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The biology of an isolated Mashona mole-rat population from southern Malawi, with implications for the diversity and biogeography of the genus Fukomys

The Mashona mole-rat, Fukomys darlingi (Thomas, 1895), is a little studied social African mole-rat (Bathyergidae) from south-astern Africa. Here, we present an integrative study characterizing the genetic diversity of populations assigned to F. darlingi with special focus on animals from Nsanje, southern Malawi. These mole-rats show pronounced differences in body mass and general appearance compared to nominate F. darlingi from Zimbabwe and Mozambique, but their taxonomic status has so far remained unclear. A genetic analysis encompassing all major lineages of the genus Fukomys suggests that this population indeed represents a deeply nested lineage within the F. darlingi clade. The karyotype of the Nsanje mole-rats also corresponds to that of the nominate form, being 2n = 54. While both nuclear and mitochondrial data agree about the assignment of the Nsanje mole-rats to F. darlingi, our analyses revealed substantial mitonuclear discordance for other branches within the Fukomys phylogenetic tree. Nsanje mole-rats are significantly larger than nominate F. darlingi and their ontogeny and reproduction closely resemble similar-sized congeneric species rather than the nominate population. The somatic growth of the Nsanje form is the slowest of all African mole-rats. The maximum life span of F. darlingi is at least 19 years. The observed differences between nominate F. darlingi and mole-rats from Nsanje may be attributed mainly to their different body mass. Our study highlights the advantages of an integrative approach for understanding the diversity of African mole-rats and emphasizes the great intraspecific variability that may be encountered in these underground-dwelling rodents.

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Comparative molecular and morphological species delineation of Halammohydra Remane, 1927 (Hydrozoa)—with the description of four new species

Whereas most cnidarians are macrofaunal, a few microscopic lineages have evolved, and some of them inhabit marine sediments. The meiofaunal genus with the most species is Halammohydra, comprising nine nominal species. Species are described with high intraspecific variability in, e.g., the number of tentacles and statocysts and the shape and length of tentacles and body, complicating morphological identification to species level. Additionally, there is not much molecular data available. This study aims to revise already described species with morphological and molecular methods, as well as, to delineate potential new species answering questions about their geographical distribution. For this, specimens were sampled at 16 locations in the Northwest Atlantic and two localities in the East Atlantic, documented with light microscopy, and fixed individually for sequencing (16S, 18S, and CO1). Herewith, morphological characters were linked to a specific sequence, enabling the testing of character variation within one molecular phylogenetic group. Phylogenetic analyses were conducted (Bayesian Interference and Maximum Likelihood) in combination with species delimitation tests (ABGD, GMYC, and bPTP). Four already described species were identified in the data sets, and all of these were found at multiple localities. Four new species are described. Overall, the combined molecular and morphological data acquisition revealed multiple new species and a high degree of sympatry in Halammohydra. This, together with the confirmed excessive intraspecific variation in morphological traits, underlines the necessity of molecular sequencing for the taxonomy and species identification of Halammohydra.

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Development and structure of the anterior nervous system and sense organs in the holopelagic annelid Tomopteris spp. (Phyllodocida, Errantia)

Tomopteridae are transparent, predatory Annelida inhabiting pelagic ocean zones. Despite being well-known for their fast metachronal swimming and species-specific bioluminescence, our knowledge of morphological adaptations in these fascinating holopelagic worms remains extremely limited. In particular, the evolutionary scenarios and adaptive changes related to the transition from putative benthic ancestors to recent free-swimming groups remain poorly investigated and understood. Therefore, we investigated different taxa and developmental stages within the holopelagic Tomopteridae. We used a comparative morphological approach, including a range of microscopic methods, in our investigations focused on the anterior nervous system and prominent sensory structures, such as nuchal organs and tentacular cirri, in early developmental and adult stages of four tomopterid species. Our data show that Tomopteridae undergo heterochronic, lecithotrophic development with early visibility of adult-like features, which is consistent with earlier investigations. Furthermore, our ultrastructural examinations of the tomopterid nuchal organ highlight the conservativism in the fine structure and development of this prominent polychaete chemosensory organ. Nevertheless, our data indicate ultrastructural differences, such as an extraordinary number of supporting cell types and a bipartite olfactory chamber, potentially related to their pelagic lifestyle. In contrast to previous assumptions, it is shown that the supporting structures in the cirrus-like appendages of the first chaetiger contain prominent intracellular skeletal elements rather than annelid chaetae. These findings highlight the need for further investigations to understand Annelida’s immense morphological diversity of organ systems. Furthermore, our data demonstrate the necessity of functional analyses to understand Annelida’s adaptive radiation of sensory and neuronal structures.

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Guess who? Taxonomic problems in the genus Eiseniella revisited by integrated approach

Eiseniella neapolitana is a semi-aquatic, diploid earthworm that for many years was related to the cosmopolitan species Eiseniella tetraedra and even considered a subspecies of it. Norealidys andaluciana was described in Spain and is usually synonymized with E. neapolitana. We collected 69 specimens from Italy, Spain, and Cyprus and studied five molecular markers (COI, 16S, 28S, 12S, and ND1) and their morphology to solve this taxonomic problem. Phylogenetic analyses reveal the possible existence of two separate genera confounded under the name Eiseniella, but the study of more molecular markers and species of the genus would be necessary to confirm this. Therefore, the synonymy between Eiseniella and Norealidys is maintained. Various genetic analyses, including species delimitation, confirm the separation between E. neapolitana and E. andaluciana (= N. andaluciana) and excluded that E. neapolitana is a subspecies of E. tetraedra. The resemblance in external appearance despite clear genetic differences of the three species could be explained by convergent adaptation to the aquatic habitat. Despite the expected low haplotype diversity based on the 28S gene, we found a surprisingly high variability in the E. andaluciana (= N. andaluciana) population in Spain. However, its stable predicted secondary structure and its high content of G + C reject the presence of a pseudogene.

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Historical biogeography highlights the role of Miocene landscape changes on the diversification of a clade of Amazonian tree frogs

The diversification processes underlying why Amazonia hosts the most species-rich vertebrate fauna on earth remain poorly understood. We studied the spatio-temporal diversification of a tree frog clade distributed throughout Amazonia (Anura: Hylidae: Osteocephalus, Tepuihyla, and Dryaderces) and tested the hypothesis that Miocene mega wetlands located in western and central Amazonia impacted connectivity among major biogeographic areas during extensive periods. We assessed the group’s diversity through DNA-based (16S rRNA) species delimitation to identify Operational Taxonomic Units (OTUs) from 557 individuals. We then selected one terminal for each OTU (n = 50) and assembled a mitogenomic matrix (~14,100 bp; complete for 17 terminals) to reconstruct a Bayesian, time-calibrated phylogeny encompassing nearly all described species. Ancestral area reconstruction indicates that each genus was restricted to one of the major Amazonian biogeographic areas (western Amazonia, Guiana Shield and Brazilian Shield, respectively) between ~10 and 20 Mya, suggesting that they diverged and diversified in isolation during this period around the Pebas mega wetland. After 10 Mya and the transition to the modern configuration of the Amazon River watershed, most speciation within each genus continued to occur within each area. In Osteocephalus, only three species expanded widely across Amazonia (< 6 Mya), and all were pond-breeders. Species with other breeding modes remained mostly restricted to narrow ranges. The spectacular radiation of Osteocephalus was probably driven by climatic stability, habitat diversity and the acquisition of new reproductive modes along the Andean foothills and western Amazonia. Our findings add evidence to the importance of major hydrological changes during the Miocene on biotic diversification in Amazonia.

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Integrative taxonomy helps to revise systematics and questions the purported cosmopolitan nature of the type species within the genus Diaforobiotus (Eutardigrada: Richtersiusidae)

Recent advances in tardigrade taxonomy have been greatly enhanced by the redescriptions of the type species for particular taxa or species groups. De novo characterisation of these key taxa now allows to describe tardigrade species diversity with improved precision and at higher rate, increasing the momentum towards resolving the taxonomic impediment in these micro-invertebrates. Since its description, Diaforobiotus islandicus (Richters, 1904) has been reported from many distinct localities around the world. This suggested, perhaps falsely, a cosmopolitan nature of the species. However, potential erroneous assignment of newly found populations to this species could be a result of the very general and superficial original description. In order to properly recognise and name species diversity within the genus, I provide here an integrative redescription of the type species (D. islandicus) with a neotype designation, a description of a new species, Diaforbiotus svalbardicus sp. nov, and dichotomous key for the genus. Both descriptions are based on detailed morphological and morphometric data associated with standard DNA sequences of four genetic markers (18S rRNA, 28S rRNA, ITS-2, and COI). The genus composition and diagnosis amendments of the family Richtersiuside are also discussed. The presented study constitutes a starting point for further systematic studies on the genus Diaforobiotus and new taxa discoveries.

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Differentiation of skull morphology and cranial kinesis in common toads

We examined the cranial morphology and cranial kinesis of the common toads Bufo bufo and B. spinosus with micro-computed tomography and geometric morphometrics and compared the results with published data for related species in a phylogenetic context. The species significantly diverge in skull shape. The skull of B. spinosus is shorter and higher, with a ventral arm of the squamosal bone and the jaw articulation point positioned perpendicular to the braincase, in comparison with a more lateral position in B. bufo. In either species, females have a shorter snout and a higher and wider skull at the jaw articulation point that is positioned more posteriorly, in comparison with conspecific males. High variation in the amount of bone ossification was recorded in both species, ranging from scarcely ossified and loosely connected bones to highly ossified and firmly connected bones. We also found that skull shape and inferred kinetic properties of the skull are highly variable across the Bufonini tribe. However, sample sizes are mostly small and intraspecific variation is high, which might compromise the analyses. Overall, the results suggest that developmental plasticity produces high variation in ossification and cranial kinesis, affecting individuals’ feeding performances. At the population level, this variation supports an efficient exploitation of the habitat and may promote morphological adaptation in a changing environment.

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Cryptic subterranean diversity: regional phylogeography of the sand termite Psammotermes allocerus Silvestri, 1908 in the wider Namib region

Psammotermes allocerus Silvestri, 1908 is the only described species representing the genus Psammotermes Desneux, 1902 in Southern Africa. The large geographical range of this subterranean termite covers both summer and winter rainfall regimes. Deadwood is the preferred food when available, but in more arid habitats, both live and dead grasses form the major dietary component. Along the Namib Desert margins, the species’ localised herbivory creates circular bare patches known as fairy circles. For a regional phylogeographic study of this species, we sampled 65 sand termite populations within drier parts of Namibia, South Africa, and Angola. Based on combined molecular and ecological data, we found considerable genetic diversification within P. allocerus. Analyses of two mitochondrial markers (COI, COII), including a Bayesian inference tree, haplotype analysis and genetic distances suggest a delineation into seven highly differentiated genetic groups. The ‘Succulent Karoo’ group is additionally characterised by unique features of the royal chamber, nest and tunnel system. In conclusion, our data suggest that P. allocerus should be not regarded as one species but as a species complex. Termites of each analysed group ‘Northern Namib’, ‘Western Kalahari Basin’, ‘Nama’, ‘Southwestern Kalahari’, ‘East Gariep’, ‘Southern Namib’ and ‘Succulent Karoo’ should be considered as distinct species. The species name P. allocerus should be used for termites of the ‘Succulent Karoo’.

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