A molecular phylogeny of Bopyroidea and Cryptoniscoidea (Crustacea: Isopoda)

Epicaridean isopods are parasitic on other crustaceans. They represent a diverse group of highly derived taxa in two superfamilies and 10 families. Little work has been done on the phylogeny of these parasites because of the difficulty in defining homologous characters for adults above the genus level. The females exhibit morphological reduction of characters and the males have few distinguishing characters. Moreover, epicarideans have only rarely been included in past studies of isopod phylogeny. Our objective was to derive a phylogeny of epicaridean taxa based on 18S rDNA, then use that phylogeny to examine the relationships of the bopyrid subfamilies, bopyroid families and epicarideans to cymothoid isopods. We tested the monophyly of the Epicaridea, evaluated hypotheses on relationships among epicaridean families and subfamilies, examined the evolution of the abdominal mode of infestation on caridean, gebiidean, axiidean and anomuran hosts and examined coevolution between epicarideans and their crustacean hosts. The molecular phylogeny indicated that Epicaridea were monophyletic with respect to Cymothooidea. Bopyroidea formed a monophyletic group without Dajidae and Entophilinae (now as Entophilidae). Both latter taxa grouped with Cryptoniscoidea, and this group was the sister taxon to the redefined Bopyroidea in all trees. The bopyrid subfamily Ioninae is the sister taxon to the other bopyrid subfamilies (except Entophilidae). Ioninae was elevated to family status but found not to be monophyletic; a new subfamily, Keponinae, was erected for all genera formerly placed in Ioninae except the type genus. The abdominal mode of parasitism appears to have evolved independently among the subfamilies. Coevolution between host and parasite phylogenies showed extensive incongruence, indicating frequent host-switching as a general pattern in Epicaridea.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:30ECFB13-2795-494E-AABE-6B5F84A57A67     

Searching factors causing implausible non-monophyly: ssu rDNA phylogeny of Isopoda Asellota (Crustacea: Peracarida) and faster evolution in marine than in freshwater habitats

This contribution addresses two questions: which alignment patterns are causing non-monophyly of the Asellota and what is the phylogenetic history of this group? The Asellota are small benthic crustaceans occurring in most aquatic habitats. In view of the complex morphological apomorphies known for this group, monophyly of the Asellota has never been questioned. Using ssu rDNA sequences of outgroups and of 16 asellote species from fresh water, littoral marine habitats and from deep-sea localities, the early divergence between the lineages in fresh water and in the ocean, and the monophyly of the deep-sea taxon Munnopsidae are confirmed. Relative substitution rates of freshwater species are much lower than in other isopod species, rates being highest in some littoral marine genera (Carpias and Jaera). Furthermore, more sequence sites are variable in marine than in freshwater species, the latter conserve outgroup character states. Monophyly is recovered with parsimony methods, but not with distance and maximum likelihood analyses, which tear apart the marine from the freshwater species. The information content of alignments was studied with spectra of supporting positions. The scarcity of signal (=apomorphic nucleotides) supporting monophyly of the Asellota is attributed to a short stem-line of this group or to erosion of signal in fast evolving marine species. Parametric boostrapping in combination with spectra indicates that a tree model cannot explain the data and that monophyly of the Asellota should not be rejected even though many topologies do not recover this taxon.     

Morphology and evolution of respiratory structures in the pleopod exopodites of terrestrial Isopoda (Crustacea, Isopoda, Oniscidea)

The morphology of the respiratory structures located in the pleopod exopodites of terrestrial Isopoda is described for representative species of different genera. Special emphasis is set on the evolution of these lungs in the context of phylogenetic relationships as revealed by other morphological characters. In the most primitive species of the Oniscidea, and still of subordinate taxa of the Crinocheta, respiration takes place in the thin ventral integument of the exopodites. The evolution of dorsal respiratory fields with a weakly wrinkled surface occurred at least six times within the Crinocheta. In five of these cases, a further development can be observed. The evolution of a partly covered respiratory field with strongly wrinkled surface may have taken place six times. It is assumed that completely internalized lungs with spiracles surrounded by a water‐repellent surface microsculpture, evolved at least six times independently within the Oniscidea: in the Tylidae, Actaecia, Aphiloscia, the Eubelidae, the Armadillidae and in a taxon probably comprising Porcellionidae plus Armadillidiidae.     

Review of methodological problems of 'Computer cladistics' exemplified with a case study on isopod phylogeny (Crustacea: Isopoda)

Using a computerized phylogenetic analysis of the Isopoda (Crustacea: Peracarida) as source of typical errors and misunderstandings, problems that may occur in computer cladistics are reviewed. It is concluded that in addition to the errors that are possible in a conventional Hennigian analysis some specific methodological problems exist in computer cladistics. It is recommended that the OTU be replaced by the groundpattern concept. Tree statistics are not useful for comparing different competing hypotheses. Arguments ought to concentrate on the hypo-thetico-deductive steps of the analysis, i.e. on character analysis. The use of computers does not add objectivity to character analysis. Single outgroup taxa should not be used in assessing the character states of ingroups. Concerning isopod phylogeny, it is argued here that the tail fan of the Isopoda can probably be derived from the eumalacostracan groundpattern and did not evolve de novo within the Isopoda.     

The ultrastructure of the heart of Oniscus asellus L. and Asellus aquaticus L. (Crustacea,Isopoda)

Summary The endocardium of Oniscus asellus L. and Asellus aquaticus L. consists of lipid cells. The epicardium consists of a layer of cells with a vesiculated cytoplasm covered by a thick extracellular fibrous sheet. The myocardium is a single layer of cells, the sarcolemma invaginates at Z disc level forming transverse tubules, and longitudinal tubules branch off from these. At the A-I level' longitudinal tubules form transverse systems, which form couplings with the sarcoplasmic reticulum. The sarcoplasmic reticulum appears as perforated sheets enveloping the myofibrils. Two types of nerve terminal are found: one is embedded in a myocardial cell process, the other lies in a myocardial cell depression. They contain clear and dense-cored synaptic vesicles.This work was supported by grants from the Norwegian Research Council for Science and the Humanities     

Morphology and evolution of the respiratory apparatus in the family Eubelidae (Crustacea,Isopoda, Oniscidea)

The morphology of the respiratory apparatus in the pleopodal lungs of the family Eubelidae was investigated. The family is a monophyletic group including more than 240 species in 53 genera (three of which are nomina dubia), mostly distributed in the Afrotropical Region (tropical Africa and Arabian Peninsula). In all the Eubelidae, except for the monospecific genus Parelumoides and two species of the genus Elumoides, the exopods of pleopods have lungs. All the pulmonary morphologies present in the entire suborder Oniscidea are found: 1) uncovered lungs, composed of a pleated respiratory surface, directly exposed to the air (Atracheodillo-type) or partially enclosed within the appendage (Synarmadilloides-type); 2) covered lungs with several spiracles and respiratory trees, housed within the appendages, with spiracles surrounded by a specialized, nonrespiratory, structure (perispiracular area) (Eubelum- and Somaloniscus-types); 3) covered lungs with only one spiracle, with or without perispiracular area, and one respiratory tree (Aethiopopactes- and Periscyphis-types), which in taxa with Periscyphis-type lung crosses the insertion of the appendage and penetrates into the pleon with bundles of respiratory tubules. The evolution of the various types of lungs is discussed. It is concluded that the two main evolutionary lines, i.e., uncovered lungs and covered lungs, originated independently from an ancestral respiratory structure-the semilunar area. A first mechanism of development of the semilunar area by folding of its surface produced the Atracheodillo-type (all folds coplanar with the surface of the exopod) and Synarmadilloides-type (folds partly coplanar and partly intraflexed inside the exopod) uncovered lungs. A second mechanism of development by tubular invagination of the cuticle of the semilunar area produced the polyspiracular Eubelum-type lungs (numerous arborescent invaginations) and the monospiracular Aethiopopactes-type lungs (only one arborescent invagination), probably passing through a common intermediate pattern. From the common pattern, both the poly- and monospiracular types would have inherited the characteristic concave cell arrangement of the perispiracular area. The Somaloniscus-type and Periscyphis-type lungs are forms specialized for arid environments, directly derived from the Eubelum-type and Aethiopopactes-type, respectively.     

Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater

The isopod crustaceans are diverse both morphologically and in described species numbers. Nearly 950 described species (∼9% of all isopods) live in continental waters, and possibly 1,400 species remain undescribed. The high frequency of cryptic species suggests that these figures are underestimates. Several major freshwater taxa have ancient biogeographic patterns dating from the division of the continents into Laurasia (Asellidae, Stenasellidae) and Gondwana (Phreatoicidea, Protojaniridae and Heterias). The suborder Asellota has the most described freshwater species, mostly in the families Asellidae and Stenasellidae. The suborder Phreatoicidea has the largest number of endemic genera. Other primary freshwater taxa have small numbers of described species, although more species are being discovered, especially in the southern hemisphere. The Oniscidea, although primarily terrestrial, has a small number of freshwater species. A diverse group of more derived isopods, the ‘Flabellifera’ sensu lato has regionally important species richness, such as in the Amazon River. These taxa are transitional between marine and freshwater realms and represent multiple colonisations of continental habitats. Most species of freshwater isopods species and many genera are narrow range endemics. This endemism ensures that human demand for fresh water will place these isopods at an increasing risk of extinction, as has already happened in a few documented cases. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Evolutionary biogeography of the Microparasellid isopod Microcharon (Crustacea) in the Mediterranean Basin

The interstitial subterranean isopod Microcharon (Crustacea) is highly diversified within the Mediterranean. The present distribution of 70 species is considered the result of the joint history of this genus and its environment. This stygobiont is derived from marine surface ancestors. Both Plate Tectonics and the two-step model of colonization and evolution, the second phase of which represents vicariance during the Tethys regressions, enable an understanding of the evolutionary history of this monophyletic group. The single extant marine littoral species was separated from its sister group by the closure of the Gibraltar Straits during the Messinian. The drift of the Apulian plate, the Alboran-Kabylian-Calabrian fragment as well as the opening of the western Mediterranean through the translation of the Corsican-Sardinia plate could explain the divergences in the western and the eastern Mediterranean. The primitive sister groups in Morocco and in Spain would have been left by the Turonian Tethys regression. Such regressions of the Tethys embayments could have played a major role in the evolution of the genus from Turanian up to Tortonian in Norht Africa, and up to Pliocene and even to Pleistocene in Italian, Iberian and Greek zones. Several eastern European species may be a product of the Paratethys regressions. Everywhere, species with plesiomorphic characters are related to old regressions and vice versa. Local geological events could have resulted in further vicariance divergences. There is a good congruence between the evolutionary history of Microcharon and other stygobiont isopod, amphipods or syncarids living in the same regions studied in the Mediterranean.Presented at the Symposium of Biogeography at Subterranean Crustaceans: the effects of different scales (June 1992; Charleston, South Carolina, U.S.A.). Manuscrips prepared for print by J. R. Holsinger.     

A Phylogenetic Analysis of Ilyocryptus Sars, 1862 (Cladocera: Ilyocryptidae)

Kotov and Štifter (2006) revised the taxonomy of the genus Ilyocryptus Sars, 1862 (Branchiopoda: Anomopoda: Ilyocryptidae) and concluded that 28 species can be recognized as valid for the world fauna. In order to test phylogenetic relationships between the species and to contribute to a better understanding of the genus, a cladistic (using branch‐and‐bound search) analysis was conducted for 25 Ilyocryptus species (two of them with two subspecies). In total, 32 morphological characters were used. An evolutionary‐morphological interpretation of the results was also made. Phylogenetic analysis revealed that Ilyocryptus elegans is an earlier derived member of the genus, while other species are separated into two main branches: agilis ‐ and sordidus ‐lines (without reference to type of moulting). Species with incomplete moulting and species with complete moulting do not form separate clusters, suggesting an independent origin of incomplete moulting within different species groups. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Structural investigation of the female genitalia and sperm-storage sites in the terrestrial isopod Armadillidium vulgare (Crustacea, Isopoda)

The cuticular genitalia of the terrestrial isopod, Armadillidium vulgare, have two distinct states during the reproductive cycle of the females. The structural differences between the reproductive and non-reproductive states, and the structure of the sperm storage sites were investigated employing electron and light microscopy. In both states the genitalia consist of a distal segment that connects to the gonopore, and a cuticular tube-like structure lining the lumen of the oviduct in the middle region of the oviduct. Sheath-like projections, apparently consisting of cuticular material, extend laterally along two sides of the cuticular tube. In the proximal region of the oviduct cuticular structures are lacking. In the non-reproductive state the distal segment consists of endo-, exo- and epicuticle. The exocuticle is three layered with unusual spongy and dense layers at the distal side. On one side the endocuticle doubles in thickness to form a cuticular bulge that fills the lumen of the distal segment leaving just a narrow U-shaped space. The cuticular tube consists of endo- and epicuticle only. In the reproductive state the distal segment is funnel-shaped and forms branched cuticular folds that increase in complexity from distal to proximal. In the cuticular tube these folds tightly fill the lumen of the oviduct. At the confluence of the oviduct with the ovary spermatozoa are stored in a seminal receptacle.     

Cirolana songkhla,a new species of brackish-water cirolanid isopod (Crustacea,Isopoda, Cirolanidae) from the lower Gulf of Thailand

Cirolana songkhla sp. n. was collected from brackish-water habitats including lagoons and estuaries in the coastal zone of the lower Gulf of Thailand. C. songkhla sp. n. is described and fully illustrated; C. songkhla sp. n. can be recognized by the presence of abundant chromatophores dorsally, lack of ornamentation on the posterior pereonites, pleonites and pleotelson, the number of robust setae on the uropodal and pleotelson margins (uropod exopod lateral margin with 12–14 RS, mesial margin with 5–8 RS; endopod lateral margin with 8–10 RS, mesial margin with 11–13 RS; pleotelson with 12–15 RS) and lack of setae on the endopods of pleopods 3–5. A dichotomous key of brackish Cirolana species in Thailand is given.     

Phylogeny and biogeography of the oil-collecting bee subgenus Centris (Wagenknechtia) (Hymenoptera,Apidae)

Centris (Wagenknechtia) is the only subgenus of centridine bees that occurs almost exclusively in the Andean Region. This study investigates the monophyly of C. (Wagenknechtia) proposing a hypothesis for the relationships among its species using 42 morphological characters of adults. The analysis resulted in one most parsimonious tree corroborating the monophyly of this group of oil-collecting bees, with the relationships among its species as follows: ((C. escomeli + C. moldenkei) (C. muralis (C. rhodophthalma + C. vardyorum) (C. cineraria + C. orellanai))). Based on these results plus available distributional records, a cladistic biogeographic analysis was performed. The resulting hypothesis of the relationships between the endemism areas in the Andean Region is as follows: (((Atacama–Puna) (Desert–Atacama–Puna)) ((Pampean–Chaco–Puna) ((Monte–Coquimban) (Coquimban (Patagonian–Valdivian Forest–Coquimban–Maule–Monte–Santiago–Magellanic Forest))))).     

The genus Coperonus Wilson, 1989 (Crustacea: Isopoda: Asellota: Munnopsidae)

This paper presents a revision of the natatory isopod genus Coperonus (Munnopsidae, Lipomerinae). A key to identify all seven known species of the genus is provided. Four new species are described and Coperonus frigida (Vanhöffen, 1914) is redescribed on the basis of the type material. The biogeography of Coperonus is described.
In dieser Arbeit wird die Gattung Coperonus (Munnopsidae, Lipomerinae), deren Vertreter sich durch gute Schwimmfähigkeit auszeichnen, revidiert. Ein Bestimmungsschlüssel für alle sieben bisher bekannten Arten ist in der Arbeit enthalten. Es werden vier neue Arten beschrieben und Coperonus frigida (Vanhöffen, 1914) wird anhand des Typenmaterials nachbeschrieben. Auβerdem wird in der Arbeit die Biogeographie von Coperonus dargestellt.     

Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Thereavidae)

Twenty-one members of the Laurasian group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hypothesis of relationships among taxa within the Gyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Anolinga , gen. nov. , Breviperna Irwin, Coleiana , gen. nov. , Crebraseta , gen. nov. , Cyclotelus Walker, Mesonana , gen.nov. , and Ozodiceromyia Bigot. In addition, three Old World genera, Ammothereva Lyneborg, Bugulaverpa , gen. nov. , and Procyclotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Brevipema-group and the Cyclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles.     

Burrow plasticity in the deep-sea isopod Bathynomus doederleini (Crustacea: Isopoda: Cirolanidae)

We investigated whether the deep-sea isopod Bathynomus doederleini has the capacity to change burrow length in response to changes in environmental conditions. We observed burrowing behavior in individuals that were placed on substrates with either simple (ST) or complex (CT) surface topographies. Individuals in the ST group (N = 10) constructed seven burrows. The mean ratio of the burrow length to body length was 1.8. The individuals in the CT group (N = 10) constructed eight burrows with a mean ratio of burrow length to body length of 2.5. Thus the burrows were significantly longer in the CT group. In addition, the isopods in the CT group often incorporated a chamber in the mid-section of the burrow. Our results may be used to infer the determinants of burrow morphology and speculate about the lifestyle of this species in the deep sea.     

The morphology of the sphaeromatid brood pouch (Crustacea: Isopoda: Sphaeromatidae)

The structure of the brood pouch and mouthparts of ovigerous female sphaeromatids is described. Taxonomically, these are shown to be important generic characters and some attempt is made to relate structure to function. Expanded maxillipeds (which produce a current of water through the marsupium) are shown to occur only in genera which have oostegites overlapping in the mid-line and reaching the mouthparts anteriorly. The reduction of the mouthparts of some genera cannot be explained, but it may conserve energy at the final moult. No platybranchiate genera (subfamily Cassidininae) have the mouthparts reduced but some have the maxilliped expanded. Some correlation between brood pouch structure and habitat is suggested. Genera with a brood pouch composed only of oostegites (and hence liable to disruption by turbulence and particle abrasion) occur sublittorally or, if intertidal, adopt a protected, crevice dwelling habit. Genera with a brood pouch composed of large opposing pockets (which are more able to resist disruption) occur intertidally or in shallow water, and may be found in wave-washed sand. Some wood-boring species of the genus Sphaeroma Latreille show a reduced number of oostegites when compared with non-boring species. This presumably relates to their burrowing habit.     

Microscopic anatomy and ultrastructure of the digestive system of Natatolana obtusata (Vanhöffen, 1914) (Crustacea, Isopoda)

The scavenging cirolanid isopod Natatolana obtusata was caught using primarily meat‐baited traps at a station in the Weddell Sea at Kap Norvegica. The hindguts of most specimens were densely stuffed with meat from the traps. The anatomy and the ultrastructure of different sections of the digestive system are described. The large and extremely expandable mouth opens into a short oesophagus which extends posteriorly into a small stomach. Lateralia and primary filter can be observed directly when the mouth opening is widened to its extreme. The anterior hindgut is extremely variable in its lumen and very distensible, depending on the nutritional condition. The cuticle of the hindgut is colonized by fungi. The length of the midgut glands varies with the nutritional conditions. Morphology and anatomy of the digestive system of Natatolana obtusata are discussed with the background of the lifestyle of the species and compared with other taxa.