Functional morphology of the mouthparts and associated structures of Pagurus rubricatus (Crustacea: Decapoda: Anomura) with special reference to feeding and grooming

Summary Pagurus rubricatus is predatory, detrivorous, macrophagous, and to a small degree, a suspension feeder. The crab searches for small invertebrates by digging shallow pits in the sediment. During this process it feeds on detritus obtained either directly from the sediment or scoured off gravel granules. Particles trapped by the dense setation of the 2nd and 3rd maxillipeds are brushed off and ingested.The distribution of the various types of setae on the mouthparts is mapped and structure of the mouthparts and their setae is correlated with function. Sediment collected by the pereiopods is brushed off by the endopodites of the 3rd maxillipeds and transferred to the inner mouthparts by the endopodites of the 2nd maxillipeds. The basipodites of the 1st maxillae form a filter screen through which particles of suitable size are pushed by the 2nd maxillae. Rejected particles are discarded by the exhalant stream via the currents generated by the exopodites of the maxillipeds. Specialized setae on the 2nd maxillae scour detritus from the surface of gravel granules applied to these appendages by the 2nd and 3rd maxillipeds. Interlocking setae from different appendages form a number of screens the main function of which is to retain material in the buccal region. The exopodite and endopodite of the 1st maxilliped and the endopodites of the 1st and 2nd maxillipeds form a channel which funnels the exhalant respiratory current away from the crab. The main grooming appendages are the endopodites of the 3rd maxillipeds, however, most of the other mouthparts have a self-cleaning function.     

Carapace movements associated with ventilation and irrigation of the branchial chambers in the semaphore crab,Heloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Summary Carapace movements in crabs are briefly reviewed. While on land and recirculating branchial water, the Australian semaphore crab Heloecius cordiformis (Decapoda: Ocypodidae), a semi-terrestrial air-breathing mangrove crab, sequentially depresses and elevates its carapace relative to its thorax (0.5–1 mm excursion) in a regular pump-like manner. In quiescent crabs each carapace-pumping cycle lasts about 4 s; carapace depression takes 3 s and elevation 1 s. Carapace movements are brought about by pressures generated within the branchial chambers by the scaphognathites, probably in combination with carapace muscles. Carapace movements are associated with bilaterally synchronised scaphognathite activity. Unilateral scaphognathite activity was not observed. During normal forward recirculation of branchial water the scaphognathites beat at about 1.5 Hz (slow-forward pumping) and the lungs (epibranchial chambers) are not ventilated. In Heloecius, the lungs are not physically separated from the gills below by an anatomical barrier. Lung ventilation is accomplished during the following sequence of events: the carapace is lowered and the scaphognathites pump in a fast-forward mode at about 2.8 Hz. This activity preferentially pumps air out of the lungs and generates suction within the branchial chambers (4–10 cm H₂O below ambient) which draws water from external body surfaces into the hypobranchial space below and around the gills. At the end of the carapace's downward travel the scaphognathites switch from fast-forward to fastreverse beating at about 4 Hz. This pumps air into the lungs and the carapace elevates. As a result, during carapace elevation the water which had previously been drawn into the branchial chambers by fast-forward pumping activity is released and flows out between the legs and into the abdominosternal cavity. When the carapace reaches its original resting or up position the scaphognathites switch from fast-reverse to slowforward beating to re-establish water recirculation through the branchial chambers. This cycle is subsequently repeated. In stationary crabs, there are 2 carapace-pumping cycles per minute, increasing to 14 per minute in active crabs (walking). When water is absent, the lungs are preferentially ventilated by slow-reverse scaphognathite pumping activity. Carapace movements do not occur in the absence of branchial water. Carapace pumping is thought to provide a mechanism which permits the scaphognathites to ventilate the lungs in the presence of recirculating branchial water, without this water interfering with lung ventilation or being lost to the environment.Abbreviations FF, FR, SF, SR fast-forward, fast-reverse, slowforward, slow-reverse scaphognathite pumping - MEA Milne Edwards aperture     

Comparative studies of the branchial morphology, gill area and gill ultrastructure of some thalassinidean mud-shrimps (Crustacea: Decapoda: Thalassinidea)

The morphology, gill area and branchial formulae of six thalassinidean decapods ( Calocaris macandreae, Jaxea nocturna, Callianassa subterranea, Upogebia stellata, U. deltaura and U. pusilla ) are reported. Additionally, the rarely-encountered Axius stirhynchus receives brief attention. Gill formulae are presented; the simplest arrangements are found in the Callianassidae and Upogebiidae. The deeper-burrowing, deposit-feeding species that are regularly exposed to prolonged periods of hypoxia, i.e. Callianassa subterranea and Jaxea nocturna , had significantly larger gill areas than thalassinideans that occupied more oxygenated burrows ( Upogebia spp., Calocaris macandreae ). The increase in gill area was a result of flattening of the trichobranchiate gill filaments giving rise to a phylloid gill form. It is suggested that the efficiency of gas transfer, and hence diffusing capacity, was enhanced in the phylloid gill by the larger gill area and, because of the reduced cuticle thickness, by the shorter water-haemolymph diffusion distance. The increased diffusion capacity of the phylloid gill is interpreted as a functional adaptation to the more severe physicochemical burrow water conditions experienced by Callianassa subterranea and Jaxea nocturna .     

The morphology of unique structures on the spines of a deep-water Hawaiian hermit crab (Crustacea: Decapoda: Paguridae)

Unique structures, which have been observed on the spines of the chelipeds in a recently described deep-water Hawaiian hermit crab, Nematopagurus spinulosensoris McLaughlin & Brock, 1974, are herein described. At present the function of these structures is unknown; however, their morphology appears to be more complicated than that of any sensory structures heretofore ascribed to decapod crustaceans.     

The population dynamics of some freshwater carideans (Crustacea: Decapoda) in Hong Kong,with special reference to Neocaridina serrata (Atyidae)

During a 15-month investigation of the population dynamics of three caridean shrimp species in the Lam Tsuen River, New Territories, Hong Kong, the numbers of the commonest species, Neocaridina serrata (Stimpson)(Atyidae), were reduced as water temperatures fell. By contrast, Caridina lanceifrons Yu became more numerous during the winter. The relatively scarce Macrobrachium hainanense (Parisi)(Palaemonidae) tended to be more abundant during the summer, and the abundance of this species was directly correlated with that of N. serrata. The population size of these two species was positively correlated with water temperatures prevailing one and two months prior to the collection of samples, but there was no significant relationship between C. lanceifrons abundance and water temperature.All species exhibited similar growth patterns with an inflection at the attainment of sexual maturity, occurring after the (presumed) 14th moult for N. serrata and in the (presumed) 13th and 18th stadium for C. lanceifrons and M. hainanensis respectively. Ovigerous shrimps of all species were only recorded when water temperatures exceeded 20 °C and the % incidence of ovigerous N. serrata was positively correlated with prevailing temperatures. It is suggested that restriction of freshwater caridean breeding periods by low temperatures may be common in the subtropics while perennial breeding is more likely to be typical of tropical regions.The mean carapace length of N. serrata populations was negatively correlated with prevailing temperatures, but more strongly correlated with temperatures recorded one and two months prior to sampling when gametogenesis may have taken place. Smallest mean sizes were recorded in late summer upon the cessation of juvenile recruitment. N. serrata has a maximum longevity of approximately 12 months and reproduction occurs in the 7th month of life or later. The hatchlings are well developed and clutch size is not related to the size of the brooding female. An annual life cycle, as seen in N. serrata, may be typical of many small freshwater carideans.Department of Zoology, The University of Hong Kong     

Spermiogenesis and sperm structure in the crabUca tangeri (Crustacea,Brachyura), with special reference to the acrosome differentiation

Summary Early spermatids of the crabUca tangeri consists of the nucleus of granular chromatin and the cytoplasm, which contains a proacrosomal vesicle in close association with membrane lamellae. In the mid spermatids an invagination of the acrosomal vesicle membrane gives rise to the formation of the perforatorium, a spindle-shaped tubule which encloses tubular membranous structures. The pair of centrioles located at the base of the acrosome is not directly involved in perforatorial differentiation. The acrosomal vesicle shows a heterogeneous content composed of the operculum, the thickened ring, and three layers of different materials concentrically arranged around the perforatorium. During the late spermatid stage the nuclear profile differentiates numerous slender arms and the chromatin arranges into fibers. Membranous tubules from the cytoplasm become incorporated into the tubular structures of the perforatorium. The mature spermatozoon has the typical structure of the branchyuran sperm, with a complex acrosome, cupped by the nucleus, and a thin cytoplasmic band intervening between the former main elements. The centrioles are degenerate. The nuclear arms are unusually numerous (more than 20) and lack microtubules or microtubular derivatives.     

Oxygen consumption and the respiratory responses to declining oxygen tension in the crab Ebalia tuberosa (Pennant) (Crustacea: Decapoda: Leucosiidae)

Oxygen consumption and its relationship to body weight and activity have been examined in Ebalia tuberosa (Pennant). Ebalia showed only very limited ability to regulate their oxygen consumption under conditions of declining P_O2. Respiratory independence was markedly affected by the level of activity of the animals. When inactive, respiratory independence was maintained down to a critical P_O2 of only about 130 mm Hg while when active, the crabs behaved as perfect conformers. Inactive Ebalia responded to a decrease in PO₂ by increasing the pumping activity of the scaphognathites and heart. This increased activity continued even at oxygen tensions lower than the P_c. The rate of reversals of the ventilatory currents did not change with decreasing P_O2. In the active crabs, the pumping activity of the scaphognathites and heart, and the rate of current reversals were higher than in the inactive animals, and a decrease in PO₂, did not bring about any further change in any of these responses. When exposed to hypoxic conditions for long periods initially inactive crabs survived longer than did active animals.     

Mechanics of branchial ventilation in the valviferan isopod Idotea wosnesenskii (Crustacea)

Idotea wosnesenskii Brandt, a valviferan isopod, has five pairs of pleopods within a branchial chamber enclosed by modified uropods (operculae). The pumping mechanism of resting isopods was videotaped to observe ventilation, and particularly to determine whether or not the swimming pleopods (first three pairs) and the gas exchange pleopods (last two pairs) have ventilatory roles. Observations on intact animals (to observe water flow into and out of the branchial chamber) and on animals with operculae propped open or with parts of their abdominal wall removed (to study the actual pumping movements of the pleopods) revealed that all five pairs of pleopods function in ventilation. The ventilatory stroke has two phases: an opening phase in which the pleopods move medially and ventrally, opening spaces between pleopods (analogous to the swimming recovery stroke), and a closing phase, wherein the pleopods move laterally and dorsally, reducing the gaps between succeeding pleopods (analogous to the swimming power stroke). Ventilatory strokes may be continuous or they may be separated by resting phases. Both the frequency and amplitude are variable: the frequency changes greatly due to the wide variation in resting phase duration. By opening the operculae the amplitude can increase so that ventilatory strokes grade into swimming strokes. The ventilatory stroke pattern at rest is generally similar to the swimming stroke pattern but differs in its slower and nearly simultaneous dorsoventral movements and its lower frequencies. Dye and particle movements around intact animals and those with exposed pleopods show that water typically takes about three strokes to pass through the branchial chamber.     

Metamorphosis of the excretory system of Paragonimus ohirai (Trematoda), with special reference to its functional significance

The fine structure of the excretory system was studied in metacercariae and juveniles of Paragonimus ohirai. The former were in vitro excysted, and the latter were collected from the abdominal cavity of a rat 24 hours postinfection. The terminal organs of the excretory system were composed of a flame cell and the first cell of a tubule. In the excysted metacercaria, there was no space between these cells to allow the passage of fluid. This suggests that the terminal organs of P. ohirai may be inactive in this stage. The excretory bladder was formed of epithelial cells which contained numerous lipid droplets and a large amount of glycogen in the cytoplasm. The bladder of the metacercaria seems to function as a storage area for nutrients and other materials. These characteristics of the metacercaria are considered to be related to the enclosed conditions created by encystment. Excretory organs had undergone cytomorphosis in the juveniles and appear to possess active excretory function. The periflagellar space in the terminal organs was formed, and lipids and concretions were excreted from a thin layer of the excretory epithelia into the lumen.     

The food of suburban foxes (Vulpes vulpes), with special reference to London

The food of foxes in suburban London was examined using stomach contents and den litter analysis. A great diversity of food was taken. Adult foxes killed nearer to the city centre were found to eat significantly fewer earthworms, domestic pets and wild mammals, but more scavenged food items, than foxes killed closer to the suburban fringe. Cubs killed closer to the city centre ate significantly fewer earthworms and insects, and more pet birds, pet mammals and scavenged food items than cubs killed nearer the periphery of the suburban area. Although seasonal trends in food selection were apparent, they were not as pronounced as those observed in studies of rural foxes. At no time of the year did a single food item predominate in the diet of adult foxes. Passeriformes were the most important single food item in the diet of young cubs, but there was no correlation between the number of Passeriformes eaten and their abundance, nor between annual changes in the abundance of small birds and mean fox birth litter size and productivity. Most foxes examined were in good body condition; there was no seasonal variation in body weight, although fat content did vary seasonally. A complementary survey in Bristol on losses of domestic pets to foxes revealed that 07% of pet cats and 80% of other pets kept outdoors were killed in a year. Damage to garden Crops was negligible.     

Evolutionary morphology of the hemolymph vascular system in hermit and king crabs (Crustacea: Decapoda: Anomala)

The morphological transformation of hermit crabs into crab‐like king crabs in the evolution of decapod crustaceans represents a remarkable case of carcinization or evolutionary shaping into a crab‐like form. In this study, we focus on internal organs such as the hemolymph vascular system and adjacent anatomical structures of several Recent hermit crab (Paguridae) and king crab (Lithodidae) species. There are various correspondences in the morphology of the arterial systems in the dorsal cephalothorax of the two taxa, especially with regard to the anterior aorta, anterior lateral arteries, and hepatic arteries. In the pleon, the posterior aorta in both taxa displays a proximal bifurcation and follows an asymmetrical course. The ventral vessel system, on the other hand, which mainly supplies the limbs, differs significantly between the taxa, with pagurids displaying the plesiomorphic condition. The pattern of the ventral vessel system in Lithodidae is influenced by morphological transformations of integumental structures during carcinization. One of these transformations was the broadening of the sternites, which resulted in a widening of the space between the endosternites. In addition, changes in the morphology of the endophragmal skeleton in Lithodidae led to an increase in the potential for intraspecific variability and interspecific variation in the arterial branching pattern. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Carcinonemertes pinnotheridophila sp. nov. (Nemertea,Enopla, Carcinonemertidae) from the branchial chambers of Pinnixa chaetopterana (Crustacea,Decapoda, Pinnotheridae): description,incidence and biological relationships with the host

A new species of Carcinonemertes, C. pinnotheridophila, is described and illustrated. The worms were found in the pinnotherid crab Pinnixa chaetopterana collected from the coasts of New Jersey, North Carolina and Florida. The anatomy of the new species is compared and contrasted with that of other members of the genus Carcinonemertes and an emended generic diagnosis is provided. The nemerteans only inhabit female hosts, crabs being infested with one or two mature female worms but with no more than one ensheathed in each branchial chamber; one or more smaller male nemerteans may be associated with each female. Sheaths are attached to the medial portion of the host branchial exoskeleton, and project through an opening in the floor of the chamber to exit via another aperture in the sternum; the anterior part of the sheath opens in the excurrent canal of the branchial chamber. Female worms cement their oval egg sacs on the pleopods to which the crab's eggs are also attached. Attachment, development and hatching of both host and symbiont eggs are synchronous. The incidence of infestation, reproductive potential of the nemertean, damage to its host and tolerance of the crab's growth cycles are described.     

The biogeography of the beachflea, Orchestia gammarellus (Crustacea, Amphipoda, Talitridae), in the North Atlantic with special reference to Iceland: a morphometric and genetic study

The beachflea [Orchestia gammarellus (Pallas)] is a trans‐Atlantic amphipod inhabiting the littoral fringe. In Iceland, its distribution is temperature‐limited; it has recently colonized Iceland's relatively warm south‐western coast, and in the cooler north‐west, several small populations inhabit isolated warm springs. We address two questions: (i) Do the warm spring populations show evidence of long‐term residence in Iceland, or of recent colonization? (ii) For the new south‐western Icelandic populations, can the source population be determined? We sequenced COI for 22 populations in Iceland, Nova Scotia, the Faroe Islands, the British Isles, Norway and Sweden. Morphometric analysis of a subset of populations assessed 16 continuous and five discrete characters. Genetically, we found a star phylogeny: a common haplotype was found at all sites except two neighbouring warm spring populations, and all haplotypes were within two base pairs of this common haplotype. Morphometrically, almost all populations examined differed significantly in some characters; however, the warm spring populations differed slightly more from each other than did other populations. Although the origins of the Icelandic populations could not be well resolved, our data are consistent with a recent European origin.     

The functional anatomy of the foregut of Porcetlana platychetes and a comparison with Galathea squamifera and Upogebia deltaura (Crustacea: Decapoda)

Muscles, ossicles and internal structures of the foregut of Porcellana platycheles are described and figured. The function of the foregut is deduced from the distribution of food particle size within its chambers. The trough of the cardio-pyloric valve is considered to be the main site of digestion and the supra-ampullary valve to have an active role in the formation of the ventral cap of faecal pellets.
The structures and function of the foregut of Galathea squamifera are similar to those of P. platycheles. Certain differences found in the foregut of Upogebia deltaura may be related to the small size of food particles ingested and the absence of a ventral cap from faecal pellets may be due to the supra-ampullary valve having no action on surrounding particles.     

The functional morphology of the feeding appendages and gut of Hippolyte varians (Crustacea: Natantia)

The musculature of the feeding appendages are described and compared with those few early accounts of other Decapoda, particularly those of Pandalus danae and Astacus fluviatilis . While the musculature is very similar to that of the latter, the relative importance of the various muscles may differ markedly and a number of previously undescribed muscles have been found, while others present in Pandulus and Astacus are not present in Hippolyte . The alimentary canal is also described, together with its associated musculature. The function of the various appendages and the parts of the gut are described from observations of living specimens.     

The structures of the tentacles of Rhabdopleura compacta (Hemichordata) with special reference to neurociliary control

Summary The tentacle of Rhabdopleura compacta (Hemichordata) consists of two layers of cells surrounding a central coelomic cavity. The two layers of cells are separated by a cell free basement lamella.The tentacles on the arms of Rhabdopleura bear three longitudinal rows of cilia. The ciliated cells are closely associated with bundles of nerve fibres, and between some of the cells and nerve fibres there are synapses. The peripheral regions of the ciliated cells are joined to one another by desmosomes. Tonofibrils join some of these desmosomes to the kinetosomes of the cilia.The nerve fibres are confined to the ectodermal layer and the muscle cells to the layer of cells within the basement lamella. In the ectodermal layer besides ciliated cells there are mucus cells, densely pigmented cells, and green bodies. The function of these last two types of cells is secretory. Most of the epithelial cells have microvilli upon their free borders.I wish to thank Professor J. Z. Young F. R. S. for enthusiastic advice and encouragement. Dr. R. Bellairs generously provided the facilities for electron microscopy. Mr. R. Moss gave excellent technical and photographic assistance. Dr. A. Stebbing of the Plymouth Marine Biological Laboratory helped me to obtain and to identify the specimens. Professor D. W. James kindly allowed me to use his facilities for interference microscopy.     

The anatomy and functional morphology of the large hermaphroditic duct of three species of Aplysia, with special reference to the atrial gland

The anatomy and functional morphology of the large hermaphroditic duct of three species of gastropod mollusc (Aplysia californica, A. dactylomela, and A. brasiliana) were examined. Each duct is composed of two parallel compartments, the red hemiduct (RHD) and the white hemiduct (WHD), which are distinguishable from the outside of the duct. Four secretory regions, all exocrine in morphology, are recognizable: the RHD secretory epithelium, the atrial gland (or atrial gland-like epithelium), the WHD secretory epithelium, and the accessory gland of the copulatory duct (AGCD). Of these regions, only the atrial gland (or atrial gland-like epithelium) contains egglaying activity and only the atrial gland (or atrial gland-like epithelium) is immunocytochemically labeled by serum antibodies generated against low molecular weight A. californica atrial gland peptides. The RHD is the functional oviduct: the egg cordon passes through a channel lined by the RHD secretory epithelium and bordered by the atrial gland (or atrial gland-like epithelium); the eggs are separated from both the WHD secretory epithelium and the AGCD by internal folds of the duct. The WHD is the functional copulatory duct: the penis, exogenous sperm, and endogenous sperm pass directly by the AGCD and in close proximity to the WHD secretory epithelium; they are separated from both the RHD secretory epithelium and the atrial gland (or atrial gland-like epithelium) by internal folds. The atrial gland (or atrial gland-like epithelium) is thus not likely to have a prostatic function or to be directly stimulated by the penis during copulation; it may play a role in oviductal function.