Simultaneous hermaphroditism in the marine shrimp Lysmata wurdemanni (Caridea: Hippolytidae): an undescribed sexual system in the decapod Crustacea

Simultaneous hermaphroditism with outcrossing, a previously unconfirmed sexual system in decapod crustaceans, is documented in the shrimp Lysmata wurdemanni (Gibbes), using time-lapse video observations on mating in pairs of “female-phase” (FP) individuals. Copulations between FPs resulted in successful spawning and development of embryos. However, female-phase hermaphrodites maintained in isolation were unable to self-fertilize spawned eggs. All smaller individuals possessed characters typical of caridean males, including male gonopores, appendices masculinae, ejaculatory ducts, and cincinnuli on the first pleopods. However, the gonad of these male-phase (MP) individuals was an ovotestis with an undeveloped ovarian portion. FPs, which spawn eggs and incubate embryos, also had male gonopores and an ovotestis terminating in ejaculatory ducts containing sperm. In FPs, male pleopod characters were absent or reduced, and a female incubatory character, expanded pleopod flanges, showed a greater relative size. Smaller size classes of the population were composed only of MPs while larger size classes consisted of both MPs and FPs. “Transitionals,” individuals with MP characters but an ovotestis containing vitellogenic oocytes, were rare and overlapped completely in size with FP and larger MP individuals. It is suggested that not all MPs transform into FP hermaphrodites. Social interactions may mediate MP change into FP hermaphrodites in L. wurdemanni. The social system of L. wurdemanni and other Lysmata spp. may determine the exact form of the sexual system in these species. Received: 26 October 1997 / Accepted: 21 April 1998     

Manipulation of food and photoperiod promotes out-of-season gametogenesis in the green sea urchin, Strongylocentrotus droebachiensis: implications for aquaculture

Green sea urchins, Strongylocentrotus droebachiensis OF Müller, collected off the coast of New Hampshire, USA, in late February 1995, were brought into the laboratory and fed an artificial diet ad libitum, and subjected to a photoperiod advanced by 4 months. During this study, temperatures and salinities for experimental urchins mirrored those recorded at the collection site. We examined the effects of changes in feeding regime and photoperiod on gametogenesis and compared the experimental urchins with those from the source population. During the 7-month period, experimental urchins showed no detectable changes in mean test height or diameter. Experimental urchins had a significantly higher gonad index (GI) in March, April and May (18 ± 6%) compared with field urchins in March (11 ± 3%). Subsequently, experimental urchins had a mean monthly GI of 25 to 30%, while the mean GI for field urchins was 11 to 13%. Gonial cell mitosis and gametogenesis occurred earlier in experimental male and female urchins compared with field urchins. Stereological and histological observations and stage–frequency data showed that the ovaries of experimental urchins were large because of the accelerated development of nutritive phagocytes, the volume fraction (V _v) of which was 89 to 90% of the gonad, while new vitellogenic primary oocytes occupied <1% V _v. In males, stereological and histological observations and stage–frequency data suggested a mobilization of materials from the nutritive phagocytes beginning between June and August, i.e. earlier than in females, and, by September, new gametes occupied a V _v of 49 ± 3% of the testes. Oocyte size–frequency distributions demonstrated that most primary oocytes were <80 μm in diameter between March and September, suggesting that cold temperatures may be needed for completion of vitellogenesis. We describe changes in the two principal cell types in the germinal epithelium of urchin gonads and indicate how knowledge of their population dynamics may be useful in aquaculture applications. Received: 21 February 1997 / Accepted: 3 June 1998     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Distribution and relative abundance of Pseudocalanus moultoni and P. newmani (Copepoda: Calanoida) on Georges Bank using molecular identification of sibling species

The sibling species, Pseudocalanus moultoni (Frost, 1989) and P. newmani (Frost, 1989), occur sympatrically on Georges Bank. Taxonomic discrimination of the species relies on subtle morphological characteristics, making routine identification of the species very difficult. DNA sequence variation of two mitochondrial genes, 16S rRNA and cytochrome oxidase I (COI), reliably discriminated P. moultoni and P.␣newmani. Levels of DNA sequence variation for both genes were consistent with those between species of calanoid copepods. A molecular systematic protocol (based on allele-specific PCR amplification) was designed from the COI sequences and used to discriminate females of the two species. The distributions and relative abundances of the two species were mapped for April 1996 based on samples of 15 to 30 females from 12 collections across Georges Bank. The results of this study indicated that P. moultoni females predominated along the northern flank of Georges Bank, while P. newmani females were common on the southern flank, deeper than the 60-m isobath. Received: 16 May 1997 / Accepted: 26 April 1998     

Effects of Trichodesmium spp. blooms on penaeid prawn larvae

The effects of blooms of the cyanobacterium Trichodesmium spp. on penaeid prawn larvae were examined using in situ and laboratory rearing experiments and plankton surveys in Albatross Bay, Gulf of Carpentaria, Australia. The in situ experiments demonstrated that, during a bloom of Trichodesmium spp., larvae of the prawn Penaeus merguiensis did not develop beyond the first protozoea stage, and survival was low compared with times when diatoms were dominant in the same study area. Laboratory experiments confirmed the in situ results. None of the prawn larvae fed Trichodesmium sp. in laboratory experiments developed beyond the first protozoeal stage. In contrast, 94% of prawn larvae fed the green flagellate Tetraselmis suecica successfully developed to the second protozoea stage. Electron microscopy of larvae gut-contents revealed that Trichodesmium spp. were ingested by larvae but were of no nutritional value, resulting in starvation. A 7 yr plankton survey, from 1985 to 1992, showed that minimum abundance of prawn larvae occurs during the annual summer blooms of Trichodesmium spp. and that maximum abundance of prawn larvae generally occurs just after the bloom. There was a negative correlation between the abundance of larvae and the abundance of Trichodesmium at individual sites, one offshore and one inshore, indicating that the blooms affect the survival of larvae. We conclude that variations in both timing and magnitude of Trichodesmium blooms are important determinants of prawn larvae abundance in Albatross Bay. Received: 28 April 1997 / Accepted: 2 April 1998     

Feeding in myodocopid ostracods: functional morphology and laboratory observations from videos

High-resolution videos, scanning electron microscopy and histology were used to study the feeding mechanism of myodocopid ostracods from the Pacific Coast of Japan, as exemplified by Vargula hilgendorfii (Müller, 1890) and a few other cypridinid species. Ostracods observed in the laboratory were attracted to a wide spectrum of natural food sources, behaving as predators of living prey (e.g. polychaete annelids), as opportunistic scavengers on dead animals (e.g. annelids, fishes, squid), and also consuming artificial food. Food sources may be detected by chemoreception. The fourth limb (endopodites with strong sclerotized setae) and the furcal lamellae (claws with teeth) act in coordination to abrade and eventually tear open the protective integument of living/dead prey such as annelids. The mandibular palps are used mainly to hold the food. Food sections and soft-body contents are transferred to the mouth by the fourth limb (endopodial “rake”) and fifth limb (exopodite with pectinate setae) and are passed to the oesophagus by the endites (mandibles, fourth and fifth limbs). Food is subsequently pumped up to the stomach by peristaltic contractions of the oesophagus (ring muscles) and stored in the stomach pouch. The upper lip of bioluminescent (V. hilgendorfii) and non-bioluminescent species of Cypridinidae often contact food, suggesting that some of the glands housed in this organ may emit digestive enzymes prior to ingestion. Ostracods are able to ingest massive quantities of food within a few minutes and to survive starvation for several weeks. In V. hilgendorfii, the midgut is a huge sac-like organ with no partition and is lined with a single layer of columnar epithelial cells. No differentiated hepatopancreas is present. The cypridinid produces a single faecal pellet wrapped in a thin reticulated, peritrophic membrane. Myodocopid ostracods exhibit a wide range of feeding strategies (detritus-feeding, comb-feeding, scavenging, predation, ectoparasitism) in both benthic and pelagic niches, and constitute a substantial source of food for many zooplankters. Adaptation of cypridinids to scavenging/predation is reflected in the morphology of their furcae, mandibles, fourth and fifth limbs, and their digestive system. Palaeontological data suggest that early Triassic cypridinids and possible late Ordovician myodocopids may have been carnivorous scavengers feeding on carcasses of cephalopods (ammonoids or orthoconic nautiloids), thus playing the same role of “recyclers” as modern representatives of the group. Received: 2 January 1998 / Accepted: 23 July 1998     

Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata

Dipolydora armata (Langerhans, 1880) is a small (4 to 5 mm) spionid polychaete found burrowing in the calcareous hydrozoan Millepora complanata Lamarck, 1816, on coral reefs at Barbados, West Indies. It excavates complex networks of interconnecting burrows and forms aggregations of worms in cavities within branches of the coral. Adult worms have a mixed feeding mode (suspension feeding and deposit feeding). Size–frequency distributions of worms in branch samples suggest that they mature in a single year and that reproduction occurs throughout the year. Burrow openings on the surface of the coral develop distinctive, erect spines caused by combined growth of worm tubes and host tissue. Millepore zooids were absent in the vicinity of tube openings and on spines, and thus the potential feeding surface of the coral will be reduced in heavily colonized branches. Burrows and openings were densest at the bases of millepore branches where weakening of the skeleton would be expected to occur. The absence of openings near the branch tips suggests difficulty in larval settlement there, amongst stinging zooids. Reproduction␣and larval development of the worms were examined, and a sequence of larval stages from one to 20 segments and a juvenile stage of 22 segments are described. Eggs are deposited in brood sacs attached to the burrow wall, and the larvae feed upon nurse eggs (adelphophagy). The presence of larvae and juveniles occurring free in the burrows suggests that larval development may be completed within the host coral as an alternative or in addition to a planktonic larval phase. Lack of provisional larval setae, early development of adult capillary setae, production of special spermatophores and a protracted breeding cycle in D. armata are all traits which would favour complete development within the host skeleton. Received: 6 March 1997 / Accepted: 25 October 1997     

A correction to: biological and oceanographic insights from larval labrid (Pisces: Labridae) identification using mtDNA sequences

Previously published mtDNA cytochrome b sequences for Xyrichtys novacula (Linnaeus) and two morphologically distinct types of Xyrichtys larvae (Pisces: Labridae) included human cytochrome b sequence, presumably due to sample contamination and/or poor preservation of sample DNA. Those sequences had been used to identify the two types of Xyrichtys larvae as X. novacula, but owing to the contamination, the identifications are invalid. Fresh specimens were collected: X.␣martinicensis (Cuvier and Valenciennes), X. novacula, X. splendens (Castelnau), and Xyrichtys sp. larvae of each of the two morphotypes (dorsal-forward eye and ventral-forward eye). The cytochrome b fragment was amplified from each specimen using the polymerase chain reaction. Comparison of the sequences with human cytochrome b sequence confirmed that the new sequences were not contaminated. The interspecific differences in Xyrichtys sequences were less than previously reported, but still greater than observed in many fish genera. Distance and parsimony analyses indicated that X. novacula and X. martinicensis were more closely related to each other than to X. splendens. This conclusion differs from our previous conclusion that X. martinicensis and X. splendens were more closely related to each other than to X. novacula. Distance and parsimony analyses also demonstrated that both larval morphotypes were X. novacula and, thus, our previous conclusions regarding larval ecology and oceanographic transport remain unchanged. Received: 12 March 1997 / Accepted: 24 October 1997     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Elemental (C, N and P) analysis of metamorphosing bonefish (Albula sp.) leptocephali: relationship to catabolism of endogenous organic compounds, tissue remodeling, and feeding ecology

Whole-body carbon (C), nitrogen (N) and phosphorus (P) content, and stable-isotope composition (¹³C:¹²C and ¹⁵N:¹⁴N), were followed during metamorphosis of bonefish (Albula sp.) larvae (leptocephali). Metamorphosing larvae depend entirely on endogenous carbon compounds (some of which contain N and P) as an energy source. Two fundamental questions are (1) Do the demands of extensive tissue remodeling during metamorphosis require the efficient retention of N and P during the catabolism of carbon compounds? (2) What effect does the lack of feeding have on stable-isotope composition? Our results showed that both C and N decreased by ～35 to 40%, reflecting the utilization of neutral lipid (triacylglycerols) and N-containing compounds (phosphatidylethanolamine and keratan sulfate glycosaminoglycan) as energy sources, and indicating that larvae have little or no capacity to retain N. Evidence suggested that collagen breakdown, measured as a loss of hydroxyproline content, also contributed to N loss. Stable-isotope ratios, expressed as ¹³C and ¹⁵N, showed no statistically significant differences in early and advanced metamorphosing larvae. In contrast to C and N, phosphorus was conserved during metamorphosis and most probably is utilized in the increased bone mineralization occurring in advanced larvae. We show, however, that advanced larvae are P-limited and that normal ossification is dependent upon a supply of exogenous P obtained after the resumption of feeding. The N:P ratio of 12.3 in early larvae decreased to 8.1 in advanced larvae owing to the conservation of P as N was lost. The mean ¹⁵N value in early metamorphic larvae (11.6‰) is consistent with results from other studies, and provides further support for the view that premetamorphic leptocephali feed at a very low trophic level.