The sensory papilla X-organ in Boreomysis arctica (Krøyer) (Crustacea Malacostraca Mysidacea)

Summary The SPX-organ in Boreomysis arctica (Krøyer) (Crustacea Malacostraca Mysidacea) was investigated light and electron microscopically. The organ consists of a group of cells (the SPX-cells) and a vesicle surrounded by a connective tissue sheath. It is situated near the base of the sensory papilla of the eye-stalk. Neurosecretory material is produced in the SPX-cells and transported in axon-like projections from these cells into the vesicle. These processes contain no neurotubuli. Numerous fibres from an afferent nerve emanating from the medulla terminalis also enter the vesicle, where they form a dense irregular meshwork. This nerve transports no neurosecretory material. There are numerous synaptic contacts between the afferent nerve fibres and the neurosecretory processes from the SPX-cells. The neurosecretory material released from them accumulates in haemocoelic spaces in the vesicle. Release is most probably effected by the afferent nerve. Acknowledgements. We are indebted to Prof. H. Brattström and the staff of Biologisk Stasjon, Espegrend for working facilities and material. Mrs B. Morawetz and Mrs L. Eriksson gave us skilled technical assistance. The investigation was made possible by grants from the Swedish Natural Science Research Council.     

Leptostraca,Mysidacea, Isopoda,and Decapoda (Anomura) (Crustacea,Malacostraca) of the Chukchi Sea and Adjacent Waters: Biogeography and Fauna Formation

Data on the distribution of 72 species of Leptostraca, Decapoda: Reptantia: Anomura, Mysidacea, and Isopoda in the northern Bering Sea, the Chukchi and East Siberian seas, and the adjacent areas of the continental slope in the Arctic basin at depths less than 500 m were used for biogeographic analysis. According to distribution, these species can be united into 15 biogeographic groups. The hydrological regime, primarily, the distribution of waters of different origin in the investigated regions, governs the distribution of different biogeographic groups of crustaceans. Pacific boreal and subtropic-boreal species mostly inhabit the southern part of the Chukchi Sea, coastal waters off Alaska to Point Barrow, and the central region of this sea to the Herald Bank. For these taxa, the boundary between the Pacific and Arctic faunas is in the Chukchi Sea crossing from Cape Serdtse Kamen and Point Barrow to the area west of the Herald Bank. Possible pathways and the major stages of formation of the fauna of the investigated crustaceans for the last 18000 years are discussed.     

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.     

Feeding appendages of the Euphausiacea (Crustacea)

The Euphausiacea comprises about 85 species and the structure of the feeding appendages of 68 of these species is discussed here. A considerable uniformity is apparent in the appendages throughout the order but generic, and even in some cases, specific differences are evident. It is concluded from the study of the morphology of the appendages and the analyses of stomach contents that the majority of species in the genera Bentheuphwsia, Thysanopoda, Meganyctiphanes, Nyctiphanes, Pseudeuphausia, Euphausia, Tessara-brachion and Thysanoessa are omnivorous; that is, they can feed by filtering material from the water and act as predators of small zooplankton, especially copepods. Species in the genera Nematoscelis, Nematobrachion and Stylocheiron can feed on bottom deposits and also by predation of zooplankton but the amount of filter-feeding which they do may be limited because their mouthparts are not so well adapted for filtering as those of the previous group of genera.
It is suggested, from the study of the appendages and several other features of the animals, that the genera Thysanopoda, Meganyctiphanes, and Euphausia are closely related to one another and that a similar relationship exists between the genera Nematoscelis, Nematobrachion and Stylocherion. The genera Nyctiphanes and Pseudeuphausia are probably more closely related to the Thysanopoda group of genera and the genera Tessarabrachion and Thysanoessa to the Nematoscelis group of genera.     

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.     

Leptostraca,Mysidacea, Isopoda,and Decapoda (Anomura) (Crustacea,Malacostraca) of the Chukchi Sea and Adjacent Waters: Hydrological Regime and Species Composition

The hydrological regime of the region is briefly outlined. The distribution of waters of Arctic and Pacific origin is discussed. Based on the original and literature data, it has been found that 73 species of the investigated orders occur in the northern part of the Bering Sea, in the Chukchi and East Siberian seas, and in the adjoining areas of the continental slope of the Arctic basin at depths down to 500 m. These are 1 leptostracan species, 11 anomurans, 22 mysids, and 39 free-living isopods. The distribution of these species by depth and in relation to the thermal and salinity characteristics of waters is analyzed.     

Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca)

Although the biology of the reptantian Decapoda has been much studied, the last comprehensive review of reptantian systematics was published more than 80 years ago. We have used cladistic methods to reconstruct the phylogenetic system of the reptantian Decapoda. We can show that the Reptantia represent a monophyletic taxon. The classical groups, the 'Palinura', 'Astacura' and 'Anomura' are paraphyletic assemblages. The Polychelida is the sister-group of all other reptantians. The Astacida is not closely related to the Homarida, but is part of a large monophyletic taxon which also includes the Thalassinida, Anomala and Brachyura. The Anomala and Brachyura are sister-groups and the Thalassinida is the sister-group of both of them. Based on our reconstruction of the sister-group relationships within the Reptantia, we discuss alternative hypotheses of reptantian interrelationships, the systematic position of the Reptantia within the decapods, and draw some conclusions concerning the habits and appearance of the reptantian stem species.     

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     

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.     

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.     

Structure of recent and fossil mysid statoliths (Crustacea,Mysidacea)

Statoliths of 61 Recent species representing all subfamilies of Mysidae were studied with special emphasis on internal structure. In addition 5 samples of fossil statoliths from Miocene deposits were examined. Species of Boreomysinae and Rhopalophthalminae show simple roughly spherical organic statoliths, with setae originating from the sensory cushion and anchored in the statolith with distal branches extending shortly below the surface. All other subfamilies possess mineralized statoliths of greater structural complexity, with differentiation in core and mantle, where each part may consist of up to three layers. Habitus is hemispherical to discoidal. External gross structures are dorsal tegmen, ventral fundus, and the ambitus forming the outer toroidal to semi-toroidal circumference. Setae penetrate the mantle through mineralic canals and insert on the surface of the core. As suggested by congeneric species of Schistomysis, there is no principal structural difference between statoliths mineralized with fluorite compared to vaterite. However, vaterite statoliths tend to be more often of moruloid appearance and are exceptional by showing a central conical hole (the hilum) or a central cavity in certain forms. These structures are typical of fossil calcite statoliths. In vaterite and fluorite statoliths, the mantle shows radially arranged (= spherulitic) crystal aggregates. Such arrangements are badly preserved in fossil calcite statoliths. In large extant statoliths, concentric structures, mainly in the form of superficial striation and/or concentric microstrata, are visible in coexistence with radial aggregates. Stratification is possibly due to stratified deposition of the nonmineralized gland product, while the spherulitic structure is indicative of subsequent radial growth of crystal aggregates. The structure of accessory fluorite statoliths in the statocyst of Mesopodopsis slabberi leads to the hypothesis that mantle material is formed by secretions of the caudal statocyst gland. After demineralization of fluorite, vaterite and calcite statoliths, an organic template remains showing most essential morphological features of the statolith. From this we conclude that the structure of the statolith is (almost) entirely matrix mediated. © 1993 Wiley-Liss, Inc.     

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 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.     

The circulatory system in Mysidacea--implications for the phylogenetic position of Lophogastrida and Mysida (Malacostraca, Crustacea)

The morphology of the circulatory organs in Mysida and Lophogastrida (traditionally combined as Mysidacea) is revisited investigating species so far unstudied. In addition to classical morphological methods, a newly developed combination of corrosion casting with micro computer tomography (MicroCT) and computer aided 3D reconstructions is used. Lophogastrida and Mysida show a highly developed arterial system. The tubular heart extends through the greater part of the thorax and is connected with the ventral vessel via an unpaired descending artery. It is suggested that a distinct ostia pattern supports the monophyly of Mysidacea. The cardiac artery system is more complex in Lophogastrida than in Mysida, consisting of up to 10 pairs of arteries that supply the viscera. In both taxa, an anterior and posterior aorta leads off the heart. In the anterior part of the cephalothorax the anterior aorta forms dilations into which muscles are internalized; these structures are called myoarterial formations. One of these myoarterial formations can also be found in all the other peracarid taxa but not in other Malacostraca.     

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