Is Syllis gracilis (Polychaeta: Syllidae) a species complex? An allozyme perspective

The genetic relationships between morphologically indistinguishable marine and brackish populations of Syllis gracilis Grube, 1840 (Polychaeta: Syllidae) were studied by means of allozyme electrophoresis. Samples of S. gracilis from marine coastal and brackish-water habitats were examined for variation at 13 presumptive loci. In addition, a sample of the closely related species S. prolifera (Krohn, 1852) was analysed. Five loci were multiallelic in at least one population of S. gracilis and eight loci in S. prolifera. Low to moderate levels of within-population genetic variability were found, with average expected heterozygosity values ranging from H = 0.068 (±0.043 SE) to 0.187 (±0.069 SE) in the populations of S. gracilis; higher values were found in S. prolifera (H = 0.325 ± 0.076). The presence of various private alleles indicated a marked genetic divergence among populations of S. gracilis, with Nei's genetic distances ranging from D = 0.000 to 0.833 and a highly significant F _ST value. Furthermore, evidence for strong genetic heterogeneity between two sympatric marine populations was found. UPGMA cluster analysis and multidimensional scaling pointed out a clear genetic divergence between brackish and marine populations. At least two genetically divergent entities occurred in marine and brackish habitats. This could be due to local adaptation of individuals coming from marine populations to brackish habitats, but more presumably to the occurrence of a species complex within S. gracilis. Received: 6 June 1999 / Accepted: 7 February 2000     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Mitochondrial DNA sequence variation and phylogeography of oceanic insects (Hemiptera: Gerridae: Halobates spp.)

Relatively few insects have invaded the marine environment, and only five species of sea skaters, Halobates Eschscholtz (Hemiptera: Gerridae), have successfully colonized the surface of the open ocean. All five species occur in the Pacific Ocean, H. germanus White also occurs in the Indian Ocean, whereas H. micans Esch- scholtz is the only species found in the Atlantic Ocean. We sequenced a 780 bp long region of the mitochondrial cytochrome oxidase subunit I gene (COI) for a total of 66 specimens of the five oceanic Halobates species. Our purpose was to investigate the genetic variation within species and estimate the amount of gene flow between populations. We defined 27 haplotypes for H. micans and found that haplotype lineages from each of the major oceans occupied by this species are significantly different, having sequences containing five to seven unique base substitutions. We conclude that gene flow between populations of H. micans inhabiting the Atlantic, Pacific, and Indian Ocean is limited and hypothesize that these populations have been separated for 1 to 3 million years. Similarly, there may be limited gene flow between H. germanus populations found in the Pacific and Indian Ocean and between H. sericeus populations inhabiting the northern and southern parts of the Pacific Ocean. Finally, we discuss our findings in relation to recent hypotheses about the influence of oceanic diffusion on the distribution and population structure of oceanic Halobates spp. Received: 29 July 1999 / Accepted: 23 November 1999     

Escape responses of four Chilean intertidal limpets to seastars

The Chilean intertidal Lottiidae limpet fauna consists of at least nine species, grouped into the monophyletic taxon Scurria and a single taxon preliminarily designated as `Lottia' orbignyi (Dall). Within the Scurria clade, the most basal species are Scurria variabilis (Sowerby), S. viridula (Lamarck), and S.zebrina (Lesson), and among the most crown groups are S. ceciliana (Orbigny) and S. araucana (Orbigny). This phylogenetic information, obtained by the analysis of molecular characters (16S mtDNA), provides a framework for a comparative study of behavioral characters. In this study, we analyze the escape responses of two basal limpet species (`L.' orbignyi and S. viridula) and two derived limpet species (S. araucana and S. ceciliana) in the presence of the limpet predatory seastar Heliaster helianthus (Lamarck) and the limpet non-predatory seastars Stichaster striatus Muller and Troschel and Patiria chilensis Lutken. Neither P.chilensis nor Stichaster striatus induced escape responses such as those observed with the predatory H. helianthus. Moreover, in the presence of H. helianthus, basal and derived limpet species differed significantly in the percentage of individuals responding, reaction time, and duration of the response. The basal species `L.' orbignyi and Scurriaviridula exhibited instantaneous and vigorous locomotor responses, whereas the derived species S. araucana and S. ceciliana displayed no locomotor responses. The same pattern was found in responses to H. helianthus homogenates. The results strongly suggest a correlation between phylogeny and anti-predator escape responses to the seastar H. helianthus. Alternative escape responses have evolved for basal and derived species within the Chilean clade. Received: 27 April 2000 / Accepted: 16 August 2000     

Contrasting life histories and secondary production of populations of Munnopsurus atlanticus (Isopoda: Asellota) from two bathyal areas of the NE Atlantic and the NW Mediterranean

The isopod Munnopsurus atlanticus occupies bathyal depths in both the Bay of Biscay (NE Atlantic; between 383 and 1022 m) and in the Catalan Sea (Northwestern Mediterranean; between 389 and 1859 m). The species was dominant in both assemblages, reaching bathymetric peaks of abundance on the upper part of the continental slope (400 m depth) in the Bay of Biscay and at ˜600 m in the Catalan Sea. Both the Atlantic and the Mediterranean populations are bivoltines. Demographic analysis of the Bay of Biscay population revealed the production of two generations per year with different potential longevity (5 mo for G₁ and 11 mo for G₂). The mean cohort-production interval (CPI) was estimated at 8 mo, and results of the demographic analysis were also used to estimate production for the Catalan Sea populations. Mean annual density (D) and biomass (B) were higher in the Bay of Biscay (D = 356.7 individuals 100 m⁻²; B = 0.803 mg dry wt m⁻² yr⁻¹) than in the Mediterranean (D = 16.3 individuals 100 m⁻²; B = 0.078 mg dry wt m⁻² yr⁻¹). Also, mean annual production was an order of magnitude higher in the Atlantic (between 4.063 and 4.812 mg dry wt 100 m⁻² yr⁻¹ depending on the method used) than in the Catalan Sea (between 0.346 and 0.519 mg dry wt 100 m⁻² yr⁻¹). M. atlanticus feeds on a wide variety of benthic and pelagic food sources. In both study areas, phytodetritus was not important in the diet of M. atlanticus. In contrast, gut-content data suggested an indirect coupling with phytoplankton production in both areas via foraminiferans. The life history and the recorded production are considered in respect to both the dynamics and levels of primary production and the total mass flux in the respective study areas. Differences in the secondary production of both populations seemed to be more consistently explained by differences in total mass flux than by differences in the primary production levels; this is also consistent with the variety of food sources exploited by M. atlanticus. Received: 22 February 1999 / Accepted: 3 February 2000     

Meiofauna response to a dynamic river plume front

 The benthic response to a plume front was studied in two areas of the northern Adriatic (Mediterranean Sea) differently influenced by the Po River freshwater input. Sediment samples were collected in June 1996 and February 1997 from 12 stations. The adopted sampling strategy was able to identify the front line in real time by satellite images and to locate sampling stations along an inner–outer plume gradient in order to cover the benthic area beneath the river plume, where enhanced biological production was expected, and open-sea sediments not directly influenced by freshwater inputs. Meiofaunal parameters were compared to the physical conditions and to phytodetritus inputs, organic matter accumulation and bacterial secondary production. The sediments of the Adriatic Sea were characterised by high concentrations of phytopigments (0.6 to 13.9 μg g⁻¹ for chlorophyll a and 1.2 to 17.7 μg g⁻¹ for phaeopigments) and biopolymeric organic carbon (0.15 to 3.02 mg g⁻¹). The plume system extended for a large sector of the northern Adriatic. In the northern area, a large and highly dynamic plume area was coupled with a sediment organic matter concentration significantly higher than in open-sea sediments. In the southern sector, where the plume area and the front line did not change markedly during the year, plume–benthic coupling was evident only in the sediments beneath the front, and corresponded to phaeopigment accumulation. Bacterial parameters and secondary production were high and significantly higher in the frontal area than at open-sea stations. Meiofauna density (1342 to 8541 ind. 10 cm⁻²) did not change either by season or between areas and was significantly correlated with phaeopigments and bacterial secondary production. Meiofauna displayed different responses to plume inputs in the two sampling areas. In the northern sector, meiofauna density was coupled with organic matter distribution and displayed highest values beneath the plume. In the southern sector, the densities of copepods, turbellarians and kinorhynchs displayed highest values under the front in summer, and the same applied to total meiofauna density in winter. Juvenile decapods and copepod nauplii significantly increased their densities in sediments beneath the front. Data presented in the present study suggest that plume inputs and frontal systems, enhancing phytodetritus accumulation and benthic bacterial response, might influence density, composition and distribution of meiofaunal assemblages. As river plumes are highly variable systems affecting the trophic characteristics of the sediments underneath, their dynamics should be considered when analysing mesoscale spatial changes of meiofaunal assemblages. Received: 30 November 1999 / Accepted: 24 May 2000     

Predation on meiofaunal and macrofaunal invertebrates by western sandpipers (Calidris mauri): evidence for dual foraging modes

Western sandpiper (Calidris mauri) predation was examined by concurrent experiments and direct observations of foraging behaviour on high intertidal mudflats of the Fraser River estuary, British Columbia. Western sandpipers foraged by either “pecking” on the surface (64% of observational time) or probing into sediment (29%). The first experiment (probe-mark method) consisted of collecting small-volume cores (21.2 cm³) of probed (experimental) and non-probed (control) sediment on the tidal flat, following a 22.5-min feeding period. The second experiment (exclosure method) involved deploying exclosures immediately prior to the feeding period and subsequent collection of cores from inside (control) and outside (experimental) the exclosures. Sediment cores were analysed for both macrofaunal and meiofaunal size fractions. Comparisons between macro- and meiofaunal invertebrate densities in experimental and control sediments revealed significant differences, attributed to shorebird predation, for both experiments. The probe-mark experiment detected the removal of large infaunal polychaetes (∼ 20 mm), while the exclosure experiment showed depletion of epifaunal harpacticoid copepods (0.063–0.5 mm). Predation on macrofaunal cumaceans was detected in both experiments. Invertebrates selected by western sandpipers neither fell within traditional infaunal size classifications (macro- vs. meiofauna; 500 μm delineation) nor corresponded to the highest densities of taxa. Rather, inference from experimental results and observations is that western sandpipers forage in two modes, by: (1) surface gleaning of epibenthic copepods and cumaceans in the macro- and meiofaunal size ranges and (2) selective probing for larger infauna, such as polychaetes. These findings were facilitated by the combination of methodologies employed. Received: 29 December 1999 / Accepted: 11 September 2000     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000