Malagasy Species Research Articles

Révision du genre Deparia Hook. & Grev. (Pteridophyta, Athyriaceae) à Madagascar: trois espèces nouvelles, une synonymie nouvelle et des hybrides présumés

Rakotondrainibe, F. & A. Jouy (2016). Revision of the genus Deparia Hook. & Grev. (Pteridophyta, Athyriaceae) from Madagascar: three new species, a new synonymy and putative hybrids. Candollea 71: 357–371. In French, French and English abstracts.The first two revisions of the genus Deparia Hook. & Grev. for Madagascar (Dryopteris Adans. sensu Christensen in 1932 and Cornopteris Nakai sensu Tardieu-Blot in 1958) were based on a limited number of specimens. Errors of identification of collections at the level of genus (between Deparia and Diplazium Sw.) and species were numerous. In the process of updating the pteridophyte flora of Madagascar, a new revision of the Malagasy species of Deparia is presented. Six species are recognized, five of which are endemic and one, Deparia boryana (Willd.) M. Kato, is paleotropical. Three species, Deparia florensiae Rakotondr., Deparia longipilosa Rakotondr., and Deparia septentrionalis Rakotondr., are newly described. Deparia forsythii majoris (C. Chr.) M. Kato is placed in synonymy under Deparia parvisora (C. Chr.) M. Kato and putative hybrids are detected. An identification key to the Malagasy species of Deparia is provided, along with illustrations and complete descriptions for each of the species.

Taxonomic revision and distribution of herbaceous Paramollugo (Molluginaceae) in the Eastern Hemisphere.

The genus Paramollugo with the type species Paramollugo nudicaulis (≡Mollugo nudicaulis) has recently been described after molecular investigations. Here we report two new endemic Malagasy species: Paramollugo simulans and Paramollugo elliotii, and transfer a forgotten New Caledonian endemic Mollugo digyna to Paramollugo (Paramollugo digyna). Consequently, the number of Paramollugo species in the Eastern Hemisphere is increased from three to six. Almost all genus representatives (except Paramollugo nudicaulis, which has a wide distribution in Southern Asia, Arabia and tropical Africa) are endemic to Madagascar, Somalia, or New Caledonia. Since the type of seed coat ornamentation is crucial for species delimitation, a diagnostic key with new taxonomically significant carpological characters and other new traits is provided for all the herbaceous Paramollugo. The distribution patterns of Paramollugo nudicaulis s.str., Paramollugo simulans and Paramollugo elliotii are presented.

A new xerophytic species of Mimosa (Mimosoideae, Leguminosae) from Madagascar

A new Malagasy species, Mimosa dupuyana, is described and illustrated. This species is closely related to M. delicatula, but they can be distinguished by the leaf formula, leaf morphology and pods. M. dupuyana has leaves with broad, flattened, and dorsiventrally compressed petioles and only 2–3 leaflet pairs, while M. delicatula has narrow, terete to subteretepetioles and 5–9 leaflet pairs. In addition, the pods of Mimosa dupuyana are constricted between seeds, coriaceous, reddish and elevated on each seeds. The latter species is quite different to the remainder known Malagasy species of Mimosa, by combination of some or all of these characters: leaf-formula, habit, form of petioles, and form and armament of pods. Together, M. delicatula and M. dupuyana resemble some lineages of South American section Batoucalon series Farinosae, which occur in seasonally dry subtropical formations, and this could be an interesting example of convergent evolution. Illustrations, keys for identification and taxonomical, distribution and ecological comments are included.

A Reconsideration of the Infrageneric Classification of Homalium Jacq. (Salicaceae)

Abstract Applequist, W.L. (2016). A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae). Candollea 71: 231–256. In English, English and French abstracts. Homalium Jacq. (Salicaceae) is widespread in tropical regions and includes considerable diversity in floral morphology. Its modern infrageneric classification is complex, including two subgenera and nine sections. Morphological variation within the genus is described, and an emended classification including ten sections is proposed. Subgenera have been defined by stamen number : subg. Homalium has stamens in fascicles of two to 12, while subg. Blackwellia (Benth.) Warb. has one stamen per petal. These subgenera are not natural groups; since the relationships of some sections are uncertain, recognition of subgenera may not be of value and is not upheld here. Sect. Eumyriantheia Warb. is polyphyletic; it is herein restricted to Malagasy species, while sect. Polyanthera Warb. is revived for fasciculate-stamened species from Asi...

Molecular phylogeny, biogeography and chromosome evolution of Malagasy dwarf geckos of the genus Lygodactylus (Squamata, Gekkonidae)

We performed a phylogenetic analysis using nuclear (RAG‐1, RAG‐2) and mitochondrial (16S) markers, a statistical Bayesian reconstruction of ancestral distribution areas and a karyological analysis on most Malagasy species of the gekkonid genus Lygodactylus. The phylogenetic analysis largely confirms major basal branching pattern of previous molecular studies, but highlights significant differences concerning both the relationships between different species groups as well as those within groups. The biogeographic analysis supports a Malagasy origin of Lygodactylus, an oversea dispersal to continental Africa and a return to Madagascar. The L. madagascariensis group (also including a new candidate species identified herein) is the most basal clade in Lygodactylus, and the sister group of a clade with all the remaining species. The second most basal clade is the L. verticillatus group, placed as the sister group of a clade comprising African and Malagasy species. The sister lineage of the L. verticillatus group originated the African radiation through an oversea dispersal out of Madagascar. Eventually, the sister lineage of the L. capensis group originated secondary dispersals from Africa to Madagascar. In Madagascar, lineage diversification in different species groups mainly occurred from southern to northern and eastern regions. Dispersal, vicariance and paleoclimatic refugia probably played a relevant role in the evolutionary history of closely related taxa and in speciation mechanisms. The cytogenetic analysis evidenced a high karyotypic variability in Lygodactylus (from 2n = 34 to 2n = 40), which is at least partly consistent with the phylogenetic relationships and the composition of the various species group. Chromosome evolution occurred independently in different lineages, mainly through a reduction in the chromosome number and starting from a putative primitive karyotype of 2n = 40 with all telocentric elements.

DNA Barcoding of Malagasy Rosewoods: Towards a Molecular Identification of CITES-Listed Dalbergia Species.

Illegal selective logging of tropical timber is of increasing concern worldwide. Madagascar is a biodiversity hotspot and home to some of the world’s most sought after tropical timber species. Malagasy rosewoods belong to the genus Dalbergia (Fabaceae), which is highly diverse and has a pantropical distribution, but these timber species are among the most threatened as a consequence of intensive illegal selective logging and deforestation. Reliable identification of Dalbergia species from Madagascar is important for law enforcement but is almost impossible without fertile plant material, which is often unavailable during forest inventories or when attempting to identify logged trees of cut wood. DNA barcoding has been promoted as a promising tool for species identification in such cases. In this study we tested whether DNA barcoding with partial sequences of three plastid markers (matK, rbcL and trnL (UAA)) can distinguish between Dalbergia from Madagascar and from other areas of its distributional range, and whether Malagasy species can be distinguished from one another. Phylogenetic analyses revealed that the Malagasy Dalbergia species studied form two monophyletic groups, each containing two subgroups, only one of which corresponds to a single species. We characterized diagnostic polymorphisms in the three DNA barcoding markers that allow rapid discrimination between Dalbergia from Madagascar and from other areas of its distribution range. Species identification success based on individual barcoding markers or combinations was poor, whereas subgroup identification success was much higher (up to 98%), revealing both the value and limitations of a DNA barcoding approach for the identification of closely related Malagasy rosewoods.

Long-distance dispersal and inter-island colonization across the western Malagasy Region explain diversification in brush-warblers (Passeriformes:Nesillas)

The present study examines the colonization history and phylogeography of the brush-warblers (Nesillas), a genus of passerines endemic to islands of the western Indian Ocean (Madagascar, Comoros, and Aldabra Atoll). The phylogeny of all recognized Nesillas taxa was reconstructed employing Bayesian phylogenetic methods and divergence times were estimated using a range of substitution rates and clock assumptions. Spatiotemporal patterns of population expansion were inferred and niches of different lineages were compared using ecological niche modelling. Our results indicate that taxa endemic to the Comoros are paraphyletic and that the two endemic species on Madagascar (Nesillas typica and Nesillas lantzii) are not sister taxa. The brush-warblers started to diversify approximately 1.6 Mya, commencing with the separation of the clade formed by two species endemic to the Comoros (Nesillas brevicaudata and Nesillas mariae) from the rest of the genus. The lineages leading to the two Malagasy species diverged approximately 0.9 Mya; each with significantly different modern ecological niches and the subject of separate demographic processes. Patterns of diversification and endemism in Nesillas were shaped by multiple long distance dispersal events and inter-island colonization, a recurring pattern for different lineages on western Indian Ocean islands. The diversification dynamics observed for Nesillas are also consistent with the taxon cycle hypothesis.

A synoptic revision of the Malagasy species ofScolopiaSchreb. (Salicaceae, Scolopieae)

ABSTRACT A synoptic revision of the Malagasy species of Scolopia is presented. Fourteen species are recognized, of which all are endemic and ten are threatened. One species, S. delphinensis Appleq. & G.E.Schatz, sp. nov., is newly described and illustrated. An identification key to the Malagasy species of Scolopia is provided.

Karyotypic Evolution in Malagasy Flying Foxes (Pteropodidae, Chiroptera) and Their Hipposiderid Relatives as Determined by Comparative Chromosome Painting

Pteropodidae and Hipposideridae are 2 of the 9 chiropteran families that occur on Madagascar. Despite major advancements in the systematic study of the island's bat fauna, few karyotypic data exist for endemic species. We utilized G- and C-banding in combination with chromosome painting with Myotismyotis probes to establish a genome-wide homology among Malagasy species belonging to the families Pteropodidae (Pteropus rufus 2n = 38; Rousettus madagascariensis, 2n = 36), Hipposideridae (Hipposideros commersoni s.s., 2n = 52), and a single South African representative of the Rhinolophidae (Rhinolophus clivosus, 2n = 58). Painting probes of M. myotis detected 26, 28, 28, and 29 regions of homology in R. madagascariensis, P. rufus, H. commersoni s.s, and R. clivosus, respectively. Translocations, pericentric inversions, and heterochromatin additions were responsible for karyotypic differences amongst the Malagasy pteropodids. Comparative chromosome painting revealed a novel pericentric inversion on P. rufus chromosome 4. Chromosomal characters suggest a close evolutionary relationship between Rousettus and Pteropus. H. commersoni s.s. shared several chromosomal characters with extralimital congeners but did not exhibit 2 chromosomal synapomorphies proposed for Hipposideridae. This study provides further insight into the ancestral karyotypes of pteropodid and hipposiderid bats and corroborates certain molecular phylogenetic hypotheses.

Une espèce nouvelle et une synonymie nouvelle dans le genre Selaginella P. Beauv. (Lycopodiophyta, Selaginellaceae) à Madagascar

Rakotondrainibe, F. & A. Jouy (2016). A new species and a new synonymy in the genus Selaginella P. Beauv. (Lycopodiophyta, Selaginellaceae) from Madagascar. Candollea 71: 143–148. In French, English and French abstracts. DOI : http//dx.doi.org/10.15553/c2016v711a17A new Malagasy species of Selaginella P. Beauv. (Lycopodiophyta, Selaginellaceae) is described and a new synonymy in the same genus is established. The new species Selaginella rasoloheryi Rakotondr. belongs to the subgenus Heterostachys Baker; it differs from all other Africano-Malagasy species of this subgenus by the main stem being prostrated throughout its entire length, without soboles, with rhizophores and ramifications unbranched, and with median leaves strongly acuminate. Selaginella rasolohery has a close affinity to Selaginella ciliaris (Retz.) Spring, an asiatic and oceanic species, but differs from it by more pronounced leaf dimorphism in both vegetative and fertile shoots, median leaves longer acuminate, strobili less compact and lateral sporophyll carina narrow and dentate (vs. carina large and ciliate in Selaginella ciliaris). The new synonymy between Selaginella fissidentoides (Hook. & Grev.) Spring f.fissidentoides and Selaginella fissidentoides f. ovata Stefanović & Rakotondr. is established after observation of numerous Malagasy specimens recently collected between 1997 and 2015 after the publication of the first Selaginella revision for Madagascar and the Comoro Islands.

New names for two Malagasy species of Canarium L. (Burseraceae)

Daly, D.C., J. Raharimampionona & S. Federman (2016). New names for two Malagasy species of Canarium L. (Burseraceae). Candollea 71 : 159–160. In English, English and French abstracts. DOI : http//dx.doi.org/10.15553/c2016v711a19New names are provided for two recently described Malagasy species of Canarium L. (Burseraceae) for which the epithets proved to be later homonyms of already published taxa. These new names are Canarium fugax Daly, Raharim. & Federman, and Canarium lobocarpum Daly, Raharim. & Federman.

Phylogenetic Reconstruction of the Myrrh Genus, Commiphora (Burseraceae), Reveals Multiple Radiations in Madagascar and Clarifies Infrageneric Relationships

Abstract Commiphora (Burseraceae) is the most species-rich genus in the frankincense and myrrh family, whose widespread and often remote distribution in seasonally dry tropical forests globally has impeded efforts to resolve its taxonomy and investigate its systematic biology. Here we focus on establishing the origin and evolution of Commiphora species native to Madagascar, all of which are endemic. Recent work has uncovered 16 new Malagasy species, bringing the total number of island endemic species to 44 and comprising nearly 25% of the genus. Previous phylogenetic studies of Commiphora have indicated that Malagasy lineages immigrated to and radiated within the island twice. We test this biogeographic hypothesis more thoroughly using a nearly comprehensive sampling of species from Madagascar, an expanded sampling of Commiphora species native to other regions, and a greater depth of DNA sequence data from the nuclear (nrETS, nrITS) and chloroplast (psbA-trnH, ndhF-rpl32, and trnD—trnT) genomes. Results f...

A Revision of the Malagasy Species of Homalium Sect. Eumyriantheia Warb. (Salicaceae)

Applequist, W.L. (2016). A revision of the Malagasy Species of Homalium Sect. Eumyriantheia Warb. (Salicaceae). Candollea 71: 33–60. In English, English and French abstracts. DOI : http//dx.doi.org/10.15553/c2016v711a7Homalium sect. Eumyriantheia Warb. (Salicaceae) is the largest and most widespread fasciculate-stamened section of Homalium Jacq. Though the section is markedly heterogeneous, the Malagasy species appear to form a natural group. A new revisionary treatment of these species is presented and an identification key is provided.Fifteen species are recognized, of which five, i.e. Homalium dorrii Appleq., Homalium pseudoboinense Appleq., Homalium randrianasoloi Appleq., Homalium ranomafanicum Appleq., and Homalium schatzii Appleq., are newly described. Occasional hybridization is observed. Most species are endangered.

No longer single! Description of female Calumma vatosoa (Squamata, Chamaeleonidae) including a review of the species and its systematic position

Calumma vatosoa is a Malagasy chameleon species that has until now been known only from the male holotype and a photograph of an additional male specimen. In this paper we describe females of the chameleon Calumma vatosoa for the first time, as well as the skull osteology of this species. The analysed females were collected many years before the description of C. vatosoa, and were originally described as female C. linotum. According to external morphology, osteology, and distribution these specimens are assigned to C. vatosoa. Furthermore we discuss the species group assignment of C. vatosoa and transfer it from the C. furcifer group to the C. nasutum group. A comparison of the external morphology of species of both groups revealed that C. vatosoa has a relatively shorter distance from the anterior margin of the orbit to the snout tip, more heterogeneous scalation at the lower arm, a significantly lower number of supralabial and infralabial scales, and a relatively longer tail than the members of the C. furcifer group. These characters are, however, in line with the species of the C. nasutum group. In addition the systematic position of C. peyrierasi also discussed, based on its morphology.

Tick (Amblyomma chabaudi) infestation of endemic tortoises in southwest Madagascar and investigation of tick-borne pathogens

Little is known about the role of endemic ticks as vectors for bacterial and protozoan pathogens for animals and humans in Madagascar and their interaction in anthropogenic habitats where humans, their livestock and native Malagasy species (vectors and hosts) come into more frequent contact than in natural forest ecosystems. The aims of the study were (1) to test whether habitat degradation is associated with increased infestation of tortoises by ticks and (2) to investigate whether ticks carried Babesia, Borrelia or Rickettsia species that might be pathogenic for humans and livestock. We studied hard ticks of two endemic Malagasy tortoises, Astrochelys radiata and Pyxis arachnoides in March and April 2013 in southwest Madagascar. Two tortoise habitats were compared, the National Park of Tsimanampetsotsa and the adjacent degraded pasture and agricultural land at the end of the wet season. Ticks were screened for protozoan and bacterial pathogens via PCR on DNA isolated from ticks using genus-specific primers. Only one out of 42 A. radiata collected from both habitats had ticks. The low prevalence did not allow further analyses of the effect of habitat degradation. Forty-two P. arachnoides were found in the anthropogenic habitat and 36 individuals in the national park. Tick infestation rates of P. arachnoides differed significantly between the two study sites. Tortoises inside the park had lower tick prevalence than outside (8 of 36 (22%) versus 32 of 42 individuals (76%)) and infected animals tended to have fewer ticks inside than outside the park. All ticks collected in both habitats were adults of the ixodid tick Amblyomma chabaudi, which is supposed to be a host-specific tick of P. arachnoides. Screening for Borrelia sp. and Babesia sp. was negative in all ticks. But all A. chabaudi ticks were infected with Rickettsia africae, known to cause spotted fever in humans. Thus, habitat degradation seems to be linked to higher infestation of tortoises with ticks with possible consequences for humans and their livestock.

The Transfer of the Malagasy Species of Strobilanthes Blume to Acanthopale C. B. Clarke (Acanthaceae)

Callmander, M. W. & P. B. Phillipson (2015). The transfer of the Malagasy species of Strobilanthes Blume to Acanthopale C. B. Clarke (Acanthaceae). Candollea 70: 253–256. In English, English and French abstracts. DOI: http//dx.doi.org/10.15553/c2015v702a12Five Acanthaceae species originally described in the genus Strobilanthes Blume are formally transferred to the genus Acanthopale C. B. Clarke, following currently accepted generic delimitation in the family. The necessary combinations: Acanthopale breviceps (Benoist) Callm. & Phillipson, Acanthopale cuneifolia (Benoist) Callm. & Phillipson, Acanthopale humblotii (Benoist) Callm. & Phillipson, Acanthopale perrieri (Benoist) Callm. & Phillipson and Acanthopale ramiflora (Benoist) Callm. & Phillipson are established.

Molecular systematics and undescribed diversity of Madagascan scolecophidian snakes (Squamata: Serpentes).

We provide an updated molecular phylogenetic analysis of global diversity of typhlopid and xenotyphlopid blindsnakes, adding a set of Madagascan samples and sequences of an additional mitochondrial gene to an existing supermatrix of nuclear and mitochondrial gene segments. Our data suggest monophyly of Madagascan typhlopids, exclusive of introduced Indotyphlops braminus. The Madagascar-endemic typhlopid clade includes two species previously assigned to the genus Lemuriatyphlops (in the subfamily Asiatyphlopinae), which were not each others closest relatives. This contradicts a previous study that described Lemuriatyphlops based on a sequence of the cytochrome oxidase subunit 1 gene from a single species and found this species not forming a clade with the other Malagasy species included. Based on our novel phylogenetic assessment we include all species in this endemic typhlopid clade in the genus Madatyphlops and in the subfamily Madatyphlopinae and consider Lemuriatyphlops as junior synonym. Within Madatyphlops, we identify several candidate species. For some of these (those in the M. arenarius complex), our preliminary data suggest sympatric occurrence and morphological differentiation, thus the existence of undescribed species. We also comment on the genus-level classification of several non-Madagascan typhlopids. We suggest that African species included in Madatyphlops (Afrotyphlops calabresii, A. cuneirostris, A. platyrhynchus, and Rhinotyphlops leucocephalus) should not be included in this genus. We furthermore argue that recent claims of Sundatyphlops, Antillotyphlops, and Cubatyphlops being "undiagnosable" or "not monophyletic" were based on errors in tree reconstruction and failure to notice diagnostic characters, and thus regard these three genera as valid.

Bird Assemblages in a Malagasy Forest-Agricultural Frontier: Effects of Habitat Structure and Forest Cover

Increasing global human population and per-capita food consumption are expected to exacerbate the already massive agricultural footprint in tropical ecosystems. Madagascar is home to exceptional levels of biodiversity and is in the midst of severe land-use change, mostly driven by slash-and-burn, smallholder agriculture. Understanding the consequences of these agricultural practices for Malagasy native species is therefore of the foremost importance for the conservation of the nation's biodiversity. We surveyed bird assemblages inside and surrounding Ranomafana National Park, southeastern Madagascar, obtaining nearly 1,000 records of more than 60 species. At each study point, habitat structure was characterized by its vegetation complexity, and forest cover was quantified within circles of radii of 100, 500 and 750 m. We found that species richness was higher in forest than in agricultural areas, and responses to land-use change were found to be guild-specific, with frugivores being especially depleted outside forest areas, whereas granivores had higher species richness in the agricultural matrix. The number of recorded species with forest affinities was highly associated with landscape-scale forest cover, while open area and generalist species responded mainly to site-scale habitat structure. Our results demonstrate a turnover from forest-associated species to open area and habitat generalist species in Madagascar's smallholder agricultural areas. Our study underscores the conservation value of landscape-scale forest cover and of site-scale vegetation complexity. A double-stranded conservation approach, in which both landscape-scale forest cover and vegetation complexity are preserved would benefit conservation of the island's forest avifauna.

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.

Catharanthus makayensis L. Allorge, Phillipson & Razakamal. (Apocynaceae), a New Species from Madagascar

Allorge, L., P. B. Phillipson & R. Razakamalala (2015). Catharanthus makayensis L. Allorge, Phillipson & Razakamal. (Apocynaceae), a new species from Madagascar. Candollea 70: 61–66. In English, English and French abstracts.Catharanthus makayensis L. Allorge, Phillipson & Razakamal. (Apocynaceae), a species new to science from Madagascar is described and illustrated. It differs from all other species in the genus by the hexagonal cross-section of its stems, and by a combination of flowering and fruiting characters. Its 5–6 mm long filiform sepals are clearly much narrower and longer than all other species with the exception of Catharanthus coriaceus Markgr., but the latter species is highly distinct with its pendulous fruits, as opposed to erect fruits of Catharanthus makayensis and all other species of the genus. Catharanthus makayensis and Catharanthus roseus (L.) G. Don are the only species to have a magenta, rather than white to pale yellow eye. The corolla lobes and the flower buds of Catharanthus makayensis are pure white or tinted pale pink whereas Catharanthus roseus typically has pink lobes. The relationships of the new species with the eight other species of Catharanthus G. Don are discussed and an identification key to the Malagasy species of the genus is provided.