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On the ontogeny of Leptodora kindtii (Crustacea, Branchiopoda, Cladocera), with notes on the phylogeny of the Cladocera.

Leptodora kindtii, a large predaceous cladoceran, is among the most deviant species of the Cladocera. Therefore, its phylogenetic position has traditionally proven difficult to determine. Its many peculiar features include, among others, long, stenopodous, forwardly directed trunk limbs, a posteriorly placed dorsal brood pouch, a tri-lobed lower lip, and a long, segmented abdomen. This study describes the ontogeny of L. kindtii (Haplopoda), including general body proportions, appendages, the carapace, and other external structures in an attempt to facilitate the comparison of its aberrant morphology to that of other branchiopods. In general, the early embryos are similar to the early embryos of other cladoceran taxa with respect to body shape and size and position and orientation of the early limb buds. Many of the unusual features of L. kindtii appear late in ontogeny. The carapace appears at an early stage as a pair of dorsolateral swellings in a position corresponding to the gap between the mandibles and the first pair of trunk limbs; it later becomes posteriorly transposed by a gradual fusion of its more anterior parts to the dorsal side of the thorax. The tri-lobed "lower lip," under the labrum of the late embryo and the adult, develops as a fusion of the first maxillae (lateral lobes) to an elevated sternal region behind the mouth (median lobe). The stenopodous, segmented trunk limbs in the adult develop from embryonic, elongate, subdivided limb buds, similar to those seen in early stages of other branchiopods. Two conflicting possibilities for the phylogeny of the Cladocera, involving two different positions of L. kindtii (Haplopoda), are discussed. Several characters support a sister-group relationship between the Haplopoda and Onychopoda. However, some characters support the Anomopoda and Onychopoda as sister groups, leaving the Haplopoda outside this clade. In contrast to recent suggestions, we prefer to retain the term "Cladocera" in its original sense as comprising the Haplopoda, Ctenopoda, Anomopoda, and Onychopoda.

Animals↗

Autapomorphies of the endophragmal system in trichodactylid freshwater crabs (Crustacea: Decapoda: Eubrachyura).

An examination of the thoracic endophragmal system in trichodactylid freshwater crabs revealed several derived features that appear restricted to this family. Trichodactylids have a highly reduced sella turcica with a rim-like overall configuration; as a consequence of sella turcica reduction the median septum is dorsoventrally diminished and exposed along its length. In addition, interosternite 7/8 is medially erased with the remnants restricted to the lateral regions of the sternum. The configurations of the sella turcica and interosternite 7/8 in trichodactylids underscore the monophyly of this family. The Dilocarcininae is characterized by having interosternites 5/6 through 7/8 restricted to the lateral sectors of the sternum, the lateral remnants of interosternite 4/5 forming prominent, rib-like ridges oriented diagonally to the longitudinal axis of the crab, and the median septum forming with interosternite 4/5 a Y-shaped pattern. These autapomorphies of the endophragmal system separate the trichodactylids from all other eubrachyurans and emphasize the morphological distinctness of the group. A description of the trichodactylid endophragmal junction plate is also provided.

Animals↗

Antennulary sensory organs in cyprids of Octolasmis and Lepas (Crustacea: Thecostraca: Cirripedia: Thoracica): a scanning electron microscopic study.

Cypris larvae of the pedunculate barnacles Octolasmis angulata (Poecilasmatidae), Lepas australis, L. pectinata, and Dosima fascicularis (Lepadidae) were studied with scanning electron microscopy, focusing on the sensory setae and the attachment disc on the antennules. The antennules of O. angulata did not exhibit any remarkable trait, but carry the same number of setae as seen in most other thoracicans. The third segment is bell-shaped and quite distinct from the second and its attachment disc is surrounded by a skirt. We found several potential synapomorphies in antennulary morphology between cyprids of the lepadid species but none of them were shared with the cyprids of Octolasmis; the list of unique lepadid characters includes: one additional, preaxial seta on the second segment; multiple similar (up to eight) postaxial setae (PS3) on the third segment, unlike all other thoracicans, where there is only a single PS3; the third segment consists almost entirely of the attachment disc, which is distended and surrounded by two parallel rows of radial setae; on the fourth segment the terminal seta E is diminutive. We found no traits in cyprids of Octolasmis that seem to be adaptations to their attachment site within the branchial chamber of swimming crabs and, in particular, no similarities with cyprids of rhizocephalan barnacles, many of which also attach in the gill chamber. The synapomorphies between cyprids of the lepadid species may be adaptations to their life in the neuston.

Animals↗

Growth and molting in epizoic pedunculate barnacles genus Octolasmis (Crustacea: Thecostraca: Cirripedia: Thoracica).

Scanning electron microscopy, light microscopy, and histology were used to study growth in species of the pedunculate barnacle genus Octolasmis (O. angulata, O. cor, O. californiana, O. mülleri). These species are epizoic in the gill chamber of portunid crabs and have highly reduced capitular shell plates, with large areas of general cuticle in between. The external integument grows by means of a system of narrow growth zones, one encircling the peduncle and a Y-shaped system on either side of the capitulum. Growth is by a regular series of molts, but shedding of old cuticle and production of new layers is entirely restricted to the growth zones. Just prior to ecdysis, the new cuticle lies in a highly folded fashion beneath the old cuticle that is about to be shed. At ecdysis, the old cuticle breaks along the margins of the growth zones and the resulting scars remain as a system of "ecdysial lines" along either side of the zone. Once exposed after ecdysis, the new cuticle remains as a part of the permanent external integument. The growth zones divide the externa into five cuticular areas, two on the peduncle and three on the capitulum. The calcareous shell plates (carina, paired scuta, and, when present, paired terga) all lie within the capitular regions and the ecdysial lines pass across, not around, these mineralized areas. The number, relative spacing, and topology of the ecdysial lines form a record of the growth history of the specimen. These and other growth patterns demonstrate that size increase is due to the formation of new cuticle by molting in the growth zones, while expansion of the shell plates by mineralization follows only after production of the new cuticle. Thus, although specialized, growth in Octolasmis still complies with the general crustacean model, complicated only by the mineralization of parts of the capitular cuticle into shell plates. The results are compared with the very scarce information on molting in other barnacles. We argue that at least the circular peduncular growth zone is omnipresent in the Cirripedia Thoracica.

Animals↗

Studies of male sexual tubes in hermit crabs (crustacea, decapoda, anomura, paguroidea). I. Morphology of the sexual tube in Micropagurus acantholepis (Stimpson, 1858), with comments on function and evolution.

The external morphology and internal structure of the male sexual tube of the hermit crab Micropagurus acantholepis, a member of the family Paguridae from Australian waters, is described in detail using histological thick sectioning and scanning and transmission electron microscopy techniques. This is the first in-depth study of a sexual tube in the Paguroidea, a group where a remarkable number of genera (55.9% in the family Paguridae) with species having these intriguing sexual structures are known. In M. acantholepis a sexual tube is present on the left side, whereas only a gonopore is present on the right side. The tube is used for the delivery of spermatophores to the female and consists of a sheath of cuticular origin surrounding an internal, functional extension of the posterior vas deferens. Pedunculate spermatophores were observed within the lumen and partially extruding from the terminal opening of the tube in preserved specimens. The tube protrudes from the left coxa of the fifth pereopod as an elongate 3-mm-long, hollow, coiled structure with a terminal opening. Exteriorly the tube consists of a conspicuous thick chitinous cuticular ridge throughout its length, and a thin chitinous cuticle with sparse, regularly arranged simple setae. Interior to the cuticle, the tube contains loose connective tissue, secretory cells, oblique muscle, circular muscle, and epithelial cells. The latter cells line a central lumen that runs the length of the sexual tube. The morphology, cellular composition, and function of the tube are discussed.

Animals↗

Comparative electron microscopic study of the stomach of Orchestia cavimana and Arcitalitrus sylvaticus (crustacea: Amphipoda).

The functional morphology of stomachs of the European semiterrestrial amphipod Orchestia cavimana and of the Australian terrestrial species Arcitalitrus sylvaticus was studied by electron microscopy. The stomach of the two amphipod species is divided longitudinally into a spacious dorsal food channel and two ventral filtration channels. Additionally, a prominent helically oriented circulation channel is situated on each lateral side of the stomach, forming a semicircular channel separated from the food channel by spines. The food channel conveys coarse food particles directly into the midgut through a funnel. The filtration channels receive fine material filtered through primary and secondary filters. Material forced through the secondary filters by the pressure of the laterally located inferolateralia eventually reaches the openings of the midgut glands. Washing of filters and soaking of ingested food items with enzymes probably is achieved by a forward stream of digestive juice from the midgut glands and conveyed through the circulatory channels. The specializations of the stomach of the two species of Amphipoda investigated are described and compared to the pertinent structures of Mysidacea and Isopoda.

Amphipoda↗

Introduction to the Rhizocephala (Crustacea: Cirripedia).

Knowledge about the Rhizocephala (parasitic barnacles) has increased exponentially over the past two decades. This introduction broadly reviews this progress, touching on rhizocephalan morphology, life-cycles, larval biology, and the effects on the crustacean hosts. Members representing both orders, the Kentrogonida and Akentrogonida, are discussed. The recent discovery regarding the vermigon stage further highlights the intricacy of the kentrogonid life-cycle. Because rhizocephalans are found in most world regions, occurring on their respective hosts from the deep ocean to freshwater, their importance is now being recognized.

Animals↗

Cypris metamorphosis, injection and earliest internal development of theRrizocephalan Loxothylacus panopaei (Gissler). Crustacea: Cirripedia: Rhizocephala: Sacculinidae.

Rhizocephala is a group of crustaceans that exclusively parasitizes other crustaceans. It is taxonomically placed within the class Cirripedia, the barnacles, with which it shares a unique larval type, the cyprid. The main objective of the cyprid is to find and irreversibly attach to a suitable substratum and initiate metamorphosis. In the presumed sister group to Rhizocephala, the true barnacles or Thoracica, metamorphosis leads to a juvenile filter-feeding version of the adult organism. In Rhizocephala the female cyprid settles on the integument of a crustacean and undergoes metamorphosis into a kentrogon that possesses a hollow cuticular-tube structure, the stylet, which penetrates the integument of the host and acts as a guide tube for the prospective internal parasite. The first, hitherto unknown endoparasitic stage of a rhizocephalan, the vermigon, was recently discovered (Glenner and Høeg [1995] Nature 377:147-150) and its migration through the hemolymph of the host, as well as its internal development, was described in Glenner et al. ([2000] Mar Biol 136:249-257). The present article provides detailed information on kentrogon and vermigon formation, the injection process, and the succeeding developmental stages up to the stage of the earliest primordium reported from the literature. The anlage of the ovary is traced back to the free-swimming cypris stage and it is implied that the mesoderm and ectoderm of the endoparasite are already differentiated in the cyprid.

Animals↗

Swimming and cleaning in the free-swimming phase of Argulus larvae (crustacea, branchiura)--appendage adaptation and functional morphology.

The free-swimming early larval stages of Argulus foliaceus (Linneaus) (Branchiura) are studied using digital video, light microscopy, and SEM. We analyze and document the mode of swimming in the hatching stage of A. foliaceus and the subsequent juvenile stages with fully developed thoracopods. We present new observations and an analysis of the functional morphology of a cleaning behavior in the first stage. This stage swims very efficiently using the large exopods of the second antennae in concert with the mandibular palp (naupliar limbs), while the subsequent stages use the now developed thoracopods for propulsion. This posterior shift in propulsion is similar to--but independent from--what is seen in other crustaceans. The hatching stage has previously been referred as a "metanauplius" but as the first and second maxillae are developed and active, and buds of all four thoracopods are present, it is too advanced to be included in the naupliar phase. The hooks of the first antennae and the distal hooks of the maxillae are demonstrated to function not only as attachment organs (to the host), but also to play a significant role in the cleaning of the naupliar swimming appendages. A digital video-based analysis of the swimming mode is provided. The larval swimming pattern is generally similar to that of other crustaceans such as Branchiopoda and Cirripedia, but autapomorphies of the Branchiura include the following: 1) While actively swimming, the naupliar appendages are almost straight during the recovery stroke and 2) they have a relatively small deflection during movement ( approximately 25 degrees or approximately 35 degrees for mandible and second antenna respectively), 3) the larval mandible has a uniramous palp which is the retained exopod. The morphological implications of the transition from the possibly nonfeeding pelagic, or free-swimming, first larval stage to the feeding, parasitic second stage are discussed and compared with other crustaceans.

Adaptation, Physiological↗

Localization of serotonin-like immunoreactivity in the eyestalk of the prawn Palaemon serratus (Crustacea, Decapoda, Natantia).

A serotonin-like substance in the organ of Bellonci in the eyestalks of embryos, larvae, and adults of the prawn Palaemon serratus was visualized by the use of two specific antisera against serotonin (5-hydroxytryptamine; 5-HT) in combination with peroxidase-antiperoxidase (PAP). The organ of Bellonci, characterized by compact onion bodies distally and degenerating onion bodies proximally, was the only site of the serotonin-like substance in adults, as well as during development in embryos and larvae. Variations in the content of the 5-HT analogue in the adult were detected during the molting cycle. There was more immunoreactivity in specimens fixed at night than in those fixed in daytime. Likewise, colchicine and nialamide injections enhanced the immunoreactivity of the serotonin-like substance. Extirpations of the medulla externa X organ (MEX), a neurosecretory cell group of the optic ganglion medulla externa, produced the same effect.

Animals↗

Spermatophore transfer in the hermit crab Clibanarius vittatus (Crustacea, Anomura, Diogenidae).

Although mating has been described in several hermit crab species, the mechanics of spermatophore transfer have not previously been demonstrated. Evidence from pleopod and gonopore morphology, video observations, and inseminated females indicates that in Clibanarius vittatus the male applies a spermatophoric mass directly onto the female via the gonopores rather than with modified pleopods 1-2 (gonopods) and/or genital papillae as in many other decapods. The single second pleopod of males of C. vittatus has a simple endopod with no apparent modifications for sperm transfer. There are no genital papillae extending from the male gonopores. The globular spermatophores are aligned in rows surrounded by a seminal secretion in the male ducts (vasa deferentia that terminate in ejaculatory ducts opening to the exterior via the gonopores). During copulation, described from time-lapse video recordings, the ventral surface of the last thoracic segment of the male, bearing the gonopores, was apposed to the ventral cephalothorax of the female. A massive amount of seminal secretion containing spermatophore ribbons, termed here the spermatophoric mass and described for the first time in a hermit crab species, was observed covering the sternites and coxae of pereopods 1-5 of a recently copulated female. It is suggested that during copulation the male emits the contents of the ejaculatory ducts directly onto the female without the aid of gonopods or genital papillae. Although spermatophore transfer is simple in C. vittatus, the presence of modified anterior pleopods or elongate genital papillae (sexual tubes) in other paguroidean species suggests the possibility of a more complex insemination process in these other hermit crabs.

Animals↗

Structural changes in sperm from the fiddler crab, Uca tangeri (Crustacea, Brachyura), during the acrosome reaction.

The acrosome reaction (AR) was induced in sperm from the brachyuran crustacean Uca tangeri either by mixing male and female gametes in filtered seawater or by treating the spermatozoa with the divalent cation ionophore A23187. This latter method provided a sufficient number of reacted spermatozoa to allow a detailed ultrastructural study of the AR. The process consists of two separate phases: a) initial release of the acrosomal vesicle contents, and b) further elongation of the acrosomal filament, which causes reversal of the rigid capsule limiting the acrosomal vesicle contents. The elongate acrosomal filament consists of an apical perforatorium and a basal columnar structure called here the proximal piece. The former derives from the perforatorium of the uninduced sperm stage with only small ultrastructural changes. The proximal piece forms from myelin-like membrane layers which are initially distributed all around the subacrosomal region and then accumulate in a column at the perforatorial base, thus promoting a sudden forward projection of the perforatorium. The AR in brachyurans is thought to be a passive mechanism that utilizes the negative pressure exerted on the nucleus--caused by emptying of the acrosomal vesicle--for an organized accumulation of membrane-rich material immediately behind the perforatorium, with the final result of the raising of a 3 microns long acrosomal filament.

Acrosome↗

Actin in the acrosome of the spermatozoa of the crab, Cancer pagurus L. (Decapoda, Crustacea).

The cytoskeletal protein actin was identified in the mature spermatozoon of the European edible crab, Cancer pagurus Linnaeus, by indirect immunofluorescence with monoclonal and polyclonal anti-actin antibodies, fluorescent phalloidin, and DNAase I. The actin was localized in two distinct concentric rings within the acrosome vesicle of the spermatozoon and appeared to correlate with the internal zonation of the vesicle. Modifications of the fluorescent pattern for actin were observed in sperm cells which were undergoing changes associated with the acrosome reaction. In these cases, fluorescent staining was observed in the nucleocytoplasm immediately subjacent to the perforatorial column and sometimes in the perforatorial column within the acrosome vesicle. Equally intense fluorescence was observed in an apical perforatorial projection. SDS-PAGE of C. pagurus sperm confirmed the presence of actin in the cells. A single band of actin (approximately 43 kDa) comigrated with rabbit muscle actin when immunoblotted onto nitrocellulose with mouse monoclonal anti-actin. The actin-associated cytoskeletal proteins alpha-actinin, tropomyosin, and spectrin were also identified within the spermatozoon of C. pagurus using specific polyclonal antibodies, but their presence was not confirmed by SDS-PAGE and immunoblotting.

Acrosome↗

Sublethal copper toxicity and food response in the freshwater crayfish Cambarus bartonii (Cambaridae, Decapoda, Crustacea).

Food response preceding and following exposure to Cu concentrations of 0.00, 0.02, and 0.2 mg/L was observed in male and female Cambarus bartonii from an uncontaminated, freshwater lake. The animals were tested in a Y-maze. Control animals demonstrated food localization, and the treated animals, either no localization or slight food avoidance. The latency for localization in the Cu-exposed males increased. Females revealed slight increases in latency. There were no significant differences between the low and the high Cu concentration groups in food detection or latency. However, there was an increase in mortality in the higher concentration group and significant difference in mortality among three copper concentrations (P<0.02). Females seem to recover from Cu-induced impairment slightly faster than males, suggesting a more efficient mechanism for detoxifying copper. Female mortality, however, remained equally high in the high Cu concentration group. The toxic effects of copper persist after removal of the toxicant and occur at concentrations much lower than reported lethal levels.

Animals↗

Stimulation of osmoregulating processes in the perfused gill of the crab Pachygrapsus marmoratus (Crustacea, Decapoda) by a sinus gland peptide.

Isolated posterior gills of the hyper-hyporegulating crab Pachygrapsus marmoratus were perfused with extracts of homologous sinus glands. Sinus gland extracts stimulated the influx of Na+ ions and increased the transepithelial potential difference in the gills in a dose-dependent and reversible fashion. The bioactivity of extracts prepared from crabs that had been acclimated to 10/1000 salinity for at least 1 week was not significantly different from that of extracts prepared from seawater (36/1000 salinity) crabs. The perfusion experiments with both extracts containing two sinus glands significantly increased Na+ influx by about 150% and transepithelial potential difference by about 45%. Sinus gland extracts also increased the Na+/K(+)-ATPase activity by 54% in incubated posterior gills. The bioactivity of extracts was reduced by pronase and trypsin, but not by heating for 10 min at 100 degrees. The molecular weight of the responsible factor(s) was > 5000 Da. Thus, the sinus gland of P. marmoratus is concluded to be involved in the neuroendocrine control of osmoregulation and to contain a peptide(s) that directly influences brachial function.

Animals↗

Neuroendocrine control of osmotic regulation in the freshwater shrimp Macrobrachium olfersii (Wiegmann) (Crustacea, decapoda): free amino acid concentrations in the hemolymph.

The participation of neuroendocrine factors present within the central nervous system in the regulation of hemolymph free amino acid (FAA) concentrations was examined in the freshwater shrimp Macrobrachium olfersii. Test shrimps were injected intramuscularly with homogenates prepared from the eyestalks (ES), ventral nerve cord (VNC), supraesophageal (SEG), or thoracic ganglia (TG) of donor shrimps previously exposed for 6 hr to a high-salinity medium (HSM, 21% salinity). After injection of the homogenate, the shrimps were maintained for up to 6 hr in either freshwater (FW) or HSM. Hemolymph was sampled by cardiac puncture and prepared for reverse phase HPLC, derivatizing the FAA with phenylisothiocyanate. An FAA profile was determined and the [FAA]:[Cl-] ratios for the four FAA present in highest concentration (Gly, Arg, Ala, and Pro for ES and VNC experiments; Glu, Leu, Ala, and Val for SEG and TG experiments) were obtained. Nonparametric analyses revealed specific, notable effects resulting from homogenate injection, e.g., ES homogenate increased [Pro]/[Cl-] ratios in FW-exposed shrimps; SEG homogenate increased [Glu]/[Cl-] and [Val]/[Cl-] ratios in HSM-exposed shrimps; and TG homogenate increased [FAA]/[Cl-] ratios for Glu, Leu, Ala, and Val in HSM-exposed shrimps. Total FAA concentrations decreased after exposure of the shrimps to HSM but were increased by the injection of ES homogenate in FW-exposed shrimps and by TG homogenate in HSM-exposed shrimps. The total [FAA]/[Cl-] ratio was also increased by TG homogenate in HSM-exposed animals. There were no clear effects on [Cl-] alone.(ABSTRACT TRUNCATED AT 250 WORDS)

Alanine↗

Occurrence of Merogony of Aggregata Frenzel 1885 (Apicomplexa) in Pleoticus muelleri and Artemesia longinaris (Crustacea: Natantia) from Patagonian Waters (Argentina)

The aim of this paper is to describe the merogonic phases of the coccidian Aggregata sp. in the natantians Pleoticus muelleri and Artemesia longinaris from Argentina. Ninety-seven specimens of P. muelleri (from San Jorge Gulf) and 49 of A. longinaris (from Rawson) were examined for intestinal parasites. Intestines were processed under standard histological conditions. The distribution of the octopuses (where gamogony and sporogony of Aggregata spp. take place) in the area considered and the importance of finding these protistans in crustaceans other than brachyurans are also explored. High prevalence values for both hosts indicate that shrimps are the primary hosts for these parasites in Argentinean waters. Copyright 1997 Academic Press. Copyright 1997 Academic Press

Journal Article↗

Morphological description of bacterial infection of digestive glands in the terrestrial isopod porcellio scaber (Isopoda, crustacea)

Morphological studies of the hepatopancreas of the terrestrial isopod Porcellio scaber revealed bacterial infection. The percentage of infected animals collected from the same site varied from 0 to 10% during the 4 years of study. Transmission electron microscopy, scanning electron microscopy, and light microscopy revealed that infected glands differed from those in healthy isopods. The most prominent sign was white spots between 100 and 200 &mgr;m in diameter along the entire gland. These spots were aggregations of vacuoles in the cells that were densely filled with bacteria in different phases of the developmental cycle that included the formation of small, dense, rod-shaped infective bacteria and much larger spherical multiplying cells filled with aggregates of polysomes and a chromatin network. Occasionally, large sphericles were filled with homogeneous electron-dense material. Bacteria were not observed in the cell nucleus. Small vacuoles of less than 5 &mgr;m were filled predominately with spherical bacteria but rod-shaped forms were also present in large numbers. Larger vacuoles of 10 to 20 &mgr;m in the main were densely filled with rod-shaped bacteria. According to the literature on the morphological characteristics of bacteria infecting invertebrates, those described in our study would be classified in the genus Rickettsiella. However, most recent investigations show that besides morphological investigations, genetic ones are also needed to define the taxonomic position of bacteria that infect invertebrates. Copyright 1999 Academic Press.

Journal Article↗