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The axiidean families Callianassidae and Ctenochelidae, sometimes treated together as Callianassoidea, are shown to represent a monophyletic taxon. It comprises 265 accepted species in 74 genera, twice this number of species if fossil taxa are included. The higher taxonomy of the group has proved difficult and fluid. In a molecular phylogenetic approach, we inferred evolutionary relationships from a maximum-likelihood (ML) and Bayesian analysis of four genes, mitochondrial 16S rRNA and 12S rRNA along with nuclear histone H3 and 18S rRNA. Our sample consisted of 298 specimens representing 123 species plus two species each of Axiidae and Callianideidae serving as outgroups. This number represented about half of all known species, but included 26 species undescribed or not confidently identified, 9% of all known. In a parallel morphological approach, the published descriptions of all species were examined and detailed observations made on about two-thirds of the known fauna in museum collections. A DELTA (Description Language for Taxonomy), database of 135 characters was made for 195 putative species, 18 of which were undescribed. A PAUP analysis found small clades coincident with the terminal clades found in the molecular treatment. Bayesian analysis of a total-evidence dataset combined elements of both molecular and morphological analyses. Clades were interpreted as seven families and 53 genera. Seventeen new genera are required to reflect the molecular and morphological phylograms. Relationships between the families and genera inferred from the two analyses differed between the two strategies in spite of retrospective searches for morphological features supporting intermediate clades. The family Ctenochelidae was recovered in both analyses but the monophyly of Paragourretia was not supported by molecular data. The hitherto well recognised family Eucalliacidae was found to be polyphyletic in the molecular analysis, but the family and its genera were well defined by morphological synapomorphies. The phylogram for Callianassidae suggested the isolation of several species from the genera to which they had traditionally been assigned and necessitated 12 new generic names. The same was true for Callichiridae, with stronger ML than Bayesian support, and five new genera are proposed. Morphological data did not reliably reflect generic relationships inferred from the molecular analysis though they did diagnose terminal taxa treated as genera. We conclude that discrepancies between molecular and morphological analyses are due at least in part to missing sequences for key species, but no less to our inability to recognise unambiguously informative morphological synapomorphies. The ML analysis revealed the presence of at least 10 complexes wherein 2–4 cryptic species masquerade under single species names.
Historically, morphological characters have been used to support the monophyly, composition, and phylogenetic relationships of scorpion families. Although recent phylogenomic analyses have recovered most of these traditional higher-level relationships as non-monophyletic, certain key taxa have yet to be sampled using a phylogenomic approach. Salient among these is the monotypic genus Caraboctonus Pocock, 1893, the type species of the family Caraboctonidae Kraepelin, 1905. Here, we examined the putative monophyly and phylogenetic placement of this family, sampling the library of C. keyserlingi Pocock, 1893 using high throughput transcriptomic sequencing. Our phylogenomic analyses recovered Caraboctonidae as polyphyletic due to the distant placement of the genera Caraboctonus and Hadrurus Thorell, 1876. Caraboctonus was stably recovered as the sister-group of the monotypic family Superstitioniidae Stahnke, 1940, whereas Hadrurus formed an unstable relationship with Uroctonus Thorell, 1876 and Belisarius Simon, 1879. Four-cluster likelihood mapping revealed that the instability inherent to the placement of Hadrurus, Uroctonus and Belisarius was attributable to significant gene tree conflict in the internodes corresponding to their divergences. To redress the polyphyly of Caraboctonidae, the following systematic actions have been taken: (1) the family Caraboctonidae has been delimited to consist of 23 species in the genera Caraboctonus and Hadruroides Pocock, 1893; (2) Caraboctonidae, previously included in the superfamily Iuroidea Thorell, 1876 or as incertae sedis, is transferred to the superfamily Caraboctonoidea (new rank); (3) the superfamily Hadruroidea (new rank) is established and the status of Hadrurinae Stahnke, 1973 is elevated to family (Hadruridae new status) including 9 species in the genera Hadrurus and Hoffmannihadrurus Fet & Soleglad, 2004 and (4) we treat Uroctonus and Belisarius as insertae sedis with respect to superfamilial placement. Our systematic actions engender the monophyly of both Iuroidea and Caraboctonidae. Future phylogenomic investigations should target similar taxon-poor and understudied lineages of potential phylogenetic significance, which are anticipated to reveal additional non-monophyletic groups.
Mimetidae is one of the three families within Araneoidea whose members do not spin foraging webs, but are unique in displaying a complex prey-capture behaviour known as aggressive mimicry. Mimetids are distributed worldwide and are most diverse in the tropics of Central and South America. Here we provide a comprehensive phylogeny of pirate spiders (Mimetidae) based on analyses that combine morphological and multigene nucleotide sequence data. We scored 147 morphological characters for 55 mimetids and 16 outgroup taxa and combined it in a total-evidence approach with the sequence data of Benavides et al. (2017) which included two nuclear ribosomal genes, 18S rRNA and 28S rRNA, two mitochondrial ribosomal genes, 12S rRNA and 16S rRNA, the nuclear protein-encoding gene histone H3 and the mitochondrial protein-encoding gene cytochrome c oxidase subunit I. We analysed the combined dataset using parsimony, maximum-likelihood and Bayesian inference methods. Our results support the monophyly of Mimetidae and of the genera Gelanor, Ero, Anansi and Australomimetus. Mimetidae is sister to Arkyidae + Tetragnathidae. Mimetus as currently circumscribed is not monophyletic under any analytical approach used, although several lineages within the genus are consistently found in our analyses. We describe, illustrate and discuss the morphological synapomorphies that support the main clades of Mimetidae. The following nomenclatural changes are proposed: Ermetus koreanus (Paik, 1967), the sole species of the genus, is transferred to Ero C. L. Koch, 1836 and thus Ermetus Ponomarev, 2008 is a junior synonym of Ero C. L. Koch, 1836 (new synonymy) and Ero koreana Paik, 1967 becomes a revalidated combination. Phobetinus sagittifer Simon, 1895, the type species of the genus, is transferred to Mimetus Hentz, 1832 and thus Phobetinus Simon, 1895 is a junior synonym of Mimetus Hentz, 1832 (new synonymy), which results in two changes: Mimetus sagittifer (Simon, 1895), new combination and Mimetus investus (Simon, 1909), new combination. Reo latro Brignoli, 1979, the type species of the genus, is transferred to Mimetus and thus Reo Brignoli, 1979 is a junior synonym of Mimetus (new synonymy), which results in the following two changes: Mimetus latro Brignoli, 1979, new combination and Mimetus eutypus Chamberlin & Ivie, 1935, revalidated combination. Arochoides integrans Mello-Leitão, 1935 is transferred to Tetragnathidae (new family placement). The type specimen of Arochoides integrans, the only species in this genus, is a subadult male of Azilia (Tetragnathidae), most likely Azilia histrio Simon, 1895. Arochoides is a junior synonym of Azilia (new synonymy).
Accurate phylogenies are important for understanding the evolutionary histories of organisms, their reproductive traits and ecological habits. The freshwater mussel order Unionida is currently thought to include six families. However, assignment of particular species to these families has been unstable, particularly for species that have been described solely on conchological characters. Unio polystictus Heude, 1877 represents such a species. Based on DNA sequence data from five genes (COI, 16S rRNA, 18S rRNA, 28S rRNA and histone H3) and complete mitochondrial genomes, we investigated the phylogenetic position and generic affinities of U. polystictus using various analytical methods. Both the five-gene and mitogenome datasets strongly supported transferring U. polystictus from Margaritiferidae to Unionidae as Aculamprotula polysticta, comb. res. Our results also supported the following intrageneric relationships: (Aculamprotula tortuosa, ((Aculamprotula polysticta, Aculamprotula scripta), (Aculamprotula fibrosa, Aculamprotula tientsinersis))). In addition, by comparing the morphological features of Aculamprotula (Unionidae, Unioninae), Lamprotula (Unionidae, Gonideinae) and Gibbosula (Margaritiferidae, Gibbosulinae) species, potential issues of relying solely on shell morphology for high-level classification of freshwater mussels are highlighted. Confirmation of classification position and genetic relationship for Aculamprotula polysticta will helpful to understand the ecological characteristics, reproductive strategies and host-fish requirements, which can be inferred from closely related taxa.
Dorvilleidae is a diverse group of annelids found in many marine environments and also commonly associated with chemosynthetic habitats. One dorvilleid genus, Parougia, currently has 11 described species, of which two are found at vents or seeps: Parougia wolfi and Parougia oregonensis. Eight new Parougia species are recognised and described in this study from collections in the Pacific Ocean, all from whale-falls, hydrothermal vents, or methane seeps at ∼600-m depth or greater. The specimens were studied using morphology and phylogenetic analyses of DNA sequences from mitochondrial (cytochrome c oxidase subunit I, 16S rRNA, and cytochrome b) and nuclear (18S rRNA and histone 3) genes. Six sympatric Parougia spp. were found at Hydrate Ridge, Oregon, while three of the Parougia species occurred at different types of chemosynthetic habitats. Two new species were found over wide geographical and bathymetric ranges. Another dorvilleid genus, Ophryotrocha, has previously been highlighted as diversifying in the deep-sea environment. Our results document the hitherto unknown diversity of another dorvilleid genus, Parougia, at various chemosynthetic environments.
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