplotypes were consistent with the Bantu haplotype. The Bantu beta s mutation is thought to have been introduced into Madagascar by Bantu-speaking immigrants (colonists or slaves) from central or east Africa. The Senegal beta s mutation may have been introduced to the island via Portuguese naval explorers. This study provides the first definitive biological evidence that a major component of Malagasy ancestry is derived from African populations, in particular, Bantu-speaking Negroids. beta A haplotypes are also consistent with the claim for a significant African contribution to Malagasy ancestry but are also suggestive of Asian/Oceanic and Caucasoid admixture within the Malagasy population.     

Field observations on some dwarf chameleons (Brookesia spp.) from rainforest areas of Madagascar, with description of a new species

Six species of Brookesia dwarf chameleons were recorded during two rainforest expeditions in Madagascar. One of these is described as a new species Brookesia valerieae based on the arrangement of the nine pairs of dorso-lateral spines, the lack of a pelvic shield, and dorsal chevron markings. Observations were made on the defence and roosting behaviours shown by these six species. Aspects concerning the evolution and conservation in Madagascar are discussed for this poorly studied group of endemic Malagasy reptiles.     

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.     

Extinction Risks and the Conservation of Madagascar's Reptiles

Background

An understanding of the conservation status of Madagascar''s endemic reptile species is needed to underpin conservation planning and priority setting in this global biodiversity hotspot, and to complement existing information on the island''s 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 island''s biodiversity.     

Large-scale phylogeny of chameleons suggests African origins and Eocene diversification

Oceanic dispersal has emerged as an important factor contributing to biogeographic patterns in numerous taxa. Chameleons are a clear example of this, as they are primarily found in Africa and Madagascar, but the age of the family is post-Gondwanan break-up. A Malagasy origin for the family has been suggested, yet this hypothesis has not been tested using modern biogeographic methods with a dated phylogeny. To examine competing hypotheses of African and Malagasy origins, we generated a dated phylogeny using between six and 13 genetic markers, for up to 174 taxa representing greater than 90 per cent of all named species. Using three different ancestral-state reconstruction methods (Bayesian and likelihood approaches), we show that the family most probably originated in Africa, with two separate oceanic dispersals to Madagascar during the Palaeocene and the Oligocene, when prevailing oceanic currents would have favoured eastward dispersal. Diversification of genus-level clades took place in the Eocene, and species-level diversification occurred primarily in the Oligocene. Plio-Pleistocene speciation is rare, resulting in a phylogeny dominated by palaeo-endemic species. We suggest that contraction and fragmentation of the Pan-African forest coupled to an increase in open habitats (savannah, grassland, heathland), since the Oligocene played a key role in diversification of this group through vicariance.