Poecilogony in Polydora hoplura (Polychaeta: Spionidae) from commercially important molluscs in South Africa

Females of the polychaete Polydora hoplura (Claparède 1869) that produced planktotrophic and adelphophagic larvae were compared genetically to determine whether the different reproductive morphs represented sibling species or poecilogony. Worms were collected from Saldanha Bay, South Africa (33°0′37.71S, 17°56′59.74E), and cultured in a laboratory setting from November 2012 to April 2013. The results based on mitochondrial and nuclear DNA (Cyt b and ATPSα, respectively) showed shared haplotypes between reproductive morphs for both markers. Additionally, variation in mtDNA sequences was significantly higher within morphs than between morphs (95.8 and 4.2 %, respectively). A comparison of developmental modes found that broods of planktotrophic larvae were larger than broods of adelphophagic larvae, while the latter had larger eggs and larvae at hatching. Developmental time from oviposition to settlement for planktotrophic and adelphophagic larvae was 40.2 and 16.6 days, respectively. Polydora hoplura represents the fifth confirmed case of poecilogony in the Spionidae.     

Molecular and morphological evidence of a single species, Boccardia proboscidea (Polychaeta: Spionidae), with multiple development modes

Most species of benthic marine invertebrates have a single mode of larval development. Poecilogonous species are those that produce more than one type of larval offspring. Reports of variable development within one species, especially in combination with widely differing ecological habitat, are frequently attributed to cryptic species. The spionid polychaete Boccardia proboscidea Hartman, 1940 exhibits development that varies both within a single brood and among broods produced by different females. Some females have planktotrophic development and produce many small larvae with a 2 week planktonic period before metamorphosis. Other females produce broods containing both planktotrophic larvae as well as nurse-egg-ingesting (adelphophagic) offspring that hatch as juveniles. Molecular analysis (RAPD-PCR) showed that a significant proportion of genetic variance is attributable to geographic origin, and not to developmental type. Adults of both developmental types showed no consistent differences in taxonomically important features (e.g. type and arrangement of chaetae, modified fifth setiger, caruncle, branchiae, pygidium) when examined with SEM. These data support the hypothesis that developmental variability in this species is a case of poecilogony, and is not attributable to cryptic species. Received: 21 April 1998 / Accepted: 20 April 1999     

Enzyme activities and allozyme polymorphism in two genetic types (or sibling species) of the genus Marenzelleria (Polychaeta: Spionidae) in Europe

Two populations of Marenzelleria spp. from the North Sea and the Baltic Sea that had already been characterized by allozyme electrophoresis and found genetically different in 1994 were again examined in 1997. The genetic identity (I = 1.0) examined in 1994 and 1997 shows that the structures of the populations of genetic Type I in the North Sea (Ringkobing Fjord) and genetic Type II in the Baltic Sea (Darss–Zingst Bodden chain) are stable. Three diagnostic gene loci that were determined additionally (citrate synthase, one locus; alanine aminotransferase, two loci) support the hypothesis of the existence of two sibling species of the genus Marenzelleria in Europe. In addition, the activities of ten electrophoretically separated enzyme systems were measured in the worms immediately after removal from their habitat and after they had been kept in the laboratory for 2 months. Under both conditions four enzymes showed significant differences between genetic types. The higher activity of the alanine aminotransferase of the North Sea worms (Type I) is discussed in the context of the different preferred salinities of the two Marenzelleria sibling species. Received: 21 January 1999 / Accepted: 13 July 1999     

Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.     

Particle-size selection by Pseudopolydora paucibranchiata (Polychaeta: Spionidae) in suspension feeding and in deposit feeding: influences of ontogeny and flow speed

Pseudopolydora paucibranchiata Okuda suspension feeds and deposit feeds at the sediment-water interface, where it is exposed to a variety of particles differing in physical characteristics and nutritional value. In flume experiments (conducted in August 1994 and May 1995) with two sizes of either suspended or deposited beads, I measured particle-size selection separately in each feeding mode. The same influences of palp width and of ambient flow speed were observed in each mode. At velocities 0.74 cm s^-1 there were no relationships between palp width and the proportion of gut contents composed of large beads. At velocities 1.8 cm s^-1 worms with narrower palps ingested relatively fewer large beads (and more small beads) than did worms with wider palps. Palp width and body length were linearly related, and results were similar when analyzed with body length as the independent variable. As flow speed increased, selectivity changed in a worm-size-specific manner: worms with a palp width -1. Assuming that in the field (1) particle size is the principle criterion for selection, and (2) the amount of digestible food component in deposited and suspended particles, respectively, is related to particle surface area and volume, I hypothesize that changes in selectivity as velocity rises can cause juveniles to experience a decreasing profitability of suspension feeding and a simultaneously increasing profitability of deposit feeding. Juveniles could maintain a diet of high food value despite flow variations by adjusting the proportion of time they spend suspension feeding relative to deposit feeding.     

Age structure and spawning of a Northumberland population of Melinna cristata (Polychaeta: Ampharetidae)

A Northumberland (England) population of Melinna cristata was studied from October, 1967 to February, 1970. M. cristata is a gregarious sedentary polychaete which lives in muddy substrates. It is dioecious, and has a very restricted breeding season. The gametes are shed solely by the two pairs of posterior nephridia, which undergo morphological changes just before the breeding season; this allows them to accommodate the mature gametes. After spawning, these nephridia ternal and, within 2 to 3 weeks of spawning, the benthic larvae metamorphose into juveniles. The juveniles grow from 3 to 10 mm during their first year, and can be distinguished from the rest of the population by their size and the complete absence of coelomic gametes. After about 12 months, the juveniles release their first batch of gametes into the coelom from the germinal epithelium, and they then cannot be distinguished from the rest of the population. The development of coelomic gametes takes 7 to 10 months to complete, so M. cristata is potentially capable of breeding for the first time when 2 years old. The majority of worms survive spawning, and M. cristata probably breeds annually for several years. In this population, not all potential breeders spawn, some resorb their mature gametes and release another batch of gametes into the coelom. The Northumberland population of M. cristata is near the southernmost limit of the species distribution, which suggests that environmental conditions for this population are not optimum. The population appears to maintain itself by producing fewer oocytes and by only part of the population spawning.     

Elysia timida (Risso, 1818) (Gastropoda, Opisthobranchia): relationship and feeding deterrence to a potential predator on the south-western Mediterranean coast

The relationship between the sacoglossan Elysia timida and the ornate wrasse Thalassoma pavo was studied in two laboratory experiments using artificial models. A feeding-preference experiment was conducted to determine whether mollusc extract deterred feeding by T. pavo, by using a "realistic" mollusc model (imitating the colour pattern of E. timida) coated with mollusc extract, and a reference model (without imitation or extract), and fishes collected from two locations. It was observed that fish approached, attacked and ate more reference models than mollusc models. A second feeding-preference experiment was designed with four different coloured models: "realistic" (W), green (G), red (R) and chequered (S) pattern. Both fish populations clearly rejected the S model, and differed in their colour preferences. Although both populations seemed to prefer the R and G models equally, the W model was clearly rejected by the fish that coexist with the mollusc at one site (Mazarrón), but was not rejected by the other population of fish which does not coexist with it (Cabo de Palos). Mazarrón fishes would identify the W model with the presence of a toxic compound during their coexistence, and therefore avoid attacking conspicuously coloured E. timida models as a response to their visual signals. Therefore, it was concluded that extract of E. timida is a deterrent for T. pavo, and its effect is sufficiently noxious that the fish tend to avoid it, so that the ability of the fishes to learn to recognise colours and identify certain colour patterns associated with obnoxious prey provides the molluscs the opportunity to survive by exhibiting a conspicuous coloration.     

Population dynamics and secondary production in an estuarine population of Caulleriella caputesocis (Polychaeta: Cirratulidae)

Between July 1978 and March 1980, the distribution, population dynamics and secondary production of Caulleriella caputesocis St. Joseph, 1864, in Southampton Water, South England, were investigated. The distribution of the polychaete was related to amount of silt and copper-content of the sediment, the highest densities occurring in sediment containing 60 to 100% silt and less than 50 ppm copper. Breeding occurred at a low level throughout the year, usually when the worms were over one year old, with peak breeding between May and September. C. caputesocis in Southampton Water is monotelic, and oocytes measured 120 to 160 m in diameter at spawning. Unshed oocytes were resorbed. Annual secondary production varied between 0.01 and 5.90 g C m^-2 yr^-1 and P:B ratio of the species ranged between 0.84 and 7.09.     

Population dynamics and secondary production in an estuarine population of Nephtys hombergii (Polychaeta: Nephtyidae)

From July 1978 to March 1980, a study was made on the distribution, population dynamics and secondary production of Nephtys hombergii Audouin et Edw. occurring in the sublittoral industrialised region of Southampton Water in south England. The distribution of the worm was related to the silt content and copper level of the sediment, the greatest densities of N. hombergii being found in sediment containing 60 to 100% silt. Breeding occurred at a low level throughout the year, with a maximum in July to September and November to January in the second year of growth. Spawning occurred when the oocytes measured 200m in diameter, and unshed gametes were resorbed. Annual production varied between 0.092 and 4.32 g C m^-2 yr^-1 (ash-free dry weight) and amounted to 1.9–39.4% of the total macrofaunal production at the sampling stations. The production:biomass (P:B) ratio of the species varied between 1.6 and 2.9.     

Effects of climate and exurban development on nest predation and predator presence in the southern Appalachian Mountains (USA)

In the eastern United States, land-use and climate change have likely contributed to declines in the abundance of Neotropical migrant birds that occupy forest interiors, but the mechanisms are not well understood. We conducted a nest-predation experiment in southern Appalachian Mountain forests (North Carolina, U.S.A.) during the 2009 and 2010 breeding seasons to determine the effects of exurban development and temperature on predator presence and the average number of days until eggs in an artificial nest were disturbed by predators. We baited artificial nests with quail (Excalfactoria chinensi) eggs and monitored them for 18 days. We used clay eggs, track plates, and motion-triggered cameras to detect and identify nest predators. The average number of days a nest was undisturbed decreased as mean temperature increased and, to a lesser extent, as the density of buildings increased. Nests on the ground were more often depredated than those in trees, likely due to increased predation by opossum (Didelphis virginiana) and other carnivores. Raccoons (Procyon lotor), opossums, corvids (Corvus brachyrhynchos and Cyanocitta cristata), chipmunks (Tamias striatus), black bears (Ursus americanus), and domestic cats (Felis catus) were the most commonly detected predators. Presence of these predators did not vary as a function of mean temperature. Domestic cats and corvids were detected more frequently in plots with high rather than low densities of buildings. Forest-interior specialists and Neotropical migrants often nest in cool, high-elevation areas with low housing density. These bird species, especially those that nest on the ground, may be most vulnerable to increased nest predation if temperature and exurban development increase at higher elevations as anticipated.     

Spatio-temporal population and reproductive responses in Perinereis gualpensis (Polychaeta: Nereididae) from estuaries under different anthropogenic influences

Reproductive and population-level responses are ecologically relevant diagnostic tools for revealing the cost of long-term tolerance to contaminants. Spatial density distribution, seasonal density, biomass, size frequency histograms and oocytes stages in Perinereis gualpensis (Polychaeta: Nereididae) from a highly anthropogenically impacted estuary (Lenga, south-central Chile) were compared with low-polluted estuaries (Tubul and Raqui). Spatial distribution showed that the highest P. gualpensis abundances occur in the central (middle) estuarine area, establishing them as suitable zones for comparisons of estuaries. Middle areas also showed differences among estuaries in terms of population and reproductive responses. Mature stage oocytes and juvenile recruitment limited to certain seasons and low adult survival led to significant low densities and biomasses in Lenga individuals (p?相似文献   

Demography of Pholoe minuta (Polychaeta: Sigalionidae) in Galway Bay,west coast of Ireland,with special reference to settlement and recruitment patterns

Pholoe minuta is a numerically dominant member of the Amphiura filiformis subcommunity in Galway Bay. A longterm (5 yr) assessment of its population structure highlighted the shortfall in information regarding juvenile stages in the post-spawning months. Additional remote and in situ sampling programmes (including the use of a Muus sampling device), covering two successive postspawning periods, were undertaken to provide information on the juvenile recruits. Estimates of density levels (130 to 680 m^-2), growth rates (doubling in size in 2 mo) and mortalities (80%) were calculated for the newly settled P. minuta. Mean monthly densities, seasonal population structure, recruitment levels (21.1%), growth (1 mm per annum) and survivorship patterns of the adult population were evaluated using the dorsal width of the fifth chaetiger, which is shown to be a suitable parameter of total body size. The success of grab sampling as a method for examining early settlement of P. minuta is critically reviewed, and a recommendation for future benthic population studies is proposed.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

Uptake,binding and clearance of divalent cadmium in Glycera dibranchiata (Annelida: Polychaeta)

The bloodworm Glycera dibranchiata Ehlers, 1968 accumulates cadmium through the general body surface and the intestine. Absorption through the gut accounts for cadmium which rapidly binds to coelomic proteins. Intracoelomic injection of ¹⁰⁹Cd demonstrates that cadmium binds readily to hemoglobin and other proteins. The degree of cadmium binding is pH-dependent. The apparent pK of binding sites in body wall and musculature homogenates is 5.39. Cadmium ions injected into the coelom at 7 g g^-1 tissue increase proline incorporation rates into the positively charged hemoglobin (cathode fraction) by 15-fold in 3 days. A 3-fold increase of proline incorporation was observed in the anode hemoglobin fraction over the same time period. Radioactivity in various protein fractions decreases at different rates after injection of ¹⁰⁹Cd. Comparisons between the function of mammalian metallothionein and the coelomic fluid proteins of G. dibranchiata as a detoxification mechanism are discussed.     

Genetic and demographic consequences of importing animals into a small population: a simulation model of the Texas State Bison Herd (USA)

The extant 40 bison (Bison bison) constituting the Texas State Bison Herd (TSBH; USA) are directly and exclusively descended from a bison herd assembled by Charles Goodnight in the 1880s, representing a historically and genetically valuable resource. The population currently suffers from low genetic variation, low heterozygosity, high calf mortality, and low natality rates compared with other closed bison populations. Population viability analysis using the VORTEX program previously indicated a 99% chance of population extinction within the next 41 years [J. Mamm. 85 (2004) in press]. We developed a stochastic simulation model to evaluate the genetic and demographic consequences of various management scenarios for the TSBH using genotypic data from 51 microsatellite loci and demographic information recorded over a 6-year period. Our results reveal that without the introduction of new genetic variation, approximately 37% of the representative microsatellite loci will become fixed as the TSBH continues to lose genetic variation at a staggering rate of 30–40% within the next 50 years. Furthermore, if the current trends in natality and mortality rates continue, our model indicates the TSBH faces a 99% chance of extinction in the next 51 years. With the importation of unrelated male bison into the TSBH, and under the assumption of increased fitness, the probability of population survival in the next 100 years increases to 100%, and the population will reach the approximate carrying capacity of 200 bison in 15–16 years. Furthermore, our model predicts increases in genetic diversity and heterozygosity of 24.7–48.4% and 17.5–36.5%, respectively, in the next 100 years following the addition of new genetic variation. We conclude that the importation of bison into the TSBH is necessary to prevent extinction and ensure long-term population survival.     

Population structure and production of Lumbrineris fragilis (Polychaeta: Lumbrineridae) in the Oslofjord (Norway) with a note on metal content of jaws

Population structure and production in Lumbrineris fragilis (O. F. Müller) based on samples collected from April 1981 to June 1983 were investigated. Based on the number of growth lines on the mandibles, the population of L. fragilis at Bjørnehode Bay in the Oslofjord could be divided into ten size classes, with the first six size classes being most commonly represented. There were strong indications that the dark bands in the mandibles were formed during winter and the light bands during summer. Neither seasonal changes in temperature nor gametogenesis were likely causes of dark band formation. A possible cause could be a reduction in feeding intensity during winter either due to changes in quantity or quality of available food. Production was calculated to be 1.32 g AFDW (ash-free dry weight) m^-2 year^-1 and the P/B ratio was 0.826. Chemical analysis of the mandibles showed that, in L. fragilis, calcium was present in highest concentrations while smaller quantities of copper, zinc, iron, phosphorous and sulphur were also recorded.     

Impact of Arenicola marina (Polychaeta) on the microbial assemblages and meiobenthos in a marine intertidal flat

The benthic microbial food web can be responsible for a large proportion of benthic carbon cycling yet there are few data on the trophic interactions between this food web and macrobenthos. A large-scale field experiment was conducted to investigate effects of eliminating the polychaete Arenicola marina on benthic microbes (prokaryotes, heterotrophic and autotrophic protists) and metazoan meiofauna in a marine intertidal flat of the North Sea, Germany. Over a period of 2 years, quantity and composition of micro- and meiobenthos from unmanipulated sites were compared to those from sites deplete of lugworms. These grazer treatments were cross-classified with different sediment characteristics (low- and mid-intertidal areas). Lugworm removal resulted in an initial increase in abundance of prokaryotes and nanoflagellates, which became less pronounced in the second year. Ciliates were not affected quantitatively, but in the absence of lugworms, diversity and the proportion of carnivorous forms increased. Meiobenthos (nematodes, ostracods and copepods) were affected only moderately. The observed changes are probably due to a combination of release from grazing/predation pressure, changes in the species composition of higher trophic levels (namely large polychaetes) and altered environmental conditions (such as depth of the oxygenated layer and sediment grain size). Spatial differences between sites of different tidal exposure/grain size appeared to be as large as temporal differences during the 2 years following the manipulation of the system. We conclude that in intertidal sediments, indirect effects due to habitat transformation are as important as direct biological interactions (grazing pressure and competition) for the dynamics of the benthic microbial food web.     

Reproduction activity of<Emphasis Type="Italic"> Exogone naidina</Emphasis> (Polychaeta: Exogoninae) in a population from the Apulian coast (Adriatic Sea)

This study reports on the reproduction activity of Exogone naidina Örsted, 1845, followed on a hard-bottom population located at a site along the coast of Bari (South Adriatic Sea), during a year of study from November 2000 to October 2001. Population structure was analysed together with the reproductive state of individuals over time. The interpretation of frequency histograms computed on the total worm length was not easy because of the almost complete lack of juvenile forms. Ripe individuals were present for most of the year, covering a large range of sizes. The reproductive season extended from April to December, with two peaks of activity: one from April to June and one in September–October. Life span was hypothesized to be up to 1 year, with generational change clearly visible only in late summer. However, the prolonged reproductive period produces an overlapping of generations. The link between population density and the reproductive cycle of the species with food availability is also discussed.Communicated by R. Cattaneo-Vietti, Genova     

Variation in protein patterns within and among polychaete sibling species of the genus<Emphasis Type="Italic"> Marenzelleria</Emphasis> (Spionidae): a preliminary survey

Differences in protein patterns of the soluble protein fraction among the sibling species Marenzelleria viridis (formerly type I) and M. neglecta (formerly type II) were investigated under common environmental conditions using two-dimensional gel electrophoresis (2D-PAGE). Protein expression was determined using general protein staining with Coomassie-blue and compared with radioactive labeling of proteins. In the well-resolved region of stained gels an average of 319 protein spots for M. viridis and 241 spots for M. neglecta could be detected. High sensitivity of radiolabeling allowed separation of an average of 517 and 496 spots for M. viridis and for M. neglecta, respectively. Differences in protein expression between both species could be attributed mainly to qualitative differences in protein patterns. Triplet pattern was used to calculate the genetic similarity of the two species. Thus, 373 protein spots were scored for this analysis; whereas 304 spots were invariant, 36 spots were specific for M. viridis, while 33 spots were specific for M. neglecta. The genetic similarity (F) of the two Marenzelleria sibling species was 0.815. Apart from presence and absence, differences between both species resulted either from slight changes in the isoelectric point or from molecular weight, but rarely from both. Genetic variability was found only among specimens of M. viridis. The experimental conditions to perform two-dimensional electrophoresis for these polychaete species were established for subsequent investigations on a proteomic level. Using 2D-PAGE we expect further insight into the evolutionary adaptation in Marenzelleria spp.Communicated by O. Kinne, Oldendorf/Luhe     

Spermatogenesis,sperm storage and comparative sperm morphology in nine species of Capitella,Capitomastus and Capitellides (Polychaeta: Capitellidae)

Light microscopy and transmission electron microscopy (TEM) were used to describe spermatogenesis and the morphology of mature sperm and sperm storage organs in five sibling species of Capitella, three species in the related genus Capitomastus, and one species in the genus Capitellides. These capitellids lack a well-developed testis, but young males have a few specialized regions of the peritoneum in the eighth setiger, where germ cells proliferate and spermatogonia are released into the coelom, and spermiogenesis is completed. Mature sperm are stored in the central regions of paired genital ducts (coelomoducts), which lie between the seventh and eighth setigers. The cells forming the walls of the coelomostome and central region of the duct are ciliated and have large glycogen deposits. The lumenal borders have extensive microvilli and there is evidence that they secrete glycogen-containing materials into the duct. All species have modified primitive sperm with a conical acrosome, elongated nucleus, and long middle piece extending along the proximal portion of the flagellum. A single ring-shaped mitochondrion encircles the centriolar region of the middle piece and the cytoplasm is filled with glycogen. The sperm of all nine species differ significantly in the lengths of their middle pieces, acrosomes and especially in their nuclear lengths. The nuclear lengths have a twofold range among the sibling species of Capitella and Capitomastus. Subtle differences in the shape and volume of the acrosomal vesicle and acrosomal space characteristic of the Capitella sibling species seem to correlate with a basic division of these species into those with diploid chromosome numbers of 20 or 26. Spermiogenesis, the number of sperm produced, and the method of sperm storage are appropriate for efficient sperm utilization in fertilization. No evidence indicates that spermatophores are formed and transferred between individuals and the method of sperm transfer is not understood. The differences in the dimensions and acrosome morphology of mature sperm, and the previously demonstrated specializations in the egg envelopes in the Capitella sibling species, are characteristic features of the reproductive isolation that exists among these capitellid species.