Some observations of factors affecting embryonic and larval growth of Pollicipes polymerus (Cirripedia: Lepadomorpha) in vitro

Observations on the effects of several environmental conditions in embryonic and larval cultures are reported for the gooseneck barnacle Pollicipes polymerus (Sowerby, 1833). When the growth rates of control embryos kept in finger bowls at ambient-temperature sea water (ca. 12°–15°C) were compared to those of embryos grown under variously modified circumstances, it was found that growth rates were faster under conditions closely simulating the adult barnacle mantle-cavity (e.g. darkness and aeration). Addition of antimicrobial drugs and reduction of egg-mass size also promoted fast growth and development. Nauplii were fed several species of algae; only three species promoted growth to Stage V or older. Naupliar growth was fastest in larvae fed the combination of Prorocentrum micans/Platymonas sp. Larvae fed Prococentrum micans/pennate diatom grew slowest, and most larvae died at Naupliar Stage III. Interactions between algal species may have affected their nutritional value for Pollicipes polymerus nauplii. Less larval activity, slower growth rates, and higher mortality rates were observed in individuals given small amounts of food. The development of embryos and larvae in culture depends greatly on the culture conditions. These conditions should be described if comparison of timetables and envents are to be made between studies.     

Fertilisation, embryogenesis and larval development in the tropical intertidal sand dollar Arachnoides placenta in response to reduced seawater pH

We examined the response of the tropical sand dollar Arachnoides placenta to reduced seawater pH in experiments spanning ca. 50 % of the planktonic larval duration. A. placenta inhabits intertidal sandy beaches where we observed a minimum in situ pH range 0.06 pH units (pH 8.10–8.16). The responses of gametes and larvae to seawater pH were tested in vitro in ambient (pH 8.14, pCO₂ = 525.7 μatm, total alkalinity = 2,651 μmol kg soln^?1) and three reduced pH seawater treatments (7.8–7.0). Percentage fertilisation decreased significantly with decreasing pH across a range of sperm/egg ratios (4:1 up to 4,000:1). A. placenta reached the advanced pluteus stage in 4 days, and during this time, we saw no difference in survival rate of larvae between the ambient (67 %) and pH 7.79 (72 %) treatments. Four-day survival was, however, reduced to 44 and 11 % in the pH 7.65 and 7.12 treatments, respectively. Larval development and morphometrics varied among pH treatments. Embryos reared in pH 7.12 exhibited arrested development. Larvae reared at pH 7.65 showed delayed development and greater mortality compared with those reared at pH 7.79 and 8.14. When larval morphometrics are compared among larvae of the same size, differences in larval width and total arm length between pH treatments disappear. These results suggest that variation in larval size among the three highest pH treatments at a given time are likely the result of slower development and apparent shrinkage of surviving larvae and not direct changes in larval shape. There were no differences in the percentage inorganic content (a proxy for calcification) in larvae reared in either an ambient or a pH 7.7 treatment. The responses of fertilisation and development to decreased pH/increased pCO₂ in A. placenta are within the range of those reported for other intertidal and subtidal echinoid species from colder latitudes.     

Habitat selection in the tropical polychaeteSpirobranchus giganteus

Total body weight, tube length, abdomen weight and branchial crown weight of individualSpirobranchus giganteus (Pallas) living on four different coral species on the bank reef on the west coast of Barbados were investigated in 1986. Worms onDiploria strigosa were larger in all size parameters than those onMontastrea annularis, which were larger than those onMontastrea cavernosa, which were larger than those onPorites porites. The differences between worms onM. annularis andM. cavernosa were not significant. Variation in worm size on the different corals could result from variation in mortality and/or variation in growth. Whichever the case, larger worms may have higher lifetime reproductive success. Abdomen weight is an index of gamete production inS. giganteus, and increases with increasing body weight for worms on all corals. Moreover, larger worms do not have lower abdomen weight at a given body size than smaller worms. This may imply that faster growth does not reduce gamete production at body size. The coral species on which worms are largest are those most preferred by worm larvae in the laboratory and most heavily colonised by adult worms in the field. The results suggest adaptive habitat selection by planktonic larvae ofS. giganteus.     

Effects of gamma-aminobutyric acid on the settlement of larvae of the black chiton Katharina tunicata

Laboratory-cultured larvae of the black chiton Katharina tunicata (Wood), collected from central California in May and June of 1979 and 1980, settled preferentially on the encrusting coralline alga Lithothamnium sp. Metamorphosis, which involves the loss of the prototrochal ciliary band, occurred within a period of 2.5 h after larvae began crawling upon the alga. In response to gamma-aminobutyric acid (GABA) at a concentration of 10^-6 M, the lecithotrophic trochophore larvae ceased swimming and settled rapidly. Developmental metamorphosis did not follow settlement in the absence of encrusting coralline alga. However, when both GABA and Lithothamnium sp. were present, larval metamorphosis occurred within 2 h of settlement. Significantly different rates of settlement exist for sibling larvae of different ages; 12 d-old larvae responded more rapily and in greater numbers to treatment with GABA than 10 d-old larvae. Differences in settling rates are attributed to the onset of metamorphic competence.     

RNA-DNA ratio: an index of larval fish growth in the sea

Data on water temperature, RNA-DNA ratio, and growth of eight species of temperate marine fish larvae reared in the laboratory were fit to the equation: $$G_{pi} = 0.93{\text{ }}\operatorname{T} + 4.75{\text{ RNA - DNA}} - 18.18$$ where G_pi is the protein growth rate in % d^-1 and T is the water temperature. Water temperature and larval RNA-DNA ratio explained 92% of the variability in growth rate of laboratory-reared larvae. The model is useful over the entire range of feeding levels (starvation to excess), temperatures (2° to 20°C) and fish species studied. Estimates of recent growth of larval cod, haddock, and sand lance caught at sea based on water temperature and RNA-DNA ratio ranged from negative to 26% d^-1. These data demonstrate the importance of food availability in larval fish mortality and suggest that short-term growth under favorable conditions may be considerably higher than expected from long-term indicators. RNA-DNA ratio analysis offers new possibilities for understanding larval growth and mortality, and their relation to environmental variability.     

Egg laying and embryonic-larval development in the snail<Emphasis Type="Italic"> Thais (Stramonita) chocolata</Emphasis> (Duclos, 1832) with observations on its evolutionary relationships within the Muricidae

Oviposition and embryonic-larval development are described for the muricacean snail Thais (Stramonita) chocolata from the Southeast Pacific coast. As with numerous other muricacean snails, this species engages in communal egg laying, with females depositing egg capsules in clusters on subtidal rocks. Each cluster of capsules contains 100–150 pedunculate, ampulliform egg capsules, with each capsule containing an average of 2,600 small (130 m) eggs. Intracapsular development was followed using light and scanning electron microscopy to describe the successive embryonic stages of the species. Free-swimming veliger larvae of about 225 m length were released from capsules after 49 days incubation at 13.6°C. The planktotrophic larvae were cultured in seawater aquaria by feeding with pure cultures of phytoplankton, recording growth and form of the larvae. Larvae reached competence after 4 months at 22°C, at 1,450–1,740 m in size, and a few larvae were observed through metamorphosis and early definitive growth. The embryonic-larval development of T. chocolata coincides with the general characteristics of the ontogeny observed in other Thais species as well as of other genera of the Rapaninae such as Concholepas. This lent support to grouping these genera into a single clade. The lack of knowledge of the development of free larvae of Thais spp. means that we do not know whether these similarities also include an extensive larval phase as generally characteristic of other members of the clade. The mode of development may be useful in characterizing some clades of this family. Thus for example, the transference of some Thais to the genus Nucella (Subfamily Ocenebrinae) is supported by differences in the mode of embryonic development, which differentiates these subfamilies. Paleobiological data reported for Neogastropoda allow postulation of primitiveness in planktotrophic larval development compared to more recent developmental strategies such as direct development of different types, which characterize various clades of this family.Communicated by O. Kinne, Oldendorf/Luhe     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

Reproductive ecology of a hippolytid shrimp,Lebbeus virentova (Caridea: Hippolytidae) at the Von Damm Vent Field,Mid-Cayman Spreading Centre

Elucidation of life-history traits is essential to understand larval dispersal and population dynamics in marine benthic assemblages. This study is the first investigation of the life history of a recently described hippolytid shrimp from a deep-sea chemosynthetic environment, Lebbeus virentova Nye, Copley, Plouviez and Van Dover, 2013 at the Von Damm Vent Field (18°22N, 81°47W, ~2,300 m depth, Mid-Cayman Spreading Centre, Caribbean), using samples collected in February and June 2013. Lebbeus virentova is gonochoric and iteroparous. The sex ratio of L. virentova was significantly female biased (1:3) in February and June. The sampled population of L. virentova had a unimodal size–frequency distribution pattern in February and June, consistent with continuous recruitment and mortality. Continuous reproduction is indicated by a lack of synchrony in oocyte size–frequency distributions within both months, and asynchronous development of embryos among females, which may result in asynchronous larval release. A large embryo size in this species (2.65 ± 0.28 mm diameter) compared with other caridean shrimps suggests possible abbreviated larval development, as described in other species of the genus from non-chemosynthetic environments. Fecundity (26–94 embryos female^?1) was lower and embryo size larger in L. virentova compared with alvinocaridids at chemosynthetic environments. This suggests that there are phylogenetic constraints on reproductive features of decapods at hydrothermal vents.     

Reproduction and developmental variability in the opisthobranchTenellia pallida

Pelagic and non-pelagic development occurs within a single population of the nudibranchTenellia pallida (Alder and Hancock) from South Carolina (USA). Both developmental types occurred simultaneously, under identical conditions, and without the aid of nurse eggs. Adults laid either ≈70 μm diameter eggs that hatched as pelagic (planktotrophic?) veligers in 68 to 122 h, or ≈100 μm diameter eggs that became well-developed veligers, metamorphosed within their egg capsules, and hatched as shelled juveniles in 140 to 190 h (15°C, 30‰ S). Adults that produced pelagic larvae laid more eggs per day and per egg mass and twice the daily egg volume produced by adults of the opposing type.     

Oceanographic habitat,growth and mortality of larval anchovy (<Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>) in the northern Aegean Sea (eastern Mediterranean)

Data from two ichthyoplankton surveys carried out during June 1995 and June 1996 were used to study the broad scale distribution patterns of anchovy eggs and larvae over the northern Aegean Sea continental shelf and the regional/inter-annual variability in growth and mortality rates of larvae. Two major spawning grounds were identified. One in the east, located in the area influenced by the Samothraki gyre (SG), in which a large amount of enriched, modified Black Sea water (BSW) is entrapped and one in the west, associated with zooplankton-rich waters in the semi-enclosed Thermaikos gulf close to several river mouths. In the NE Aegean, anticyclonic gyres generated over the continental shelf and fed by the circulating stream of BSW (like the SG) may act as retention areas for larval anchovy. In the west, the high enclosure of the Thermaikos Gulf contributes to reducing offshore dispersal. Major changes were observed in egg and larval abundance as well as larval mortality between June 1995 and June 1996 in both the western and eastern part of the continental shelf. Mean abundance of eggs and early larvae was >5 times higher in 1996 than in 1995, when waters were significantly cooler, fresher and richer in mesozooplankton. Larval survival decreased from 79 to 69% day⁻¹ in the east and from 89 to 74% day⁻¹ in the west between 1995 and 1996. Hence increased egg production was coupled with higher larval mortality during June 1996. Furthermore, a highly significant positive relationship between larval mortality (Z) and mean egg abundance (A) emerged (Z = −0154 + 0.205 log[A], r ² = 0.96, n = 7) when data from this study and a similar study in the NW Mediterranean were regressed. Mean growth rate of anchovy larvae in the study area (∼0.5 mm day⁻¹) did not differ significantly between areas/years. A marked ontogenetic change was observed in the otolith size/recent otolith growth-on-fish size relationships, which exhibited significant inflection points at ∼6 mm formalin preserved length. This change seems to coincide with performance (e.g., catchability) and behavioral changes (e.g., onset of vertical migrations) in European anchovy associated with the development of the caudal fin (the flexion stage).     

Biochemical changes during larval development of the xanthid crab Rhithro panopeus harrisii. II. Nucleic acids

Deoxyribonucleic acid (DNA) and ribonucleic acid (RNA) contents were measured daily during the zoeal development of the crab Rhithropanopeus harrisii (Gould). DNA concentration (per unit protein) decreased as larvae increased in size. Total DNA content per larva showed an abrupt increase at the second molt and increased steadily during the third and fourth zoeal stages. Ratios of fresh weight to DNA were highest during the first two zoeal stages, dropped sharply at the second molt, then increased during the third and fourth zoeal stages. RNA:DNA ratios showed cyclical activity apparently related to the molt cycle. Assuming that fresh weight: DNA ratio reflects cell size and that DNA content reflects cell number, growth during the first two stages is due primarily to increase in cell size and during the last two stages to a combination of increase in cell size and number. Assuming further that RNA:DNA ratios reflect protein-synthesis activity, such activity in these larvae follows a rhythm related to the molt cycle.     

The influence of irradiance on the biochemical composition of three phytoplankton species and their nutritional value for larvae of the Pacific Oyster (Crassostrea gigas)

Three species of phytoplankton grown at high (HL) or low light (LL) were fed as saturating rations to laboratory-reared larval Crassostrea gigas. Larval C. gigas fed diets of HL grown Chaetoceros gracilis and HL grown Isochrysis aff. galbana grew faster than those fed LL grown cells of the same phytoplankton species. Faster growth of C. gigas larvae was consistently associated with increases in the percent composition of short chain saturated fatty acids (FA) 14:0+16:0 in the HL grown cells. There were no consistent and significant differences between HL and LL grown phytoplankton cells in their content of carbon, nitrogen, protein, lipid or carbohydrate. Intraspecific increases in percent composition of essential fatty acids (EFAs), 20:53 and 22:63, in the phytoplankton were not associated with improvements in the growth or survival of the oyster larvae. Oyster larvae fed diets of Phaeodactylum tricornutum with a relatively high proportion of EFAs grew more slowly than those fed C. gracilis. In this experiment the proportion of dietary EFA 20:53 was negatively correlated with oyster growth rates. The faster growing oyster larvae contained relatively more of the FAs 14:0+16:0 which may be useful as measures of larval oyster condition. After a diet of one phytoplankton species for ca. 10 d, oyster larvae acquired distinctive FA profiles resembling that of their phytoplankton prey.     

Spawning,larval development and settlement behaviour of Gibbula cineraria (Gastropoda: Prosobranchia) with a reappraisal of torsion in gastropods

Cultures of developing larvae of Gibbula cineraria (L.) were obtained from adults spawning in the laboratory, and these cultures were reared to settlement of the larvae at 9 days. Dispersal of the outer jelly coat of the egg appeared to be delayed in the presence of spermatozoa. Early cleavage followed the typical spiral pattern, and gastrulation was by epiboly. The trochophore larvae hatched at about 28 h, before shell-formation began. The first 90° of torsion was completed between 48 and 56 h. The second part of torsion was completed within 4 days, and preliminary attempts to retract into the shell were made at 76h. After 96 h, although torsion was complete, the larvae were still unable to retract fully into the shell. The larvae remained in a stage of swimming-attempted creeping until the 4th or 5th day after torsion was complete, but were unable to pull the shell upright until the end of this time. Throughout this stage, the velum was reduced and was finally shed. The animals all died after metamorphosis, which occurred at about 9 days, when the larval shell comprised 1 1/2 whorls, with a breadth of 250 to 300m. Settlement appeared to be passive, i.e., governed by wave action and tides. In the light of these observations, theories concerning torsion are reappraised. The advantage to the larva of torsion is not clear, as the larvae are not able to retract into the shell until after metamorphosis, yet it is clear from previous authors' observations that the first part of torsion in most gastropods involves only larval structures. This makes it difficult to suggest that torsion is entirely of advantage to the adult snails. It is thus proposed that the two components of torsion may have evolved independently. The first, relatively rapid component of torsion, resulting in 90° of displacement of the mantle cavity in relation to the visceral mass, may be of advantage to the swimming larva. The final, slower component of torsion may be of advantage to the newly-metamorphosed benthic snail, and is the only component of torsion found in those gastropods which have no free-swimming larva.     

Distribution and abundance of fish larvae off the coast of West Pakistan

The material was collected during 4 different cruises on the shelf off the coast of W. Pakistan in November–December, 1964 and in March, 1967 and 1968. The hydrography as well as the zooplankton-biomass distribution are briefly described. A total of 5777 larval fish were taken in 33 positive vertical hauls by an Indian Ocean Standard Net. The number of larvae in positive hauls ranged from 2 to 1262 larvae haul^-1. The most productive area of the ichthyoplankton was the waters south of Karachi, covered during the Machera Cruise, where 828 to 1262 larvae haul^-1 were obtained. The larvae were identified to species, genera or family. Sardinella sindensis (specific identification uncertain) larvae dominated in abundance. The other abundant larvae belonged to Benthosema spp., Amentum commersonii, Vinciguerria spp. and Diaphus spp. Larval distributions and abundances are described. Aggregation and spawning of adult Sardinella (sindensis) in the waters off the coast of W. Pakistan in November/December, 1964, are discussed. Presumably, the waters south of Karachi represent good feeding grounds for s. (sindensis) larvae. Displacement volume, length and weight relationships of various larval size groups have been studied and a high rate of larval mortality (at a length of 5.0 to 8.5 mm) observed.     

Effects of temperature on oxygen consumption,growth, and development of embryos and yolk-sac larvae of Siganus randalli (Pisces: Siganidae)

Eggs from laboratory spawnings of the coralreef fish Siganus randalli Woodland were incubated at two temperatures (27 and 30 °C). Eggs and larvae were sampled until larval starvation, while changes in oxygen consumption, growth, yolk utilization, and development were monitored. Oxygen consumption, which peaked at hatching, was higher for embryos incubated at 30 °C than at 27 °C. Rates of oxygen consumption (nl h^-1 individual^-1) at hatching were similar to those for other temperate and tropical species. Rates of oxygen consumption by yolk-sac larvae were highly variable, and these data suggest that larval oxygen consumption prior to yolk-sac absorption may not be significantly influenced by temperature. Rates of yolk depletion were higher for larvae at the higher temperature. After an initial rapid increase in length, length of larvae at 30 °C decreased with age. Egg size, egg weight, and maximum notochord length of larvae differed significantly between spawns. Age-specific oxygen consumption rates by the embryos varied between spawns, but regressions describing oxygen consumption as a function of age did not differ significantly. The initiation and completion of eye pigmentation were used as developmental markers to calculate the amount of yolk remaining for larvae at the different temperatures. Larvae maintained at 30 °C completed eye pigmentation approximately 3 h sooner than those maintained at 27 °C, but had less endogenous reserves. This finding indicates a trade-off between rapid development and efficient utilization of the endogenous reserves. The completion of eye pigmentation in larvae incubated at the higher temperature occurred at midnight and, depending on the amount of time that the larvae have to initiate feeding prior to the point-of-no-return, the timing of completion of eye pigmentation could influence larval survival.     

Prey selection by larval weakfish (Cynoscion regalis): the effects of prey size,speed, and abundance

We examined feeding by larval weakfish, Cynoscion regalis (Bloch and Schneider), in laboratory experiments conducted during the 1991 spawning season. under natural conditions weakfish larval development is ca. 3 wk, and we ran separate experiments with larvae of five different ages (5, 8, 11, 14, and 17 d post-hatching). We used two different size classes of rotifers (Brachionus plicatilis) and brine shrimp nauplii (Artemia sp.) as prey organisms. Contrary to results of previous research, weakfish larvae did not select prey based on size alone. When prey abundance was above 100 itemsl^-1 weakfish, larvae always chose large rotifers (length = 216 m) over small rotifers (length = 160 m). At 11 d post-hatching, larvae switched their diet from large rotifers to small brine shrimp nauplii (length = 449 m); however, when fed small rotifers and small brine shrimp nauplii the change in diet occurred at 14 d post-hatching. This pattern of selectivity was maintained in each larval age class. Early-stage larvae (5 and 8 d post-hatching) did not feed selectively when prey abundance was less than 100 itemsl^-1. Late-stage larvae (17 d post-hatching) fed selectively at abundances ranging from 10 to 10000 items^-1. Lwimming speeds of prey items, which ranged from 1 to 6 mms^-1, had no consistent effect on prey selection. These results suggest that weakfish larvae are able to feed selectively, that selectivity changes as larvae age, and that selectivity is also influenced by prey abundance.     

Feeding of laboratory-reared larvae of the sea bream Archosargus rhomboidalis (Sparidae)

Feeding by larvae of the sea bream Archosargus rhomboidalis (Linnaeus) was investigated from late September, 1972 to early May, 1973 using laboratory-reared larvae. Fertilized eggs were collected from plankton tows in Biscayne Bay, and the larvae were reared on zooplankton also collected in plankton nets. Techniques were developed to estimate feeding rate, food selection, gross growth efficiency, and daily ration. Daily estimates of these were obtained through 16 days after hatching at rearing temperatures of 23°, 26°, and 29°C. Feeding rate increased exponentially as the larvae grew, and increased as temperature was raised. At 23°C larvae began feeding on Day 3, at 26° and 29°C larvae began feeding on Day 2. Feeding rates at initiation of feeding and on Day 16 were, respectively: 23°C, 7.16 food organisms per larva per hour (flh) and 53.78 flh; 26°C, 7.90 flh and 168.80 flh; 29°C, 17.62 flh and 142.07 flh. Sea bream larvae selected food organisms by size. At initiation of feeding they selected organisms less than 100 m in width. As larvae grew they selected larger organisms and rejected smaller ones. The major food (more than85% of the organisms ingested) was copepod nauplii, copepodites, and copepod adults. Minor food items were barnacle nauplii, tintinnids, invertebrate eggs, and polychaete larvae. Mean values for gross growth efficiency of sea bream larvae ranged from 30.6% at 23°C to 23.9% at 29°C. Mean values for daily ration, expressed as a percentage of larval weight, ranged from 84% at 23°C to 151% at 29°C and tended to decline as the larvae grew.This paper is a contribution from the Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida, USA     

Reproductive isolation between intertidal and subtidal<Emphasis Type="Italic"> Scoloplos armiger</Emphasis> (Polychaeta,Orbiniidae) indicates sibling species in the North Sea

Two distinct modes of development in the common polychaete Scoloplos armiger (O. F. Müller, 1776) occur in the North Sea region: holobenthic development in egg cocoons and pelagic larvae hatching from suspended eggs. In the northern Wadden Sea near the island of Sylt, we observed that egg cocoons are produced intertidally while pelagic larvae originate from the adjacent subtidal zone. A previous genetic comparison between these subtidal and intertidal populations revealed distinct gene pools, suggesting that reproductive differences are not phenotypic but heritable. In this study, crossbreeding experiments show that intertidal and subtidal populations are reproductively isolated. Couples with males and females from different habitats had no offspring. Production of egg cocoons is determined by female origin from the intertidal zone. Pelagic larvae occurred only in couples with subtidal females and subtidal males. Intertidal males have spermatozoa with heads twice as long as those from subtidal males and a significantly shorter flagellum. We suspect that deviating sperm morphology may cause the reproductive breakdown at the fertilization stage. Juveniles hatching from cocoons have shorter anal cirri compared to juveniles that metamorphosed from pelagic larvae. We conclude there to be two sympatric sibling species in S. armiger: 'type I' in intertidal areas, which have egg cocoons, no pelagic larvae, elongated sperm heads, shortened sperm flagella and anal cirri; and a subtidal 'type S', lacking egg cocoons but with pelagic larvae, short sperm heads, long sperm flagella and anal cirri.Communicated by O. Kinne, Oldendorf/Luhe