Fine structure and ecdysis of mandibular sensilla associated with the lacinia mobilis in Neomysis integer (Leach, 1814) (Crustacea,Malacostraca, Peracarida)

The external and internal structures of adult Neomysis integer mandibles were studied using light and electron microscopy with special reference to the lacinia mobilis, a highly specialized appendage on the gnathal edge of many crustaceans. The right and left lacinia mobilis are equipped with ciliary primary sensory cells revealing that both laciniae are also mechanosensory organs in addition to their mechanical function during mastication. A detailed character analyses indicated that the right lacinia is probably a highly derived sensory seta, whereas two alternative interpretations are considered for the left lacinia; it could be a sensillar appendage equipped with two mechanosensory units, or it could be a movable appendage of the incisor process containing two sensilla deprived of external appendages. The ecdysis of the lacinia mobilis corresponds very well to type I sensillar ecdysis, suggesting classification as a sensillar appendage. These features support a possible homology of the right lacinia mobilis in Peracarida and Decapoda, tracing them to an origin as a member of the setal row. Whether the left lacinia mobilis is a sensillum or an appendage with sensilla cannot be resolved presently.     

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 lacinia mobilis and Similar Structures – a Valuable Character in Arthropod Phylogenetics?

Homology between the lacinia mobilis of peracarid crustaceans (Malacostraca) and movable appendages of the mandibular edge in other crustaceans, hexapods, and myriapods has been advocated in classical as well as recent phylogenetic studies, and in some cases this feature has been attributed major significance in arthropod systematics. A comparative SEM survey of the lacinia mobilis in Peracarida and its alleged homologues (prostheca of Hexapoda, internal tooth of Diplopoda, ‘lacinia mobilis’ of Symphyla and Remipedia) rejects the primary homology of these varied structures. The lacinia mobilis of Peracarida can be characterized precisely by asymmetry on the left and right mandibles, as a strong tooth-like structure on the left mandible which is oriented at a right angle to the remaining edge and as a stalked, spine-like structure on the right mandible. A fundamental difference to other Malacostraca is that the peracarid lacinia mobilis characterizes the adult mandibles. Supposed homologues of the peracarid lacinia mobilis are instead a modified pectinate lamella (Diplopoda, Chilognatha) or a movable appendage that is associated with the pars molaris rather than the pars incisivus (Symphyla), or an inner separation of the incisor process (Remipedia). The detailed structure of the prostheca in Campodeoida and Ephemeroptera weakens interpretations of its origin as a separation of the incisor process.     

The foregut of the Mysida (Crustacea,Peracarida) and its phylogenetic relevance

The foreguts of the mysids Antarctomysis maxima, A. ohlinii, Hansenomysis antarctica, Heteromysis formosa, Mesopodopsis slabberi, Neomysis integer, Paramysis kessleri, Praunus flexuosus, and Siriella jaltensis were examined by maceration methods, histological techniques, and scanning electron microscopy. Their morphology, their connection with the midgut glands, and probable function are described and summarized. Previous stomach investigations on mysids and the results of the present study are tabulated; a list of foregut characters, common to all Mysida, is presented. The phylogenetic relevance of these characters within the Malacostraca, especially within the Peracarida, is discussed. Most features are inherited from the ground pattern of the Malacostraca or Eumalacostraca. The bulbous cardia with its dorsal fold, the armature of the lateralia, and the construction of the funnel region are apomorphies for the Mysida. The results suggest that characters of mysidan and other peracaridan foreguts might also be useful in the elucidation of the phylogeny of the Mysida and Peracarida, respectively.     

28S rDNA evolution in the Eumalacostraca and the phylogenetic position of krill

The Malacostraca are an ancient and morphologically diverse class of Crustacea. The phylogenetic position of one order within this class, the Euphausiacea ("krill," subclass Eumalacostraca) was investigated using 28S rDNA sequences from representatives of several malacostracan orders. Phylogenies for these sequences were estimated by maximum-likelihood and maximum-parsimony analysis. The results of these analyses produced a new scheme for evolution within the Eumalacostraca. The new phylogenies suggested that Euphausiacea are most closely related to the Mysida and not the Decapoda, as is generally thought. Furthermore, the Mysida were found not to be closely related to the Lophogastrida, which are often considered their sister taxon. These hypotheses were tested against the hypotheses of monophyly for the Eucarida, Mysidacea, and Peracarida and found to be significantly better on the basis of the 28S rDNA data.     

Suprabenthic Peracarida (Crustacea, Malacostraca) sampled at 75°N off East Greenland

During an expedition with RV Polarstern in 1994 samples were taken at 75°N off Greenland, by means of an epibenthic sledge, which was equipped with an epi- and an additional suprabenthic net with an opening 1–1.33 m above the seafloor. Within the Peracarida (Crustacea, Malacostraca) sampled, 12 species were more frequent in the suprabenthic net, while 8 species were only sampled in this upper net. Received: 9 August 1996 / Accepted: 1 October 1996     

Structural Plans as Functional Models Exemplified by the Crustacea Malacostraca

Abstract
Dahl, E. (Zoological Institute, University of Lund, S-223 62 Lund, Sweden.) Structural plans as functional models exemplified by the Crustacea Malacostraca . Zool. Scr. 5 (3–4): 163–166, 1976.—The structural plans of higher taxa also represent integrated functional model systems which may be of value in dealing with evolutionary and phylogenetic problems. The model concept is tested on the Crustacea Malacostraca. Their origin and interrelationships are discussed, and a new alternative for their derivation is presented for criticism and comparison with previous hypotheses.     

Evolutionary morphology of the circulatory system in Peracarida (Malacostraca; Crustacea)

We demonstrate that by formulating guidelines for evolutionary morphology the transparency, reproducibility, and intersubject testability of evolutionary hypotheses based on morphological data can be enhanced. The five main steps in our concept of evolutionary morphology are (i) taxon sampling, (ii) structural analysis, (iii) character conceptualization, (iv) phylogenetic analysis, and (v) evolutionary interpretation. We illustrate this concept on the example of the morphology of the circulatory organs in peracarid Malacostraca. The analysis is based on recently published accounts in which detailed structural analyses were carried out, and on the older literature. Detailed conceptualizations of 22 characters of the circulatory system are given for 28 terminals. In a further step these characters are included in a recently revised matrix, resulting in 110 characters. The resulting parsimony analysis yielded a single most parsimonious tree with a length of 309 steps. The most significant results are that Peracarida is monophyletic, Amphipoda is the sister taxon to the Mancoida sensu stricto, the relict cave‐dwelling taxa Thermosbaenacea, Spelaeogriphacea, and Mictocarididae form a monophylum and Tanaidacea is the sister group to a monophylum comprising Cumacea and Isopoda. The evolutionary analysis shows that the ground pattern features of the circulatory organs in Peracarida are a tubular heart extending through the whole thorax, a posterior aorta with lateral arteries, and a ventral vessel system. Important features within the Peracarida are the backward shift of the anterior border of the heart, the reduction of the ventral vessel system, and two patterns of cardiac arteries, one common to the amphipod and tanaidacean terminals, and one to the cumacean and isopod terminals. © The Willi Hennig Society 2009.     

Global diversity of mysids (Crustacea-Mysida) in freshwater

In this article we present a biogeographical assessment of species diversity within the Mysida (Crustacea: Malacostraca: Peracarida) from inland waters. Inland species represent 6.7% (72 species) of mysid diversity. These species represent three of the four families within the Mysida (Lepidomysidae, Stygiomysidae, and Mysidae) and are concentrated in the Palaearctic and Neotropical regions. The inland mysid species distributional patterns can be explained by four main groups representing different freshwater invasion routes: (1) Subterranean Tethyan relicts (24 spp.); (2) Autochthonous Ponto-Caspian endemics (20 spp.); (3) Mysis spp. ‘Glacial Relicts’ (8 spp.); and (4) Euryhaline estuarine species (20 spp.). The center of inland mysid species diversity is the Ponto-Caspian region, containing 24 species, a large portion of which are the results of a radiation in the genus Paramysis. 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 and K. Martens Freshwater Animal Diversity Assessment     

Assembly of a transmembrane b-type cytochrome is mainly driven by transmembrane helix interactions

Folding, assembly and stability of alpha-helical membrane proteins is still not very well understood. Several of these membrane proteins contain cofactors, which are essential for their function and which can be involved in protein assembly and/or stabilization. The effect of heme binding on the assembly and stability of the transmembrane b-type cytochrome b'559 was studied by fluorescence resonance energy transfer. Cytochrome b'559 consists of two monomers of a 44 amino acid long polypeptide, which contains one transmembrane domain. The synthesis of two variants of the b'559 monomer, each carrying a specific fluorescent dye, allowed monitoring helix-helix interactions in micelles by resonance energy transfer. The measurements demonstrate that the transmembrane peptides dimerize in detergent in the absence and presence of the heme cofactor. Cofactor binding only marginally enhances dimerization and, apparently, the redox state of the heme group has no effect on dimerization.     

The Amphipod Functional Model and Its Bearing upon Systematics and Phylogeny

The peculiar habitus of Amphipods with lateral compression, deep coxal plates 1-4, broad bases of peraeopods 5-7, and enlarged pleura (epimeral plates) of abdominal segments 1-3 is seen as a functional system closely dependent upon the currents produced by the incessant beating of the pleopods. The derivation and position of branchiae and oostegites is discussed. The morphology of more or less aberrant groups, including the suborders Caprellidea and Ingolfiellidea, and its correlations with the mode of life and the basic functional model is analyzed. The possible bearing of the functional model system upon the position of the Amphipoda among the Peracarida is briefly dealt with.     

Germ band differentiation in the stomatopod Gonodactylaceus falcatus and the origin of the stereotyped cell division pattern in Malacostraca (Crustacea)

We analysed aspects of the embryonic development of the stomatopod crustacean Gonodactylaceus falcatus focusing on the cell division in the ectoderm of the germ band. As in many other malacostracan crustaceans, the growth zone in the caudal papilla is formed by 19 ectoteloblasts and 8 mesoteloblasts arranged in rings. These teloblasts give rise to the cellular material of the largest part of the post-naupliar germ band in a stereotyped cell division pattern. The regularly arranged cells of the genealogical units produced by the ectoteloblast divide twice in longitudinal direction. The intersegmental furrows form within the descendants of one genealogical unit in the ectoderm. Hence, embryos of G. falcatus share some features of the stereotyped cell division pattern with that in other malacostracan crustaceans, which is unique among arthropods. In contrast to the other malacostracan taxa studied so far, stomatopods show slightly oblique spindle direction and a tilted position of the cells within the genealogical units. The inclusion of data on Leptostraca suggests that aspects of stereotyped cell divisions in the germ band must be assumed for the ground pattern of Malacostraca. Moreover, Stomatopoda and Leptostraca share the lateral displacement of cells during the mediolateral divisions of the ectodermal genealogical units in the post-naupliar germ band. The Caridoida within the Eumalacostraca apomorphically evolved the strict longitudinal orientation of spindle axes and cell positions, reaching the highest degree of regularity in the Peracarida. The phylogenetic analysis of the distribution of developmental characters is the prerequisite for the analysis of the evolution of developmental patterns and mechanisms.     

Muscle precursor cells in the developing limbs of two isopods (Crustacea, Peracarida): an immunohistochemical study using a novel monoclonal antibody against myosin heavy chain

In the hot debate on arthropod relationships, Crustaceans and the morphology of their appendages play a pivotal role. To gain new insights into how arthropod appendages evolved, developmental biologists recently have begun to examine the expression and function of Drosophila appendage genes in Crustaceans. However, cellular aspects of Crustacean limb development such as myogenesis are poorly understood in Crustaceans so that the interpretative context in which to analyse gene functions is still fragmentary. The goal of the present project was to analyse muscle development in Crustacean appendages, and to that end, monoclonal antibodies against arthropod muscle proteins were generated. One of these antibodies recognises certain isoforms of myosin heavy chain and strongly binds to muscle precursor cells in malacostracan Crustacea. We used this antibody to study myogenesis in two isopods, Porcellio scaber and Idotea balthica (Crustacea, Malacostraca, Peracarida), by immunohistochemistry. In these animals, muscles in the limbs originate from single muscle precursor cells, which subsequently grow to form multinucleated muscle precursors. The pattern of primordial muscles in the thoracic limbs was mapped, and results compared to muscle development in other Crustaceans and in insects. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Observations of the mating behavior and dentition of the round stingray,Urolophus halleri

Synopsis The mating behavior and dentition ofUrolophus halleri, the round stingray was examined. Males frequently bite females during the mating period but most male biting does not result in copulation. In bites that do not lead to copulation, males bite the posterior (or occasionally the medial) portion of the females' disc but females often free themselves from these bites. In bites that precede copulation, males bite the anterior portion of the females' disc and females do not struggle to free themselves. Thus, females may exert some form of choice when they are bitten. Mature males have sexually dimorphic dentition that may aid in holding females. A principal component analysis (PCA) showed that in juvenile males, the relative size of the teeth decrease while the relative thickness of the disc increases as body size enlarges; adult males displayed no clear pattern. In adult females, there is a relative decrease in tooth size and increase in relative disc thickness as body size enlarges. The relative increase in females disc thickness in areas where they are bitten may function to minimize the amount of damage due to non-copulatory biting. There is no indication that biting functions to induce female receptivity but it may allow females and males to acquire information about potential mates. Thus, copulatory biting functions to maintain contact during copulation while the function of non-copulatory biting is less clear.     

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).     

A comparison of the spermatozoa of OratosquiMa stephensoni and Squilla mantis (Crustacea, Stomatopoda) with comments on the phylogeny of the Malacostraca

Each stomatopod sperm, aflagellate and obovoid, is surrounded by an electron dense coat. A spermatophore is absent. The discus-shaped acrosome vesicle is penetrated and underlain by a straight, slender acrosome rod (perforatorium) ensheathed, below the vesicle, in subacrosomal material. Feulgen-positive granular material, indicating chromatin, fills most of the length of the cell but there is no certain nuclear membrane. Two centrioles, consisting of doublets each with a radial “foot” as in decapods and peracarids, occur near the acrosome and like it are embedded in the chromatin. Myelin-like membranes are associated with degenerating mitochondria in the posterior region of the cell. Thiéry-positive granules are aggregated as a glycogen body posteriorly in the cell. Spermatozoa1 ultrastructure confirms monophyly of the mysid–amphipod–isopod– cumacean section of the Peracarida but affinities of the tanaids are uncertain. A closer phylogenetic relationship of Phyllocarida to the Branchiopoda than to the Malacostraca is suggested. A weak similarity (synapomorphy?) of syncarids and peracarids is the filiform perforatorium bipassing the nucleus. Stomatopods resemble decapods in their diffuse sperm chromatin but are placed below the syncarid–peracarid–decapod assemblage.     

Molecular characterization of the principal substrate binding site of the ubiquitous folding catalyst protein disulfide isomerase

Disulfide bond formation in the endoplasmic reticulum of eukaryotes is catalyzed by the ubiquitously expressed enzyme protein disulfide isomerase (PDI). The effectiveness of PDI as a catalyst of native disulfide bond formation in folding polypeptides depends on the ability to catalyze disulfide-dithiol exchange, to bind non-native proteins, and to trigger conformational changes in the bound substrate, allowing access to buried cysteine residues. It is known that the b' domain of PDI provides the principal peptide binding site of PDI and that this domain is critical for catalysis of isomerization but not oxidation reactions in protein substrates. Here we use homology modeling to define more precisely the boundaries of the b' domain and show the existence of an intradomain linker between the b' and a' domains. We have expressed the recombinant b' domain thus defined; the stability and conformational properties of the recombinant product confirm the validity of the domain boundaries. We have modeled the tertiary structure of the b' domain and identified the primary substrate binding site within it. Mutations within this site, expressed both in the isolated domain and in full-length PDI, greatly reduce the binding affinity for small peptide substrates, with the greatest effect being I272W, a mutation that appears to have no structural effect.     

Functional incorporation of chimeric b subunits into F1Fo ATP synthase

F(1)F(o) ATP synthases function by a rotary mechanism. The enzyme's peripheral stalk serves as the stator that holds the F(1) sector and its catalytic sites against the movement of the rotor. In Escherichia coli, the peripheral stalk is a homodimer of identical b subunits, but photosynthetic bacteria have open reading frames for two different b-like subunits thought to form heterodimeric b/b' peripheral stalks. Chimeric b subunit genes have been constructed by substituting sequence from the Thermosynechococcus elongatus b and b' genes in the E. coli uncF gene, encoding the b subunit. The recombinant genes were expressed alone and in combination in the E. coli deletion strain KM2 (Deltab). Although not all of the chimeric subunits were incorporated into F(1)F(o) ATP synthase complexes, plasmids expressing either chimeric b(E39-I86) or b'(E39-I86) were capable of functionally complementing strain KM2 (Deltab). Strains expressing these subunits grew better than cells with smaller chimeric segments, such as those expressing the b'(E39-D53) or b(L54-I86) subunit, indicating intragenic suppression. In general, the chimeric subunits modeled on the T. elongatus b subunit proved to be more stable than the b' subunit in vitro. Coexpression of the b(E39-I86) and b'(E39-I86) subunits in strain KM2 (Deltab) yielded F(1)F(o) complexes containing heterodimeric peripheral stalks composed of both subunits.     

Formal biting in stumptailed macaques (Macaca arctoides)

Form and function of a specific kind of biting were studied in a captive group of stumptailed macaques (Macaca arctoides) composed of 1 adult male and 10 females. Behavior patterns were recorded with all occurrence and scan sampling techniques. Formal biting consisted of slowly gripping with teeth a part of the body of an apparently willing partner, most often the forearms, lips, or eyebrows. It may occur either following an agonistic interaction or outside the context of aggression. Such biting was directed from dominants towards subordinates, and appeared especially frequently between individuals having frequent agonistic interactions. It is concluded that this behavior represents a ritualized interaction expressing formal dominance between partners.     

Carbon 13 NMR studies of saturated fatty acids bound to bovine serum albumin. II. Electrostatic interactions in individual fatty acid binding sites

13C NMR chemical shift results as a function of pH for a series of carboxyl 13C-enriched saturated fatty acids (8-18 carbons) bound to bovine serum albumin (BSA) are presented. For octanoic acid bound to BSA (6:1, mol/mol), the chemical shift of the only FA carboxyl resonance (designated as peak c), plotted as a function of pH, exhibited a complete sigmoidal titration curve that deviated in shape from a corresponding theoretical Henderson-Hasselbach curve. However, FA carboxyl chemical shift plotted as a function of added HCl yielded a linear titration curve analogous to those obtained for protein-free monomeric fatty acid (FA) in water. The apparent pK of BSA-bound octanoic acid was 4.3 +/- 0.2. However, the intrinsic pK (corrected for electrostatic effects resulting from the net positive charge on BSA) was approximately 4.8, a value identical to that obtained for monomeric octanoic acid in water in the absence of protein. For long-chain FA (greater than or equal to 12 carbons) bound to BSA (6:1, mol/mol), chemical shift titration curves for peak c were similar to those obtained for octanoic acid/BSA. However, the four additional FA carboxyl resonances observed (designated as peaks a, b, b', and d) exhibited no change in chemical shift between pH 8 and 3. For C14.0 X BSA complexes (3:1 and 6:1, mol/mol) peaks b' and a exhibited chemical shift changes between pH 8.8 and 11.5 concomitant with chemical shift changes in the epsilon-carbon (lysine) resonance. In contrast, peaks c and d exhibited no change and peak b only a slight change in chemical shift over the same pH range. We conclude: the carboxyl groups of bound FA represented by peaks a, b, b', and d were involved in ion pair electrostatic interactions with positively charged amino acyl residues on BSA; the carboxyl groups of bound FA represented by peak c were not involved in electrostatic interactions with BSA; the similarity of the titration curves of peak c for BSA-bound octanoic acid and long-chain FA suggested that short-chain and long-chain FA represented by peak c were bound to the same binding site(s) on BSA; bound FA represented by peaks b' and a (but not d or b) were directly adjacent to BSA lysine residues. We present a model which correlates NMR peaks b, b', and d with the putative locations of three individual high-affinity binding sites in a three-dimensional model of BSA.