Reproductive strategies and life histories in the cheilostome marine bryozoans Chartella papyracea and Bugula flabellata

Seasonal life history phenomena were monitored through 1978 for two sublittoral cheilostome bryozoans from souther Britain: Chartella papyracea (Ellis and Solander), a non-placental brooder, and Bugula flabellata (Thompson in Gray), which is placental. The intracolonial relationships between growth by zooid budding, polypide recycling, and sexual reproduction, were analysed for both species — quantitatively in the case of C. papyracea. Although both species grow only in summer, their life histories differ. C. papyracea is more of a K-strategist, producing perennial fronds that release larvae in several successive years, most being liberated in winter months. The reduced need for maximization of the larval productivity rate is reflected in the intracolonial configuration of polypide and sexual recycling. B. flabellata is more of an r-strategist. The larvae are produced by the ephemeral fronds of two successive colony generations each summer. Colonies are dormant in winter. The need to maximize the larval productivity rate in this species is reflected in the polypide/sexual recycling configuration. It is believed that embryonic placentation contributes to the maximization of the larval productivity rate in this and other placental brooders.     

Competitive interactions between related clades: evolutionary implications of overgrowth interactions between encrustin cyclostome and cheilostome bryozoans

Overgrowth interactions between encrusting cyclostome (eight species) and cheilostome (over 50 species) colonies were examined in subtidal communities in the northern Adriatic Sea off Rovinj, Croatia, in May–June 1988 and September–October 1990. Cheilostome colonies occupied ca. 80% of the available substrate space, whereas the cyclostome colonies occupied <5%. Out of 210 recorded interactions, cheilostomes overlapped cyclostomes in 164, while cyclostomes overlapped cheilostomes in only 16; 30 encounters resulted in mutual overlap or at least temporary growth termination along the line of contact. Most of the recorded interactions were for the cyclostomes Diplosolen obelia (Johnston) and Plagioecia patina (Lamarck). Both species elevated their colony margins on contact with a cheilostome in some instances, but D. obelia only overgrew competing cheilostomes in 12/120 encounters, and P. patina never prevailed. I propose that the cheilostomes have a key evolutionary innovation, lacking in cyclostomes, consisting of more rapid ontogenetic development of zooids along colony margins so that encrusting cheilostomes have substantially higher colony margins. This results in an outflow of filtered water along the edge of encrusting cheilostome colonies, while cyclostomes take water in at colony margins. These contrasting feeding currents apparently give a competitive advantage to the cheilostomes where colony margins approach and make contact with those of cyclostomes.     

Population genetic consequences of larval dispersal mode and hydrography: a case study with bryozoans

Genetic analysis of the marine bryozoans Celleporella hyalina and Electra pilosa using the RAPD technique revealed population structuring corresponding to the contrasting modes of larval dispersal. Samples of C. hyalina exhibited genetic differentiation over distances as small as 10 m, concordant with the limited dispersal predicted by a simulation model, based on the short pelagic phase of the lecithotrophic larvae and the local hydrography. In contrast, E. pilosa showed high levels of genetic heterogeneity only over much larger spatial scales, commensurate with its production of comparatively long-lived planktotrophic larvae. The population differentiation observed between samples of E. pilosa, collected from sites 70 km apart, is reconcilable with coastal water currents and frontal systems that restrict the exchange of water masses between the two sites. Hydrographic conditions and discontinuous distribution of suitable substrata therefore are seen to constrain gene flow, creating opportunities for local genetic differentiation despite the high dispersal potential of pelagic larvae. Received: 9 August 2000 / Accepted: 18 November 2000     

Effects of contrasting modes of larval development on the genetic structures of populations of three species of prosobranch gastropods

 In south-eastern Australia, the prosobranch gastropods Morula marginalba (Blainville), Cominella lineolata (Lamarck) and Bedeva hanleyi (Angas) have similar fine-scale distributions, but appear to possess very different dispersal capabilities due to contrasting modes of larval development. M.marginalba produce planktonic larvae, whereas C. lineolata and B. hanleyi undergo direct development in benthic egg capsules and emerge as crawling juveniles. To test for possible effects of contrasting life histories on levels of genetic variation within and among populations, a survey was conducted of allozyme variation at six polymorphic loci in 8 to 9 local populations of each species. Collections of snails were made between June 1992 and November 1993. Sampling ranges spanned between 162 and 180 km of coast. Regardless of larval type, proportions of single-locus genotypes in each collection were consistent with the recruitment of offspring which had been generated through random mating. However, genotypic diversity was lower in those species that undergo direct development. Loci surveyed in C. lineolata and B. hanleyi were polymorphic (i.e. frequency of most common allele <95%) in fewer populations than those examined for M.␣marginalba (P <0.001) and, where polymorphisms occurred, also possessed significantly fewer alleles (P <0.001). Consequently, average levels of expected heterozygosity were greater in populations of M. marginalba than in those of either of the other species (P <0.001). Genetic variation among populations, expressed as the standardised variance in allele frequencies (F _ST ), was inversely related to expected larval dispersal capability. The nine collections of M. marginalba showed little overall differentiation (F _ST  = 0.017; P <0.001), reflecting the ability of planktonic larvae to interconnect local populations, and so limit divergence due to drift and natural selection. In contrast, there were high levels of allelic heterogeneity among the nine collections of C. lineolata (F _ST  = 0.523; P <0.001) and eight collections of B. hanleyi (F _ST  = 0.140; P <0.001). These data imply that for species which undergo direct development, local populations are effectively closed and evolve largely independent of one another. Received: 3 May 1996 / Accepted: 12 July 1996     

降尘对棉花叶片解剖结构的影响

根据模拟降尘的盆栽试验,通过显微技术,研究了受自然降尘影响和未受影响的棉花（Gossypium hirsutumL.）叶片解剖结构的特点。结果表明：受降尘影响的棉花叶片栅栏组织排列不规则,部分栅栏组织细胞收缩、变短,甚至弯曲。栅栏组织细胞之间空隙增大,海绵组织细胞中内含物减少;但在出现伤害症状的叶片中,维管束仍保持原状。用石英砂替代降尘处理的和未受降尘影响的棉花叶片的组织结构保持正常。     

Population structure and energetics of the shallow-water antarctic sea star Odontaster validus in contrasting habitats

Individuals and populations of Odontaster validus Koehler differed markedly among different habitats, as revealed in a study from October 1984 through January 1986 in McMurdo Sound, Antarctica. At McMurdo Station, individual sizes (wet weight) and population biomass (g wet wt m^-2 and kJ m^-2) decreased significantly with increasing depth. Individuals from shallow (10 to 20 m) habitats were in superior nutritional condition to those from deeper water (30 and 165 m), as shown by higher gonad and pyloric cecum indexes, and by higher lipid and energetic levels in the pyloric ceca. Moreover, gonadal output (reproductive output) was higher in shallow-water individuals. Higher levels of chlorophyll in the pyloric ceca and richer yellow to red coloration of the body wall in the shallow-water individuals indicate that they utilize the higher levels of primary production at shallow depths. At East Cape Armitage, where nearly permanent, thick, snow-covered ice most of the year resulted in very low levels of benthic primary production, the lowdensity sea stars were all very small and nutritionally similar to the deep-water individuals at McMurdo Station. At Cape Evans, where the generally snow-free sea-ice that broke up in mid-summer resulted in a luxurient benthic cover of diatoms and macroalgae, the sea stars were smaller than at McMurdo Station at comparable depths, but population densities were higher, resulting in 4 to 9 times greater biomass. Growth rates of sea stars fed in the laboratory were very low, especially compared to laboratory-reared temperate and tropical species; well-fed individuals need about 9 yr to reach 30 g wet weight, near the mean size of shallowwater individuals at McMurdo Station. No growth was detected in individuals caged at McMurdo Station for one year, suggesting even lower growth rates in the field. The stable size-frequency distributions at the different sites and depths throughout the year-long study suggest highly stable populations with low temporal variability in recruitment, migration and mortality. These data indicate that individuals and populations of O. validus quantitatively and qualitatively reflect the general level of productivity of a habitat. Differences noted in size, coloration, nutrition, and reproductive effort may be the result of long-term integration of local levels of primary production. These ubiquitous sea stars may serve as a biotic indicator of productivity in localized habitats around the continental shelf of Antarctica.     

Distribution of encrusting bryozoans and other epifauna on the subtidal bivalveChlamys opercularis

The shells of living queen scallops,Chlamys opercularis (L.), are often completely colonised with epifauna. In this study, the distribution of the major epifaunal groups across the surfaces of left (upper) and right (lower) valves is examined using samples from two commercial queen scallop beds off the Isle of Man. The composition of the epifauna differs across and between the valves and also varies with the age of the scallop. YoungerC. opercularis are commonly encrusted by sponges, but on older shells other epifauna colonise the surfaces. The encrusting bryozoan assemblage found on livingC. opercularis is not as abundant or speciose as on some other substrata such as stones and dead shell. This may be a result of intense competition from other phyla. The distribution of different bryozoan species between the surfaces is examined. Most species are more abundant on the lower valves.     

多效唑对水仙观赏效果的影响

水仙对多效唑比较敏感,无论浸泡法或叶面喷施法都具有明显的矮化效应,但当浓度≥200mg·l-1时又会产生药害,也有碍观赏;叶面喷施法对水仙矮化效应不如浸泡法,开花期亦迟10·15d;浸泡法的最佳处理浓度是50 mg·l-1;叶面喷施法以浓100-200mg·l-1效应最好,但此法费工费时,操作不便,生产上最好不采用。     

Influence of food concentration,temperature and salinity on the larval development ofBalanus amphitrite

Influence of food concentration (0.5, 1 and 2 x 10⁵ cell ml^–1 ofSkeletonema costatum), temperature (20 and 30°C) and salinity (15, 25 and 35) on the larval development ofBalanus amphitrite (Cirripedia: Thoracica) was examined. The mortality rate at 20°C was lower than at 30°C in general. Increase in food concentration from 0.5 to 1 x 105 cells ml^–1 improved the survival rate, but this was not evident when food concentration was increased to 2 x 105 cells ml^–1. The results indicate that food availability and temperature jointly determine the energy allocation for metamorphic progress. It was observed that the influence of the tested variables varied with instar. At 20 °C the mean duration of the second instar exceeded 3 d and was much longer than other instar durations. The fourth, fifth and sixth instars and the total naupliar period showed that the effect of different salinities at given food concentrations was negligible at 20°C, while at 30°C there was a marked decrease in duration with increasing salinity.     

Life-history variation in three coexisting cheilostome bryozoan species of the genus Stylopoma in Panamá

Life-history variation was studied in three sympatric species of Stylopoma Levinsen [S. spongites (Pallas), S. projecta Canu and Bassler and Stylopoma n. sp. 15] in Panamá. Bryozoan colonies were collected from 27 reefs along 300 km of the Caribbean coast of Panamá. The distribution and abundance of each species were very patchy, but with broad overlap in occurrence among localities and depths. Nevertheless, species differed considerably in colony size, size at first reproduction and numbers of brooding larvae; implying that interspecific differences in life-history traits may contribute to their coexistence. To examine closely this variation in life-history patterns, we grew, in a common garden experiment, the offspring of the two most common species that were obtained from parent colonies collected from several reefs. There were highly significant differences in growth rates and the timing and extent of sexual reproduction which corresponded well to patterns observed in individuals from the field. Other factors, including size of larvae, extent of secondary zooidal calcification and numbers of avicularia were also correlated with differences in life histories. Despite this additional complexity, however, ecological consequences of trade-offs in life history among modular species such as Stylopoma spp. appear very similar to those among unitary species.     

土地利用方式对土壤动物多样性的影响

为了解不同利用方式下土壤动物多样性状况及其演变,对广州市东部郊区的水稻田、蔬菜地、果园旱地和林地4种土地利用类型0～5、5～10、10～15、15～20cm土层进行了土壤动物取样调查,共获得土壤动物24683只,分别隶属于4门10纲23类。统计分析结果表明,土壤动物多样性受土地利用方式的影响明显。果园旱地和林地的个体数显著高于蔬菜地和水稻田,但果园旱地与林地、蔬菜地与水稻田之间无显著差异;果园旱地和林地的土壤动物类群数显著高于水稻田,但果园旱地与林地、林地与蔬菜地、蔬菜地与水稻田之间的差异不显著;林地和果园旱地的复杂性指数显著高于蔬菜地,但林地与果园旱地和水稻田、水稻田与蔬菜地之间的差异不显著（P〈O．05）。土壤动物的个体数和类群数量随着土壤深度的增加而明显减少,但不同土地利用方式下各层土壤动物的丰富度及其随土层加深而递减的程度则有明显不同,其中,林地和果园旱地的土壤动物在不同土层中较丰富,垂直变化比较和缓,水稻田和蔬菜地的类群数和个体数则随土层的加深而急剧减少。     

Vertical structure of very nearshore larval fish assemblages in a temperate rocky coast

Small-scale vertical patterns of larval distribution were studied at a very nearshore larval fish assemblage, during the spring–summer period of several years, at two depth strata (surface and bottom) using sub-surface and bottom trawls. A total of 4,589 larvae (2,016 from surface samples and 2,573 from bottom samples) belonging to 62 taxa included in 22 families were collected. Most larvae belonged to coastal species. Although inter-annual variations in larval density and diversity could be found, total larval abundance was always higher near the bottom whereas diversity was higher at the surface. A marked distinction between the structure of surface and bottom assemblages was found. Sixteen taxa explained 95% of the similarity among surface samples. Larvae which contributed most to this similarity included species like clupeiformes, sparids and serranids, and also blenniids, tripterygiids and some labrids. In the bottom samples, fewer species were present, with only six taxa, almost exclusively from species which lay demersal eggs, contributing to 95% of the similarity between samples. Larvae present at the surface were significantly smaller than at the bottom. For some of the most abundant species found at the bottom, only small larvae occurred at the surface while the whole range of sizes was present at the bottom, indicating that larvae may be completing the entire pelagic phase near the adults’ habitat. These results indicate that larval retention near the reefs probably occurs for these species, although for others dispersal seems to be the prevailing mechanism.     

The influence of per offspring investment (POI) and starvation on larval developmental plasticity within the palaemonid shrimp,Palaemonetes varians

At the inter-specific level, per offspring investment (POI), degree of abbreviated development, and lecithotrophic potential all increase with increasing latitude and freshwater penetration among crustaceans. These traits are considered adaptations to conditions of decreasing growth potential. We hypothesise that this relationship between POI and abbreviated development also occurs at the intra-specific level. We studied the caridean shrimp, Palaemonetes varians, to investigate the hypothesis that under food-limited conditions, higher POI enables development through fewer larval instars. Under starvation stress, larvae from broods of greater POI (measured as hatchling brood average dry weight, DW) generally developed through fewer larval instars. With increasing starvation period, larval development time increased, whilst larval growth rate, juvenile DW, juvenile carbon mass, and juvenile carbon:nitrogen (C:N) ratio all decreased. Larval development time generally decreased with increasing brood average dry weight. In contrast, larval growth rate, juvenile DW, juvenile carbon mass, and juvenile C:N ratio all increased with increasing larval brood average DW. The relationship between POI and larval instar number (abbreviation of development) reported here is consistent with that at the inter-specific level and supports the concept that macro-ecological trends in development modes at the inter-specific level may be driven by selection occurring on POI at the intra-specific level.     

Influence of three sediment types on copper toxicity to the polychaete Neanthes arenaceodentata

Adult male Neanthes arenaceodentata were exposed to 0.10±0.015 mg l^-1 copper in the seawater of a continuous-flow bioassay system in the presence of a sand, a mud, a mixture of the sand and mud, and no sediment, to assess the influence of sediment type on Cu-induced mortality. The sediment type did influence mortality. The time to 50% mortality was 7.8 days without sediment, 36.5 days with sand, 54.5 days with the mixture, and 50.0 days with mud. There was no threshold Cu body burden that caused death. The mean Cu concentration per gram of N. arenaceodentata (24 h after death) without sediment was 270 g, in sand 994 g, in the mixture 1047 g, and in mud 1464 g. The differences in the toxic responses are discussed.     

Genetic structure of populations of two closely related brittle stars with contrasting sexual and asexual life histories,with observations on the genetic structure of a second asexual species

Ophiocoma pumila Lütken andOphiocomella ophiactoides (H. L. Clark) are morphologically similar brittle stars with contrasting life histories, the former obligately sexual, the latter fissiparous (capable of both sexual reproduction and asexual proliferation by binary fission). Electrophoretic analysis of five polymorphic enzymes was used to assess the genetic consequences of these differing life histories and provide a genetic perspective on the taxonomic relationship between the two species. Genotypic diversity ofOphiocoma pumila collected at Discovery Bay, Jamaica, in 1985 conformed to expectations for a sexually reproducing population. In contrast, genotypic diversity ofOphiocomella ophiactoides at this site was significantly lower than expected for a sexually reproducing population, due largely to the predominance of clonal proliferation over larval recruitment. Large variation in clonal composition over a short (50 m) distance emphasized the very localized scale of clonal mixing in this species. Allozymic data are indicative of a close sibling species relationship betweenOphiocoma pumila andOphiocomella ophiactoides which suggests that the present generic separation of the two species should be re-examined. Electrophoretic analysis was also used to examine the genetic structure of sponge- and alga-dwelling populations of a second fissiparous brittle star,Ophiactis savignyi (Müller & Troschel), which was also collected at Jamaica in 1985. Striking differences in the allelic composition of sponge- and alga-dwellingO. savignyi were observed. Genotypic diversity ofO. savignyi in sponges was very low, each sponge being dominated by a single genotype. Genotypic diversity ofO. savignyi in algae was higher, although still significantly lower than expectations for a sexually reproducing population. In the light of the highly clonal composition of fissiparous brittle-star populations, the adaptive significance of clonal growth may be related to an increase in the overall fitness of dispersed clones (genets), compared to individuals of strictly sexual counterparts, through greater genotypespecific biomass and, hence, fecundity.Contribution No. 476 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Mode of larval development and genetic population structure inNodilittorina africana knysnaensis (Prosobranchia: Littorinidae)

We examined electrophoretic variation in the supralittoral gastropod,Nodilittorina africana knysnaensis, along the coast of South Africa in 1989. There was little allozyme variation in nine samples collected over 1 500 km for three highly polymorphic loci,Est, Gpi, andPgm, and Wahlund's effect was absent in the pooled sample. A gene diversity analysis showed that 98.1% of total genic diversity, on average, was contained within populations, and that only 1.9% was due to allozyme frequency differences among populations. Since the mode of larval development has not yet been described for this species, we suggest that the lack of strong allozyme frequency differences over long distances indicates thatN. a. knysnaensis has a planktonic mode of larval development.     

Spatial autocorrelation analysis of small-scale genetic structure in a clonal soft coral with limited larval dispersal

The philopatric larval dispesal and small effective population sizes characteristic of many clonal species should promote the development of significant small-scale genetic structure within populations as a result of isolation-by-distance. We used spatial autocorrelation statistics to detect genetic structure, arising from both clonal reproduction and philopatric dispersal of sexual propagules, for five allozyme loci within populations of the soft coral Alcyonium sp. In a population on Tatoosh Island, Washington, USA, sampled in 1991/1992, we found significant positive spatial autocorrelation at all loci among individuals separated by <40 cm, reflecting the presence of significant smallscale genetic structure due to associations among clonemates. For 4 of 5 loci, however, we detected no significant spatial autocorrelation among the different clones within this population over distances of 1 to 40 m. Analysis of soft-coral populations from six additional, topographically diverse sites in the north-east Pacific also did not reveal significant spatial autocorrelation among clones at any loci. This general lack of spatial autocorrelation of genotypes among clones suggests that significant small-scale genetic structure has not arisen in populations of Alcyonium sp. as a consequence of isolation-by-distance.     

Influence of variability in larval development on recruitment success in the euphausiid Euphausia pacifica : elasticity and sensitivity analyses

Simple life-history models were employed to identify those larval stages of Euphausia pacifica (Hansen) with the greatest potential contribution to variability in recruitment success. Using stage-specific vital parameters (i.e. stage duration and specific mortality) the models explored the influence of developmental variability without ascribing the variability to any particular cause or forcing. Through such a non-mechanistic modelling approach, the critical periods in larval development were determined. The results led to inferences about larval population responses to temperature/food heterogeneity in the natural environment. Peaks in recruitment-success elasticity were evident for variability imposed during the egg and Furcilia I and II stages. The high elasticity of the egg stage suggested the importance of female nutrition and the timing and location of egg release in determining their viability and subsequent recruitment success. Peak elasticity during the Furcilia I and II stages supported the notion that indirect developmental pathways of pleopod development during these stages may be associated with diminished recruitment success. Sensitivity analyses further highlighted the influence of these early furcilia stages in determining recruitment success, and also suggested the importance of the Furcilia III and VII stages. The robustness of elasticity and sensitivity results with respect to the life-history structure utilized is discussed. A criterion is presented for the identification of life-history structures which might obscure the true elasticity relationships of a developmental sequence. Received: 27 April 1998 / Accepted: 14 September 1998     

Influence of irradiance on the nutritional value of two phytoplankton species fed to larval Japanese scallops (Patinopecten yessoensis)

Japanese scallop (Patinopectin yessoensis Jay) larvae grew faster and were larger after 18 d when fed a diet of high-light(HL)-grown Chaetoceros simplex or HL Pavlova lutheri relative to diets of the same phytoplankton species grown at low light (LL). When provided as saturating rations to larval scallop, these diets could be ranked: HL C. simplex>LL C. simplex>HL P. lutheri>LL P. lutheri. In both phytoplankton species, HL-grown cells contained more of the short-chain saturated fatty acid (FA), 16:0 than LL-grown cells. Scallop growth rates were a significant function of the amounts (mg g^-1 dry wt) and the proportions (as percentage of total FAs) of the FAs 14:0 and 14:0+16:0 (total saturated FAs) in their diet. The proximate biochemical composition of HL- versus LL-grown phytoplankton showed no significant differences in protein, total lipid, carbon, carbohydrate or nitrogen per cell which were consistently associated with the greater nutritional value of HL cells. In spite of this high variability in proximate composition, the larval growth rate was a significant function of the average carbon content, nitrogen content and cell volume of the phytoplankton cells. Increased amounts of the essential polyunsaturated FAs 20:5 3 and 22:6 3 in the phytoplankton were negatively correlated with larval scallop growth rates. Thus HL-grown phytoplankton cells were nutritionally superior to LL-grown cells. This nutritional superiority seems to be determined by the fatty acid composition of the cells which, in turn, is controlled by variation in irradiance. The general tendency of predator FA profiles to resemble that of their prey was not observed in larvae fed P. lutheri. The much greater amounts of 18:4 3, 20:5 3, and 22:6 3 FA in P. lutheri relative to C. simplex were not evident in the scallop larvae fed these cells.