No abstract

Highly diverse and so far apparently untouched by emergent diseases, Malagasy frogs nevertheless are threatened by ongoing habitat destruction, making pro-active conservation actions especially important for preserving this unique, pre-decline, amphibian fauna.

The black-spined toad, Duttaphrynus melanostictus, is widespread in South and South-East (SE) Asia, although recent molecular analyses have revealed that it represents a species complex (here called the D. melanostictus complex). Invasive populations of this toad have been detected in Madagascar since, at least, 2014. We here trace the origin of this introduction based on mitochondrial DNA sequences of 340 samples. All 102 specimens from Madagascar have identical sequences pointing to a single introduction event. Their haplotype corresponds to a lineage occurring in Cambodia, China, Laos, Thailand, Vietnam, and some locations of eastern Myanmar and northern Malaysia, here named the SE Asian lineage. Within this lineage, specimens from one location in Cambodia and three locations in Vietnam have the same haplotype as found in Madagascar. This includes Ho Chi Minh City, which has a major seaport and might have been the source for the introduction. Species distribution models suggest that the current range of the Madagascan invasive population is within the bioclimatic space occupied by the SE Asian lineage in its native range. The potential invasion zone in Madagascar is narrower than suggested by models from localities representing the full range of the D. melanostictus complex. Thus, an accurate taxonomy is essential for such inferences, but it remains uncertain if the toad might be able to spread beyond the potential suitable range because (1) knowledge on species-delimitation of the complex is insufficient, and (2) the native range in SE Asia might be influenced by historical biogeography or competition.

The Asian toad, Duttaphrynus melanostictus, was accidentally introduced to Toamasina (Eastern Madagascar) around 2010, and since then has spread at a substantial rate across a larger area. This study documents the expansion of the invasive range of this species, calculates the invasion spread rate, and it further estimates the toad abundance and habitat preferences. Updates of the distribution range revealed a fivefold increase of the invaded area during three years, and a doubling of the rate of spread, showing a shift of the invasion towards the North-West, most probably because of the absence of ecological barriers. We used N-mixture models to estimate toad's abundance on the basis of repeated count data of six areas in Toamasina and its surrounding countryside. Toad distribution shows heterogeneous density across the distribution range, with an average abundance of 184 toads ha -1 (95% CI, 132-263). The toad's abundance was highest in sites with the presence of organic waste, and was negatively related to the density of road networks in the proximity of study sites. The rapid expansion of the Asian toad in the Toamasina region suggests that this toad is an increasing threat for Madagascar. We identify immediate management actions that could limit the spread of alien toads in this megadiverse country.

We propose a method of analysing genetic data to obtain separate estimates of the size (N(p)) and migration rate (m(p)) for the sampled populations, without precise prior knowledge of mutation rates at each locus ( micro(L)). The effects of migration and mutation can be distinguished because high migration has the effect of reducing genetic differentiation across all loci, whereas a high mutation rate will only affect the locus in question. The method also takes account of any differences between the spectra of immigrant alleles and of new mutant alleles. If the genetic data come from a range of population sizes, and the loci have a range of mutation rates, it is possible to estimate the relative sizes of the different N(p) values, and likewise the m(p) and the micro(L). Microsatellite loci may also be particularly appropriate because loci with a high mutation rate can reach mutation-drift-migration equilibrium more quickly, and because the spectra of mutants arriving in a population can be particularly distinct from the immigrants. We demonstrate this principle using a microsatellite data set from Mauritian skinks. The method identifies low gene flow between a putative new species and populations of its sister species, whereas the differentiation of two other populations is attributed to small population size. These distinct interpretations were not readily apparent from conventional measures of genetic differentiation and gene diversity. When the method is evaluated using simulated data sets, it correctly distinguishes low gene flow from small population size. Loci that are not at mutation-migration-drift equilibrium can distort the parameter estimates slightly. We discuss strategies for detecting and overcoming this effect.