A Family Level Analysis of Tardigrade Phylogeny

In the present study a character data set suitable for cladistic analysis at the family level was developed. A data matrix consisting of 50 morphological characters from 15 families of tardigrades was analyzed by maximum parsimony. Kinorhynchs, loriciferans, and gastrotrichs were used as outgroups. The results agree with the currently accepted hypothesis that Eutardigrada and Heterotardigrada are distinct monophyletic groups. Among the eutardigrades, Eohypsibiidae was found to be a sister group to Macrobiotidae+Hypsibiidae, while Milnesiidae was the basal eutardigrade family. The basal heterotardigrade family was found to be Oreellidae. Echiniscoideans grouped with some traditional Arthrotardigrada (Renaudarctidae, Coronarctidae+Batillipedidae) suggesting that the arthrotardigrades are not monophyletic. The 18S rRNA gene sequence of Batillipes mirus Richters, 1909 and Calohypsibius schusteri Nelson & McGlothlin, 1996 were obtained and their addition to a previously published dataset supports the monophyly of Heterotardigrada and of Parachela versus Apochela within the Eutardigrada.     

An upgraded comprehensive multilocus phylogeny of the Tardigrada tree of life

Providing accurate animals’ phylogenies rely on increasing knowledge of neglected phyla. Tardigrada diversity evaluated in broad phylogenies (among phyla) is biased towards eutardigrades. A comprehensive phylogeny is demanded to establish the representative diversity and propose a more natural classification of the phylum. So, we have performed multilocus (18S rRNA and 28S rRNA) phylogenies with Bayesian inference and maximum likelihood. We propose the creation of a new class within Tardigrada, erecting the order Apochela (Eutardigrada) as a new Tardigrada class, named Apotardigrada comb. n. Two groups of evidence support its creation: (a) morphological, presence of cephalic appendages, unique morphology for claws (separated branches) and wide‐elongated buccopharyngeal apparatus without placoids, and (b) phylogenetic support based on molecular data. Consequently, order Parachela is suppressed and its superfamilies erected as orders within Eutardigrada, maintaining their current names. We propose a new classification within the family Echiniscidae (Echiniscoidea, Heterotardigrada) with morphological and phylogenetic support: (a) subfamily Echiniscinae subfam. n., with two tribes Echiniscini tribe n. and Bryodelphaxini tribe n.; (b) subfamily Pseudechiniscinae subfam. n., with three tribes Cornechiniscini tribe n., Pseudechiniscini tribe n. and Anthechiniscini tribe n.; and (c) subfamily Parechiniscinae subfam. n., with two tribes Parechiniscini tribe n. and Novechiniscini tribe n. Reliable biodiversity selection for tardigrades in broad phylogenies is proposed due to biased analyses performed up to now. We use our comprehensive molecular phylogeny to evaluate the evolution of claws in the clawless genus Apodibius and claw reduction across the Tardigrada tree of life. Evolutionary consequences are discussed.     

A molecular approach to arthrotardigrade phylogeny (Heterotardigrada,Tardigrada)

The marine order Arthrotardigrada (class Heterotardigrada, phylum Tardigrada) is known for its conspicuously high morphological diversity and has been traditionally recognized as the most ancestral group within the phylum. Despite its potential importance in understanding the evolution of the phylum, the phylogenetic relationships of Arthrotardigrada have not been clarified. This study conducted molecular phylogenetic analyses of the order encompassing all families except Neoarctidae using nuclear 18S and 28S rRNA fragments. Data from two rare families, Coronarctidae and Renaudarctidae, were included for the first time. The analyses confirmed the monophyly of Heterotardigrada and inferred Coronarctidae as the sister group to all other heterotardigrade taxa. Furthermore, the results support a monophyletic Renaudarctidae + Stygarctidae clade, which has been previously suggested on morphology. Our data indicated that two subfamilies currently placed in Halechiniscidae are only distantly related to this family. We propose that these taxa are each elevated to family level (Styraconyxidae (new rank) and Tanarctidae (new rank)). The morphology of tardigrades is discussed in the context of the inferred phylogeny.     

Molecular phylogeny of Arthrotardigrada (Tardigrada)

Tardigrades are microscopic ecdysozoans with a worldwide distribution covering marine, limnic and terrestrial habitats. They are regarded as a neglected phylum with regard to studies of their phylogeny. During the last decade molecular data have been included in the investigation of tardigrades. However, the marine arthrotardigrades are still poorly sampled due to their relative rarity, difficult identification and minute size even for tardigrades. In the present study, we have sampled various arthrotardigrades and sequenced the 18S and partial 28S ribosomal subunits. The phylogenetic analyses based on Bayesian inference and maximum parsimony inferred Heterotardigrada (Arthrotardigrada + Echiniscoidea) and Eutardigrada to be monophyletic. Arthrotardigrada was inferred to be paraphyletic as the monophyletic Echiniscoidea is included within the arthrotardigrades. The phylogenetic positions of Stygarctidae and Batillipedidae are poorly resolved with low branch support. The Halechiniscidae is inferred to be polyphyletic as the currently recognized Styraconyxinae is not part of the family. Archechiniscus is the sister-group to the Halechiniscidae and Orzeliscus is placed as one of the basal halechiniscids. The phylogeny of the included eutardigrade taxa resembles the current molecular phylogenies. The genetic diversity within Arthrotardigrada is much larger (18S 15.1–26.5%, 28S 7.2–20.7%) than within Eutardigrada (18S 1.0–12.6%, 28S 1.3–8.2%). This can be explained by higher substitution rates in the arthrotardigrades or by a much younger evolutionary age of the sampled eutardigrades.     

Phylum Tardigrada: an “individual” approach

Phylum Tardigrada consists of ～1000 tiny, hardy metazoan species distributed throughout terrestrial, limno‐terrestrial and oceanic habitats. Their phylogenetic status has been debated, with current evidence placing them in the Ecdysozoa. Although there have been efforts to explore tardigrade phylogeny using both morphological and molecular data, limitations such as their few morphological characters and low genomic DNA concentrations have resulted in restricted taxonomic coverage. Using a protocol that allows us to identify and extract DNA from individuals, we have sequenced 18S rDNA from 343 tardigrades from across the globe. Using maximum parsimony and Bayesian analyses we have found support for dividing Order Parachela into three super‐families and further evidence that indicates the traditional taxonomic perspective of families in the class Eutardigrada are nonmonophyletic and require re‐working. It appears that conserved morphology within Tardigrada has resulted in conservative taxonomy as we have found cases of several discrete lineages grouped into single genera. Although this work substantially adds to the understanding of the evolution and taxonomy of the phylum, we highlight that inferences gained from this work are likely to be refined with the inclusion of further taxa—specifically representatives of the nine families yet to be sampled. © The Willi Hennig Society 2008.     

Trehalose and anhydrobiosis in tardigrades--evidence for divergence in responses to dehydration

To withstand desiccation, many invertebrates such as rotifers, nematodes and tardigrades enter a state known as anhydrobiosis, which is thought to require accumulation of compatible osmolytes, such as the non-reducing disaccharide trehalose to protect against dehydration damage. The trehalose levels of eight tardigrade species comprising Heterotardigrada and Eutardigrada were observed in five different states of hydration and dehydration. Although many species accumulate trehalose during dehydration, the data revealed significant differences between the species. Although trehalose accumulation was found in species of the order Parachela (Eutardigrada), it was not possible to detect any trehalose in the species Milnesium tardigradum and no change in the trehalose level has been observed in any species of Heterotardigrada so far investigated. These results expand our current understanding of anhydrobiosis in tardigrades and, for the first time, demonstrate the accumulation of trehalose in developing tardigrade embryos, which have been shown to have a high level of desiccation tolerance.     

Robust support for tardigrade clades and their ages from three protein-coding nuclear genes

Abstract. Coding sequences (5,334 nt total) from elongation factor-1α, elongation factor-2, and the largest subunit of RNA polymerase II were determined for 6 species of Tardigrada, 2 of Arthropoda, and 2 of Onychophora. Parsimony and likelihood analyses of nucleotides and amino acids yielded strong support for Tardigrada and all internal nodes (i.e., 100% bootstrap support for Tardigrada, Eutardigrada, Parachela, Hypsibiidae, and Macrobiotidae). Results are in agreement with morphology and an earlier molecular study based on analysis of 18S ribosomal sequences. Divergence times have been estimated from amino acid sequence data using an empirical Bayesian statistical approach, which does not assume a strict molecular clock. Divergence time estimates are pre-Vendian for Tardigrada/Arthropoda, Vendian or earlier for Eutardigrada/Heterotardigrada, Silurian to Ordovician for Parachela/Apochela, Permian to Carboniferous for Hypsibiidae and Macrobiotidae, and Mesozoic for Isohypsibius/Thulinia (both within Hypsibiidae) and Macrobiotus/Richtersius (both within Macrobiotidae).     

Reconstructing the phylogeny of the Sipuncula

Sipunculans are marine spiralian worms with possible close affinities to the Mollusca or Annelida. Currently 147 species, 17 genera, 6 families, 4 orders and 2 classes are recognized. In this paper we review sipunculan morphology, anatomy, paleontological data and historical affiliations. We have conducted cladistic analyses for two data sets to elucidate the phylogenetic relationships among sipunculan species. We first analyzed the relationships among the 45 species of Phascolosomatidea with representatives of the Sipunculidea as outgroups, using 35 morphological characters. The resulting consensus tree has low resolution and branch support is low for most branches. The second analysis was based on DNA sequence data from two nuclear ribosomal genes (18S rRNA and 28S rRNA) and one nuclear protein-coding gene, histone H3. Outgroups were chosen among representative spiralians. In a third analysis, the molecular data were combined with the morphological data. Data were analyzed using parsimony as the optimality criterion and branch support evaluated with jackknifing and Bremer support values. Branch support for outgroup relationships is low but the monophyly of the Sipuncula is well supported. Within Sipuncula, the monophyly of the two major groups, Phascolosomatidea and Sipunculidea is not confirmed. Of the currently recognized families, only Themistidae appears monophyletic. The Aspidosiphonidae, Phascolosomatidae and Golfingiidae would be monophyletic with some adjustments in their definition. The Sipunculidae is clearly polyphyletic, with Sipunculus nudus as the sister group to the remaining Sipuncula, Siphonosoma cumanense the sister group to a clade containing Siphonosoma vastumand the Phascolosomatidea, and Phascolopsis gouldi grouping within the Golfingiiformes, as suggested previously by some authors. Of the genera with multiple representatives, only Phascolosoma and Themiste are monophyletic as currently defined. We are aiming to expand our current dataset with more species in our molecular database and more detailed morphological studies.     

New molecular data for tardigrade phylogeny, with the erection of Paramacrobiotus gen. nov.

Up to few years ago, the phylogenies of tardigrade taxa have been investigated using morphological data, but relationships within and between many taxa are still unresolved. Our aim has been to verify those relationships adding molecular analysis to morphological analysis, using nearly complete 18S ribosomal DNA gene sequences (five new) of 19 species, as well as cytochrome oxidase subunit 1 (COI) mitochondrial DNA gene sequences (15 new) from 20 species, from a total of seven families. The 18S rDNA tree was calculated by minimum evolution, maximum parsimony (MP) and maximum likelihood (ML) analyses. DNA sequences coding for COI were translated to amino acid sequences and a tree was also calculated by neighbour-joining, MP and ML analyses. For both trees (18S rDNA and COI) posterior probabilities were calculated by MrBayes. Prominent findings are as follows: the molecular data on Echiniscidae (Heterotardigrada) are in line with the phylogenetic relationships identifiable by morphological analysis. Among Eutardigrada, orders Apochela and Parachela are confirmed as sister groups. Ramazzottius (Hypsibiidae) results more related to Macrobiotidae than to the genera here considered of Hypsibiidae. Macrobiotidae and Macrobiotus result not monophyletic and confirm morphological data on the presence of at least two large groups within Macrobiotus. Using 18S rDNA and COI mtDNA genes, a new phylogenetic line has been identified within Macrobiotus , corresponding to the ' richtersi-areolatus group'. Moreover, cryptic species have been identified within the Macrobiotus ' richtersi group' and within Richtersius . Some evolutionary lines of tardigrades are confirmed, but others suggest taxonomic revision. In particular, the new genus Paramacrobiotus gen. n. has been identified, corresponding to the phylogenetic line represented by the ' richtersi-areolatus group'.     

Are the supportive structures of the tardigrade pharynx homologous throughout the entire group?

Based on the differences in position, structure and function of the cuticular supportive structures in the pharynx of representatives of two major groups of tardigrades, the Heterotardigrada and Eutardigrada, it is postulated that these structures are not homologous throughout the Tardigrada. Hithero, these structures have been termed placoids in both classes (fused placoids in Heterotardigrada versus macro- and microplacoids in Eutardigrada). It is proposed here that there are two different kinds of fused placoids within the Heterotardigrada and that only one of these is homologous to the macro- and microplacoids of the Eutardigrada. It is furthermore proposed that the plesiomorphic condition for the tardigrade pharynx is without reinforced cuticular supportive structures and not the presence of encrusted fused placoids as found in Heterotardigrada.     

Evolutionary relationships within the protostome phylum Sipuncula: a molecular analysis of ribosomal genes and histone H3 sequence data

The phylogenetic relationships of the members of the phylum Sipuncula are investigated by means of DNA sequence data from three nuclear markers, two ribosomal genes (18S rRNA and the D3 expansion fragment of 28S rRNA), and one protein-coding gene, histone H3. Phylogenetic analysis via direct optimization of DNA sequence data using parsimony as optimality criterion is executed for 12 combinations of parameter sets accounting for different indel costs and transversion/transition cost ratios in a sensitivity analysis framework. Alternative outgroup analyses are also performed to test whether they affected rooting of the sipunculan topology. Nodal support is measured by parsimony jackknifing and Bremer support values. Results from the different partitions are highly congruent, and the combined analysis for the parameter set that minimizes overall incongruence supports monophyly of Sipuncula, but nonmonophyly of several higher taxa recognized for the phylum. Mostly responsible for this is the split of the family Sipunculidae in three main lineages, with the genus Sipunculus being the sister group to the remaining sipunculans, the genus Phascolopsis nesting within the Golfingiiformes, and the genus Siphonosoma being associated to the Phascolosomatidea. Other interesting results are the position of Phascolion within Golfingiidae and the position of Antillesoma within Aspidosiphonidae. These results are not affected by the loci selected or by the outgroup chosen. The position of Apionsoma is discussed, although more data would be needed to better ascertain its phylogenetic affinities. Monophyly of the genera with multiple representatives (Themiste, Aspidosiphon, and Phascolosoma) is well supported, but not the monophyly of the genera Nephasoma or Golfingia. Interesting phylogeographic questions arise from analysis of multiple representatives of a few species.     

Phylogenetic analysis based on 18S ribosomal RNA gene sequences supports the existence of class polyacanthocephala (acanthocephala)

Members of phylum Acanthocephala are parasites of vertebrates and arthropods and are distributed worldwide. The phylum has traditionally been divided into three classes, Archiacanthocephala, Palaeacanthocephala, and Eoacanthocephala; a fourth class, Polyacanthocephala, has been recently proposed. However, erection of this new class, based on morphological characters, has been controversial. We sequenced the near complete 18S rRNA gene of Polyacanthorhynchus caballeroi (Polyacanthocephala) and Rhadinorhynchus sp. (Palaeacanthocephala); these sequences were aligned with another 21 sequences of acanthocephalans representing the three widely recognized classes of the phylum and with 16 sequences from outgroup taxa. Phylogenetic relationships inferred by maximum-likelihood and maximum-parsimony analyses showed Archiacanthocephala as the most basal group within the phylum, whereas classes Polyacanthocephala + Eoacanthocephala formed a monophyletic clade, with Palaeacanthocephala as its sister group. These results are consistent with the view of Polyacanthocephala representing an independent class within Acanthocephala.     

A modern approach to rotiferan phylogeny: combining morphological and molecular data

The phylogeny of selected members of the phylum Rotifera is examined based on analyses under parsimony direct optimization and Bayesian inference of phylogeny. Species of the higher metazoan lineages Acanthocephala, Micrognathozoa, Cycliophora, and potential outgroups are included to test rotiferan monophyly. The data include 74 morphological characters combined with DNA sequence data from four molecular loci, including the nuclear 18S rRNA, 28S rRNA, histone H3, and the mitochondrial cytochrome c oxidase subunit I. The combined molecular and total evidence analyses support the inclusion of Acanthocephala as a rotiferan ingroup, but do not support the inclusion of Micrognathozoa and Cycliophora. Within Rotifera, the monophyletic Monogononta is sister group to a clade consisting of Acanthocephala, Seisonidea, and Bdelloidea-for which we propose the name Hemirotifera. We also formally propose the inclusion of Acanthocephala within Rotifera, but maintaining the name Rotifera for the new expanded phylum. Within Monogononta, Gnesiotrocha and Ploima are also supported by the data. The relationships within Ploima remain unstable to parameter variation or to the method of phylogeny reconstruction and poorly supported, and the analyses showed that monophyly was questionable for the families Dicranophoridae, Notommatidae, and Brachionidae, and for the genus Proales. Otherwise, monophyly was generally supported for the represented ploimid families and genera.     

A molecular phylogeny of Heterodonta (Bivalvia) based on small ribosomal subunit RNA sequences

Within Heterodonta, phylogenesis has so far been studied almost exclusively on the basis of morphological data. Results have often been discordant, and an exhaustive molecular approach has not yet been attempted. The present study was undertaken to clarify the phylogenetic relationships obtaining among Heterodonta families through the analysis of 18S rRNA gene. To do this, the whole sequence of this gene was analyzed in 29 species of eight superfamilies of the order of Veneroida (Arcticoidea, Cardioidea, Galeommatoidea, Mactroidea, Solenoidea, Tellinoidea, Tridacnoidea, and Veneroidea) and in two superfamilies of Myoida (Pholaloidea and Myoidea). The study was extended by constructing phylogenetic trees using partial sequences. This strategy made it possible to include 11 additional species by introducing three further superfamilies: Chamoidea, Corbiculoidea, and Hiatellinoidea. At variance with the conclusions reached on the basis of morphological features, the molecular data clearly show that the Myoida species included in this study belong to Veneroida, thus undermining the legitimacy of the division of Heterodonta into two orders, and that considerable differences in the phylogenetic relationships obtain among superfamilies.     

Localization of chitin in the cuticle of tardigrada using wheat germ agglutinin—Gold conjugate as a specific electron-dense marker

Wheat germ agglutinin, a lectin with binding sites specific for N-acetyl-glucosamine residues, was coupled with colloidal gold as an electron-dense marker and used for the electron microscopical localization of chitin in the cuticles of Echiniscus testudo (Heterotardigrada, Echiniscoidea), Macrobiotua hufelandi (Eutardigrada, Parachela) and Milnesium tardigradum (Eutardigrada, Apochela). In all species investigated the cuticular layer next to the epidermis, the procuticle, was labelled, and in Milnesium tardigradum additionally the so-called intracuticle. In Echiniscus testudo only parts of the dorsal intracuticle showed some additional labelling. The other cuticular layers (ventral intracuticle in Echiniscus testudo and epicuticles of all species investigated) remained unlabelled. Specificity of binding was controlled by competitive inhibition with triacetyl chitotriose. The results are discussed with regard to a chitin-containing procuticle and the occurrence of the intracuticle in Tardigrada.     

Molecular phylogeny of the Byrrhoidea–Buprestoidea complex (Coleoptera,Elateriformia)

The superfamilies of Elateriformia have been in a state of flux since their establishment. The recent classifications recognize Dascilloidea, Buprestoidea, Byrrhoidea and Elateroidea. The most problematic part of the elateriform phylogeny is the monophyly of Byrrhoidea and the relationships of its families. To investigate these issues, we merged more than 500 newly produced sequences of 18S rRNA, 28S rRNA, rrnL mtDNA and cox1 mtDNA for 140 elateriform taxa with data from GenBank. We assembled an all‐taxa (488 terminals) and a pruned data set, which included taxa with full fragment representation (251 terminals); both were aligned in various programs and analysed using maximum‐likelihood criterion and Bayesian inference. Most analyses recovered monophyletic superfamilies and broadly similar relationships; however, we obtained limited statistical support for the backbone of trees. Dascilloidea were sister to the remaining Elateriformia, and Elateroidea were sister to the clade of byrrhoid lineages including Buprestoidea. This clade mostly consisted of four major lineages, that is (i) Byrrhidae, (ii) Dryopidae + Lutrochidae, (iii) Buprestoidea (Schizopodidae sister to Buprestidae) and (iv) a clade formed by the remaining byrrhoid families. Buprestoidea and byrrhoid lineages, with the exception of Byrrhidae and Dryopidae + Lutrochidae, were usually merged into a single clade. Most byrrhoid families were recovered as monophyletic. Callirhipidae and Eulichadidae formed independent terminal lineages within the Byrrhoidea–Buprestoidea clade. Paraphyletic Limnichidae were found in a clade with Heteroceridae and often also with Chelonariidae. Psephenidae, represented by Eubriinae and Eubrianacinae, never formed a monophylum. Ptilodactylidae were monophyletic only when Paralichas (Cladotominae) was excluded. Elmidae regularly formed a clade with a bulk of Ptilodactylidae; however, elmid subfamilies (Elminae and Larainae) were not recovered. Despite the densest sampling of Byrrhoidea diversity up to date, the results are not statistically supported and resolved only a limited number of relationships. Furthermore, questions arose which should be considered in the future studies on byrrhoid phylogeny.     

Molecular phylogeny of the Calanoida (Crustacea: Copepoda)

The order Calanoida includes some of the most successful planktonic groups in both marine and freshwater environments. Due to the morphological complexity of the taxonomic characters in this group, subdivision and phylogenies have been complex and problematic. This study establishes a multi-gene molecular phylogeny of the calanoid copepods based upon small (18S) and large (28S) subunits of nuclear ribosomal RNA genes and mitochondrial encoded cytochrome b and cytochrome c oxidase subunit-I genes, including 29 families from 7 superfamilies of the order. This analysis is more comprehensive than earlier studies in terms of number of families, range of molecular markers, and breadth of taxonomic levels resolved. Patterns of divergence of ribosomal RNA genes are shown to be significantly heterogeneous among superfamilies, providing a likely explanation for disparate results of previous studies. The multi-gene phylogeny recovers a monophyletic Calanoida, as well as the superfamilies Augaptiloidea, Centropagoidea, Bathypontioidea, Eucalanoidea, Spinocalanoidea and Clausocalanoidea. The phylogeny largely agrees with previously-published morphological phylogenies, including e.g., enlargement of the Bathypontioidea to include the Fosshageniidae.     

Congruence between molecular phylogeny and cuticular design in Echiniscoidea (Tardigrada,Heterotardigrada)

Although morphological characters distinguishing echiniscid genera and species are well understood, the phylogenetic relationships of these taxa are not well established. We thus investigated the phylogeny of Echiniscidae, assessed the monophyly of Echiniscus, and explored the value of cuticular ornamentation as a phylogenetic character within Echiniscus. To do this, DNA was extracted from single individuals for multiple Echiniscus species, and 18S and 28S rRNA gene fragments were sequenced. Each specimen was photographed, and published in an open database prior to DNA extraction, to make morphological evidence available for future inquiries. An updated phylogeny of the class Heterotardigrada is provided, and conflict between the obtained molecular trees and the distribution of dorsal plates among echiniscid genera is highlighted. The monophyly of Echiniscus was corroborated by the data, with the recent genus Diploechiniscus inferred as its sister group, and Testechiniscus as the sister group of this assemblage. Three groups that closely correspond to specific types of cuticular design in Echiniscus have been found with a parsimony network constructed with 18S rRNA data. © 2013 The Linnean Society of London     

A molecular phylogeny of the Percidae (Teleostei, Perciformes) based on mitochondrial DNA sequence

The family Percidae is among the most speciose families of northern hemisphere fishes with > 178 178 North American species and 14 Eurasian species. Previous phylogenetic studies have been hampered by a lack of informative characters, inadequate taxonomic sampling, and conflicting data. We estimated phylogenetic relationships among 54 percid species (9 of 10 genera and all but one subgenus of darters) and four outgroup taxa using mitochondrial DNA data from the 12S rRNA and cytochrome b genes. Four primary evolutionary lineages were consistently recovered: Etheostomatinae (Ammocrypta, Crystallaria, Etheostoma, and Percina), Perca, Luciopercinae (Romanichthys, Sander, and Zingel), and Gymnocephalus. Except Etheostoma and Zingel, all polytypic genera were monophyletic. The Etheostoma subgenus Nothonotus failed to resolve with other members of the genus resulting in a paraphyletic Etheostoma. The subfamily Percinae (Gymnocephalus and Perca) was not recovered in phylogenetic analyses with Gymnocephalus sister to Luciopercinae. Etheostomatinae and Romanichthyini were never resolved as sister groups supporting convergent evolution as the cause of small, benthic, stream-inhabiting percids in North American and Eurasian waters.     

Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification

Grapsoid and ocypodoid crabs receive a lot of attention in the literature due to their predominance and important role as primary and secondary consumers in intertidal as well as supratidal marine habitats. They are especially species‐rich in the tropics, where they have been found to repeatedly invade terrestrial and freshwater habitats. However, the systematics of the crabs belonging to these two superfamilies is still not settled, despite recent steps clarifying phylogenetic relationships and introducing new taxa. In this study, a molecular phylogeny of grapsoid crabs primarily based on East African representatives is constructed based on DNA sequences of the mitochondrial small and large ribosomal subunits (12S and 16S rRNA), thus complementing previous molecular taxonomic studies that had been carried out with the American and East Asian fauna. In addition, selected representatives of all ocypodoid families and subfamilies were included. The monophyly of Grapsidae, Ocypodidae (sensu stuctu), Sesarmidae and Varunidae is well confirmed, if the genera Cyclograpsus, Helice are considered Varunidae and Euchirograpsus a Plagusiidae, as previously suggested. The monophyly of the family Gecarcinidae cannot be supported with our data. The family Plagusiidae in its present composition is polyphyletic. Special attention was given to the large family Sesarmidae, which has many endemic genera in the Indo‐West Pacific. According to this study, two of the most speciose genera, Chiromantes and Parasesarma, are not monophyletic and need to be redefined. On the higher taxonomic level, it becomes evident that both superfamilies, Grapsoidea and Ocypodoidea, are not monophyletic in their current composition, as exemplified by a proposed sister group relationship of Varunidae and Macrophthalmidae. These results confirm those from previous molecular studies and we therefore propose to refrain from the traditional use of the Grapsoidea and Ocypodoidea as monophyletic superfamilies and treat the constituent families separately.