Phylogenetic relationships and biogeography of Ptilophora (Gelidiales,Rhodophyta) with descriptions of P. aureolusa,P. malagasya,and P. spongiophila from Madagascar

The genus Ptilophora currently includes 16 species occurring mostly in subtidal habitats of the Indo‐Pacific Ocean, but its global diversity and biogeography are poorly understood. We analyzed mitochondrial cox1, plastid rbcL and plastid psbA sequences from specimens collected in southern Madagascar during the 2010 Atimo Vatae expedition and studied their morphologies. Both morphological and molecular data sets demonstrated the presence of five species in southern Madagascar: Ptilophora hildebrandtii, P. pterocladioides, and three new species described here, P. aureolusa, P. malagasya, and P. spongiophila. Ptilophora aureolusa is distinguished by its compound pinnae with uniformly spaced pinnules. Ptilophora malagasya has an indistinct midrib and irregularly spaced pinnules. Ptilophora spongiophila, heavily coated with sponges, has cylindrical to flattened main axes, lateral and surface proliferations, and spatulate tetrasporangial sori. The species of Ptilophora found in Madagascar are endemic, except P. hildebrandtii, which also occurs in eastern Africa. Ptilophora comprises four phylogenetic groups that map to eastern Australia, Japan, western Australia/Southeast Asia/Madagascar/eastern Africa, and Madagascar/eastern Africa/Aegean Sea. Biogeographical analysis revealed that the ancestor of Ptilophora originated in Australia, but most of the species radiated from Madagascar.     

Riverbeds demarcate distinct conservation units of the radiated tortoise (<Emphasis Type="Italic">Geochelone radiata</Emphasis>) in southern Madagascar

The radiated tortoise (Geochelone radiata) is an endangered species endemic to Madagascar. It inhabits the semiarid spiny forest of the southern part of the island, an ecosystem heavily affected by habitat destruction. Furthermore, illegal harvesting greatly threatens this species. The main objective of our study was to acquire better knowledge of its genetic structure, in order to take appropriate management decisions concerning, for instance, the reintroduction of confiscated individuals. Our hypothesis was that rivers represent effective barriers to tortoise dispersal despite the fact that they are dry most of the year. We used 13 polymorphic microsatellite markers to compare samples from six populations across the range of the species. All analyses (Fisher’s exact tests, F _ST values, AMOVA) indicated that the radiated tortoise exhibits moderate levels of genetic structure throughout its range. In addition, we used a multiple regression approach that revealed the importance of rivers to explain the observed structure. This analysis supported the role of the Menarandra and Manambovo Rivers as major barriers to the dispersal of most radiated tortoises, but Markov chain Monte Carlo simulations revealed that low levels of recurrent gene flow may explain why F _ST values were not higher. We identified three distinct conservation units with relatively high assignments rates (87%), which should be valuable for the management of the species. This is the first study to report the genetic structure of a species sampled throughout the Malagasy spiny forest.     

Characterization of 21 microsatellite marker loci in the ring-tailed lemur (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Ring-tailed lemur (Lemur catta) is the only species in the Genus Lemur, distributed in the deciduous and spiny forests of southwestern Madagascar. This species is listed as endangered due to the loss and fragmentation of its natural habitat, a consequence of deforestation. Twenty-one nuclear microsatellite loci were isolated from a genomic DNA derived from a free-ranging ring-tailed lemur population from the Tsimanampetsotsa National Park, Madagascar. We report various parameter estimates and measures to establish the utility of this marker suite as screened among individuals this single forest fragment.     

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.     

Characterization of 18 microsatellite marker loci in the white-collared lemur (Eulemur cinereiceps)

The white-collared lemur (Eulemur cinereiceps) is found in southeastern Madagascar. This species is endangered due to the loss and fragmentation of its natural habitat and hunting. Eighteen nuclear microsatellite loci were isolated from genomic DNA derived from an animal from Mahabo Forest in Madagascar. Population genetic parameters were estimated from a population of white-collared lemur sampled at Vevembe Forest 7 years apart to evaluate the potential utility of this marker suite.     

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.     

Characterization of 20 microsatellite marker loci in Coquerel's sifaka (Propithecus coquereli)

The Coquerel's sifaka (Propithecus coquereli) is one of the species of Propithecus, living in the dry forest of northwest Madagascar. This species is endangered due to the loss and fragmentation of its natural habitat, consequential of deforestation. Twenty nuclear microsatellite loci were isolated from genomic DNA derived from a free ranging Coquerel's sifaka from the Mariarano Classified Forest, Madagascar. Population genetic parameters were estimated as baseline values from samples collected in that forest.     

Characterization of 26 microsatellite marker loci in the fossa (Cryptoprocta ferox)

The fossa (Cryptoprocta ferox) is one of eight carnivore species endemic to Madagascar. This species lives in the forests of Madagascar and is classified as vulnerable by the IUCN due to the loss and fragmentation of its natural habitat and the hunting pressure of local people. Twenty-six nuclear microsatellite loci were isolated from genomic DNA derived from a free-ranging fossa from Betampona, Madagascar. Population genetic parameters were estimated on 28 individuals from nine European zoos establishing a baseline to study the genetic health and diversity of captive fossas to support the European Breeding and Conservation Program (EEP). The marker suite averaged 4.4 alleles per locus with an average polymorphic information content of 0.478.     

Characterization of 20 microsatellite markers in the plowshare tortoise,Astrochelys yniphora

The plowshare tortoise, Astrochelys yniphora, or Angonoka is one of Madagascar’s land tortoises, living in the vicinity of Baly Bay bamboo scrub in northwest Madagascar. Primary threats to this endangered species are due to poaching for pet trade and the loss and fragmentation of its natural habitat. The number of wild Angonoka is estimated to be ~400. Twenty polymorphic nuclear microsatellite loci were isolated from a genomic DNA on individuals from Andrafiafaly in Baly Bay National Park, Madagascar. Results from this study will help for future studies of the social structure and population dynamics of this species as well as identification of confiscated individuals from the illegal pet trade.     

Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers

We studied mitochondrial divergence in 27 individuals of colubrid snakes of the genus Madagascarophis Mertens from most of its distribution area in Madagascar. Combined analyses of 16S rRNA and cytochrome b sequences identified six major clades which only partly agreed with previously proposed classifications. Analysis of nuclear DNA sequences of the c-mos gene as well as of ISSR fingerprints revealed consistent differences only among three clades which we consider as distinct species: a widespread Madagascarophis colubrinus (Schlegel), with M. citrinus (Boettger) as a junior synonym, a southern M. meridionalis Domergue, and a presumably undescribed species from the extreme north of Madagascar. The species M. ocellatus Domergue was not available for our study. Within M. colubrinus there are two populations from the north-west, each showing two divergent haplotypes with pairwise divergences of up to 5.2% in the cytochrome b gene. Maximum divergence in this gene within M. colubrinus was 7.1%. These high values emphasise that caution needs to be applied before genetic distance values are used for species delimitation. Phylogeographically, most of the genetic variation in M. colubrinus is found in northern Madagascar, indicating that the species might have originated in this region. Later one haplotype clade colonised western and eastern Madagascar, with a putative secondary introgression into north-western populations.     

Characterization of 20 microsatellites marker loci in the golden-crowned sifaka (<Emphasis Type="Italic">Propithecus tattersalli)</Emphasis>

The golden-crowned sifaka (Propithecus tattersalli) is one of the most critically endangered species of Propithecus endemic to a limited area in the Daraina region of north-eastern Madagascar. This species is endangered due to habitat loss and fragmentation, a consequence of deforestation. Twenty nuclear microsatellite loci were isolated from genomic DNA derived from a Propithecus tattersalli from the Daraina area in north-eastern Madagascar. Population genetic parameters were estimated on 20 individuals from two Daraina forest fragments to determine the potential utility of this marker suite for future studies on the golden-crowned sifaka.     

The first report of <Emphasis Type="Italic">Spondias</Emphasis> native to Madagascar: <Emphasis Type="Italic">Spondias tefyi</Emphasis>, sp. nov. (Anacardiaceae)

Spondias represents a genus new to Madagascar’s native flora. Like Campnosperma, it is now known from both American and Asian tropics and Madagascar but not from continental Africa. The new species Spondias tefyi is easily distinguished from all of its Asian congeners by having the stamens shorter than the pistil and fruits brown and lenticellate at maturity (vs. greenish, yellow, orange or red, and relatively smooth). The new species is one of several Anacardiaceae whose fruits are eaten by lemurs in the Analavelona Forest, highlighting the importance of conserving this threatened subhumid forest remnant in southern Madagascar.     

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.     

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.     

The subfossil occurrence and paleoecological implications of Macrotarsomys petteri (Rodentia: Nesomyidae) in extreme southeastern Madagascar

Remains of a large-bodied species of endemic nesomyid rodent, Macrotarsomys petteri Goodman and Soarimalala, 2005, were identified from subfossil deposits recovered from Andrahomana Cave in extreme southeastern Madagascar. This recently described extant species was previously only known from a single specimen collected at a site about 450 km northwest of Andrahomana and with distinctly different forest habitat than currently found in the vicinity of the cave. Radiocarbon dating of remains of M. petteri from the cave site indicates that it persisted in the region subsequent to human settlement. Previous reports of a large member of Macrotarsomys from other Quaternary sites in southern Madagascar are almost certainly of M. petteri. It is proposed that this species once had a broad distribution across the southern portion of the island during a more mesic period and subsequent aridification of the region has led to its local extirpation across most of its former range. To cite this article: S.M. Goodman et al., C. R. Palevol 5 (2006).     

Novelties in Adenia (Passifloraceae): Four new species, a new combination, a vegetative key, and diagnostic characters for known Madagascan species

Four new species and one new combination ofAdenia are presented, along with a vegetative key and diagnostic characters of the Madagascan species. Additional notes are provided about unusual specimens and field observations.Adenia kigogoensis from Tanzania is shown to be distinct fromA. stenodactyla (its putative closest relative) andA. digitata based on anther connation and other floral traits. The remaining new taxa are from Madagascar.Adenia litoralis has been observed from one coastal locality in northern Madagascar. It is distinctive in fruit size and leaf form.Adenia metamorpha is the only Madagascan taxon with a narrow cylindrical trunk and large napiform tuber; it is also known from only one locality in Madagascar in the Ankarana Reserve.Adenia mcdadiana is a robust liana with highly reduced glands and leaves that appear to be neotenic compared to its closest putative relative,A. sphaerocarpa. Finally, the position ofA. stylosa has been clarified. This species was once treated asA. firingalavensis var.stylosa, and prior to that asA. epigea var.stylosa, but molecular and morphological data suggest it is separate from these species.     

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.     

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.     

How common is common? Rapidly assessing population size and structure of the frog Mantidactylus betsileanus at a site in east‐central Madagascar

Population‐level data are urgently needed for amphibians in light of the ongoing amphibian extinction crisis. Studies focused on population dynamics are not only important for rare species but also for common species which shape ecosystems to a greater degree than those that are rare. Some of the greatest global amphibian species diversity is found in Madagascar, yet there are few studies on the ecology of frog species on the island. We carried out a mark‐recapture study on the widespread frog Mantidactylus betsileanus (Mantellidae: Mantellinae: Mantidactylus) at two adjacent rainforest sites in east‐central Madagascar to assess its population size and structure. To do so, we validated and implemented an individual identification protocol using photographs of the ventral patterns of frogs and identified individuals with photographic‐matching software. Using this rapid, non‐invasive survey method, we were able to estimate a density of 26 and 28 frogs per 100 m² at each of the two sites sampled. Our results show the rainforests near the village of Andasibe, Madagascar support remarkably high amphibian abundance, helping illustrate the significant ecological role of frogs in this ecosystem. Further, individual frog markings allowed us to develop more precise estimates than traditional survey methods. This study provides a blueprint to augment existing population studies or develop new monitoring programs in Madagascar and beyond.     

Isolation of 22 new Haliaeetus microsatellite loci and their characterization in the critically endangered Madagascar fish‐eagle (Haliaeetus vociferoides) and three other Haliaeetus eagle species

We isolated a total of 22 microsatellite loci from two Haliaeetus species: the Madagascar fish‐eagle (Haliaeetus vociferoides) and the bald eagle (Haliaeetus leucocephalus). Five loci were monomorphic in both the Madagascar fish‐eagle (n = 24–43) and the bald eagle (n = 2–8) but were found to be polymorphic in other Haliaeetus species. Haliaeetus loci have proved useful for investigating gene flow in Haliaeetus and Aquila eagles. Ten loci were polymorphic in the critically endangered Madagascar fish‐eagle and will be used to investigate the genetic population structure and mating system in this species.