Amphi-Atlantic phylogeography of direct-developing lineages of Lasaea, a genus of brooding bivalves

Direct-developing intertidal Lasaea spp. occur in the North Atlantic as both continental margin and oceanic island populations. We conducted a molecular phylogenetic analysis of representative populations in order to test colonization hypotheses for North Atlantic oceanic islands. Thirty individuals each were collected in 1995 and 1996 from two continental putative source populations (Florida, Iberia) and two oceanic island populations (Bermuda, Azores). They were sequenced for a 462 nucleotide portion of the mitochondrial large ribosomal subunit (16S) gene. No amphi-Atlantic genotypes were detected: Bermudan lineages co-clustered exclusively with Floridian congeners, and Azorean samples formed an exclusive clade with Iberian haplotypes. Our data indicate that geographical proximity to continental source populations is a better predictor of phylogenetic relationships in North Atlantic Lasaea spp. than present-day oceanic surface circulation patterns. The phylogenetic trees generated are not consistent with colonization of oceanic islands by indirect-developing ancestral lineages or by truly trans-oceanic rafting events. However, they are consistent with predicted topologies resulting from limited (≤ 2000 km), long-distance colonization by rafting (against present-day circulation patterns in the case of the Azores) and from anthropogenic introductions. Received: 17 December 1998 / Accepted: 7 June 1999     

Reproductive ecology of the brooding,hermaphroditic clam Lasaea subviridis

The reproductive ecology of a small intertidal clam was studied for two years (September 1980-November 1982) at two mid-intertidal sites on the northern shore of Monterey Bay, California. Lasaea subviridis is hermaphroditic and incubatory; the young are released as juveniles. Brooding adults are found throughout the year. All embryos in a clam are at the same stage of development, but the number and stage of embryos varies among clams at any one time. Individuals are iteroparous and produce small numbers of oocytes and sperms simultaneously. All stages of gonad development are found in different individuals at any one time. There was no evidence of seasonality. Young reared in the laboratory after being dissected from the parent reached brooding size (shell length of 1.7 mm) within 6 mo. Laboratory experiments indicated that juveniles crawl away soon after release from the parent. These reproductive characteristics may contribute to this taxon's ability to colonize new environments rapidly.     

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.     

Evolution of reproductive features and larval development in the genus Amphiglena (Polychaeta: Sabellidae)

Larval development in Amphiglena mediterranea is direct in the parental tube, and larvae crawl away as young juveniles with a radiolar crown capable of feeding. The non-feeding larvae of A. nathae are initially brooded in the parental tube, and then have a swimming phase before settling and developing a radiolar crown. Developmental data and reproductive features such as sperm structure are combined with other morphological data for an assessment of the cladistic relationships of the species in the genus Amphiglena. Two species of Laonome are included as part of the ingroup. Six genera, representing a sister group to the Laonome/Amphiglena clade, are used as an outgroup. Two most-parsimonious cladograms were found, and the evolution of reproductive features are discussed. The monophyly of Amphiglena is indicated by features previously identified, as well as two features based on sperm morphology. At present a plesiomorphic larval form cannot be identified for the genus Amphiglena. Received: 16 June 1997 / Accepted: 29 January 1998     

Characterization of a sex pheromone in a simultaneous hermaphroditic shrimp, Lysmata wurdemanni

Olfactory chemical cues have been described to play important roles in the control of mate recognition in many decapod crustaceans. However, we still know very little about the chemical characteristics of the cues that coordinate pre-copulative behaviour. In this study, we partially characterized a waterborne sex pheromone of a marine shrimp, Lysmata wurdemanni. Female moulting water was collected and ultrafiltered using 1,000 and 500-Da membranes, respectively, and analysed using HPLC with a Lichrosphere™ RP18 (C18) column. The sex pheromone is likely to be a molecule between 500 and 1,000 Da in size because behavioural bioassays showed that males responded to the supernatant of 500-Da and to the 1,000-Da filtrate, but did not respond to the supernatant of 1,000-Da or to the 500-Da filtrate. There was only one dominant peak (2.86 min) detectable in HPLC chromatograms of the supernatant of the 500-Da filtration. This peak showed a UV absorbance maximum at 274 nm, similar to the recently identified shore crab sex pheromone Uridine-di-phosphate (UDP). Behavioural bioassays confirmed that this peak is a bioactive component of a potential pheromone bouquet, but is different from UDP, which showed no bioactivity in Lysmata wurdemanni. Our results lay the foundation for future studies to purify and eventually identify this sex pheromone.     

The paradox of Rockall: why is a brooding gastropod (Littorina saxatilis) more widespread than one having a planktonic larval dispersal stage (L. littorea)?

In benthic invertebrates dispersal of planktotrophic larvae is generally considered more effective than is, for example, the rafting of adults or egg masses. It is certainly true that over short distances, viz., in the range of tens of kilometres or less, a moderately long-lived planktotrophic larva represents an effective mechanism of dispersal. However, turbulent mixing and mortality will decrease the concentration of planktotrophic larvae, and at some distance from the ancestral population the density of settlers may be too low to enable future matings between adults of low mobility. On the other hand, adults, juveniles or benthic egg masses drifted over long distances may colonize new habitats. The crucial point is the type of larval development of the organism. If the founder group belongs to a species with direct development or which produces very short-lived planktonic larvae, the low mobility of all life-stages will maintain a population within a restricted area so that mates will be likely to encounter each other even in a small population. Even if transport of benthic stages happens very rarely, this may be more influential than larval dispersal over long distances. To show that this may be true the detailed geographical distribution of two intertidal gastropod species with contrasting modes of development is presented and further support from the literature for this hypothesis is discussed.     

Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Ontogenetic development of digestive enzyme activities in larval walleye pollock,Theragra chalcogramma

Activities of digestive enzymes trypsin, amylase and lipase in laboratory-reared walleye pollock, Theragra chalcogramma, were measured from hatching to Day 39 (just before notochord flexion) in 1993. All measurements were conducted individually or semi-individually (groups of two larvae of the same standard length). Close relationships between digestive enzyme activities and morphological development of digestive organs were observed. Activities of trypsin and lipase were low during the transition period from endogenous to exogenous energy. Amylase activity was constant with large variance during the same period. Specific enzyme activities of trypsin and amylase indicated high values with large variance during the early period. All three enzyme activities increased with age afterthe transition period, and the specific enzyme activities became constant. The existence of two types of lipase was suggested. One lipase showed a peak of specific activity at Day 4 and might be related to yolk-sac absorption. The activity of the other lipase increased with age after Day 14 and might be related to digestion of prey lipid. Our results suggest that digestive enzymes included in food organisms supplement larval pollock digestive enzymes.     

Filtration rate,using a new indirect technique,in thirteen species of suspension-feeding bivalves

A method for determining filtration rates in undisturbed suspension-feeding bivalves is described. Concentrations of particulate matter in the water collected in the inhalant (C _i) and exhalant (C _e) currents were estimated with an electronic particle counter. The clearance was calculated as , where Fl=flow rate through the tube collecting exhalant water. Only above critical levels of water flow (Fl) were clearances representative of filtration rates. At 10° to 13°C, the filtration rates (F, 1 h^-1) within one or two orders of magnitude of dry weight (w, g), in Cardium echinatum L., C. edule L., Mytilus edulis L., Modiolus modiolus (L.) and Arctica islandica (L.) followed the allometric equations: 4.22w ^0.62, 11.60w ^0.70, 7.45w ^0.66, 6.00w ^0.75 and 5.55w ^0.62, respectively. Five species of bivalves [Spisula subtruncata (da Costa), Hiatella striata (Fleuriau), Cultellus pellucidus (Pennant), Mya arenaria L. and Venerupis pullastra (Montagu)] filtered with the same rates as individuals of Cardium echinatum and A. islandica of equivalent soft weight. In Pecten furtivus and P. opercularis filtration rates were about twice the rates measured in individuals of Mytilus edulis of comparable body weight. The gill area in M. edulis increases with size at the same rate as the filtration rate.     

Patterns of larval and early juvenile growth in a semiterrestrial crab, Armases angustipes (Decapoda: Sesarmidae): comparison with a congener with abbreviated development

In a semiterrestrial and estuarine tropical crab, Armases angustipes Dana (Grapsoidea: Sesarmidae), changes in biomass (measured as dry mass, W; carbon, C; nitrogen, N; and hydrogen, H; per individual) and relative elemental composition (C, N, H, in percent of W; C:N mass ratio) were studied during development from an early egg stage through hatching, the complete larval phase, metamorphosis and the first juvenile crab stage (CI). In the megalopa and CI, growth was measured also within the moulting cycle, and biomass and elemental composition were determined in cast exuviae. From an early egg stage to the freshly hatched larva, A. angustipes lost about 20% of W, 29% of C, 5% of N and 32% of H. Proportionally higher losses in C than in N were reflected also in a significantly decreasing C:N mass ratio (from 5.02 to 3.74). These results indicate that lipids mobilised from yolk reserves represented the principal metabolic substrate for embryonic energy production, while proteins were catabolised at a much lower rate. The present data of growth and exuviation are compared with previously published data from a congener, A. miersii Rathbun, which has an abbreviated and facultatively lecithotrophic mode of larval development (with three instead of four zoeal stages; stages I and II in principle independent of food). When growth is measured as an increase in the final (premoult) biomass of successive developmental stages, both species show an exponential pattern. Within the moulting cycles of the megalopa and the first juvenile, both species show parabola-shaped growth curves, with a rapid biomass increase in postmoult and intermoult stages, and losses in the premoult phase. Thus, the two Armases species show, in general, similar patterns of larval and early juvenile growth. However, the initial size of eggs and larvae is about four times larger in A. miersii, and its biomass remains higher throughout the period of larval and early juvenile development. A. angustipes is able to partially make up for this difference, as it has an additional zoeal stage, and its megalopa and CI stages show higher relative biomass increments (in percent of initial values). Due to this compensatory growth pattern, A. angustipes reaches in its CI stage about half the biomass of a juvenile A. miersii. When exuvial losses of megalopae and juveniles are compared between these two species, A. miersii shows higher biomass losses per individual (corresponding with its larger size), but lower relative losses (C, N, H, in percent of late premoult body mass or in percent of previously achieved growth increments). Differences in larval and early juvenile growth and in the exuvial losses of megalopae and juveniles of these two congeners are discussed in relation to their differential ecology, life history and reproductive strategy.     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type=Italic>Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.     

Evolution, radiation and chemotaxonomy of Lamellodysidea, a demosponge genus with anti-plasmodial metabolites

Sponges of the family Dysideidae (Dictyoceratida) are renowned for their diversity of secondary metabolites, and its genus Lamellodysidea, particularly Lamellodysidea herbacea, is the most studied taxon biochemically. Despite its importance, the taxonomic status of L. herbacea—whether it is a distinct species or a species complex—has never been assessed. Recent biochemical profiling revealed anti-plasmodial activity of brominated compounds in Lamellodysidea of the Pacific. Here, we present a comparative chemotaxonomic and molecular analysis of selected Dysideidae from the Pacific and the Indian Ocean (New Caledonia, Great Barrier Reef, Fiji, Mayotte, Guam, Palau). We investigated the phylogenetic relationships between the populations and assessed their bioactive (PBDE) compounds in order to unravel the taxonomic status of this commercially important group of sponges and assessed patterns of dispersal and biochemical variation. The molecular phylogeny was based on the internal transcribed ribosomal spacer and compared against a PBDE phylogeny for several specimens. Molecular data revealed a diversity of Indo-Pacific L. herbacea populations, also reflected by different PBDE compound profiles. Molecular and biochemical data also revealed a Lamellodysidea species new to science. Several specimens misidentified as Lamellodysidea were detected based on their position on different clades in the molecular phylogeny and their production of different halogenated compounds (brominated vs. chlorinated). The direct comparison of molecular and biochemical data also provided evidence for the occurrence of a host switch event and support for the theory that abiotic factors, such as sedimentation, affect the chemical constituents produced in L. herbacea.     

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     

Male mating preference is associated with risk of pre-copulatory cannibalism in a socially polymorphic spider

We performed male attraction experiments and staged courtship sequences to test for non-random mating with respect to social behavioral phenotype in the comb-footed spider, Anelosimus studiosus. While asocial behavior is the dominant phenotype in all A. studiosus populations examined to date, a social phenotype approaches a frequency of 15% in colder environments. We collected test subjects from higher latitude polymorphic populations and scored all individuals as to their behavioral phenotype prior to their use in these trials. Males of both phenotypes differentially approached and courted social females over asocial females and no-spider controls. By offering males different numbers of females of one type vs. the other in subsequent trials, we determined that the difference in attractiveness between the two phenotypes social/asocial is 1.5/1. Both the web produced by a female and a female that has been removed from its web attract males. We suggest that the male attracting pheromone is present on females and is also attached to silk threads. Staged encounters completed between males and females of the respective phenotypes demonstrated that courting males suffer significantly less pre-copulatory sexual cannibalism with social females than with asocial ones, and thus, female social tendency is phenotypically linked to sexual aggression. We propose that the male preference for social females is adaptive because of the observed asymmetry in courtship success.     

Environmental factors associated with the formation of larval anchovy fishing grounds in the coastal waters of southwest Taiwan

Daily commercial catches, school sizes and school densities of larval anchovy, and oceanographic and meteorological variables for the fishing seasons (January to June) from 1988 to 1990 were analyzed by means of principal component analysis to identify causative factors for the aggregation and abundance of larval anchovy, which make up the grounds in the coastal waters of southwest Taiwan. The results suggested that the primary factors associated with the formation of fishing grounds were surface water temperature that related to the occurrence of Engraulis japonicus, surface water salinity that was affected by influex of offshore water and river discharge, and phosphate content and solar radiation that affected plankton production.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Breeding success of a brood parasite is associated with social mating status of its host

Reproductive success of brood parasites varies considerably both among and within host species, mainly due to differences in host egg-rejection rates and survival of parasitic chicks. Here, we investigated the breeding success of the cuckoo (Cuculus canorus) in one of its major hosts, the great reed warbler (Acrocephalus arundinaceus), with respect to host social mating status. In this passerine, polygynous males provide less parental care to their young per nest than monogamous males. Consequently, their less-assisted females may fledge lower numbers of nestlings than monogamous females. This may be especially true for secondary females, which often receive limited or no paternal help with young at all. Based on these findings, we expected higher cuckoo reproductive success in nests of socially monogamous than polygynous great reed warbler males. More specifically, we predicted lower fledging success of cuckoo young in nests of secondary than primary or monogamous females. In line with the prediction, we found higher cuckoo fledging success in nests of monogamous than polygynous males, monogamous nests being more than twice as successful as secondary nests. We detected, however, only a tendency to lower cuckoo success in primary compared to monogamous nests and no differences between primary and secondary nests. Moreover, neither parasitism nor host egg-rejection rates differed among the nests of different status. Our results show, for the first time, that the social mating status of a host may influence the overall reproductive success of a brood parasite and thus should be considered in further studies.     

Correlations between weather and recruitment of Nassarius pauperatus,an intertidal gastropod with a planktotrophic larval stage

Recruitment of the intertidal gastropod Nassarius pauperatus (Lamarck) was estimated annually for more than a decade at three sites in South Australia. Recruitment varied temporally, but was relatively synchronous amongst sites, even though these were at least 10 km apart. The density of recruits at each site was negatively correlated with evaporation and also with several indices of temperature for February. We postulate that either high temperatures or desiccation cause the death of recently settled individuals, which may be unable to avoid these extremes by burrowing deeply into the substratum.     

Hydrography and distribution dynamics of larval and juvenile fishes in the coastal waters of the Tanshui River estuary,Taiwan, with reference to estuarine larval transport

Distribution dynamics of fish larvae and juveniles in the coastal waters of the Tanshui River, Taiwan was studied fortnightly using surface horizontal tows with a larval net in daytime during the period from early April through early June 1991. Environmental factors, including water temperature, salinity, dissolved oxygen, pH, transparency and depth at sampling stations, were also monitored. A total of 10737 fish eggs and 1387 individuals, representing 43 families and 93 species, was collected during five cruises from 12 stations in the coastal waters. Most fish were estuarine-dependent marine species. Liza macrolepis, Ambassis gymnocephalus, Terapon jarbua, Mullidae and Gobiidae were the most dominant, making up 64.7% of the total catch. Early life stages, including egg, preflexion, flexion and postflexion larvae were abundant in surface samples. However, yolk-sac larvae were absent in the surface water, probably due to an ontogenetic behavioral shift as a consequence of a change in specific weight during early development. The species composition of fish larvae and juveniles was related to the microhabitats found in the coastal waters. The physico-chemical conditions, along with ontogenetic behavior, played an important role in larval fish distribution in the coastal waters.