Phylogenetic analysis of the Malacostraca (Crustacea)

The Malacostraca comprises about 28 000 species with a broad disparity in morphology, anatomy, embryology, behaviour and ecology. The phylogenetic relationships of the major taxa are still under debate. Is the Leptostraca the sister group of the remaining Malacostraca, or is this taxon more closely related to other Crustacea? Does the Stomatopoda or the Bathynellacea represent the most basal taxon within the remaining taxa? Is the Peracarida monophyletic or are some peracarid taxa more closely related to other ‘caridoid’ taxa? Is the Thermosbaenacea part of the Peracarida or its sister group, and how much support is there for a taxon Amphipoda + Isopoda? To answer these questions a phylogenetic analysis of the Malacostraca combining different phylogenetic approaches was undertaken. In a first step, the monophyly of the Malacostraca including the Leptostraca is shown using the ‘Hennigian approach’. A computer cladistic analysis of the Malacostraca was carried out with NONA and PEE ‐WEE , based on 93 characters from morphology, anatomy and embryology. Nineteen higher malacostracan taxa are included in our analysis. Taxa whose representatives are exclusively fossils were not included. The Leptostraca was used as an operational out‐group. The present analysis supports the basal position of the Stomatopoda. Syncarida and Peracarida (including Thermosbaenacea) are supported as monophyletic, the Eucarida is not. Instead a sister‐group relationship is suggested between Euphausiacea and Peracarida (including Thermosbaenacea), with the Syncarida as the sister group to both taxa. Certain embryonic characters are interpreted as support for the monophyly of the Peracarida (without Thermosbaenacea) because convergences or reversals of these characters seem implausible. Within the Peracarida, the Mysidacea (Lophogastrida + Mysida) represents the sister group to the remaining taxa. A sister‐group relationship between Amphipoda and Isopoda is not supported.     

The disunity of "Mysidacea" (Crustacea)

New studies on malacostracan relationships have drawn attention to issues concerning monophyly of the order Mysidacea, manifested in recent crustacean classifications that treat the taxon as two separate orders, Lophogastrida and Mysida. We present molecular phylogenies of these orders based on complete sequences of nuclear small-subunit ribosomal DNA (18S rRNA), and morphological evidence is used to revise the classification of the order Mysida to better reflect evolutionary history. A secondary structure model for 18S rRNA was constructed and used to assign putative stem and loop regions to two groups of partitions for phylogenetic analyses. Phylogenies were estimated by maximum-likelihood, Bayesian inference, and maximum-parsimony. The analyses gave strong support for three independently derived lineages, represented by three monophyletic groups, Lophogastrida, Stygiomysida, and Mysida. The family Petalophthalmidae is considered as sister group to the family Mysidae, and Boreomysinae and Rhopalophthalminae are the most early derived of the Mysidae. The tribes contained in the current classification of the subfamily Mysinae are not well-supported by either molecular data or morphology.     

Spermiogenesis in Anaspides tasmaniae (Thomson) (Crustacea,Malacostraca, Syncarida)

Spermiogenesis of the syncarid Anaspides tasmaniae (subclass Eumalacostraca) was investigated by transmission electron microscopy. The spermatozoan of Anaspides is an ovoid cell with an acrosome covering the anterior pole and a lobulated nucleus and mitochondria occupying the rest of the cell. A long subacrosomal filament bypasses the nucleus and forms a spiral that supports a thin extension of the posterior cytoplasm, giving the spermatozoan a bell-shaped appearance. No flagellum is present at any stage. The immobile spermatozoans are embedded in a hard capsule, secreted by the cells of the wall of the vas deferens.     

Ommochrome pigments in the eyes of Euphausia superba (Crustacea,Euphausiacea)

Summary Ommin and a xanthommatin-like ommochrome pigment have been extracted from the eyes of Euphausia superba, and identified by their absorbance spectra and redox reactions invarious solvents. The absorbance spectrum of ommin is relatively stable, but that of the xanthommatin pigment is sensitive to light, pH, and hydroxylamine. Both pigments are readily soluble in 2% digitonin. They are therefore likely contaminants of visual pigment extracts, and may consequently introduce artifacts into visual pigment characterizations. Microspectrophotometric measurements show that both pigments are components of the distal and proximal screening pigment granules in E. superba ommatidia.This paper is dedicated to the memory of Professor Mary Alice McWhinnie, to whom I owe a great debt of gratitude for her patient tutelage and the opportunity to participate in her Antarctic research, for the encouragement to pursue my own interests, and for her generous friendship. It was a cherished association which ended sadly with her untimely death in March, 1980. She will always be fondly remembered     

Comparative analysis of the circulatory system in Amphipoda (Malacostraca, Crustacea)

We present data on the haemolymph vascular system (HVS) in four representatives of the major amphipod lineages Gammaridea, Hyperiidea and Caprellidea based on corrosion casting and three‐dimensional reconstructions of histological semi‐thin sections. In all these species the HVS comprises a dorsal pulsatile heart, which is continued in the body axis by the anterior and posterior aortae. The heart is equipped with three pairs of incurrent ostia. The number of cardiac arteries that lead off the heart varies among species: in the studied Gammaridea four pairs occur, in Hyperia galba only the three posterior pairs of cardiac arteries occur, while in Caprella mutica cardiac arteries are absent. In all the studied species the posterior aorta leads as a simple tube into the pleon attached to the dorsal diaphragm. The anterior aorta runs from its origin in the anterior part of the second thoracic segment into the cephalothorax. Both pairs of antennae have an arterial supply off the anterior aorta. An overview of previously studied species including our present findings shows the amphipod HVS to be relatively uniform and the gammarid form is discussed as being closest to the ground pattern of Amphipoda.     

Comparative morphology and actuopalaeontology of mysid statoliths (Crustacea,Mysidacea)

Summary A comprehensive and comparative study of the external statolith morphology of the family Mysidae is presented. The study covers 48 species from major systematic groups occupying a large number of habitats in different biogeographical zones of the globe. Statoliths generally show high morphological diversity. The traditional classification scheme of subtaxa and the correlation of statolith characters with segmentation patterns of body appendages suggest that the organic composition and the nearly spherical structure of the statoliths of Boreomysinae and Rhopalophthalminae are plesiomorphic compared with the more complex mineralized statoliths found in all other subfamilies. During ontogenetic development the number of sensorial setae and associated pores and pore groups on the statolith increase with body size and statolith diameter. Although patterns of caudal pores are highly specific for some genera, the high intraspecific variance of pore numbers strongly reduces the diagnostic value of this feature in most species. Statolith characters can be successfully used for identification of subfamilies, tribes, and especially genera. For future palaeontological applications a proper diagnosis of fossil mysid statoliths is essential. Therefore, we provide a key to subfamilies and tribes based exclusively on statolith characters.     

DNA barcodes for species identification of euphausiids (Euphausiacea, Crustacea)

Many species of euphausiids (Euphausiacea, Crustacea) are distinguishedby subtle or geographically variable morphological characters,and erroneous identification of euphausiid species may be morefrequent than currently acknowledged. DNA barcodes (short DNAsequences that discriminate species and aid in recognition ofunknown species) are of use for this group. A 650 bp regionof mitochondrial cytochrome oxidase I (mtCOI) was sequencedfor 40 species of 10 euphausiid genera: Bentheuphausia, Euphausia,Meganyctiphanes, Nematobrachion, Nematoscelis, Nyctiphanes,Stylocheiron, Tessarabrachion, Thyssanoessa and Thysanopoda.mtCOI sequence variation discriminated all species; pairwisedifferences averaged 16.4% (range 7–24%); mean generalizedtime reversible (GTR) genetic distance was 26.7%. mtCOI reliablyidentified euphausiid species: variation within species wastypically < 1% and GTR distance was typically < 2%. Atlanticand Pacific Ocean populations of Euphausia brevis differed by13% (GTR genetic distance = 28%) and may deserve status as distinctspecies. mtCOI gene trees were reconstructed for five generausing maximum parsimony, maximum likelihood and Bayesian algorithms;best-fit models of nucleotide evolution were determined foreach genus. The mtCOI gene tree for 20 species of Euphausiareproduced one of three morphologically defined species groups.mtCOI resolved relationships among closely related species ofmost genera, usually in accord with morphological groupings.A comprehensive DNA barcode database for euphausiids will helpensure accurate species identification, recognition of crypticspecies and evaluation of taxonomically meaningful geographicvariation.     

Limb ontogeny and trunk segmentation in Nebalia species (Crustacea, Malacostraca, Leptostraca)

The ’egg-larval’ development of two species of Nebalia has been examined with SEM. Various details concerning limb ontogeny and trunk segmentation are described. The most important of these are the following. The tripartite state of the peduncle of antenna 2 in the adult of Nebalia species is derived from the fusion of the third and fourth podomeres, present in late larvae. The proximal portion of the mandible in the adult of Nebalia brucei, carrying the ’coxal process’, is, based on the ontogenetic evidence, interpreted as the combined basis and coxa, and the bipartite palp is interpreted as the endopod. The early development of the thoracopods and the three anteriormost pleopods is identical. They all start as laterally directed, biramous limb buds. This suggests that tagmatisation of the trunk of the Leptostraca (and other Malacostraca) has been developed from an ancestor with an undivided trunk region with serially similar limbs. Certain early stages reveal an extra, ’eighth’, limbless pleon segment, as compared with the normal number of seven pleomeres of adult Leptostraca. The presence of a row of ventral, sternitic, triangular processes between the bases of the thoracopods, as they are found in certain stages of a species of Nebalia, is suggested as a possible ground pattern for the Malacostraca. Accepted: 1 February 2000     

Limb articulation in caridoid crustaceans revisited - new evidence from Euphausiacea (Malacostraca)

The propodial articulation of thoracopods in Malacostraca is revisited. Two major joints at the base of the limb, a thorax-coxa joint and a coxa-basis joint permit promotion-remotion and abduction-adduction, respectively. In representatives of Decapoda, Anaspidacea and Euphausiacea, the coxa forms proximally a dicondylic articulation with the thorax, permitting promotion-remotion, and distally another dicondylic joint with the basis, permitting abduction-adduction. In Lophogastrida and Mysida, the thorax-coxa hinge line is antero-posteriorly oriented, as is the coxa-basis hinge line. Promotion-remotion in Mysida and Lophogastrida is possible because of the presence of an intrabasal joint which is also present in Euphausiacea and Anaspidacea. In Mysida, Lophogastrida and Euphausiacea, the intrabasal joint is only present anteriorly, just distally of the anterior coxa-basis joint between a small, triangular proximal part of the basis and a larger distal part. In Anaspidacea, the intrabasal joint is also present posteriorly and permits abduction-adduction. Homology with the intrabasal joint of the other taxa seems doubtful. Limb articulation in Anaspidacea shows, nevertheless, correspondences with that in Euphausiacea, Lophogastrida and Mysida: the coxa is posteriorly invaginated and has an open ring-like structure very different from the solid coxa in decapods. Despite the high level of structural correspondence between the intrabasal joint in Euphausiacea and that in Lophogastrida and Mysida, their different functional roles make homology implausible. In Lophogastrida and Mysida the intrabasal joint is thought to replace the promotion-remotion movement of the thorax-coxa articulation, which in these taxa permits abduction-adduction only, probably in connection with the evolution of the marsupium. In Euphausiacea, the intrabasal joint might play a role in feeding mechanisms. Neither the feeding basket nor a marsupium can reasonably be suggested for any common ancestor of Euphausiacea and Mysidacea (or Peracarida).     

Eye function of Mysidacea (Crustacea) in the northern Baltic Sea

Eye spectral sensitivity, [S(lambda)], was measured in seven northern Baltic mysid species using an electroretinogram technique. Their S(lambda) curves were compared with the spectral distribution of underwater light at their normal habitats. In the littoral species Neomysis integer, Praunus flexuosus and Praunus inermis, the S(lambda) maxima, [S(lambda)(max)], were in the wavelength-bands of 525-535, 505-515 and 520-530 nm respectively. The neoimmigrant species Hemimysis anomala had a S(lambda)(max) around 500 nm and high sensitivity at 393 nm, possibly indicating UV-sensitivity. S(lambda) of the pelagic species Mysis mixta and Mysis relicta sp. II was at about 505-520 nm. M. relicta sp. I from Pojoviken Bay and fresh water humic Lake P??j?rvi had S(lambda)(max) at approximately 550 nm and 570 nm respectively. This is in accordance with a similar long-wavelength shift in light transmittance of the respective waters. The eyes of the latter population were also damaged by strong light. The pontocaspian neoimmigrant H. anomala is clearly adapted to waters transmitting more blue light.     

On the occurrence and biology of some Antarctic Mysidacea (Crustacea)

Summary 10 species of Mysidacea were sampled during the Polarstern and Walther Herwig SIBEX cruises to the Antarctic Peninsula region from 1983 to 1986. The commonest were Antarctomysis maxima, A. ohlini and Mysidetes posthon. For these species the size and maturity stage composition as well as the length-weight relationships are given. The species considered are growing slowly and attain a long life span, M. posthon reached age class 3+, A. ohlini at least lived as long as 5 years, while A. maxima was found to be of age 5+ in the Peninsula area and probably 6+ in the Weddell Sea. The generation time of Antarctomysis species was 4 years and 3 years in Mysidetes posthon. Fecundity was low, mean number of offspring was about 13 for M. posthon and 21 for A. maxima. Rematuration was observed for the species A. maxima, indicating several spawning events during its life span. Biomass production is low for A. maxima, shown by a P/B-index of 0.98. Mortality of this species was estimated to be Z=1.1, which indicates that 33% of the specimens of an age group survive until the next year. The distribution and spatial separation of the two Antarctomysis species is discussed.     

Cleavage and gastrulation of the dendrobranchiate shrimp Penaeus monodon (Crustacea,Malacostraca, Decapoda)

The cleavage pattern of the black tiger shrimp Penaeus monodon was analyzed from the first division until gastrulation. Observations were based on microscopy combined with the use of fluorescent dyes, histological techniques, and computer based three-dimensional reconstructions. Early cleavage is holoblastic and follows a stereotypic pattern, which largely corresponds to what is known from other dendrobranchiate decapods. However, for the first time in this group, we report the presence of an intracellular structure throughout early development. This intracellular body (icb) marks the lineage of one of the two enlarged and division-delayed mesendoderm cells that initiate gastrulation. The identity of the icb as a natural marker and putative determinant of the germ line and its implications on the establishment of the body axes are discussed. The icb as a landmark reveals that the same stereotypic cell division pattern can lead to different fates of individual cells. Hence, the results of this study permit an additional approach to study the relation between cell lineage pattern and the identity of cell lineages.     

Cell Death in the Epithelia of the Intestine and Hepatopancreas in Neocaridina heteropoda (Crustacea,Malacostraca)

The endodermal region of the digestive system in the freshwater shrimp Neocaridina heteropoda (Crustacea, Malacostraca) consists of a tube-shaped intestine and large hepatopancreas, which is formed by numerous blind-ended tubules. The precise structure and ultrastructure of these regions were presented in our previous studies, while here we focused on the cell death processes and their effect on the functioning of the midgut. We used transmission electron microscopy, light and confocal microscopes to describe and detect cell death, while a quantitative assessment of cells with depolarized mitochondria helped us to establish whether there is the relationship between cell death and the inactivation of mitochondria. Three types of the cell death were observed in the intestine and hepatopancreas–apoptosis, necrosis and autophagy. No differences were observed in the course of these processes in males and females and or in the intestine and hepatopancreas of the shrimp that were examined. Our studies revealed that apoptosis, necrosis and autophagy only involves the fully developed cells of the midgut epithelium that have contact with the midgut lumen–D-cells in the intestine and B- and F-cells in hepatopancreas, while E-cells (midgut stem cells) did not die. A distinct correlation between the accumulation of E-cells and the activation of apoptosis was detected in the anterior region of the intestine, while necrosis was an accidental process. Degenerating organelles, mainly mitochondria were neutralized and eventually, the activation of cell death was prevented in the entire epithelium due to autophagy. Therefore, we state that autophagy plays a role of the survival factor.     

A new species of Gastrosaccus (Crustacea,Mysidacea) from beaches in Madagascar

Gastrosaccus madagascariensis sp.nov. is common in thesurfzone of sandy beaches along the east and south coasts ofMadagascar. Morphologically, it is characterised by having sevenlateral spines along each side of the telson; with spines 2–4 andthe penultimate spine being dorso-lateral in origin. One or twospinules are interspersed between the fifth and terminal spines.The fourth exopod segment of pleopod 3 in the male is bulbous, bearing twoterminal barbed unequal spines. A robust and naked seta is alsopresent on the same segment in the male.     

Angustidontidae (Crustacea: Malacostraca) from the Upper Devonian of Ukraine and Belarus

Previously unknown specimens of angustidontid crustaceans from the Frasnian and Famennian of Ukraine and from the Famennian of Belarus are described.     

Aspects of the biology of Antarctomysis maxima (Crustacea: Mysidacea)

Summary Population structure and aspects of the reproductive biology of Antarctomysis maxima from South Georgia and the South Orkney Islands were examined. Size frequency analysis indicated that the generation time is two years at South Georgia, but may approach four years at the South Orkneys. Egg production is greater for a given size of female at South Georgia but by virtue of a larger egg, brood production in terms of ash-free dry weight is slightly greater at the South Orkneys. Young are brooded over winter at both sites and are released in spring. Females are clearly capable of breeding twice at South Georgia and it is suggested that this is also the case at the South Orkneys. These results are discussed in relation to other studies on cold water mysids and to environmental adaptations exhibited by other Antarctic marine invertebrates.     

Euphausiid visual pigments. The rhodopsins of Euphausia superba and Meganyctiphanes norvegica (Crustacea, Euphausiacea)

The rhabdoms of Euphausia superba contain one digitonin-extractable rhodopsin, lambda max 485 nm. The rhodopsin undergoes unusual pH- dependent spectral changes: above neutrality, the absorbance decreases progressively at 485 nm and rises near 370 nm. This change is reversible and appears to reflect an equilibrium between a protonated and an unprotonated form of the rhodopsin Schiff-base linkage. Near neutral pH and at 10 degrees C, the rhodopsin is partiaLly converted by 420-nm light to a stable 493-nm metarhodopsin. The metarhodopsin is partially photoconverted to rhodopsin by long-wavelength light in the absence of NH2OH; in the presence of NH2OH, it is slowly converted to retinal oxime and opsin. The rhodopsin of Meganyctiphanes norvegica measured in fresh rhabdoms by microspectrophotometry has properties very similar to those of the extracted rhodopsin of E. superba. Its lambda max is 488 nm and it is partially photoconverted by short wavelength irradiation to a stable photoconvertible metarhodopsin similar to that of E. superba. In the presence of light and NH2OH, the M. norvegica metarhodopsin is converted to retinal oxime and opsin. Our results indicate that previous determinations of euphausiid rhodopsin absorbance spectra were incorrect because of accessory pigment contamination.     

Fine Structure of the Organ of Bellonci (SPX) in Boreomysis arctica (Kroyer) (Crustacea, Mysidacea)

The organ of Bellonci (oB) in Boreomysis arctica (Krøyer) is described. The fine structure of the organ is found to agree with that of the oB among other investigated peracarideans. The difficulties met by Chaigneau in 1971 when homologizing the organs in B. arctica and in the Isopoda have been eliminated. Into the cavity of the oB, (probably) sensory neurones protrude. The perikarya of the neurones are found in the wall of the oB. On the dendrites there are subterminal dendritic swellings from which dendritic branches and cilia arise. The branching dendrites are characteristic of the oB in B. arctica. Also the cilia branch, thus increasing the amount of sensory membranes in the organ.