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The Great Barrier Reef is the largest coral reef ecosystem on the planet and supports a diverse community of marine fishes, as well as the organisms that parasitise them. Although the digenetic trematodes that parasitise fishes of the Great Barrier Reef have been studied for over a century, the species richness and diversity of many trematode lineages is yet to be explored. Trigonocephalotrema, gen. nov. is proposed to accommodate three new species, Trigonocephalotrema euclidi, sp. nov., T. hipparchi, sp. nov. and T. sohcahtoa, sp. nov., parasitic in fishes of Naso Lacepède and Zebrasoma Swainson (Acanthuridae) in the tropical Pacific. Species of Trigonocephalotrema are characterised with morphological and molecular data (18S rRNA, ITS2 and 28S rRNA). Species of Trigonocephalotrema are morphologically distinguished from all other haplosplanchnid lineages by having terminal, triangular, plate-like oral suckers. With the inclusion of the new molecular data, Bayesian inference and maximum likelihood analyses of the Haplosplanchnidae Poche, 1926 recovered identical tree topologies and demonstrated Trigonocephalotrema as a well-supported monophyletic group. Although species of Trigonocephalotrema are differentiated from all other haplosplanchnid lineages on the basis of morphology, species within the genus are morphologically cryptic; thus, accurate species identification will require inclusion of host and molecular data. Species of Trigonocephalotrema cannot be assigned to a recognised subfamily within the Haplosplanchnidae using either morphological or molecular data and would require the erection of a new subfamily to accommodate them. However, we find little value in the use of subfamilies within the Haplosplanchnidae, given that there are so few taxa in the family, and herein propose that their use be avoided.
Psammodrilidae constitutes a family of understudied, nearly completely ciliated, small-sized annelids, whose systematic position in Annelida remains unsettled and whose internal phylogeny is here investigated for the first time. Psammodrilids possess hooked chaetae typical of macroscopic tube-dwelling semi-sessile annelids, such as Arenicolidae. Yet, several minute members resemble, with their conspicuous gliding by ciliary motion and vagile lifestyle, interstitial fauna, adapted to move between sand grains. Moreover, psammodrilids exhibit a range of unique features, for example, bendable aciculae, a collar region with polygonal unciliated cells, and a muscular pumping pharynx. We here present a combined phylogeny of Psammodrilidae including molecular and morphological data of all eight described species (two described herein as Psammodrilus didomenicoi, sp. nov. and P. norenburgi, sp. nov.) as well as four undescribed species. Ancestral character state reconstruction suggests the ancestor of Psammodrilidae was a semi-sessile larger form. Miniaturisation seems to have occurred multiple times independently within Psammodrilidae, possibly through progenesis, yielding small species with resemblance to a juvenile stage of the larger species. We find several new cryptic species and generally reveal an unexpected diversity and distribution of this small family. This success may be favoured by their adaptive morphology, here indicated to be genetically susceptible to progenesis.
The decline of morphologically based taxonomy is mainly linked to increasing species redundancy, which probably contributed to a worldwide disinterest in taxonomy, and to a reduction of funding for systematic biology and for expertise training. The present trend in the study of biodiversity is integrated taxonomy, which merges morphological and molecular approaches. At the same time, in many cases new molecular techniques have eclipsed the morphological approach. The application of Standardised Integrative Taxonomy, i.e. a rigorous, common method of description based on the integration between ecological and morphological characteristics, may increase the precision, accessibility, exploitability and longevity of the collected data, and favour the renaissance of taxonomy by new investments in biodiversity exploration.
Peronina Plate, 1893 is a genus of onchidiids that live on the mud in mangrove forests. Peronina can be identified in the field by the lung opening at the margin between the ventral hyponotum and the dorsal notum, and by the distinctive scalloped notum edge. This genus was previously known only from the holotype of the type species, Peronina alta Plate, 1893, from eastern India. Onchidium tenerum Stoliczka, 1869 is moved to Peronina and applies to the same species as Peronina alta. Peronina species are described using an integrative approach (natural history, comparative anatomy and DNA sequences). Mitochondrial COI and 16S sequences and nuclear ITS2 and 28S sequences are used to independently test species boundaries. Mitochondrial sequences yielded three units separated by a large barcode gap, but nuclear sequences yielded two units. Because these two units are congruent with differences in the male copulatory apparatus, they are accepted as species. Explanations for highly divergent COI haplotypes within one species are discussed. Peronina tenera (Stoliczka, 1869) is distributed in the Bay of Bengal and the Strait of Malacca, while P. zulfigari Goulding & Dayrat, sp. nov. is endemic to the Strait of Malacca. The two species differ internally but are cryptic externally.
Our study reports on the occurrence of a new species of Leucetta (Calcinea, Calcarea) from the Southern Ocean, Leucetta giribeti, sp. nov., collected in the shallow waters (15 m depth) of Deception Island, South Shetland Islands. This new taxon is described based on a combination of morphological and molecular data, including the description of oocytes, embryos, larvae and sperm found in the choanosome. While female reproductive elements showed great similarities with those of other calcineans, sperm is reported here for the first time in the whole Calcinea subclass. Sperm cells are flagellated and possess a typical spermatic mid-piece, which is usually observed in cnidarians. In our phylogenetic analyses, we recovered Leucetta giribeti, sp. nov. as sister species of a clade formed by species of the genera Leucetta, Pericharax and Leucettusa. Although the clade in which Leucetta giribeti, sp. nov. is placed is supported by molecular and morphological features, we cannot propose a new genus due to uncertainties regarding the type species of the genus, Leucetta primigenia Haeckel, 1872. Our study reinforces the relevance of integrative approaches in the description of new taxa and contributes to resolving the poorly known reproductive patterns of Antarctic sponge species.
Gonzalo Giribet, Rebecca S. Buckman-Young, Cristiano Sampaio Costa, Caitlin M. Baker, Ligia R. Benavides, Michael G. Branstetter, Savel R. Daniels, Ricardo Pinto-da-Rocha
Onychophorans, or velvet worms, are cryptic but extremely charismatic terrestrial invertebrates that have often been the subject of interesting biogeographic debate. Despite great interest, a well resolved and complete phylogeny of the group and a reliable chronogram have been elusive due to their broad geographic distribution, paucity of samples, and challenging molecular composition. Here we present a molecular phylogenetic analysis of Onychophora that includes previously unsampled and undersampled lineages and we analyse the expanded dataset using a series of nested taxon sets designed to increase the amount of information available for particular subclades. These include a dataset with outgroups, one restricted to the ingroup taxa, and three others for Peripatopsidae, Peripatidae and Neopatida (= the Neotropical Peripatidae). To explore competing biogeographic scenarios we generate a new time tree for Onychophora using the few available reliable fossils as calibration points. Comparing our results to those of Cyphophthalmi, we reconsider the hypothesis that velvet worms reached Southeast Asia via Eurogondwana, and conclude that a more likely scenario is that they reached Southeast Asia by rafting on the Sibumasu terrane. Our phylogenetic results support the reciprocal monophyly of both families as well as an early division between East and West Gondwana, also in both families, each beginning to diversify between the Permian and the Jurassic. Peripatopsidae clearly supports paraphyly of South Africa with respect to southern South America (Chile) and a sister group relationship of the Southeast Asian/New Guinean Paraperipatus to the Australian/New Zealand taxa. The latter includes a clade that divides between Western Australia and Eastern Australia and two sister clades of trans-Tasman species (one oviparous and one viviparous). This pattern clearly shows that oviparity is secondarily derived in velvet worms. Peripatidae finds a sister group relationship between the Southeast Asian Eoperipatus and the West Gondwanan clade, which divides into the African Mesoperipatus and Neopatida. The latter shows a well supported split between the Pacific Oroperipatus (although it is unclear whether they form one or two clades) and a sister clade that includes the members of the genera Peripatus, Epiperipatus, Macroperipatus and representatives of the monotypic genera Cerradopatus, Plicatoperipatus and Principapillatus. However, Peripatus, Epiperipatus and Macroperipatus are not monophyletic, and all the species from the monotypic genera are related to geographically close species. The same goes for the type species of Macroperipatus (from Trinidad, and sister group to other Trinidad and Tobago species of Epiperipatus) and Epiperipatus (from French Guiana, and related to other Guyana shield species of Epiperipatus and Peripatus). Geographic structure within Neopatida is largely obscured by an unresolved backbone, but many well supported instances of generic non-monophyly challenge the current taxonomic framework, which has often relied on anatomical characters that are untested phylogenetically.
The higher classification of the mirid subfamily Bryocorinae has received comparatively little attention. It is not highly species-rich in comparison with other mirid subfamilies but does exhibit extraordinary morphological heterogeneity. In this work we provide a synthesis of the subfamily on a global basis, providing a new key and updated diagnoses of supraspecific taxa. Five tribes are recognised: Bryocorini, Dicyphini, Eccritotarsini, Felisacini and Monaloniini. The genus Campyloneura Fieber is transferred from the tribe Dicyphini to the Eccritotarsini. Analysis of distributional patterns and a survey of host plant associations are provided. Available data on distribution of the main bryocorine lineages are summarised in tabular form and evaluated using UPGMA methods, and geographically structured patterns were detected. The synthesis will enable users to identify bryocorines to tribal level with confidence and provides a classificatory framework for future revisionary and phylogenetic studies.
The target species of this work is a large tritonid nudibranch, relatively common in Galician waters (north-western Spain), where it has been regularly observed since the year 2000. To date, this species has been misidentified as Tritonia hombergii, a species from northern latitudes, because of its remarkable size. On the other hand, the external anatomy of the specimens from north-western Spain resembles that of Marionia blainvillea, a species common in the Mediterranean. To untangle the real taxonomic identity of the species, a detailed anatomical study was performed, revealing several morphological differences that has led us to consider the Galician specimen as new to science. A phylogenetic analysis was conducted for both 16S and COI genes to try to establish its systematic position in relation with other taxa. Marionia gemmii, sp. nov. differs from all known Tritoniidae in the size, colour pattern, number of velar processes, presence of stomach plates, radular formula, the shape of spawning masses and even in its feeding habits. The phylogenetic results support the comprehensive morphological analysis and also make it clear that most tritoniid genera currently recognised are not monophyletic and that a major systematic revision of the family is needed.
The restricted distribution of viviparous onychophorans in Tasmania has long been a subject of discussion, but their evolutionary history remains unclear. We applied morphological, molecular and karyological methods to assess the taxonomy and phylogenetic relationships of the four viviparous species reported from Tasmania, including Tasmanipatus barretti, T. anophthalmus and two undescribed species previously referred to as ‘Tasmania’ sp. 1 and sp. 2. We demonstrate that all four species can be unambiguously distinguished based on independent character sets. The two ‘Tasmania’ species, which were previously thought to be cryptic, proved to exhibit a set of distinct morphological characters. Molecular phylogenetic analyses revealed that the four species belong to a major clade that includes Peripatoides from New Zealand, and that species from the two landmasses show reciprocal monophyly within this clade. Within the Tasmanian clade, T. anophthalmus is more closely related to the two ‘Tasmania’ species than to T. barretti. Based on this relationship and the lack of morphological and/or karyological characters supporting the Tasmanian viviparous clade, we erect two new genera to accommodate the two ‘Tasmania’ species (Diemenipatus, gen. nov.) and T. anophthalmus (Leucopatus, gen. nov.). An emended diagnosis followed by a redescription of T. barretti is provided and ‘Tasmania’ sp. 1 and sp. 2 are formally described as D. taiti, gen. et sp. nov. and D. mesibovi, gen. et sp. nov., respectively.
Forest-dwelling myrmeleontids are often inadequately known and poorly represented in collections, impeding the study of their affinities and ecology. An exceptional example is the Neotropical species Ripalda insignis (Rambur, 1842), described on a single specimen and never observed again for more than 170 years. The recent rediscovery of this species in Paraguay allowed us to include it in a broader phylogenetic context of the antlion tribe Nemoleontini. Our morphology-based phylogenetic reconstruction, including both adult and larval characters, supports the synonymisation of the genus Araucaleon Banks, 1938 with Ripalda Navás, 1915, as the members of these genera form a well-supported clade. Ripalda appears to be related to the diverse genus Eremoleon Banks, 1901 and the small genera Sericoleon Esben-Petersen, 1933, Navasoleon Banks, 1943 and Elachyleon Esben-Petersen, 1927. The genus is also revised in light of a new cladistic framework, redescribing the three previously known species, R. insignis (Rambur, 1842), R. withycombei (Esben-Petersen, 1927) and R. inca (Banks, 1938) and naming a new species, R. wayana, sp. nov. from French Guyana. The larva of R. withycombei is also described for the first time. Larval morphology is broadly consistent with Eremoleon, also sharing a similar ecological niche.
An integrative approach to investigate the species-level diversity in Oxynoe (Mollusca, Heterobranchia, Sacoglossa) revealed the existence of 11 distinct taxa. Oxynoe viridis (Pease, 1861) and Oxynoe antillarum Mörch, 1863 are redescribed; Oxynoe natalensis E. A. Smith, 1903 and Oxynoe azuropunctata Jensen, 1980 are regarded as valid. Species originally described from empty shells are regarded as nomina dubia. Seven new species are described, four from the tropical Indo-West Pacific: Oxynoe kylei, sp. nov., Oxynoe neridae, sp. nov., Oxynoe jordani, sp. nov. and Oxynoe jacksoni, sp. nov.; and two from the tropical Atlantic: Oxynoe struthioe, sp. nov. and Oxynoe ilani, sp. nov. The name Oxynoe panamensis Pilsbry & Olsson, 1943 has been applied to eastern Pacific specimens, but was introduced based on material collected from the Caribbean; therefore, the new name Oxynoe aliciae, sp. nov. is introduced for eastern Pacific specimens. Species are delineated using molecular and morphological traits, as well as algal host and reproductive biology. Results from morphological comparisons are concordant with molecular and integrative species delimitation analyses, providing robust evidence for species hypotheses. As Oxynoe is one of the few groups specialised to feed on the green algal genus Caulerpa, which includes highly invasive species, clarifying the taxonomy of Oxynoe may inform efforts to predict community response to disruptive algal invasions.
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