Genetic heterogeneity among blacktip shark,<Emphasis Type="Italic"> Carcharhinus limbatus</Emphasis>, continental nurseries along the U.S. Atlantic and Gulf of Mexico

Genetic population structure of the blacktip shark, Carcharhinus limbatus, a commercially and recreationally important species in the southeast U.S. shark fishery, was investigated using mitochondrial DNA control region sequences. Neonate blacktip sharks were sampled from three nurseries, Pine Island Sound, Terra Ceia Bay, and Yankeetown, along the Gulf of Mexico coast of Florida (Gulf) and one nursery, Bulls Bay, on the Atlantic Ocean coast of South Carolina (Atlantic). Sequencing of the complete mitochondrial control region of 169 neonates revealed 10 polymorphic sites and 13 haplotypes. Overall haplotype diversity and percent nucleotide diversity were 0.710 and 0.106%, respectively. Haplotype frequencies were compared among nurseries to determine if the high mobility and seasonal migrations of adult blacktip sharks have maintained genetic homogeneity among nurseries in the Atlantic and Gulf. Chi-square analysis and AMOVA did not detect significant structuring of haplotypes among the three Gulf nurseries, P(²)=0.294, _ST=–0.005 to –0.002. All pairwise AMOVA between Gulf nurseries and the Atlantic nursery detected significant partitioning of haplotypes between the Gulf and Atlantic (_ST=0.087–0.129, P<0.008), as did comparison between grouped Florida Gulf nurseries and the Atlantic, _CT=0.090, P<0.001. Based upon the dispersal abilities and seasonal migrations of blacktip sharks, these results support the presence of philopatry for nursery areas among female blacktip sharks. Our data also support the treatment of Atlantic and Gulf blacktip shark nursery areas as separate management units.Communicated by P.W.Sammarco, Chauvin     

Population genetic and phylogeographic structure of wahoo,<Emphasis Type="Italic"> Acanthocybium solandri</Emphasis>, from the western central Atlantic and central Pacific Oceans

The wahoo, Acanthocybium solandri (Cuvier, 1832), is a pelagic, highly migratory, scombroid fish, distributed worldwide throughout tropical and warm temperate seas. To evaluate population genetic and phylogeographic structure against a null hypothesis of panmixia, the entire mitochondrial DNA control-region (~890 base pairs) was sequenced for 231 wahoo. Samples were collected from 1997 to 2001 from seven sites: North Carolina (NC; n=23), east central Florida (CF; n=30), Bimini, Bahamas (BB; n=40), southern tip of Florida (SF; n=21), Cayman Islands (CI; n=23), northern Gulf of Mexico (NG; n=54), and Hawaii (HI; n=40). Inter-annual samples were obtained from four of these locations (NC, BB, SF, NG). Seventeen haplotypes were shared by individuals within and among samples; 187 singleton haplotypes were observed. Within-sample haplotype diversities ranged from 0.995 to 1.000 (overall h=0.999) and within-sample nucleotide diversities ranged from 0.049 to 0.055 (overall =0.053). A neighbor-joining tree based on inter-haplotypic distances revealed two monophyletic lineages differing by 13.6% nucleotide divergence. Nested within each major lineage were several, well-supported subclades. There was no evidence of temporal heterogeneity in haplotype distributions. Partitioning mtDNA variation, 99.75% of the variance was within samples and 0.25% (P=0.307) between samples; the fixation index (_ST=0.0025) was not significant. Likewise, pairwise _ST values were low or negative, and none were significant on a table-wide basis. Exact tests for sample differentiation in haplotypes were also non-significant. All population analyses were consistent with the null hypothesis of panmixia. However, analytical power was limited by sample size. Mismatch distributions were inconsistent with expected distributions based on sudden-expansion and static-growth models. Wahoo exhibit concurrently high haplotype and nucleotide diversities, presumably a consequence of secondary contact between historical subpopulations rather than a long, stable evolutionary history. Given the level of geographic and individual sampling, wahoo thus far represent the sole example of a scombroid or xiphioid fish exhibiting coarse-grain genetic homogeneity across a broad, inter-oceanic range despite a deeply coalescing genealogical structure. Accordingly, cooperative fishery management on a broad, inter-ocean scale may be warranted.Communicated by J.P. Grassle, New Brunswick     

Chemical defense: incorporation of diet-derived pyrrolizidine alkaloid into the integumental scales of a moth (<Emphasis Type="Italic">Utetheisa ornatrix</Emphasis>)

Summary. Evidence is presented that pyrrolizidine alkaloid acquired by Utetheisa ornatrix (Lepidoptera, Arctiidae) as a larva from Crotalaria foodplants is incorporated in part into the scales of the adult. A single forewing of a male or female moth may contain in the order of 6 to 13 g monocrotaline in its scale cover or about 1 to 2% of the moths systemic monocrotaline content. Based on estimates of the number of scales per forewing, the monocrotaline content of individual scales is calculated to be in the order of 0.1 and 0.2 ng monocrotaline per male and female scale, respectively. This amounts to concentrations of about 1 and 3%, values roughly at a par with the average systemic concentration (0.5-0.6%) previously determined for monocrotaline in Utetheisa. It is argued that the presence of pyrrolizidine alkaloid in the moths scale coating could account for the promptness with which adult Utetheisa are rejected by spiders. It is suggested further that chemical impregnation of scales with substances deterrent to predators may be more widespread among insects than generally assumed.     

Respiration rates of<Emphasis Type="Italic"> Beroe ovata</Emphasis> in the Black Sea

Metabolic rates of the ctenophore Beroe ovata within the length range from 0.4 mm (newly hatched larvae) to 60 mm were investigated. At 20° the respiration rates (Q, µg O₂ ind.^–1 h^–1) of individuals with wet weights (W, mg) less than or greater than 100 mg changed according to the equations Q=0.093W^0.62 and Q=0.016W^0.99, respectively. The weight-specific respiration rate of the juvenile ctenophores with wet body weights of 0.021–100 mg diminished approximately 20-fold (from 0.35 to 0.017 µg O₂ mg^–1 h^–1, respectively), but did not change within the range from 100 to 30,000 mg. The difference in the slope of the regression lines seems to be attributable to the ontogenetic changes in B. ovata metabolism. For the tested temperature range of 10–28°, the mean Q₁₀ coefficient was equal to 2.17±0.5. The basal metabolism of B. ovata narcotised by chloral hydrate was 4.5±0.9 times lower than total metabolism. Such a metabolic range is considered to be characteristic of aquatic invertebrates with high levels of locomotory activity.Communicated by O. Kinne, Oldendorf/Luhe     

Inter-oceanic genetic differentiation among albacore (<Emphasis Type="Italic">Thunnus alalunga</Emphasis>) populations

Albacore (Thunnus alalunga) is a highly migratory pelagic species distributed in all tropical and temperate oceans. Recent analyses using both mitochondrial DNA (mtDNA) and nuclear DNA markers have demonstrated genetic subdivision within and between Atlantic and Pacific populations. However, although numerous biological differences have been reported for Atlantic and Mediterranean albacore, the genetic differentiation for these basins has not been demonstrated. We characterized 373 base pairs of nucleotide sequence from the mitochondrial DNA control region of 134 individuals collected in the Pacific (n=30), the northeast (NE) Atlantic (n=54) and the Mediterranean (n=50). Analysis of molecular variance (AMOVA) revealed a small, but highly significant, proportion of genetic variation separating these three regions (_st =0.041; P=0.009), a pattern also supported by pairwise comparisons. These results demonstrate for the first time the genetic distinctiveness of the Mediterranean albacore from the NE Atlantic population giving support to the current management practices based on separate units. This outcome is concordant with reported migratory movements related to reproductive behavior between the NE Atlantic and the Mediterranean Sea. Additionally, the phylogenetic analyses of DNA sequences revealed the presence of a shallow genetic discontinuity with no geographic association. These two phylogroups are more likely the result of the demographic history of this species (i.e. long demographic stable history) as opposed to historical vicariance as has been proposed for other highly migratory fishes.Communicated by P.W. Sammarco, Chauvin     

Precision and accuracy of stable isotope signals in otoliths of pen-reared cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) when sampled with a high-resolution micromill

A high-resolution sampling technique for stable-isotope measurements in otoliths is now available, but little is known about the precision and accuracy of the results when using this technique. Otolith samples of pen-reared cod (Gadus morhua L.) with well-defined temperature history were obtained using this high-resolution sampling technique. Samples from different-aged cod otoliths were collected in two yearly increments corresponding to the calendar years 1994 and 1995, and the stable-oxygen (¹⁸O) and -carbon (¹³C) isotope compositions of the samples were measured. Otolith ¹⁸O values had a clear seasonal pattern in response to the seasonal water temperature. Otolith material deposited in the same calendar years of different-aged fish showed similar ¹⁸O values. High precision of the temperature estimate using otolith ¹⁸O values was obtained when comparing results from different otoliths. The accuracy of the temperature estimates, relative to measured mean monthly water temperature, was also high, but reduced sampling resolution in the otoliths significantly reduced the accuracy due to attenuation of the ¹⁸O signals. We found that the otolith ¹³C values had a cyclic pattern, roughly in phase with the ¹⁸O values. This was most likely caused by temperature-increased metabolism. Indications of age-specific otolith ¹³C values are also presented.Communicated by M. Kühl, Helsingør     

Molecular genetic structure suggests limited larval dispersal in grass rockfish,<Emphasis Type="Italic"> Sebastes rastrelliger</Emphasis>

The grass rockfish (Sebastes rastrelliger Jordan and Gilbert, 1880) is a non-migratory, benthic, near shore species distributed along rocky reefs and sea grass beds. It occurs from Baja California, Mexico, to Oregon, USA, spanning the Oregonian and Californian biogeographic provinces. In California this fish receives intense fishing pressure from an expanding and loosely regulated live-fish fishery. It is not known whether or by what mechanism larvae and juveniles are retained locally or dispersed widely during the early life-history phase. Tissue samples of S. rastrelliger were collected between 1996 and 2001 from 405 adult fish at eight sites (42.70°N; 124.50°W to 32.67°N; 117.25°W) spanning the species range. Individuals were surveyed for polymorphism at six microsatellite loci. Allele frequency heterogeneity was not significant among all sites (F_ST=0.001; P=0.18), nor in pairwise comparisons, but a clear correlation of genetic and geographic distance was detected (P=0.019). Fit of genetic and geographic distance was stronger within biogeographic provinces than at a range-wide scale, suggesting that populations north and south of the Point Conception biogeographic boundary are not in equilibrium with respect to migration and genetic drift. Estimates of mean coastal dispersal distances associated with the isolation-by-distance relationship are on the order of 10 km generation^–1. Such limited dispersal in a species with a pelagic early life history suggests active retention mechanisms near the shore. This has important implications for coastal management zones and design of marine reserves.Communicated by J.P. Grassle, New Brunswick     

Molecular genetic variation in tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis> Valenciennes) in the northern Atlantic Ocean

The tarpon (Megalops atlanticus) is a highly valued game fish and occasional food fish in the eastern and western Atlantic Ocean. Tarpon have a high capacity for dispersal, but some regional biological differences have been reported. In this study we used two molecular genetic techniques—protein electrophoresis of nuclear DNA loci, and restriction fragment length polymorphism analysis of the mitochondrial DNA (mtDNA)—to assess this species population genetic structure in the eastern (coastal waters off Gabon and Sierra Leone, Africa) and western (coastal waters off Florida, Caribbean Sea) Atlantic Ocean north of the equator. Genetic differentiation was observed between tarpon from Africa and tarpon from the western Atlantic Ocean. A unique allele and haplotype, significant differences in allozyme allele and mtDNA haplotype frequencies between the African and western Atlantic samples, and significant F_ST analyses suggest that levels of gene flow between tarpon from these two regions is low. Among the western Atlantic Ocean collections, genetic diversity values and allele and haplotype frequencies were similar. AMOVA analyses also showed a degree of genetic relatedness among most of the western Atlantic Ocean collections: however, some significant population structuring was detected in the allozyme data. A regional jackknifed F_ST analysis indicated the distinction of the Costa Rica population from the other western Atlantic populations and, in pairwise analyses, F_ST values tended to be higher (i.e., genetic relatedness was lower) when the Costa Rican sample was paired with any of the other western Atlantic samples. These data suggest that Costa Rican tarpon could be partially isolated from other western Atlantic tarpon populations. Ultimately, international cooperation will be essential in the management of this species in both the eastern and western Atlantic Ocean.Communicated by P.W. Sammarco, Chauvin     

Photoperiodic activity pattern in the brittle star<Emphasis Type="Italic"> Amphiura filiformis</Emphasis>

Diurnal activity pattern of suspension-feeding arms of the infaunal brittle star Amphiura filiformis was recorded in situ by time-lapse photography at 30 m water depth in the Gullmarsfjord, west Sweden. Activity was clearly related to the photoperiod with high activity at night and low or no activity during daytime. In laboratory experiments, the activity of the arms of A. filiformis changed in relation to new manipulated photoperiods, demonstrating exogenous circadian rhythms. The time difference for the greatest change in activity of the arms between control and treatment was correlated with the photoperiodicity. The recently found microlenses and associated neural photoreceptors in ophiuroids may apply also to A. filiformis. The trade-off between efficient feeding and predator avoidance is discussed.Communicated by M. Kühl, Helsingør     

Triterpenoid and caffeic acid derivatives in the leaves of ragweed, <Emphasis Type="Italic">Ambrosia artemisiifolia</Emphasis> L. (Asterales: Asteraceae), as feeding stimulants of <Emphasis Type="Italic">Ophraella communa</Emphasis> LeSage (Coleoptera: Chrysomelidae)

Summary. The leaf beetle Ophraella communa infests almost exclusively Ambrosia artemisiifolia in the fields of Japan, even though it normally feeds on several Asteraceous plants. A filter paper bioassay showed that the feeding of O. communa is strongly stimulated by methanolic extracts of A. artemisiifolia. The feeding stimulants for O. communa have been isolated from methanolic extracts of A. artemisiifolia. -Amyrin acetate, -amyrin acetate, 5-caffeoylquinic acid (chlorogenic acid) and 3,5-dicaffeoylquinic acid from A. artemisiifolia have been identified as feeding stimulants for O. communa. Triterpenoid derivatives (-amyrin acetate or -amyrin acetate) and caffeic acid derivatives (3, 5-dicaffoylquinic acid or 5-caffeoylquinic acid) showed feeding stimulant activity when mixed together.     

Relationship of daily and circatidal activity rhythms of the fiddler crab,<Emphasis Type="Italic"> Uca princeps</Emphasis>, to the harmonic structure of semidiurnal and mixed tides

Intertidal organisms may employ circatidal rhythms to track the tidal cycle, but tidal patterns may vary within a species range and necessitate adaptation to the local tides. Circatidal rhythms were examined in populations of the eastern Pacific fiddler crab Uca princeps (Smith) from four sites with differing tidal characteristics, La Paz (24°10N; 110°21W), San Blas (21°33N; 105°18W) and Manzanillo (19°6N; 104°24W), Mexico (lower amplitude, mixed semidiurnal tides) and Mata de Limon, Costa Rica (9°55N; 84°43W) (high-amplitude, semidiurnal tides). Local tides were characterized by harmonic constants of M₂, S₂, K₁, and O₁, partial tides that largely determine their semidiurnal and diurnal features. Rhythmic structure in continuously recorded locomotor activity of individual crabs held under laboratory conditions was described by cosinor and periodogram methods of time-series analysis. Both daily and circatidal rhythms were found in crabs studied in light–dark cycles set to local conditions at the time of collection. Crabs at all four sites shared a tendency toward bimodality, with a mid-morning activity peak and varying degrees of nocturnal activity. Circatidal rhythms closely matching the period of the 12.42-h M₂ partial tide were consistently present at all sites except Manzanillo. At Mata de Limon, the circatidal rhythm clearly dominated locomotor activity, but was strongly modulated by a daily rhythm in a repeating pattern at a semilunar interval. In contrast, the amplitude of the daily rhythm was higher than that of the circatidal rhythm in crabs from the three mixed tide sites on the Mexican coast, where the tidal pattern is dominated by a diurnal inequality arising from the diurnal K₁ and O₁ partial tides. These results suggest that populations of U. princeps use both daily and circatidal timing systems to track local forms of the tide generated by their M₂, S₂, K₁, and O₁ geophysical counterparts.Communicated by J.P. Grassle, New Brunswick     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.^–1 day^–1) were calculated for this species from laboratory estimates of energy losses due to routine (R_R), active (R_A), and feeding (R_SDA) oxygen consumption rates (J ind.^–1 h^–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to R_R and R_A increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q₁₀=3.01) than between 10°C and 14°C (Q₁₀=2.85). Rates of R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.^–1 day^–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Stability of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) depth limits: influence of habitat type

Seagrass meadows are generally considered stable although few studies have specified and tested this statement. On the basis of a large monitoring dataset from Danish coastal waters, we aimed to test whether the stability of deep eelgrass populations changes along a eutrophication gradient extending from inner bays over outer bays to open coastal waters. We defined stability in terms of the stability properties of constancy, resilience, and/or persistence. Our data allowed us to investigate the stability property constancy expressed as temporal variability in eelgrass depth limits over a time scale of 10 years. We hypothesised that colonisation depths were large and relatively constant along open coasts because of low and relatively constant levels of nutrients and turbidity. Conversely, colonisation depths were hypothesised to be low and variable in protected bays due to higher and more variable levels of nutrients and turbidity. We found that depth limits increased from inner bays towards open coastal waters, matching declines in nutrient concentration and increases in water clarity and oxygen concentration. Stability expressed as constancy of depth limits did not differ significantly between habitat types, and neither did stability of physicochemical variables. However, when data from all habitat types were analysed together, they showed that eelgrass populations at the depth limit were significantly more constant and thus, in this respect, more stable when occurring in deep waters as compared to shallow waters. Areas of good water quality may thus obtain the double benefit of deeper-growing and more stable eelgrass populations. The most likely reason why this pattern did not appear at habitat-type level is that the habitat types studied represented wide spatial variation in water quality and depth limit. We conclude that the question of whether eelgrass populations are stable depends on the stability property and the ecological situation in question. Populations may be considered unstable in terms of inter-annual variation but stable in terms of long-term persistence. Therefore, the common statement that eelgrass populations are stable is not universally true.Communicated by M. Kühl, Helsingør     

Reproduction and sex reversal of the solitary vermetid gastropod<Emphasis Type="Italic"> Serpulorbis arenarius</Emphasis>

The reproductive biology of the Mediterranean vermetid gastropod Serpulorbis arenarius, a subtidal sessile solitary species that lives in shady rocky communities, was studied at two localities south-east of the Iberian Peninsula. Its population density is usually low, with specimens sometimes sparsely distributed. Its shell often appears to be covered by epibiotic organisms. The sex ratio is biased toward males (63%) in the populations of SE Spain, and the results point towards hermaphroditism with alternating sexuality. The results obtained suggest that all specimens are first males, change to females under some as yet unknown condition and, after the ovogenesis cycle has been completed, revert to the original sexual state (male). The possible adaptive forces that favour hermaphroditism in this species are discussed, and we hypothesise that the seizure of a spermatophore by males may stimulate sex change to female. Brooding females (51% of females studied) were found from March to December. In males, histological study of testes revealed that spermatogenesis seems to occur throughout the year. Females brooded between 5 and 28 (=15) egg capsules, which were attached by stalks to the inner surface of the shell. It is noteworthy that the gonads of two of the specimens brooding egg capsules were developing testes. The newly laid egg capsules were filled with an organic intracapsular matrix, in which 40–153 (=94) eggs were embedded. The mean diameter of the uncleaved ova was 352.5 µm. The organic intracapsular matrix and some nurse eggs were used as an extra-embryonic food source. Hatching occurred at an advanced veliger stage, immediately followed by metamorphosis. Therefore, the larval development of S. arenarius is of a lecithotrophic intracapsular type, with a short, free-swimming/crawling phase.Communicated by O. Kinne, Oldendorf/Luhe     

Population genetics of the swimming crab<Emphasis Type="Italic"> Callinectes bellicosus</Emphasis> (Brachyura: Portunidae) from the eastern Pacific Ocean

The population genetics and historical demography of the swimming crab Callinectes bellicosus from the eastern Pacific were assessed using mitochondrial DNA (mtDNA) sequences from portions of the cytochrome c oxidase subunit I (COI) and cytochrome b (Cytb) genes. Analysis of molecular variance of sequence data from crabs collected from nine localities, ranging from the upper to lower Gulf of California and the outer coast of the Baja California peninsula, revealed an absence of population structure, suggesting a high level of gene flow over a wide geographic area. Maximum-likelihood estimates of long-term effective population size obtained with the program FLUCTUATE, in addition to highly significant values obtained from neutrality tests (Tajimas D, Fu and Lis D, and Fus F_S) and mismatch analysis, are consistent with a population expansion dating to the Pleistocene epoch. Phylogenetic analysis of C. bellicosus sequences using both neighbor-joining and Bayesian methods revealed a widely distributed subclade (clade II) cryptically embedded at low frequency in the main (clade I) population. Although sequence divergence between the two clades was low (1.1% COI; 0.6% Cytb), statistical support for the split was high. The Kimura-2-parameter genetic distance between C. bellicosus and the sympatric and morphologically similar C. arcuatus was high (d=0.17) and similar to the genetic distance between C. bellicosus and the allopatric C. sapidus from the western Atlantic (d=0.18), suggesting an ancient (Miocene) divergence of C. bellicosus and C. arcuatus.Communicated by P.W. Sammarco, Chauvin     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Sibling species in interstitial flatworms: a case study using<Emphasis Type="Italic"> Monocelis lineata</Emphasis> (Proseriata: Monocelididae)

The genetic relationships between morphologically indistinguishable marine and brackish-water populations of Monocelis lineata (O.F. Müller, 1774) (Proseriata: Monocelididae) were analysed by means of allozyme electrophoresis. Fifteen samples of M. lineata (13 from the Mediterranean and two from the Atlantic) from coastal marine and brackish-water habitats were examined for variation at 18 loci. Eleven loci were polymorphic in at least one population of M. lineata. Low levels of within-population genetic variability were found, with average observed and expected heterozygosity values ranging from H_o=0.015±0.015 to 0.113±0.044, and from H_e=0.028±0.028 to 0.138±0.054, respectively. The occurrence of a number of private alleles indicated a marked genetic divergence among populations of M. lineata, with Rogers genetic distances ranging from D_R=0.003 to 0.676 and a highly significant F_ST value (0.918±0.012, P<0.001). UPGMA (unweighted pair-group method using arithmetic average) cluster analysis and multidimensional scaling showed a clear genetic divergence between marine and brackish-water populations. Moreover, Atlantic and Mediterranean populations were sharply separated. Our results suggest that M. lineata is a complex of sibling species.Communicated by R. Cattaneo-Vietti, Genova     

Competitive interactions between two fucoid algae with different growth forms,<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> Himanthalia elongata</Emphasis>

Competitive interactions between two fucoid algae with different growth forms, Fucus serratus L. and Himanthalia elongata (L.) S.F. Gray were examined both in the laboratory and on a shore of the Isle of Man, Irish Sea. The growth of germlings of both species declined with increasing density, irrespective of whether they were with cohorts or rival species, indicating that intra- and interspecific competition occurred between germlings. H. elongata suppressed the performance of F. serratus at the germling stage by virtue of its larger initial size, and at the mushroom stage by forming a miniature canopy with the caps of the adjacent plants. In a field experiment, the mortality of H. elongata juveniles generally increased in mixtures with F. serratus and was highest when F. serratus were 50% of the plants. At the juvenile stage, the negative effect of F. serratus on H. elongata was more severe than the other way round. This was because F. serratus grows predominantly upwards, whereas H. elongata had already begun to expand laterally at the distal end. If F. serratus survives in sparse mixed stands with H. elongata juveniles, it can overgrow them and inhibit their subsequent survivorship and growth, probably by both shading and physical sweeping. H. elongata and F. serratus maintain their discrete monospecific stands because of the varying outcomes of mutual competitive exclusion resulting from their differing growth patterns. Thus it is possible for them to co-occur at a similar shore height.Communicated by O. Kinne, Oldendorf/Luhe     

Proclivity and effectiveness in gall defence by soldiers in five species of gall-inducing thrips: benefits of morphological caste dimorphism in two species (<Emphasis Type="Italic">Kladothrips intermedius</Emphasis> and<Emphasis Type="Italic"> K. habrus</Emphasis>)

The essence of eusociality is a trade-off between producing ones own offspring and helping collateral kin via such activities as defence, foraging and brood rearing. This trade-off often involves morphological differences between helper and reproductive castes, but the advantages, correlates and phylogenetic context of morphological caste differentiation have seldom been analysed. Six species of Australian gall-inducing thrips on Acacia show morphological polymorphism. One morph, referred to as a soldier, has reduced wings and antennae but greatly enlarged fore-femora, which are thought to be adaptations for gall defence. The other, dispersing morph, has fully developed wings and relatively slight fore-femora. Here, we quantify the defensive behaviour of soldier morphs, and compare soldier and foundresses, using behavioural assays designed to measure proclivity to attack kleptoparasites (specialised invaders in the genus Koptothrips) and effectiveness in killing them. In all five species investigated, soldiers were able to kill Koptothrips. Moreover, the effectiveness of soldiers was relatively high in the more-derived species in the phylogeny of the clade, Kladothrips intermedius, K. habrus and K. waterhousei. Soldiers of K. intermedius and K. habrus also killed kleptoparasites more effectively than did foundresses, and K. habrus soldiers exhibited higher proclivity to attack than did foundresses. Data from naturally invaded galls demonstrate that soldiers in the field do kill Koptothrips, and vice versa. These results show that soldiers of Australian gall thrips are motivated and effective for gall defence.Communicated by D. Gwynne