Dorippids are Heterotremata: Evidence from ultrastructure of the spermatozoa ofNeodorippe astuta (Dorippidae) andPortunus pelagicus (Portunidae) Brachyura: Decapoda

Ultrastructural comparison between the sperm of the dorippid crabNeodorippe astuta (Fabricius, 1793) and the portunidPortunus pelagicus (Linnaeus, 1766) from Queensland, Australia, supports placement of dorippids with portunids and their relatives in the heterotreme section of the Eubrachyura (the Heterotremata - Thoracotremata or the Oxyrhyncha - Cancridea - Brachygnatha assemblage) and not withRanina ranina (in the Archaeobrachyura or the Oxystomata). Similarities between spermatozoa ofN. astuta and ofP. pelagicus (and other Eubrachyura) andR. ranina include: the large spherical, multi-layered, capsule-bound acrosome vesicle; the electron-dense operculum capping the vesicle; an invaginated core, or perforatorium; concentric zonation of the contents of the vesicle; a layer of cytoplasm, between the acrosome vesicle and the nucleus, which contains mitochondria (mostly degenerating) and lattice-like lamellar complexes or membrane remnants; a diffuse nucleus which is bounded externally by a combined nuclear and plasma membrane and cups the scanty cytoplasm and the large acrosome vesicle; and lateral arms into which the chromatin extends. Characteristic eubrachyuran features of theN. astuta sperm absent fromR. ranina are the long perforatorium (short and conical with a unique subacrosomal chamber inR. ranina) extending almost to the operculum; presence in the perforatorium of longitudinally arranged convoluted tubules; a zone of acrosomal rays forming the outer part of an inner dense zone; the presence of a thickened ring surrounding the basal part of the perforatorium; and, basally, two centrioles (absent fromR. ranina but also from some eubrachyurans). The sperm ofN. astuta is more similar to that ofP. pelagicus than to that of other investigated Brachyura. A heterotreme status ofN. astuta is thus unequivocally supported. Both species lack the posterior median process seen in the nucleus of majids andR. ranina.     

Ultrastructure of the mature spermatozoon of the coconut crabBirgus latro (Coenobitidae: Paguroidea: Decapoda)

The spermatozoon ofBirgus latro (Linnaeus, 1767) is approx 14µm in length. It is composed of a large multi-layered oblong-ovoid acrosome which is capped by a conical operculum and lies anterior to a small ring of cytoplasm and an amorphous nucleus which is drawn out into a series of arms or extensions. Originating from the cytoplasmic area are three further long microtubular arms. The sperm ofB. latro is very similar to the sperm of the only other genus in the Coenobitidae,Coenobita, of whichC. clypeatus is a representative species. They share a suite of ultrastructural characters including: a long, cylindrical, capsule-bound acrosome containing an inner acrosome core, a large acrosome ray zone and a thin outer acrosome zone; an apical operculum anterior to a subopercular zone divided into two areas of differing density; an invaginated perforatorial zone with a bipartite granular matrix; microvillus-like extensions of the inner acrosome core projecting into the perforatorial invagination; a ring of cytoplasm, around the base of the acrosome, containing numerous mitochondria, extensive lamellar systems and the bases of three microtubular arms; granular nuclear material forming irregular arms; and at the posterior portion of the cell membrane a combination of nuclear and plasma membranes. Some ultrastructural characters which separate the two genera are: a domed operculum inC. clypeatus as opposed to a conical one inB. latro; inB. latro there is some residual cytoplasm external to the operculum and centrioles are absent from the mature spermatozoon; inC. clypeatus the inner acrosome core does not appear to invest the perforatorium and a series of dense rods are found subjacent and internal to the operculum. Similarities between the two coenobitids are greater than those shared with the paguridEupagurus bernhardus. In this latter species, the acrosome is more ovoid than cylindrical and the acrosome zones are less conspicuous; the operculum is absent from the mature sperm (although present during spermiohistogenesis); no subopercular zone is present; the perforatorium contains longitudinally arranged microtubules and extends the full length of the acrosome; and the nuclear material does not form separate nuclear arms. Sperm ultrastructure supports monophyly of the Paguroidea, while distinguishing coenobitids from pagurids within this superfamily.     

Strontium/Calcium ratios in statoliths of the neon flying squid, Ommastrephes bartrami (Cephalopoda), in the North Pacific Ocean

Strontium to calcium ratios were observed along longitudinal sections of statoliths of nine neon flying squid, Ommastrephes bartrami (LeSueur, 1821), including three mature females (422 to 454 mm mantle length, ML; 207 to 306 d old) obtained from the North Pacific (27–35°N; 144–150°E) during winter and six immature males and females (187 to 226 mm ML; 126 to 164 d old) collected from 39°N; 145°E and 39°N; 169°W during summer. The distances between the nucleus (core) and the edge of the dorsal dome were approximately 660 to 690 μm in mature females and 450 to 510 μm in the immature squid. Sr/Ca ratios were determined at intervals of 30 μm between the nucleus and edge of the dorsal dome. Sr/Ca ratios were higher in areas near the nuclei and peripheral portions of the dorsal dome than in the middle portions of the statoliths (270 to 420 μm from the nuclei, corresponding to ages of 60 to 90 d) in mature females; thus a U-shaped pattern was evident. Sr/Ca ratios in the six immature squid decreased from nucleus to the dorsal dome; in three squid the ratios slightly increased toward the dorsal dome edge. The observed Sr/Ca ratios in immature squid were considered to represent younger portions of the U-shaped pattern. In the present study we discuss this pattern in relation to environmental and biological conditions of O. bartrami, which undertakes seasonal migrations between spawning grounds in the Subtropical Domain and feeding grounds in the Subarctic Domain and Transitional Zone in the North Pacific Ocean. Although Sr/Ca ratios are potentially affected by ambient water temperature and ontogenetic conditions, including somatic growth and statolith growth, it was impossible to evaluate each environmental and biological effect separately, as variations in these factors are complicated and effects could be interdependent. Received: 11 April 1997 / Accepted: 27 December 1997     

Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

The relationship between allele frequency and tidal height in a mussel hybrid zone: a test of the differential settlement hypothesis

The blue mussels Mytilus edulis L. and M. galloprovincialis Lmk. hybridize in western Europe. Within hybrid populations nuclear alleles specific to M. galloprovincialis increase in frequency with age and size. This relationship changes with tidal height; alleles from M. galloprovincialis occur more frequently high in the intertidal zone, while M. edulis alleles predominate in the low intertidal zone. We tested the hypotheses that larvae with M. galloprovincialis alleles tend to settle higher in the intertidal zone, or that mussels redistribute themselves with respect to tidal height after initial larval settlement. We sampled recently metamorphosed mussels every 2 weeks in a hybrid mussel population at Whitsand Bay in southwest England throughout the summer of 1996. We observed four cohorts of newly settled mussels. There was no evidence of differential settlement of mussels with different genotypes in connection with tidal height, or into shaded versus unshaded microsites. Therefore, we rejected the preferential settlement hypothesis. There was substantial movement of juvenile mussels in the first 4 weeks following initial settlement, but this “secondary settlement” did not result in genetic differentiation with respect to tidal height. Further, significant differences in allele frequencies were found between primary and secondary spat. This allele frequency change was in the opposite direction of that seen in the adult population, suggesting newly settled larvae may be experiencing different selective pressures than adults. We propose that the genetic structure of hybrid mussel populations with respect to tidal height is the consequence of differences in selection intensity. Received: 30 April 1999 / Accepted: 5 May 2000     

Sperm polymorphism and genetic divergence in the mussel Perumytilus purpuratus

For the first time in a marine bivalve species, geographic intraspecific variation in sperm morphology in the Chilean mussel, Perumytilus purpuratus, is reported. Samples were obtained in December 2010 from the rocky intertidal at three locations: Antofagasta (23o44′01.08′′S, 70o26′26.11′′W), Valparaíso (32o57′23.07′′S, 71o33′01.86′′W), and Valdivia (39°50′44.26′′S, 73°25′51.99′′W). Scanning electron microscopy was used to describe sperm traits, and genetic data were obtained by sequencing a 16S mtDNA 427-bp fragment. The results showed geographic intraspecific variation with two statistically significant patterns of genetically differentiated sperm morphologies. One genetic clade was found for Antofagasta and Valparaíso, displaying a unique sperm morphotype with a short acrosome accounting for 26 % of the head length, and the other from Valdivia, with a larger acrosome occupying 47 % of the head length. These results will lead to new research to distinguish among alternative hypotheses: the presence of cryptic species, incipient speciation, or a polymorphism in sperm traits of P. purpuratus.     

Reproductive biology and ecology of a deep-demersal fish, alfonsino Beryx splendens, over the seamounts off New Caledonia

The reproductive biology of the alfonsino Beryx splendens was studied by histological examinations, gonadosomatic index and macroscopic scales of maturation of a large sample of gonads. Alfonsino is a gonochoric species. The size-frequency distribution of the sex ratio was bimodal and considered to be due to size dimorphism. In New Caledonia, the breeding period of this species occurs during the southern summer, with a peak in December to January. The spawning stage is attained at a minimum fork length of 28 cm for females and 33 cm for males. The size at which 50% of the population attain sexual maturity (FL₅₀) is 33.2 cm for females and 34.5 cm for males. Maximum potential fecundity is estimated to lie between 270 000 to 675 000 eggs for fish between 34 and 40 cm in fork length. It was possible to differentiate vegetative zones, in which juvenile alfonsino grow until they reach maturity, from reproductive zones (fishing grounds) which are inhabited by mature individuals. The larvae and juveniles could be carried from the reproductive zone to the vegetative zone by currents in an oceanic eddy system. Received: 26 April 1996 / Accepted: 20 September 1996     

Acrosome differentiation and the acrosome reaction in ascidian spermatozoa:<Emphasis Type="Italic"> Ascidiella aspersa</Emphasis> and<Emphasis Type="Italic"> Ascidia mentula</Emphasis> with some implications for tunicate phylogeny

The spermatozoa of both Ascidiella aspersa and Ascidia mentula have architectural features characteristic of ascidian spermatozoa that have previously been described. They have an elongated head (7 µm long for A. aspersa and 4 µm long for A. mentula), a single mitochondrion that is applied laterally to the nucleus and lacks a midpiece. The acrosome of A. aspersa spermatozoa is a flattened vesicle, about 200 nm×100 nm×40 nm (length, width and height). The acrosome of A. mentula spermatozoa consists of multiple vesicles; they are about 50 nm×50 nm×40 nm (length, width and height). During spermiogenesis in both species, several proacrosomal vesicles (50–70 nm in diameter) appear in a blister at the future apex of the spermatid. In A. aspersa, these vesicles fuse with each other to form a single acrosomal vesicle, while in A. mentula these vesicles do not fuse with each other, and form multiple acrosomal vesicles. In A. aspersa spermatozoa, calcium ionophore A23187 induces the acrosome reaction in which membrane fusion between the acrosomal apical membrane and the overlying sperm plasma membrane occurs along the peripheral margin of the acrosome, resulting in the release of a hybrid, membrane-bound, small vesicle. In A. mentula, multiple acrosomal vesicles disappear by releasing small vesicles after treatment with the calcium ionophore A23187; this also appears to be an acrosome reaction. This paper discusses the way in which acrosome structure and function may have changed during the evolution of the Tunicata.Communicated by T. Ikeda, Hakodate     

Fine structure of spermatids and sperm of Dolioletta gegenbauri and Doliolum nationalis (Tunicata: Thaliacea): implications for tunicate phylogeny

In Dolioletta gegenbauri and Doliolum nationalis, collected in 1987 in the bay of Villefranche-sur-mer (French Mediterranean Sea), spermiogenesis is essentially the same. Early spermatids have a round head, a flagellum arising from a single centriole with short microtubules at 45° to its base, several mitochondria, and an acrosome 50 nm thick and 250 nm long with its long axis parallel to the plasma membrane. The acrosomal contents are dense, with a central denser plate. The nuclear envelope next to the acrosome is thickened and concave. In elongating nuclei, strands of chromatin become oriented parallel to the length of the nucleus and then twist helically. Although the mitochondria surround the nucleus, they remain relatively short and do not fuse into a single mitochondrion as in sperm of other tunicates. In very late doliolid spermatids, the acrosome undergoes exocytosis, and exposes fibrous material that stays associated with the tip of the sperm; no acrosomal tubule forms. Exocytosis at this stage may be triggered by fixation. If so, exocytosis probably occurs naturally at some time before fusion of sperm and egg. Sperm have elongate heads (1 m×10 m), with the anterior two-thirds of the nucleus surrounded by mitochondria. Spermiogenesis in doliolids, compared to that in other tunicates, is most like that in solitary members of the class Ascidiacea, except that in the latter the sperm mitochondria fuse and the acrosome appears incapable of exocytosis. In contrast, previous work has shown that salps (class Thaliacea) and colonial didemnid ascidians have an acrosomeless sperm with a spiral mitochondrion, while the class Appendicularia has a sperm with a midpiece, a compact head and an acrosome capable of exocytosis and acrosomal tubule formation. By outgroup comparison with echinoderms and acraniates, appendicularian sperm are plesiomorphic within the Tunicata. Thus, gamete morphology indicates that (1) solitary ascidians and doliolids had a common ancestor, (2) the popular idea that doliolids gave rise to appendicularians is incorrect, and (3) the Thaliacea are polyphyletic, doliolids having arisen very early from the ascidian lineage and salps having arisen later.     

Differential increment-deposition rate in embryonic statoliths of the loliginid squid Loligo vulgaris

 Apart from one study that reported growth of less than one increment per day in statoliths of the squid Alloteuthis subulata, most studies so far have presumed that one increment was laid down per day in the statoliths of the squid species they examined. The present study provides evidence of differential daily growth rates in embryonic statoliths of the squid Loligo vulgaris Lamarck, 1798, thus confirming a previous report for A. subulata. Incremental growth rates of L. vulgaris statoliths differ as a function of temperature. Squid embryos were incubated in the laboratory at three temperatures (12.0, 15.5 and 21.1 °C), and tetracycline staining was used to follow statolith growth. This growth slowed in squid exposed to the lowest temperature, but recovered when the squid were returned to warm conditions, indicating statolith adaptation. Statolith growth rate after incubation at 12 °C was 1.3% d⁻¹ and reached 6.1% d⁻¹ for squids exposed to 21.1 °C. Statoliths from embryos incubated at 15.5 °C yielded a rate of 1 increment d⁻¹ and a mean daily growth of 2.2 μm in the dorsal dome area of the statolith. In contrast, the slow growth of statoliths incubated at 12 °C yielded a mean daily growth of only 0.9 μm in the dorsal dome and the readings resulted in a less-than-daily increment-deposition rate. Received: 9 October 1999 / Accepted: 30 March 2000     

Ultrastructure of the vitelline coat in the ascidians Phallusia mammillata, Ascidia mentula and Ciona intestinalis : new aspects revealed by freeze-substitution and deep-etching

The microstructure of the vitelline coat (VC) of the three ascidian species Phallusia mammillata, Ascidia mentula and Ciona intestinalis was analyzed by transmission electron microscopy (TEM) using freeze-substitution and deep-etching. In addition, eggs were examined by light microscopy and prepared by conventional chemical fixation methods for TEM in order to illustrate the advantages of cryopreparation for the preservation of the VC ultrastructure. In conventionally fixed eggs of P. mammillata and A. mentula the VC appeared as a tripartite structure, consisting of a thin outer fibrous layer (OFL), a central dense layer (CDL) and a relatively wide inner fibrous layer (IFL). In C. intestinalis a sparse OFL is made up by fine tufts of filaments but an IFL is lacking. In all three species the test cells were located in the area between the inner VC boundary and the egg cell membrane. Using our modified freeze-substitution method a clear improvement in preservation of the VC matrix was obtained. In all three species the OFL as well as the IFL in P. mammillata and A. mentula appeared as extended fibrous meshworks. Furthermore, the area of the perivitelline space, empty after conventional TEM preparation, was found to be filled by an additional matrix, designated here as perivitelline fibrous matrix (PVFM). Quick-freeze/deep-etch/rotary-shadowing enabled us to confirm the overall architecture of the egg matrix and allowed a further detailed investigation of the VC fine structure in P. mammillata. Accordingly, the vitelline coat of P. mammillata and A. mentula is a multilayered matrix composed of four distinct subunits (OFL, CDL, IFL, PVFM), whereas in C. intestinalis an IFL is lacking. Received: 7 November 1996 / Accepted: 5 December 1996     

Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region

Carbon consumption and nitrogen requirements were estimated for populations of the sandy beach bivalve Donax serra on nine beaches of the west coast of South Africa. Subtidal populations composed mainly of adult clams were responsible for the bulk of standing stock (3538 g C m⁻¹), annual carbon consumption (13 444 g C m⁻¹ yr⁻¹), faeces production (6478 g C m⁻¹ yr⁻¹ ) and nitrogen regeneration (2525 g N m⁻¹ yr⁻¹). Kelp detritus, bacteria and kelp consumers' faeces available in the water column surpass several times the carbon and nitrogen requirements of intertidal and subtidal clam populations. Individual Donax serra pop ulations, in turn, may regenerate up to 3.2% of the total nitrogen requirements of all primary producers from kelp beds and 14% of the requirements of phytoplankton. These high standing stocks of clams are presumably supported mainly by organic matter originating from kelp which, in contrast to phytoplankton, is in constant supply and comprises the largest proportion of the annual production of particulate organic matter on this coast. Wide and shallow continental shelves with gentle slopes probably limit the penetration of upwelled waters to the nearshore waters, decreasing the influence of external inputs and increasing the importance of internal flows of nutrients and carbon within the nearshore zone. In this context, sandy beaches, rocky shores and kelp beds may be more closely interlinked compartments of a larger ecosystem encompassing the whole nearshore than traditionally thought. Received: 28 August 1996 / Accepted: 7 October 1996     

Description and comparison of major foregut ossicles in hydrothermal vent crabs

Major foregut (gastric mill) ossicles, including the dorsal median tooth, lateral teeth, accessory lateral teeth, and cardiopyloric valve, of hydrothermal vent crabs were dissected and examined during the summer of 1996 from specimens housed at the Natural History Museum of Los Angeles County. Ossicles are described for two species of hydrothermal vent crabs (family Bythograeidae Williams, 1980). The western Pacific Austinograea williamsi Hessler and Martin has an unusual dorsal median tooth. The surrounding cuticular flange is scalloped and bears spinulose setae at the tip of each of the protruding edges, a condition perhaps unique in the Brachyura. The lateral teeth are mostly unremarkable, bearing the typical large anterior denticles and deep serrations seen in other crab families, but with a higher number of serrations than is known for any species previously described. The accessory lateral teeth bear flattened, plate-like spines that are widest basally and that taper to a cylindrical tip. The basic armature of the foregut of Bythograea thermydron Williams, known only from vents in the eastern Pacific, is very similar. Scalloping of the median tooth borders is less pronounced, however, and the shape of the tooth itself and of the plate from which it arises is slightly different. The lateral teeth bear fewer and more widely spaced grooves, and the cardiopyloric valve entrance appears less setose at its extremity. Comparison with foregut ossicles in other crab families based on earlier studies, most of which have not employed SEM, reveals some similarities between bythograeids and some xanthids, but does not clarify the phylogenetic position of the bythograeids. Because of the paucity of other SEM studies of the brachyuran foregut, it is difficult to ascertain whether some of the many spine and setal types in the bythograeid foregut are unique or even unusual compared to those of other crab families. Nothing about the foregut of the vent crabs is indicative of their unusual habitat. Anecdotal observations of feeding in vent crabs indicate that they are opportunistic scavengers and omnivores, which is in keeping with the non-specialized nature of the foregut. The debate between adaptation vs phylogeny as determinants of the form of the gastric mill components is briefly discussed. Received: 10 December 1996 / Accepted: 13 November 1997     

Spermatogenesis inPyrosoma atlanticum (Tunicata: Thaliacea: Pyrosomatida): Implications for tunicate phylogeny

Colonies ofPyrosoma atlanticum were collected by submersible in October 1988 in the Caribbean Sea, and testes were studied by electron microscopy. Spermatogonia, spermatocytes and early spermatids have two centrioles. The proximal centriole subsequently disappears, its remains apparently persisting in the spermatozoon as dense material adjacent to the distal centriole, which gives rise to the axoneme. At the tip of early spermatids are several 50 nm proacrosomal vesicles, which disappear leaving no trace in early elongating spermatids. The spermatozoon lacks an acrosome and has a head 35µm long. The head is differentiated into a bulbous posterior portion 5µm long × 1µm wide, a thinner anterior portion 25µm long tapering from a width of 0.7µm to a width of 0.4µm, and a very thin anterior extension 5µm long × 0.5µm wide. At the start of elongation, the anterior extension begins to form just lateral to the proacrosomal vesicles as a spiral projection comprising part of the nucleus, covered by a thin sheath of cytoplasm. This sheath of cytoplasm undergoes a complex differentiation. Ultimately, the nucleus in the anterior extension is overlain by two membrane-bound sheaths of cytoplasm connected by a spiral flange of cytoplasm. Between these two sheaths is a spiral space, open to the exterior through a subterminal pore near the sperm tip. In early spermatids the mitochondria fuse into a single mitochondrion, which remains lateral to the nucleus. The cristae become modified late in spermatogenesis. Throughout elongation of the spermatid there are patches of dense material between the nucleus and mitochondrion. A manchette of microtubules transiently encircles the thin anterior portion of the nucleus during the last phase of elongation. A manchette is not present during most of elongation. In the spermatozoon the mitochondrion, which has reticulate cristae, spirals a few times about the nucleus and extends from the junction between the bulbous portion and the thinner anterior portion of the nucleus to the junction between the thinner anterior portion and the nuclear extension. Spermatogenesis inP. atlanticum, compared to that in other tunicates, most closely resembles that in colonial ascidians, and supports the majority view that pyrosomes arose from aplousobranch ascidians that lost their attachment to the substratum. Pyrosome sperm are more highly derived than doliolid sperm, which have an acrosome that is probably capable of exocytosis. When salp and pyrosome sperm are compared, both are highly derived, but neither shares any apomorphies with the other that it does not share with at least one other tunicate order. Thus, sperm morphology does not support the majority view that pyrosomes gave rise to doliolids and neither confirms nor denies the idea that pyrosomes are intermediate between aplousobranch ascidians and salps. Therefore, it is likely that the class Thaliacea is polyphyletic, with doliolids arising very early from the ascidian lineage and with salps and pyrosomes arising somewhat later.     

Morphology of the reproductive system and spermatozoa of Mesodasys adenotubulatus (Gastrotricha: Macrodasyida)

Mesodasys adenotubulatus Hummon, Todaro and Tongiorgi 1993 is a simultaneous hermaphrodite. Two club-shaped testes extend along the sides of the anterior intestine and show spermatogenetic stages maturing in a caudo-cephalic and centripetal direction. Sperm ducts join together dorsally to the intestine before opening into a pear-like caudal organ. An unpaired, dorsal ovary lies anteriorly to the confluence of sperm ducts. Oocytes mature in a caudo-cephalic direction; no oviduct nor female pore is present. The spermatozoa are fusiform, with a very short tail. The anterior end of the sperm is a small, conic hood which corresponds to the acrosome. The following, spindle-shaped corkscrew region contains a nuclear helix surrounding a central axis made of a single, giant mitochondrion. The 9+2 axoneme is enclosed by a bilayered, obliquely striated cylinder only in the proximal tract. The morphology of the reproductive system and spermatozoa of Mesodasys species is discussed, and the hypothesis of hypodermic impregnation in this genus is supported. Received: 20 November 1998 / Accepted: 6 July 1999     

Sex and reproductive aspects in Serranus cabrilla (Osteichthyes: Serranidae): macroscopic and histological approaches

The gonads of Serranus cabrilla (Linnaeus, 1758) from the Canary Islands were studied between September 1992 and July 1993. Macroscopic classification of the maturity stages was compared with histolog ical classification of gonad characteristics, and the effect␣of the classification method on the determination of the spawning season and size at maturity was investigated. Analysis of gonad organization and development confirmed that this species is synchronously hermaphroditic. Ovarian-tissue development is asynchronous, and testicular tissue consists of continuous spermatogonia. Six stages of oogenesis and five of spermatogenesis are described, based on differences in staining and in size and on differences in the nucleus and cytoplasm structure as viewed through a light microscope. Two types of atretic degeneration of eggs are also described. Agreement between macroscopic and histological staging was low, although both methods provided similar results in spawning-season determination and size at maturity. The spawning season of S. cabrilla is from February to July, with a peak in May. Fish size at first maturity (size at which 50% of all fish sampled are in relevant maturity stage) is 152 mm standard length (SL), and size at mass maturity (size at which 95% of all fish sampled are in relevant maturity stage) is 167 mm SL. Received: 21 May 1996 / Accepted: 9 August 1996     

Megabenthic decapod crustacean assemblages on the Galician continental shelf and upper slope (north-west Spain)

The structure of megabenthic decapod crustacean assemblages on the Galician (north-west Spain) continental shelf (100 to 200 depth) and upper slope (200 to 500 m) was analyzed based on surveys carried out in autumn and spring, from 1980 to 1987. Forty species belonging to 19 families were caught. The portunid crab Polybius henslowii, a species with pelagic phases, was the most abundant species, but displayed strong spatial and temporal fluctuations. Other dominant species were the Norway lobster Nephrops norvegicus, the portunid Liocarcinus depurator, the galatheids Munida intermedia and M. sarsi, and the shrimps Solenocera membranacea, Plesionika heterocarpus, Pasiphaea sivado and Dichelopandalus bonnieri. Total abundance and biomass (average values excluding Polybius henslowii = 255 individuals and 2.06 kg/30 min tow) and species richness and diversity, H′ (6.85 species and H′ = 1.45 per tow) displayed a significant positive correlation with depth, and strong interannual fluctuations. The factors determining community organization were depth and, to a lesser extent, spatial structure. There was clear evidence of bathymetric zonation, differentiating between species characteristic of the slope (D. bonnieri and Pasiphaea sivado), shelf-slope edge (Macropipus tuberculatus, Pontophilus spinosus, Munida sarsi, S. membranacea, Processa spp.) and shelf (L. depurator, Macropodia tenuirostris, Paguridae and Chlorotocus crassicornis). The spatial zonation was related to changes in oceanography and sediment along the continental margin. Goneplax rhomboides, N. norvegicus, C. crassicornis and Alpheus glaber are benthic species which generally exhibit burrowing behaviour, and they were found mainly in the southern area where there are fine sediments due to the outwelling from the Rías Baixas. Different benthopelagic shrimps (Pontophilus spinosus, Plesionika heterocarpus,Processa spp. and Pasiphaea sivado) were typical of the zone just north of Fisterra, characterized by a convergence of water masses bringing about an increase in productivity due to upwelling. The benthic anomuran and brachyuran crabs Munida intermedia, M.␣sarsi, L.␣depurator and Macropipus tuberculatus were characteristic of the northwestern zone between Fisterra and Estaca, where the infauna reaches high biomass despite coarser sediments with a lower concentration of organic material than in the southern area. Lastly, both the Paguridae and Macropodia tenuirostris were species typically found in the waters in the northern shelf. Based on interannual changes in assemblage structure, two periods could be distinguished: between 1980 and 1984, when Polybius henslowii, D. bonnieri and Pasiphaea sivado had abundance peaks; and another period from 1985 to 1987 when L. depurator, Munida intermedia, M.␣sarsi and Macropipus tuberculatus increased in abundance. Received: 21 May 1996 / Accepted: 9 August 1996     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. IV. Pseudodiaptomus marinus, the egg-carrying calanoid

Population dynamics and production of the egg-carrying calanoid copepod Pseudodiaptomus marinus were studied for a year in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large numerical peak in June and small peaks in September/October and November/December. During the study period, at least 11 generations could be detected. For each generation, the stage-specific survival from egg to Copepodite Stage (C) V was determined; it was very high during early life stages (egg to NIII), and gradually decreased beyond. On average, 94% of eggs recruited into NIII, which is strongly contrasted with very high (>ca. 90%) mortality during the corresponding stages for free-spawning copepods, i.e. Acartia omorii, Centropages abdominalis and Paracalans sp. This demonstrates that the egg-carrying strategy has a great advantage to reduce mortality in egg stage. The biomass of this species showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous somatic growth rate increased linearly with temperature. The population production rate was estimated as the sum of somatic growth of larval stages and egg production of adult females; the annual integration was 51.0 mg C m⁻³ yr⁻¹ or 0.38 g C m⁻² yr⁻¹. Received: 11 November 1996 / Accepted: 7 December 1996     

Effects of municipal piers on the growth of juvenile fishes in the Hudson River estuary: a study across a pier edge

The growth rates of two fish species, the winter flounder Pseudopleuronectes americanus (Walbaum) (19.3 to 42.6 mm total length, TL) and the tautog Tautogaonitis (Linnaeus) (23.9 to 55.9 mm TL), were used to evaluate habitat quality under and around municipal piers in the Hudson River estuary, USA. Growth rates were measured in a series of 10 d field caging-experiments conducted at two large piers in the summers of 1996 and 1997. Cages (0.64 m²) were deployed along␣transects that stretched from underneath the piers to beyond them, encompassing the pier edge (the transitional zone between the pier interior and the outside). Growth in weight (G _w ) was determined at five locations along the transect, 40 m beneath the pier, 20 m beneath the pier, at the pier edge, 20 m beyond the pier edge, and 40 m beyond. Under piers, mean growth rates of winter flounder and tautogs were negative (xˉG _W = −0.02 d⁻¹), and rates were comparable to laboratory-starved control fishes (xˉG _W = −0.02 d⁻¹). In contrast, mean growth rates at pier edges and in open waters beyond piers were generally positive (xˉG _W ranged from −0.001 to +0.05 d⁻¹), with growth at pier edges often being more variable and less rapid than at open-water sites. Analyses of stomach contents upon retrieval of caged fishes revealed that dry weights of food were generally higher among fishes caged at open-water stations (xˉ range = 0.02 to 0.72 mg dry wt) than at pier-edge (xˉ range = 0.01 to 0.54 mg) or under-pier (xˉ range = 0.03 to 0.11 mg) stations, although it was apparent that benthic prey were available at all stations on the transect. Our results indicate poor feeding conditions among fishes caged under piers, and suboptimal foraging among fishes caged at pier edges. Inadequate growth rates can lead to higher rates of mortality, and, based on these and other earlier experiments, we conclude that under-pier environments are poor-quality habitats for some species of juvenile fishes. Received: 12 March 1998 / Accepted: 9 November 1998     

Benthic fauna of a shallow-water gaseohydrothermal vent area in the Aegean Sea (Milos, Greece)

The benthic community of shallow-water hydrothermal vents (10 m water depth) in a sandy bay on the south coast of Milos, Greece was studied. The macro- and meiofauna, as well as environmental factors such as temperature, salinity, sulfide concentration, grain size composition, ATP concentration, organic carbon, chlorophyll a and phaeopigments were evaluated. Samples were taken along transects from a seagrass meadow into a hydrothermally active area. Hydrothermally active spots were distinguished from the surrounding sediment by their black sediment coloration and an overlying white bacterial mat. For comparison, a control transect from a seagrass meadow into bare sand in a non-vent area in the same bay was studied. Overall, we found decreasing faunal diversity from the seagrass bed towards the hydrothermally active area. Along this gradient, four different zones could be distinguished. (a) The seagrass area had a relatively low diversity compared to the control stations, but much higher diversity than in the areas directly influenced by hydrothermal activity. (b) The border zone between the seagrass and the hydrothermal area, with moderate sulfide concentrations, pH values, seawater salinity and temperatures, was inhabited by an impoverished community dominated by the polychaete Capitella capitata, the nematodes Oncholaimus campylocercoides and Sabatieria sp., and an as yet undescribed chromadorid nematode (Chromadorina sp.). (c) The transition zone at the border of the hydrothermal area proper, with temperatures and sulfide concentrations higher than in the border zone, was characterized by high salinity (60‰) and low pH values. Along with the gastropod Cyclope neritea, single specimens of C. capitata and O. campylocercoides occurred. (d) The hydrothermal area proper, characterized by high sulfide concentrations, had high sediment temperatures and high salinities. The pH was as low as 6. Here, the gastropod Cyclope neritea was the only species occurring in considerable numbers. The gaseohydrothermal activity seems to largely exclude the surrounding fauna in a graded zonal pattern. The increasing hydrothermal stress entrains a change of the ambient biota towards a depauperate consortium of tolerant, opportunistic species, e.g. Capitella capitata (Polychaeta) and Oncholaimus campylocercoides (Nematoda). Although the hydrothermal area proper was characterized by a thick bacterial mat, the benthic community was characterized not by bacteria feeders but by scavenging species. Received: 13 September 1996 / Accepted: 24 October 1996