Karyological evolution and systematics of Malagasy microhylid frogs

Microhylid frogs are a group of largely unresolved phylogeny, and diverse data sets are needed to improve the evolutionary understanding of these amphibians. We here report karyological data for 22 species of this family, belonging to the Malagasy genera Anodonthyla, Cophyla, Platypelis, Plethodontohyla, Rhombophryne, and Stumpffia (Cophylinae); Scaphiophryne and Paradoxophyla (Scaphiophryninae); and Dyscophus (Dyscophinae); and the Asian genera Calluella and Ramanella (Microhylinae). All species studied have 2n=26 chromosomes, most of which are metacentric or submetacentric. Chromosome morphology, banding pattern, and position of the nucleolar organizer regions (NORs) provide relevant characters for the understanding of the phylogeny and systematics of these frogs. The species of the Cophylinae are characterized by a subtelocentric or telocentric fourth chromosome pair (submetacentric only in Anodonthyla), which can be seen as a synapomorphy for this subfamily. Shifts in NOR position within the Cophylinae are frequent and agree with recent mitochondrial DNA data, corroborating the non-monophyly of the genus Plethodontohyla. Changes of NOR position and chromosome morphology (i.e., occurrence of subtelocentric and telocentric elements) were also common in this subfamily, possibly being related to their faster mitochondrial substitution rate and high species diversity. The ninth chromosome pair of the examined specimens of Dyscophus guineti, all juveniles, is heteromorphic. In this pair, one of the two chromosomes is longer due to the addition of two heterochromatic segments, raising the possibility that one chromosome of this pair may be a sex chromosome.     

Diversification rates have declined in the Malagasy herpetofauna

The evolutionary origins of Madagascar''s biodiversity remain mysterious despite the fact that relative to land area, there is no other place with consistently high levels of species richness and endemism across a range of taxonomic levels. Most efforts to explain diversification on the island have focused on geographical models of speciation, but recent studies have begun to address the island''s accumulation of species through time, although with conflicting results. Prevailing hypotheses for diversification on the island involve either constant diversification rates or scenarios where rates decline through time. Using relative-time-calibrated phylogenies for seven endemic vertebrate clades and a model-fitting framework, I find evidence that diversification rates have declined through time on Madagascar. I show that diversification rates have clearly declined throughout the history of each clade, and models invoking diversity-dependent reductions to diversification rates best explain the diversification histories for each clade. These results are consistent with the ecological theory of adaptive radiation, and, coupled with ancillary observations about ecomorphological and life-history evolution, strongly suggest that adaptive radiation was an important formative process for one of the most species-rich regions on the Earth. These results cast the Malagasy biota in a new light and provide macroevolutionary justification for conservation initiatives.     

Early Holocene chronology and environment of Ampasambazimba,A Malagasy subfossil lemur site

According to a traditional but hitherto untested reconstruction of Madagascar’s Holocene environment, continuous forest preceded the monotonous grassland formations that now cover most of the island’s interior. Preliminary analyses of pollen samples collected near¹⁴C-dated horizons at Ampasambazimba (central Madagascar) indicate that a mosaic of woodlands, bushlands, and savanna existed close to this important vertebrate subfossil site around 7000–8000 BP. Although most members of Madagascar’s recently extinct “subfossil” fauna are thought to have been forest dwellers, several may have preferred a more open habitat like the one inferred for the region of Ampasambazimba. Dry savanna-woodland burns more readily than does dense forest and may have been more severely affected by the forces which transformed the Malagasy environment during the later Holocene. If so, the loss of savanna-woodland, not dense forest, may have been an important factor in the subfossil extinctions.     

Mitochondrial evidence for distinct phylogeographic units in the endangered Malagasy poison frog Mantella bernhardi

Mantella bernhardi is an endemic species of Malagasy poison frog threatened by loss and fragmentation of its natural habitat and collection for the pet trade. It is classified as threatened according to the International Union for Conservation of Nature and Natural Resources (IUCN) categories and included in Appendix II of the Convention on the International Trade of Endangered Species (CITES). A recent survey has increased the known distributional range of the species from one to eight populations across southeastern Madagascar, but little is known about its biology and genetic diversity. Here we estimate inter- and intrapopulation mitochondrial genetic variation of four populations. Populations from the northern and southern parts of the distributional range showed a high degree of divergence (maximum of 11.35% in cytochrome b) and were recovered as reciprocally monophyletic groups. Nine haplotypes were detected in the northern and 12 in the southern populations. The population from Ranomafana National Park showed the lowest number of haplotypes and nucleotide diversity, and shared its most common haplotype with the second northern population from Tolongoina. All the other detected haplotypes were unique to each of the four populations. This suggests the existence of important barriers to gene flow, pre-dating human colonization of Madagascar at about 2000 years ago, in distinct contrast to other Mantella species that show a high degree of haplotype sharing throughout their range. The continued habitat fragmentation within the distribution range of M. bernhardi prevents any connection between its populations. Our data indicate the existence of at least two different management units for conservation in this species, corresponding to the North and South of its distribution range, and highlight the existence of strong regional endemism in southeastern Madagascar.     

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.     

Ontogenetic correlates of diet in Malagasy lemurs

There is a well-documented relationship between development and other life-history parameters among anthropoid primates. Smaller-bodied anthropoids tend to mature more rapidly than do larger-bodied species. Among anthropoids of similar body sizes, folivorous species tend to grow and mature more quickly than do frugivorous species, thus attaining adult body size at an earlier age. This pattern conforms to the expectations of Janson and van Schaik's "ecological risk aversion hypothesis," which predicts that rates of growth and maturation should vary in inverse relation to the intensity of intraspecific feeding competition. According to the ecological risk aversion hypothesis (RAH), species experiencing high intraspecific feeding competition will grow and mature slowly to reduce the risk of mortality due to food shortages. Species experiencing low levels of intraspecific feeding competition will shorten the juvenile period to reduce the overall duration of this high-risk portion of the life cycle. This paper focuses on development and maturation in lemurs. We show that folivorous lemurs (such as indriids) grow and mature more slowly than like-sized frugivorous lemurs (e.g., most lemurids), but tend to exhibit faster dental development. Their dental developmental schedules are accelerated on an absolute scale, relative to craniofacial growth, and relative to particular life-history landmarks, such as weaning. Dental development has a strong phylogenetic component: even those lemurids that consume substantial amounts of foliage have slower dental development than those indriids that consume substantial amounts of fruit. Implications of these results for the RAH are discussed, and an explanation for this hypothesis' failure to predict lemur growth schedules is offered. We propose that the differing developmental schedules of folivorous and frugivorous lemurs may reflect different solutions to the ecological problem of environmental instability: some rely on a strategy of low maternal input and slow returns, while others rely on a strategy of high maternal input and fast returns.     

Core and Satellite Species in Degraded Habitats: an Analysis Using Malagasy Tree Communities

Core-satellite theory predicts that, via the “rescue effect”, widespread, abundant species should have reduced risk of local extinctions. We test this hypothesis in southeastern Malagasy littoral forest using data on distribution and abundance of trees and woody understory vegetation in tropical forest fragments along a disturbance gradient. We partition the mortality risk into two kinds of extinction factors, separately operating at demographic (local) and landscape (regional) scales, contrary to core-satellite predictions, for both trees and woody understory vegetation, that the relative number of core (abundant) species declined significantly with increasing disturbance. In the least-degraded forest fragments there was a strong mode of core species, while in the moderately- and severely-degraded fragments the species distributions were essentially log-normal, lacking a substantial core mode. While the rescue effect mitigates one kind of extinction risk, namely local environmental and demographic stochasticity, it may not counterbalance widespread pervasive sources of mortality. The amount of internal forest fragmentation appears to have a much greater effect on species richness and diversity than either fragment size or shape.     

Hawk-moth scale analysis and pollination specialization in the epilithic Malagasy endemic Aerangis ellisii (Reichenb. fil.) Schltr. (Orchidaceae)

The pollination biology of the epilithic endemic Aerangis ellisii (Reichenb. fil.) Schltr. (Orchidaceae) was studied on an inselberg in the Antananarivo region, central Madagascar. The flowers exhibit long nectariferous spurs indicative of hawk-moth pollination, the mechanism evidently involving pollinia transfer via the frons and palpi of the vectors. Analysis of hawk-moth scales on naturally pollinated stigmas showed that the principal pollinators were Agrius conuolvuli (Linnaeus) and Panogena lingens (Butler). Proboscis morphology of long-tongued Malagasy Sphingidae in relation to nectar position and spur morphology in A. ellisii also indicated that A. conuolvuli and P. lingens were best suited to interact with the plant's floral adaptation. Aerangis ellisii seems to display a relatively moderate specialization versus the Malagasy hawk-moth guild since its nectar is accessible even to those long-proboscis hawk-moth species which are not able to act as pollinators.     

Sociality in a Malagasy allodapine bee, Macrogalea antanosy, and the impacts of the facultative social parasite, Macrogalea maizina

Social parasitism has been researched extensively in many taxa of social insects, including ants, wasps and bees. However, little research has been done on allodapine bees, a taxon that has numerous independent origins of social parasitism. This study looks at two species of Macrogalea from Madagascar, one of which was previously believed to be a social parasite. Macrogalea is an important genus to study as it is the sister clade to all other allodapine genera, and the species of Macrogalea in Madagascar diverged recently, meaning that the study of a social parasite in this genera would provide insights into the very early stages of social parasite evolution. Macrogalea maizina was determined to be facultatively parasitic based on the presence of many traits that are common to other allodapine social parasites. The host, Macrogalea antanosy, was found to be quasisocial, with most females within a colony being able to reproduce. This has unique consequences for a parasitic strategy, as any invading parasite has no need to remove a queen or suppress the reproduction of the other colony members, a strategy that has been commonly observed for facultative parasites in other taxa. Received 10 May 2005; revised 22 July 2005; accepted 24 August 2005.     

Exploring the potential of life-history key innovation: brook breeding in the radiation of the Malagasy treefrog genus Boophis

The treefrog genus Boophis is one of the most species-rich endemic amphibian groups of Madagascar. It consists of species specialized to breeding in brooks (48 species) and ponds (10 species). We reconstructed the phylogeny of Boophis using 16S ribosomal DNA sequences (558 bp) from 27 species. Brook-breeders were monophyletic and probably derived from an ancestral pond-breeding lineage. Pond-breeders were paraphyletic. The disparity in diversification among pond-breeders and brook-breeders was notable among endemic Malagasy frogs, although it was not significant when considering Boophis alone. Sibling species which have different advertisement calls but are virtually indistinguishable by morphology were common among brook-breeders; genetic divergence between these species was high (modal 8% total pairwise divergence). Substitution rates in brook-breeding species were significantly higher than in pond-breeders. Speciation of pond-breeders may be hindered by their usually more synchronous reproduction and a higher vagility which enhances gene flow, while a higher potential of spatial segregation and speciation may exist along brooks.     

Molecular phylogeny and biogeography of Malagasy frogs of the genus Gephyromantis

The genus Gephyromantis is a clade within the Malagasy-Comoroan family Mantellidae composed of rainforest frogs that live and breed to varying degrees independently from water. Based on DNA sequences of five mitochondrial and five nuclear genes we inferred the phylogeny of these frogs with full taxon coverage at the species level. Our preferred consensus tree from a partitioned Bayesian analysis of 5843 base pairs of 51 nominal and candidate species supports various major clades within the genus although the basal relationships among these remain unresolved. The data provide strong evidence for the monophyly of the subgenera Gephyromantis (after exclusion of Gephyromantis klemmeri), Laurentomantis, Vatomantis, and Phylacomantis. Species assigned to the subgenus Duboimantis belong to two strongly supported clades of uncertain relationships. G. klemmeri, previously in the subgenus Gephyromantis, was placed with high support sister to the Laurentomantis clade, and the Laurentomantis + G. klemmeri clade was sister to Vatomantis. A reconstruction of ancestral distribution areas indicates a diversification of several subgenera in the northern biogeographic regions of Madagascar and the dispersal out of northern Madagascar for several clades.     

Improved recruitment of a lemur-dispersed tree in Malagasy dry forests after the demise of vertebrates in forest fragments

The objective of this study was to examine how the processes of seed dispersal and seed predation were altered in forest fragments of the dry forest of Madagascar, where the usual seed dispersers and vertebrate seed predators were absent, using a lemur-dispersed tree species (Strychnos madagascariensis; Loganiaceae) as an example. We then assessed how the changes in vertebrate community composition alter the regeneration pattern and establishment of this tree species and thus, ultimately, the species composition of the forest fragments. By using size-selective exclosures, data from forest fragments were compared with results from continuous forest where vertebrate dispersers and predators were abundant. Visits to the exclosures by mammalian seed predators were monitored with hair traps. In the continuous forest up to 100% of the seeds were removed within the 7 days of the experiments. A substantial proportion of them was lost to seed predation by native rodents. In contrast, practically no predation took place in the forest fragments and almost all seeds removed were dispersed into the safety of ant nests by Aphaenogaster swammerdami, which improves chances of seedling establishment. In congruence with these findings, the abundance of S. madagascariensis in the forest fragments exceeded that of the continuous forest. Thus, the lack of vertebrate seed dispersers in these forest fragments did not lead to a decline in regeneration of this animal-dispersed tree species as would have been expected, but rather was counterbalanced by the concomitant demise of vertebrate seed predators and an increased activity of ants taking over the role of seed dispersers, and possibly even out-doing the original candidates. This study provides an example of a native vertebrate-dispersed species apparently profiting from fragmentation due to flexible animal-plant interactions in different facets, possibly resulting in an impoverished tree species community.     

Revision of the Malagasy genus Trichoteleia Kieffer (Hymenoptera, Platygastroidea, Platygastridae)

The species of the genus Trichoteleia Kieffer (Hymenoptera: Platygastridae) are revised: 42 species are recognized, of which two were previously named and are redescribed: Trichoteleia afo Talamas, sp. n., Trichoteleia albidipes Kieffer, Trichoteleia bicolor Talamas, sp. n.; Trichoteleia bidentata Talamas sp. n.; Trichoteleia carinata Talamas, sp. n.; Trichoteleia cincta Talamas & Masner, sp. n.; Trichoteleia delilah Talamas, sp. n.; Trichoteleia eburata Talamas, sp. n.; Trichoteleia echinata Talamas, sp. n.; Trichoteleia fisheri Talamas & Masner, sp. n.; Trichoteleia funesta Talamas, sp. n.; Trichoteleia halterata Talamas & Masner, sp. n.; Trichoteleia hemlyae Talamas & Masner, sp. n.; Trichoteleia irwini Talamas & Masner, sp. n.; Trichoteleia janus Talamas, sp. n.; Trichoteleia jiro Talamas, sp. n.; T. ketrona Talamas, sp. n.; Trichoteleia levii Talamas & Johnson, sp. n.; Trichoteleia longiventris Talamas & Masner, sp. n.; Trichoteleia minima Talamas, sp. n.; Trichoteleia nify Talamas & Masner, sp. n.; Trichoteleia oculea Talamas, sp. n.; Trichoteleia orona Talamas & Masner, sp. n.; Trichoteleia parvipennis Talamas & Masner, sp. n.; Trichoteleia pauliani (Risbec); Trichoteleia picturata Talamas, sp. n.; Trichoteleia prima Talamas, sp. n.; Trichoteleia prolixa Talamas, sp. n.; Trichoteleia quazii Talamas, sp. n.; Trichoteleia ravaka Talamas, sp. n.; Trichoteleia rugifrons Talamas & Masner, sp. n.; Trichoteleia solocis Talamas, sp. n.; Trichoteleia sphaerica Talamas, sp. n.; Trichoteleia subtilis Talamas & Masner, sp. n.; Trichoteleia tahotra Talamas & Masner, sp. n.; Trichoteleia takariva Talamas, sp. n.; Trichoteleia tezitra Talamas, sp. n.; Trichoteleia tigris Talamas, sp. n.; Trichoteleia tonsa Talamas, sp. n.; Trichoteleia warreni Talamas & Masner, sp. n.; Trichoteleia xantrox Talamas, sp. n.; Trichoteleia zuparkoi Talamas & Masner, sp. n. A neotype is designated for Trichoteleia albidipes and a lectotype is designated for Trichoteleia pauliani.     

Pair-specific usage of sleeping sites and their implications for social organization in a nocturnal Malagasy primate, the Milne Edwards' sportive lemur (Lepilemur edwardsi)

Safe sleeping sites may be a limited resource crucial for survival. In order to investigate their potential significance for social organization in nocturnal primates, we analyzed the spatial distribution of daily sleeping sites, their characteristics, their usage, and sleeping group compositions in the nocturnal Milne Edwards' sportive lemur during a 6-month field study in the dry deciduous forest of northwestern Madagascar. Sexes did not differ either in body size or in body mass. Sleeping sites were used almost exclusively by adult male-female pairs. Individuals showed a high sleeping-site fidelity limited to 2-3 different sleeping sites in close vicinity during the whole study period. Most females showed a higher fidelity to one distinct sleeping site than their male partners. Sleeping groups consisted of one adult male and one adult female and remained stable in composition over the whole study period. Exclusive pair-specific usage of sleeping sites suggests sleeping site related territoriality of male-female pairs, perhaps influenced by inter- and intrasexual resource competition. Results give first insights into the distribution patterns and social organization of this species. They imply dispersed monogamy for the Milne Edwards' sportive lemur, with sleeping sites as a potentially restricted and defendable resource.     

The roles of birds, lizards, and rodents as hosts for the western black-legged tick Ixodes pacificus.

We compared the infestation by ixodid ticks of lizards, rodents, and birds collected simultaneously within areas representing common habitat types in Mendocino County, CA. Lizards were infested only by Ixodes pacificus Cooley and Kohls, birds by I. pacificus and Haemaphysalis leporispalustris (Packard), and rodents by I. pacificus, I. spinipalpis Hadwen and Nuttall, I. woodi Bishopp, Dermacentor occidentalis Marx, and D. variabilis (Say). Infestation by I. pacificus larvae and nymphs of lizards (Sceloporus occidentalis Baird and Girard; Elgaria spp.) and western gray squirrels (Sciurus griseus Ord) (means of 9-35 larvae and 5-6 nymphs per animal) was several times greater than for Neotoma fuscipes Baird woodrats, Peromyscus spp. mice, and birds (means of 0.9-3.5 larvae and 0-0.3 nymphs). Overall, Borrelia-refractory lizards accounted for 84% of I. pacificus larvae and 91% of nymphs collected from animals in dense woodlands. Bird species frequently utilizing tick-questing substrates such as leaf litter (guild I birds) were more heavily infested by I. pacificus subadults (5.2 larvae and 1.0 nymphs per bird) than guild IV birds with minimal perceived contact with tick-questing substrates (0.08 larvae and 0.06 nymphs per bird). Notably, guild I birds carried similar larval loads and at least 20-fold higher nymphal loads relative to woodrats and mice. Only guild IV birds carried as few I. pacificus nymphs as did these rodents. The ratios of larvae to nymphs suggest that, relative to birds, lizards, and squirrels (infested by 1.3-6.0 larvae per nymph), nocturnally active ground-dwelling rodents such as woodrats and mice are underutilized by the nymphal stage (69 to >100 larvae per nymph). The western gray squirrel and guild I-II birds (e.g., the dark-eyed junco, Junco hyemalis [L.]) were the only potential reservoirs of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt, and Brenner (the causative agent of Lyme disease in North America) that were frequently infested with both I. pacificus larvae and nymphs and commonly utilized dense woodland habitats.     

Plants as alternative hosts for Salmonella

Recent findings show that many human pathogenic bacteria can use multiple host organisms. For example, Salmonella Typhimurium can use plants as alternative hosts to humans and other animals. These bacteria are able to adhere to plant surfaces and actively infect the interior of plants. Similarly to the infection of animal cells, S. Typhimurium suppresses plant defense responses by a type III secretion mechanism, indicating that these bacteria possess a dedicated multi-kingdom infection strategy, raising the question of host specificity. In addition, evidence is accumulating that the interaction of Salmonella with plants is an active process with different levels of specificity, because different Salmonella serovars show variations in pathogenicity, and different plant species reveal various levels of resistance towards these bacteria.