Poecilogony in an estuarine opisthobranch: planktotrophy, lecithotrophy, and mixed clutches in a population of the ascoglossan Alderia modesta

A San Diego population of the opisthobranch mollusc Alderia modesta (Lovén, 1844) exhibits poecilogony, the presence of two development modes within a single species. In spring, half of the adults spawned masses containing ∼300 eggs with a mean diameter of 68 μm. After 3 d, these egg masses hatched planktotrophic veligers with a maximum shell dimension of 116 μm. The remaining adults spawned masses containing ∼30 eggs with a mean diameter of 105 μm. These egg masses hatched after 5 to 6 d, releasing lecithotrophic larvae with a maximum shell dimension of 186 μm. About 1% of field-collected adults produced mixed clutches containing a continuum of larval sizes, spanning the size extremes of planktotrophy and lecithotrophy and hatching larvae with a mean maximum shell dimension of 152 μm. Adults producing planktotrophic and lecithotrophic larvae were interfertile, and no hybrid breakdown was observed through the F3 generation. When starved, adults which previously produced only lecithotrophic larvae switched to producing planktotrophic larvae or mixed clutches with both planktotrophic and lecithotrophic larvae. Sequence-polymorphisms from a portion of the mitochondrial cytochrome c oxidase I gene support the conclusion that the two reproductive morphs represent a single species. Most of the lecithotrophic larvae and a small percentage of the larvae from mixed clutches were metamorphically competent within 3 d of hatching. A. modesta is the only molluscan species as yet known to have both planktotrophic and pelagic lecithotrophic development within a single natural population. Received: 14 August 1997 / Accepted: 11 April 1998     

Reproduction and development of Sabella media,a sabellid polychaete with extratubular brooding

Known reproductive patterns in the polychaete family Sabellidae include: (1) broadcasting of gametes, (2) depositing of benthic egg masses, (3) brooding outside the lip of the tube, and (4) brooding within the tube. There is little information for sabellids in the third category; there-fore we have studied the reproductive biology of Sabella (=Demonax) media (Bush), one of the few species known to brood its early developmental stages outside the tube. A total of 32 adults of S. media with egg cocoons were collected from the surfaces of compound ascidians (Aplidium sp. and Didemnum sp.) and from holdfasts of a brown alga (Hedophyllum sessile) at Eagle Cove, San Juan Island, Washington, USA, from June 1981 through June 1982, between tidal levels of 0.0 and 0.6 m. The donutshaped cocoons, each containing over 1 000 eggs, are deposited just outside the tube aperture from April through September. Larval development takes place within the cocoon through the 3-setiger stage, at which time the larvae emerge (about 8 d after egg deposition). The newly hatched larvae are demersal and swim near the bottom of the culture dish for 1 d before settlement. Metamorphosis takes nearly a week, and initial formation of the tube is also gradual. The reproductive patterns within the family Sabellidae are discussed, and the adaptive significance of extratubular brooding considered.     

Effects of self-fertilisation and other factors on the early development of the scallop Pecten maximus

Adult Pecten maximus (L.) were dredged off north-east Anglesey, Wales, UK, during 1981. A 2×5 factorial mating was carried out involving self- and cross-fertilisation and the use of stripped spermatozoa. Assessments of yield, normality, and larval size were made at the D larva stage and larval size and mortality were measured after a fortnight's growth of the veligers. Underlying genetic variation was evident at all stages, with egg and sperm generally having a significant interactive effect. Cultures sired with stripped spermatozoa had fewer larvae, with more abnormality, grew slower and suffered higher mortality than most other cultures. Larvae from selfed cultures grew significantly slower than all other larvae. Data from past larval cultures also show that selfed larvae have a reduced growth rate. it is suggested that stripped spermatozoa may interfere with egg and sperm interaction at fertilisation, thus reducing the viability of larvae. On the other hand, the poor growth rate of selfed larvae is probably due to overall reduced heterozygosity compared to outbred larvae.     

Distribution pattern, reproductive traits, and molecular analysis of two coexisting vermetid gastropods of the genus Petaloconchus: a Caribbean endemic and a potential invasive species

Vermetid gastropods are characterized by complex taxonomy and unusual ecology. A survey of the fouling community in the intake channel at the Planta Centro Power Plan in Puerto Cabello, Venezuela, found massive colonies of vermetid gastropods of the genus Petaloconchus. We arbitrarily named two prevalent varieties as black and brown-orange morphs, distinguishing based on the color of their soft bodies. Spatial distribution was different for the morphs. The black morph was present along the jetty, with higher average densities in the shallower intertidal area (410 vs. 143 ind m⁻²), while the brown-orange morph was only present at the initial part of the channel (μ = 83 ind m⁻²). Both produced small eggs (142 vs. 180 μm diameter on average), with the orange-brown eggs being slightly larger, but the morphs differed in other reproductive aspects. The brown-orange morph produced significantly fewer capsules (up to nine simultaneously per female) with fewer embryos (average of 27 viable embryos/capsule) that hatched at a larger size (577 μm on average), allocating about 17.64% of the initial egg production to nurse eggs. In contrast, the black morph produced up to 14 capsules simultaneously, hatched an average of 178 veligers of 212 μm, and produced no nurse eggs. The intra-capsular development reached a more advanced stage in the brown-orange morph than in the black one. Several cytochrome subunit I (COI) and 16S rRNA (16S) haplotypes were found for the black morph, when compared to only one haplotype for both genes present for the brown-orange morph. Preliminary phylogenetic analyses separated the morphs into different clades, supported by robust bootstrap values and posterior probabilities (>98). Our results indicate that the morphs are two different species: the black morph was identified as Petaloconchus cf. varians and the brown-orange morph as a non-described species, Petaloconchus sp. (orange). The first is endemic to the Caribbean, while the second is potentially an introduced species.     

Diapause embryos in the neustonic copepodAnomalocera patersoni

The neustonic copepodAnomalocera patersoni Templeton, collected from the Gulf of Naples from January to May 1988, laid subitaneous and diapause eggs that were of equal size and dark and smooth in appearance. In January and February, most eggs were subitaneous and hatched within 2 to 3 d at room temperature. Conspicuous numbers of diapause eggs were first obtained in February when several clutches contained a second egg type which did not hatch in a subitaneous manner. By March and April, most clutches contained only diapause eggs. Transmission electron microscope studies of subitaneous and diapause eggs revealed striking morphological differences. Subitaneous eggs had a thin vitelline coat covering the plasma membrane, whereas diapause eggs were enveloped by a complex four-layer structure which is assembled after spawning and which probably serves as a protective shell during dormancy. No major morphological differences were discernible in the oogenic cycle of females spawning either of the two egg types. Diapause eggs were shown to be embryos in early stages of development, and segmentation of these embryos was arrested for the entire 6-mo period of investigation. Diapause eggs hatched after 7 mo of dormancy.     

Reproductive success of parasitized males in a marine reef fish

Parasitism is hypothesized to reduce reproductive success in heavily parasitized males because females may preferentially mate with less parasitized males (parasite-mediated sexual selection) or parasites may compromise male competitiveness. In marine systems, this hypothesis is largely unexplored. This paper provides the first confirmed record of a copepod ectoparasite (Caligus buechlerae Hewitt 1964) on the common triplefin (Forsterygion lapillum) and evaluates the hypothesis that males parasitized with C. buechlerae experience lower reproductive success than unparasitized males (as determined by the presence and area of eggs within male nests). We found that 38 % of males we surveyed were infected with at least one C. buechlerae, with a median of two individuals per infected male. About 32 % of males were defending eggs, with 62.5 % of those males infected with at least one parasite. Males of greater total length (TL) were both more likely to be infected and more likely to be defending eggs. However, when statistically accounting for the effects of TL, parasite infection had no effect on the probability of defending eggs, or the average surface area of eggs when present. Positive covariation in fish length, the presence of eggs and parasite infection observed here potentially suggest that the importance of parasitic infection on reproductive success may depend upon the strength of selection for larger male body size. Our study is one of the few studies to investigate the effects of ectoparasites on reproductive success in reef fish and also provides a quantitative measure of infection for a widespread species within New Zealand.     

Description of the early development of the halibut Hippoglossus hippoglossus and attempts to rear the larvae past first feeding

Rearing experiments on the halibut Hippoglossus hippoglossus (L.) were carried out using gametes from parents caught at a depth of 600 to 700 m off the Norwegian coast in February 1980. After fertilization, the average egg diameter was 3.08 mm, average dry weight 1 038 μg and neutral buoyancy was at 36.5‰ S. The eggs hatched after 20 d at 4.7°C, 18 d at 5°C and 13 d at 7°C. Survival to hatching was better when antibiotics were used. At hatching the larvae were 6.4 mm long, there were no functional eyes or mouth, but prominent neuromast organs were present. Resorption of yolk lasted 50 d at 5.3°C; the eyes and mouth were then functioning and the larva was about 11.5 mm long. The larvae were offered zooplankton as food, but with little success in initial feeding. A few larvae fed and grew in 2 500-litre plastic bags, one reaching a length of 24 mm after 90 d.     

Resting eggs of zooplankton (Copepoda and Cladocera) from the Kiel Bay and adjacent waters (southwestern Baltic)

Resting eggs of four species of calanoid copepods and three species of cladocerans were collected from sediments up to 5cm depth from the Kiel Bay and adjacent waters in the southwestern Baltic Sea during April–May 1994. All but one species of cladoceran was successfully hatched/reared in the laboratory. In the Kiel Bay, egg abundances varied from 1.8x10⁵ to 7.4x10⁵ m^-2. Hatching success of copepod eggs collected from all depths was high (49 to 94%), but was 0 to 79% for cladoceran eggs. Darkness did not seem to affect hatching. Eggs found in the 4 to 5 cm layer of sediment were estimated to be about 15 yr old, showing the presence of an egg bank in the Baltic. Formation of resting eggs may be a genetic trait acquired during the ice ages.     

Water and salt balance of halibut eggs and larvae (Hippoglossus hippoglossus)

Eggs of halibut [Hippoglossus hippoglossus (L.)] have a negative buoyancy in sea water of 35 S, in contrast to eggs of most flatfish species. The cause of this was investigated. The osmolality of the yolk is 350–420 mOsm during embryonic development. This is within the range for marine teleost serum and for yolk of pelagic eggs. Concentrations of major inorganic ions are comparable with those of pelagic plaice eggs [Pleuronectes platessa (L.)]. The values for Na⁺, K⁺ and Cl^- are 6, 85, and 64 mmol · (1 H₂O)^-1 after fertilization, and at the time of hatching the corresponding values are 17, 11, and 80. Large amounts of other inorganic constitutents are excluded for osmotic reasons. Malfunction in the regulation of osmolality or of inorganic constituents is thus unlikely to be the cause of negative buoyancy. The relative dry weight of the chorion (egg shell) in halibut eggs is less than in several pelagic egg types, excluding the chorion as the main contributor of negative buoyancy. It is concluded that a high content of organic matter in the rest of the egg is the cause of the negative buoyancy.     

Precise sex ratio in a parasitic wasp: the result of counting eggs

Summary The parasitic wasp Nesolynx albiclavus maintains a very skewed offspring sex ratio (ca. 4% males). We studied proximate, causal factors of the son-daughter choice mechanism to obtain insight into the precision of the procedures and the underlying rules. We analyzed the number of males in clutches produced during sessions of 12 or 24 h. The results show that the timing of the next male egg depends on the time elapsed and the number of eggs laid since the previous one. The influence of the passage of time per se during non-laying periods was reduced in an experiment with 4-h periods. The data show that the first male egg is produced by a fairly rigid process: about the sixth egg in a run will be left unfertilized. No further male egg will appear in the next 10–20 eggs if they are laid in rapid succession.     

Ecophysiological studies on the developing eggs and embryos of the European lobster Homarus gammarus

During the course of its embryonic development, the European lobster Homarus gammarus Linnaeus exhibits progressive increases in content of water (54.0 to 83.1%), ash (2.7 to 16.7%), protein (47.4 to 50.9%) and non-protein nitrogen (1.0 to 2.4%), and steady decreases in content of fat (43.8 to 25.4%) and energy (6343 to 5431 cal/g dry organic substance). Cumulative yolk utilization efficiency during the total development is 81.8% for dry weight; the corresponding value for energy is 60.1, for protein 75.6 and for fat 47.4%. Energy content of a single egg is 10.49 cal. Of 4.20 cal expended for metabolic processes of the embryo, only 13.3% energy is drawn from protein oxidation; fat oxidation supplies as much as 87.7% energy, that of carbohydrate only 2.3%. Embryonic development results in a remarkable decrease in net yolk utilization efficiency, which falls from 85.5% in the early developmental stages, to less than 70% in later developmental stages. The mean dry weight of a single egg membrane increases from 38 g (2.2% of egg weight) in a freshly laid and attached egg, to 81 g in an egg with an almost completely developed embryo. This result supports the earlier observation of Cheung (1966) that the formation of the inner chitinous egg membrane occurs after the egg is laid and attached to the setum. Protein seems to be the major constituent of the egg membrane (4049 cal/g dry weight), which has the following composition: protein 70.4%, non-protein nitrogen 0.13%, ash 2.83%. Initial permeability of the egg membrane to water (about 6% of the total water requirement is let in) is followed by a period during which the egg membrane is almost impermeable to water (stages I to III); the egg membrane becomes permeable to water again and lets in 85% of the total water requirement (the rest, i.e. 9%, is metabolic water) at a relatively advanced stage of development. These assumed changes in egg membrane permeability appear to be indicative of variations in the egg's osmoconcentration leading to shiftings in net transport of water. Rates of water and salt uptake during embryonic development are essentially parallel (Fig. 1). The egg membrane remains permeable to salts throughout development; salt intake almost doubles after the egg passes through stage III. A single egg, weighing 3.7 mg requires 4.9 mg water for successful completion of embryonic development. The imbibition of water by the developing marine demersal egg seems to (1) serve in osmotic hatching; (2) float the hatched larva by means of specific gravity reduction; (3) aid the larva to quickly adjust its body temperature. The simple osmotic hatching mechanism, proposed by previous workers, seems to be inadequate to account for the events and timing of the hatching process in the lobster. It is suggested that hatching time is determined not solely by increased internal pressure caused by inflow of water and salts, but also by some unknown internal factor. In the lobster egg, as well as in many other marine demersal eggs, protein metabolism is suppressed to a considerable extent, and fat metabolism is geared up Thus, the non-cleidoic lobster eggs exhibit metablic properties which are typical of cleidoic eggs. This finding is discussed in the light of Needham's (1950) concept of cleidoicterrestrial and non cleidoic-aquatic eggs.Dedicated to Professor P. Kochukutta Menon, Montreal, on his 62nd birthday, June 3, 1970.     

Yolk utilization and hatching time in the Canadian lobster Homarus americanus

During the course of its embryonic development, the Canadian lobster Homarus americanus Milne-Edwards exhibits steady increases in water content (56.2 to 86.8%) and ash (5.8 to 21.2%), and a progressive decrease in energy content from 6636 to 4292 cal/g dry weight. Mean dry weight of a single egg is 965 g, equivalent to 6.4 cal; a freshly hatched egg. The lobster hatches about 1,500 larvae per night over a period of 4 to 5 days. Dry weight, ash and calorific contents of larvae hatched on different days show considerable variations. After larvae hatch on the first day, continuous salt absorption by eggs to hatch on subsequent days leads to a steady increase in ash content from 143 g/larva hatched on the first day to 255 g/larva hatched on the fourth day, and consequently, to an increase in dry weight from 854 to 956 g/larva. Metabolism of embryos (0.1 cal/day), which are yet to be hatched on subsequent days, depletes the calorific content per unit weight (from 4637 to 3837 cal/g dry weight) as well as per larva (from 3.98 to 3.67 cal).Dedicated to my colleague K.-H. Schumann (Biologische Anstalt Helgoland) who died on December 12, 1969 due to a diving accident.     

Development of the pericalymma larva of Solemya reidi (Bivalvia: Cryptodonta: Solemyidae) as revealed by light and electron microscopy

Solemya reidi Bernard 1980 is a gutless protobranch bivalve known to possess intracellular chemoautotrophic bacterial symbionts in its gill. A light and electron microscope study on the embryology and larval development of S. reidi provides data for the bivalve Subclass Cryptodonta. S. reidi spontaneously spawned large eggs (271 m in diameter), which developed within individual gelatious egg capsules. The first several cleavages were equal and a distinct molluscan cross was formed at the animal pole of the embryo, features previously unreported in bivalve development. Lecithotrophic pericalymma larvae (similar to the larvae of paleotaxodont protobranch bivalves and aplacophoran molluscs) hatched at 18 to 24 h and remained in the water column for a further 5 d at 10°C. At hatching, larvae measured from 360 to 440 m in length and from 225 to 265 m in cross-sectional diameter. Definitive adult structures developed within an epithelial locomotory test entirely covered with compound cilia. The test histolysed at metamorphosis and was ingested throught the mouth into the perivisceral cavity. Length and height of the shell following metamorphosis was 433 m (±42 m, n=16) and 282 m (± 29 m, n=13), respectively. Primary data and data from the literature show that the type of larval development in both paleotaxodont and cryptodont bivalves cannot be reliably estimated from egg or prodissoconch sizes.     

Reproductive periodicity of meiobenthic copepods: Seasonal of continuous?

Much of the literature on meiofauna reproductive periodicity suggests continuous reproduction throughout the year. Field data collected over a two-year period in the North Inlet area, South Carolina (USA) indicate that the dominant meiobenthic copepods at each station sampled are in a reproductive state all year whereas less abundant species have distinct seasonal reproductive periods. Reproduction throughout the year is not restricted to, nor does it encompass all of, the interstitial copepods with a low number of eggs. In those species where seasonality is evident, there is a temporal separation of the most closely allied copepods, and when one species is present the other is absent. Alternation of reproductive cycles in these species reduces competition and allows for maximum resource utilization.     

Ecology of the northern Red Sea crinoids and their Epi- and Endozoic fauna

Fourteen crinoid species are found along the coral reefs of the northern Red Sea that form aggregations of different species at various depths. The shallowest aggregation consists partly of Lamprometra klunzingeri and Capillaster multiradiatus, occurring subtidal to a depth of 2 to 5 m. Of this group, Heterometra savignii inhabits depths down to 12–15 m. Further down, another group of 10-armed crinoids occurs, dominated by Decametra chadwicki and Oligometra serripinna. The deepest aggregation of crinoids occurs at 45 m; its most common members are Colobometra arabica and Comaster distinctus. Feeding on micro- and nanoplankton, shallow-water crinoids show a circadian activity rhythm; in deeper aggregations (10 to 12 m), this behaviour changes with decreasing illumination, to a diurnal activity rhythm. The symbiotic animals living on crinoids comprise 27 taxons, among them Copepoda (6 species), Mollusca (2 species), Polychaeta (11 species, especially Myzostomida); Ophiurida (1 species), Crustacea, Decapoda (6 species); and the clingfish Lepadichthys lineatus. These symbiotic animals form food webs at various depths, according to the distribution of their crinoid host. The occurrence of Indo-West Pacific and Mediterranean commensals on the same crinoid in the Gulf of Aqaba is of special interest.     

Development and metamorphosis of the planktotrophic larvae of Rostanga pulchra (Mollusca: Nudibranchia)

Rostanga pulchra MacFarland, a small (1 to 2 cm) dorid nudibranch, lays an average of 7000 eggs in the laboratory during a period of 30 days in the summer. The veligers hatch 15 to 16 days after oviposition and it takes another 35 to 40 days to become competent for metamorphosis at a temperature of 10° to 15°C. Larval cultures were maintained initially at a concentration of 500 veligers per 100 ml of filtered sea water (antibiotics added). During the planktotrophic phase of development, the veliger grows from 150 to 300 m in shell length. Although the veligers are generalists in their food preference, the best result (faster growth) was achieved by feeding them with a combination of Monochrysis lutheri and Isochrysis galbana. The concentration of food cells was kept at 10⁴ cells per ml of culture media and was supplied every 2 to 3 days. A veliger which is competent to metamorphose is identifiable morphologically by its propodium, eyespots, rhinophores, and spiculated dorsal papillae. The entire metamorphic process lasts 24 h when a suitable substrate such as the food sponge Ophlitaspongia pennata is provided. The competent veliger is able to delay metamorphosis for at least 3 weeks. Juveniles were kept in the laboratory for 70 days and, during this period, grew to a length of 4.5 mm.     

Developmental types of shallow-water asteroids of McMurdo Sound,Antarctica

The mode of development was ascertained for 14 of the 16 species of sea stars known to occur in shallow waters of McMurdo Sound, Antarctica (77°51S; 166°40E). The species were collected between September 1984 and December 1985. Females of three species,Odontaster validus, O. meridionalis andPorania antarctica, spawn small to moderate eggs (0.17 to 0.55 mm), have a high fecundity, and produce feeding larvae. Females of an undescribedPorania species spawn a few eggs (150 to 310) that are 0.55 mm in diameter and develop into demersal non-feeding larvae. Females ofDiplasterias brucei andNotasterias armata produce a few (<300) large eggs (2.8 to 3.5 mm) and brood their young. Females of the remaining eight species have moderate fecundity and produce pelagic non-feeding larvae, as determined from egg type (buoyant, 0.54 to 1.28 mm diam) and direct observations of spawning and development. The high incidence (11 out of 14 species; 79%) of non-feeding development is consistent with predictions that environmental conditions in high-latitude regions are unfavorable for planktotrophic development. Nonetheless, most of the species surveyed (11 out of 14) had pelagic larvae, which contradicts inferences of unusual selection for benthic development in the Antarctic.     

Production, hatching success and surface ornamentation of eggs of calanoid copepods during a winter at 57°N

Close to 50 species of marine Calanoid copepods have been reported to produce diapause eggs (Engel and Hirche in J Plankton Res 26:1083–1093, 2004); eggs that are viable but require a refractory phase before they hatch, sometimes after months. Diapause eggs are often described as morphologically different with respect to egg membrane ultrastructure and having a thicker egg shell with surface ornamentation as opposed to the smooth shell found in subitaneous eggs that hatch within days (Belmonte in J Mar Syst 15:35–39, 1998; Chen and Marcus in Mar Biol 127:587–597, 1997; Castro-Longoria in Crustaceana 74:225–236, 2001). Egg production rates, egg surface ornamentation, and hatching success were monitored in large aquaculture fish enclosures during winter with close to zero water temperatures (N57°). Surprisingly, all female copepods (Acartia spp.—presumably A. tonsa, and Centropages hamatus) produced eggs all through the winter with no obvious pattern with respect to light, temperature and food availability, and no diapause eggs were observed. However, individual females produced several categories of eggs with or without surface spines even within the same egg batch as evidenced by scanning electron microscopy (SEM). Four egg categories were distinguishable: ‘no spines’, smooth eggs; ‘short spines’, 5–15 μm long; ‘truncated spines’, with the spine tips cut-off <10 μm long; and ‘long spines’, up to 30 μm long. All egg categories remained unchanged with respect to surface structures from when we took them out of the incubation bottles until they hatched. In general, the frequency of ‘no spines’ was 10–40%, and most eggs were ornamented with ‘short-’ or ‘long spines’. Further, a given egg can be ornamented with all types of surface spines simultaneously, which might even be a fifth egg category. The different egg categories were all able to hatch within days when exposed to normoxic conditions suggesting that they were subitaneous.