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      Resurrection and re-description of Plethodontohyla laevis (Boettger, 1913) and transfer of Rhombophryne alluaudi (Mocquard, 1901) to the genus Plethodontohyla (Amphibia, Microhylidae, Cophylinae)

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          Abstract

          The systematics of the cophyline microhylid frog genera Plethodontohyla and Rhombophryne have long been intertwined, and their relationships have only recently started to become clear. While Rhombophryne has received a lot of recent taxonomic attention, Plethodontohyla has been largely neglected. Our study is a showcase of just how complex the taxonomic situation between these two genera is, and the care that must be taken to resolve taxonomic conundrums where old material, multiple genus transitions, and misattribution of new material obfuscate the picture. We assessed the identity of the historic names Dyscophus alluaudi (currently in the genus Rhombophryne), Phrynocara laeve and Plethodontohyla laevis tsianovohensis (both synonyms of Rhombophryne alluaudi) based on an integrative taxonomic approach harnessing genetics, external morphology, osteological data obtained via micro-Computed Tomography (micro-CT) and bioacoustics. We show that (1) the holotype of Dyscophus alluaudi is a member of the genus Plethodontohyla; (2) the Rhombophryne specimens from central Madagascar currently assigned to Rhombophryne alluaudi have no affinity with that species, and are instead an undescribed species; and (3) Phrynocara laeve and Dyscophus alluaudi are not synonymous, but represent closely related species, whereas Plethodontohyla laevis tsianovohensis is tentatively confirmed as synonym of D. alluaudi. We resurrect and re-describe Plethodontohyla laevis, and re-allocate and re-describe Plethodontohyla alluaudi on the basis of new and historic material.

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          Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers.

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            Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory.

            Amphibians are in decline worldwide. However, their patterns of diversity, especially in the tropics, are not well understood, mainly because of incomplete information on taxonomy and distribution. We assess morphological, bioacoustic, and genetic variation of Madagascar's amphibians, one of the first near-complete taxon samplings from a biodiversity hotspot. Based on DNA sequences of 2,850 specimens sampled from over 170 localities, our analyses reveal an extreme proportion of amphibian diversity, projecting an almost 2-fold increase in species numbers from the currently described 244 species to a minimum of 373 and up to 465. This diversity is widespread geographically and across most major phylogenetic lineages except in a few previously well-studied genera, and is not restricted to morphologically cryptic clades. We classify the genealogical lineages in confirmed and unconfirmed candidate species or deeply divergent conspecific lineages based on concordance of genetic divergences with other characters. This integrative approach may be widely applicable to improve estimates of organismal diversity. Our results suggest that in Madagascar the spatial pattern of amphibian richness and endemism must be revisited, and current habitat destruction may be affecting more species than previously thought, in amphibians as well as in other animal groups. This case study suggests that worldwide tropical amphibian diversity is probably underestimated at an unprecedented level and stresses the need for integrated taxonomic surveys as a basis for prioritizing conservation efforts within biodiversity hotspots.
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              The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice.

              Vocalizations of anuran amphibians have received much attention in studies of behavioral ecology and physiology, but also provide informative characters for identifying and delimiting species. We here review the terminology and variation of frog calls from a perspective of integrative taxonomy, and provide hands-on protocols for recording, analyzing, comparing, interpreting and describing these sounds. Our focus is on advertisement calls, which serve as premating isolation mechanisms and, therefore, convey important taxonomic information. We provide recommendations for terminology of frog vocalizations, with call, note and pulse being the fundamental subunits to be used in descriptions and comparisons. However, due to the complexity and diversity of these signals, an unequivocal application of the terms call and note can be challenging. We therefore provide two coherent concepts that either follow a note-centered approach (defining uninterrupted units of sound as notes, and their entirety as call) or a call-centered approach (defining uninterrupted units as call whenever they are separated by long silent intervals) in terminology. Based on surveys of literature, we show that numerous call traits can be highly variable within and between individuals of one species. Despite idiosyncrasies of species and higher taxa, the duration of calls or notes, pulse rate within notes, and number of pulses per note appear to be more static within individuals and somewhat less affected by temperature. Therefore, these variables might often be preferable as taxonomic characters over call rate or note rate, which are heavily influenced by various factors. Dominant frequency is also comparatively static and only weakly affected by temperature, but depends strongly on body size. As with other taxonomic characters, strong call divergence is typically indicative of species-level differences, whereas call similarities of two populations are no evidence for them being conspecific. Taxonomic conclusions can especially be drawn when the general advertisement call structure of two candidate species is radically different and qualitative call differences are thus observed. On the other hand, quantitative differences in call traits might substantially vary within and among conspecific populations, and require careful evaluation and analysis. We provide guidelines for the taxonomic interpretation of advertisement call differences in sympatric and allopatric situations, and emphasize the need for an integrative use of multiple datasets (bio-acoustics, morphology, genetics), particularly for allopatric scenarios. We show that small-sized frogs often emit calls with frequency components in the ultrasound spectrum, although it is unlikely that these high frequencies are of biological relevance for the majority of them, and we illustrate that detection of upper harmonics depends also on recording distance because higher frequencies are attenuated more strongly. Bioacoustics remains a prime approach in integrative taxonomy of anurans if uncertainty due to possible intraspecific variation and technical artifacts is adequately considered and acknowledged.
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                Author and article information

                Journal
                Zoosystematics and Evolution
                ZSE
                Pensoft Publishers
                1860-0743
                1435-1935
                February 02 2018
                February 02 2018
                : 94
                : 1
                : 109-135
                Article
                10.3897/zse.94.14698
                318d8dd4-9784-4059-b3a8-56d765671d78
                © 2018

                http://creativecommons.org/licenses/by/4.0/

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