Food webs and fishing affect parasitism of the sea urchin Eucidaris galapagensis in the Galápagos

In the Galápagos Islands, two eulimid snails parasitize the common pencil sea urchin, Eucidaris galapagensis. Past work in the Galápagos suggests that fishing reduces lobster and fish densities and, due to this relaxation of predation pressure, indirectly increases urchin densities, creating the potential for complex indirect interactions between fishing and parasitic snails. To measure indirect effects of fishing on these parasitic snails, we investigated the spatial relationships among urchins, parasitic snails, commensal crabs, and large urchin predators (hogfish and lobsters). Parasitic snails had higher densities at sites where urchins were abundant, probably due to increased resource availability. Commensal crabs that shelter under urchin spines, particularly the endemic Mithrax nodosus, preyed on the parasitic snails in aquaria, and snails were less abundant at field sites where these crabs were common. In aquaria, hogfish and lobsters readily ate crabs, but crabs were protected from predation under urchin spines, leading to a facultative mutualism between commensal crabs and urchins. In the field, fishing appeared to indirectly increase the abundance of urchins and their commensal crabs by reducing predation pressure from fish and lobsters. Fished sites had fewer snails per urchin, probably due to increased predation from commensal crabs. However, because fished sites also tended to have more urchins, there was no significant net effect of fishing on the number of snails per square meter. These results suggest that fishing can have complex indirect effects on parasites by altering food webs.     

Diet, movement, and covering behavior of the sea urchin Toxopneustes roseus in rhodolith beds in the Gulf of California, México

The density, diet, movement, and covering behavior of Toxopneustes roseus (Agassiz) were investigated in rhodolith beds in the Gulf of California. Densities varied from a mean of 0.4 to 1.8/20 m² with most urchins occurring in aggregations. Spatial patterns of urchins varied with depth, with greatest abundance at intermediate depths (7.5–9.4 m) in the middle of the rhodolith bed. Urchins ate rhodoliths and nongeniculate coralline algal crusts almost exclusively, despite the availability of other algae. The mean amounts ingested were 3.87 and 7.96 g carbonate/individual per day. Even when food was abundant, animals were highly mobile, moving an average of 6.6–11.7 cm/h depending on site and time of day. Diel movement may be a behavioral adaptation to avoid surge, which is greatest during the day. Covering behavior may also be related to surge, as the ratio of covering material:body weight and the percent cover of material held were highest at the site with the most surge. While an urchin consumed rhodoliths, its movement spread the grazing impact over large areas. Bioturbation resulting from urchin feeding, movement, and covering activity probably benefits the rhodoliths by turning them, which maintains rhodolith integrity, prevents fouling, and contributes to bed persistence. Received: 22 February 1999 / Accepted: 3 July 2000     

Interplay between proteases and protease inhibitors in the sea fan—Aspergillus pathosystem

Environmental conditions greatly impact the dynamics of host–pathogen relationships, affecting outbreaks and emergence of new diseases. In corals, epizootics have been a main contributor to the decline of reef ecosystems and rising sea surface temperatures associated with climate change are thought to exacerbate virulence of pathogens and/or compromise the immune responses of coral hosts. Among pathogens strategies, protease secretion is a common virulence factor that promotes colonization of the host. As such, protease inhibitors play an invaluable role to the host for protection against pathogen invasion. This study describes changes in activity of proteases secreted by the fungal pathogen, Aspergillus sydowii, and protease inhibitors of Gorgonia ventalina under ambient and elevated temperatures and health conditions (disease and healthy). G. ventalina colonies were collected from Florida Keys, Florida (24° 34.138N, 81° 22.905W) or La Parguera, Puerto Rico (17° 56.091N, 67° 02.577W) in 2007 and 2012, respectively. At elevated temperatures, protease activity was significantly higher than at ambient temperatures in A. sydowii. Temperature stress did not induce a change in protease inhibitor activity, but in healthy G. ventalina colonies, inhibitor activity against proteases was higher than in diseased individuals. Healthy colonies appear capable of resisting proteolytic activity, but diseased individuals are still likely affected by pathogen infections. These data suggest that rising sea surface temperatures due to climate change may increase levels of virulence (protease activity) of A. sydowii, while immunity (protease inhibitor) of G. ventalina may not be affected.     

Phylogeography of California and Galápagos sea lions and population structure within the California sea lion

We investigate the phylogeography of California (Zalophus californianus) and Galápagos (Z. wollebaeki) sea lions and describe within-population structure for the California sea lion based on mitochondrial DNA. Fifty control-region haplotypes were found, 41 from Z. californianus and 9 from Z. wollebaeki, with three fixed differences between the two species. Ranked population boundaries along the range of Z. californianus were defined based on the Monmonier Maximum Difference Algorithm, resulting in five genetically distinct populations, two in the Pacific Ocean and three inside the Gulf of California. A Minimum Spanning Network showed a strong phylogeographic signal with two well-defined clusters, Z. californianus and Z. wollebaeki, separated by six base-pair differences, supporting the existence of two genetically distinct species with an estimated divergence time of ~0.8 Ma. Results are discussed in the context of the historical geologic and paleoceanographic events of the last 1 Ma in the eastern Pacific.     

Early reproduction and development of dominant calanoid copepods in the sea ice zone of the Barents Sea—need for a change of paradigms?

Reproduction and growth of the dominant copepods Calanus finmarchicus, C. glacialis, C. hyperboreus and Pseudocalanus minutus were studied on transects across the sea ice zone of the northern Barents Sea in May and June 1997. C. glacialis and C. finmarchicus were numerically dominant and also the largest component of the biomass. C. hyperboreus was rather rare. Moderate levels of phytoplankton and eventually high concentrations of ice algae supported maximum egg production rates of 53.6 and 48.5 eggs female^–1 day^–1 of C. glacialis in May and June, respectively. Results of incubation experiments were supported by a tremendous abundance of C. glacialis eggs in the water column ranging from 7×10³ to 4.4×10⁴ m^–2 in May and from 9.8×10³ to a maximum of 9.7×10⁴ m^–2 in June. In contrast, C. finmarchicus spawned only in the vicinity of the ice edge, at a maximum rate of 30 eggs female^–1 day^–1. Egg sacs of P. minutus were often observed in the preserved samples, but contained only few eggs, which may be due to loss during sampling. The presence of considerable concentrations of young stages in May and June indicated successful recruitment of C. glacialis and P. minutus. Back calculation using published stage duration estimates indicates March/April as the begin of the reproductive and growth period for these species under the first-year ice of the Barents Sea. Hence, secondary production in the study area starts at the same time as in open water regions and polynyas in the northern North Atlantic. Although the role of ice algae in the nutrition of copepods was not clarified here, the significant relationship between phytoplankton chlorophyll and egg production of C. glacialis suggests that high reproductive activity has already been achieved at moderate food concentrations.Communicated by O. Kinne, Oldendorf/Luhe     

Fine structure of the “button setae” in the deep-sea pelagic copepods of the genus Euaugaptilus (Calanoida: Augaptilidae)

 Many species of the carnivorous copepod family Augaptilidae, a primarily meso- and bathypelagic group, possess specialized “buttons” on the setae of their maxilla and maxilliped, which have been thought to be a simple cuticular outgrowth. The fine structure of the button setae in three species of Euaugaptilus was examined by light- and electron microscopy from samples collected in the eastern Indian Ocean and the subtropical western North Pacific. The buttons are arranged in two rows along the inner surface of the setae. There are differences in the shape, size, and arrangement of buttons among the three species, Euaugaptiluslaticeps, E.longimanus, and E.magnus. The button setae have an elaborate internal structure consisting of “setal shaft”, “stalk”, “disc”, and “outer membrane”, with microtubule bundles in the setal lumen, but have no muscular tissues. The disc has only pin-point contact to the core of the stalk, but has connection over a wider area with the fibrous tissue surrounding the core, and the contact area between the stalk and the setal shaft is also small. These structures suggest that the disc will move freely on the apex of the core of the stalk and the stalk may be movable at its connection to the setal shaft. All these components of the button setae may function in concert to fit the surface of the discs firmly to the surface of a prey, and to absorb the energy generated by the struggling prey. Received: 30 November 1999 / Accepted: 19 May 2000     

Does the composition of the demersal fish assemblages in temperate coastal waters change with depth and undergo consistent seasonal changes?

The aim of this study was to determine whether the composition of the demersal fish fauna in coastal marine waters in temperate Australia changes markedly with increasing water depth and distance from the shore and whether the composition of the fish fauna in water depths of 5 to 35 m undergoes cyclic, seasonal changes. Samples of demersal fishes were therefore collected by trawling over the predominantly sandy substrate at nine sites located in water depths of 5 to 15 m or 20 to 35 m and within 20 km of the shore in four regions along ∼200 km on the lower west coast of Australia. The sampling regime involved trawling for fishes at each site at night in seven consecutive seasons between the summer of 1990/1991 and winter of 1992. A total of 72 435 fishes, representing 77 families, 143 genera and 172 species was caught. The compositions of the fish faunas in offshore waters with depths of 5 to 35 m were shown to differ markedly from those previously recorded for nearshore marine waters in the same regions. However, as some species, such as Sillago burrus, S. vittata, S. bassensis and Rhabdosargus sarba, increase in size, they move out from their nursery areas in nearshore waters into deeper and more offshore waters, where spawning occurs. Ordination showed that, in each of the four regions, the composition of the fish fauna in depths of 5 to 15 m differs from that in depths of 20 to 35 m. This difference is attributable to the fact that some species, such as  S. burrus, S. vittata and Upeneichthys lineatus, are far more abundant in depths of 5 to 15 m, whereas other species, such as S. robusta, U. stotti and Lepidotrigla modesta, occur predominantly in depths of 20 to 35 m. However, the samples collected from the single site that was inshore but in deeper water demonstrate that the composition of the fish fauna is influenced by distance from shore as well as by water depth. The compositions of the fish faunas differed with latitude, largely due to the fact that some subtropical species, such as Polyspina piosae, S. burrus and  S. robusta, did not extend down into the more southern regions. Ordination also showed that the composition of the fish faunas at all but one of the nine sites underwent pronounced and consistent cyclic, seasonal changes. This seasonal cyclicity at the different sites was attributable to sequential patterns of immigrations and emigrations by a number of fish species during the course of the year. These seasonal migrations involved, inter alia (1) movements of certain species from their nursery areas into these deeper waters, e.g.  S. bassensis and Scobinichthys granulatus; (2) migrations into and off the sandy areas of the inner continental shelf, e.g. Arnoglossus muelleri; (3) migrations to spawning areas, e.g. Sillago robusta; and (4) movements into areas where detached macrophytes accumulate in winter, e.g. Cnidoglanis macrocephalus and Apogon rueppellii. Received: 21 August 1998 / Accepted: 9 February 1999     

Metabolic transformations of acquired lucibufagins by firefly “femmes fatales”

Summary. Female Photuris fireflies sequester defensive steroidal pyrones (lucibufagins) from male fireflies of the genus Photinus. Lucibufagin analyses of Photuris females and Photinus ignitus males show that the lucibufagin mixtures of predator and prey differ in their composition. Analyses of whole body extracts showed that P. ignitus males contain a mixture of eight non-glycosylated lucibufagins, composed mostly of compounds with two oxygenated positions in the steroidal A-ring (C-3, C-5). After feeding on P. ignitus males, Photuris females contain six major lucibufagins. Three of these compounds are not present in the prey, including the novel lucibufagin glycoside 5β,11α-dihydroxy-12-oxo-3β-O-β-D-xylopyranosylbufalin, and two other lucibufagins with a trioxygenated A-ring (C-2, C-3, C-5). These results indicate that Photuris females transform the sequestered lucibufagins both by glycosylation and oxidation, which could affect the systemic transportability of these compounds due to an increase in their polarity. Received 18 February 1999; accepted 19 April 1999.     

Two distinct forms of the chitin-degrading enzyme N-acetyl-β-d-glucosaminidase in the Antarctic krill: specialists in digestion and moult

In the Antarctic krill Euphausia superba two forms of the chitinolytic enzyme N-acetyl-β-d-glucosaminidase (NAGase, EC 3.2.1.52) have been described, previously identified as NAGase B and NAGase C. Here, we demonstrate the organ-specific distribution and physiological relevance of both forms using a polyclonal antibody preparation which allows them to be distinguished immunologically. While NAGase B was localized in the integument and displayed a pattern of activity related to the moult cycle, the activity of NAGase C was independent of the moult cycle and was predominantly found in the gastrointestinal tract. Accordingly, NAGase B played a significant role in chitin degradation during the krill's moult, whereas NAGase C participated in the digestion of chitin-containing dietary components. Chromatographic elution profiles of isolated organs confirmed the immunological results by displaying characteristic organ-specific patterns in NAGase activity. The molecular characteristics of the moulting form, NAGase B, may further indicate a vesicular transport of moulting enzymes from the epidermis into the ecdysial space. Based on our results we develop a hypothesis explaining the concurrent processes of simultaneous chitin degradation and chitin synthesis occurring during moult. Received: 30 December 1998 / Accepted: 23 April 1999     

An analysis of the developmental differences in Borneo’s Southeastern and Northwestern Coastal Zones

In this research, Landsat TM images were used to extract the land use information for Borneo’s coastal zones (0 to 35 km) during three time periods (1990, 2000, and 2010). Then, in combination with the DEM and shoreline types, the developmental differences under the influences of the geographical boundaries of the Borneo coastal zones were examined from such perspectives as the spatial distribution characteristics of the land development, sea-land gradient variation characteristics, and comparative analyses of the different regions. The results indicated that the geographic boundaries, as well as the different countries, had great influences on the development of the coastal zones. The regions with the most active development in Borneo’s coastal zones were mainly located in the estuary bay, onshore with land on both sides, and the near-shore alluvial plain with DEM?<?50 m. The northwestern coast zone showed relatively flat topography, and was close to an important international waterway. Therefore, the development in this area was found to be more intense than in the southeastern coastal zone, and which was characterized by a long developmental history. On one side of the geographical boundary, the development was also found to differ among the different countries. Malaysia was observed to be dominated by industrial plantations. Meanwhile, the industrial plantations, agriculture, and aquaculture in Indonesia were significantly increased, which was closely linked with the national economic policies. Furthermore, the development of Borneo’s coastal zones has led to vegetation degradation towards a constant inland development, plantation enlargements, agricultural development, and cleared and burnt areas, which have resulted in the land use transfer of 22,000 km² of forest land within the 0 to 35 km range.     

Toxicity models of metal mixtures established on the basis of “additivity” and “interactions”

Observed effects of metal mixtures on animals and plants often differ from the estimates, which are commonly calculated by adding up the biological responses of individual metals. This difference from additivity is commonly referred to as being a consequence of specific interactions between metals. The science of how to quantify metal interactions and whether to include them in risk assessment models is in its infancy. This review summarizes the existing predictive tools for evaluating the combined toxicity of metals present in mixtures and indicates the advantages and disadvantages of each method. We intend to provide eco-toxicologists with background information on how to make good use of the tools and how to advance the methods for assessing toxicity of metal mixtures. It is concluded that statistically significant deviations from additivity are not necessarily biologically relevant. Incorporation of interactions between metals in a model does not on forehand mean that the model is more accurate than a model developed based on additivity only. It is recommended to first use a relatively simple method for effect prediction of uninvestigated metal mixtures. To improve the reliability of toxicity modeling for metal mixtures, further efforts should focus on balancing the relationship between the significance of statistics and the biological meaning, and unraveling the toxicity mechanisms of metals and their mixtures.

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Abundance and variability of microorganisms and transparent exopolymer particles across the ice–water interface of melting first-year sea ice in the Laptev Sea (Arctic)

The distribution and abundance of transparent exopolymer particles (TEP) was determined in and below pack ice of the Laptev Sea from July to September 1995. Samples were collected from the lowermost 10 cm of ice floes and at 10 cm below the ice–water interface. Abundance of bacteria, protists and TEP was determined, and the sea ice–water boundary layer was characterized using temperature, salinity and molecular viscous shear stress. TEP, with a distinct size distribution signal, were found in highest concentrations inside the sea ice, ranging from not detectable to 16 cm² l⁻¹ (median: 2.9 cm² l⁻¹). In the water, concentrations were one order of magnitude lower, ranged from below detection to 2.7 cm² l⁻¹ (median: 0.2 cm² l⁻¹) and decreased after the middle of August, whereas abundances of autotrophic flagellates (AF), diatoms, heterotrophic flagellates (HF) and ciliates increased. The abundance of TEP decreased with its size in all samples following a power law relationship. The relation of TEP to the microbial community differed between the sea ice and water, being positively correlated with bacteria and diatoms in the ice and negatively correlated with HF in the sea water. The presence of a pycnocline significantly influenced the abundance of organisms, diatom composition and TEP concentrations. Pennate diatoms dominated by Nitzschia frigida were most abundant inside the ice. Though bacteria have the potential to produce exopolymeric substances (EPS), the results of this study indicate that the majority of TEP at the ice–water interface in first-year Arctic summer pack ice are produced by diatoms. Received: 19 August 1999 / Accepted: 4 July 2000     

Investigation of the sources of aliphatic hydrocarbons in the mussel Mytilus edulis from north sea oil production platforms by Capillary glc and CGCMS†

Aliphatic hydrocarbons isolated from mussels collected over a 20 month period from three North Sea Forties field oil production platforms have been examined by capillary gas chromatography and gas chromatography‐mass spectrometry.

The biological hydrocarbons consist predominantly (300–700 μh g^‐1 lipid) of C₃₁ and C₃₃ n‐alkenes with 2, 3 and 4 double bonds probably derived from a dietary intake of Emiliania huxleyi, a microscopic coccolithophorid alga. In some of the mussels C₁₈, C₂₀ and C₂₂ n‐alkanes are present in unusually high abundance compared to their odd carbon number homologues. Possible reasons for this distribution are discussed.

The concentrations of fossil fuel hydrocarbons (mainly unresolved complex mixtures) in the mussels ranged from 330 to 5,298 μg g^‐1 of lipid. No significant increase in values was detected two months after the start of discharge of treated Forties production water from the Forties D platform.

To determine the pollutant hydrocarbon sources, the sterane and terpane distributions of hydrocarbon fractions isolated from the mussels and from a number of fossil fuels, including Forties crude oil, were examined by mass fragmentography. The steranes in the mussels from the Forties C and D platforms contained higher proportions of regular 14α(H), 17α(H) components than Forties oil in which the steranes were mainly 13ß(H), 17α(H) diasteranes. In addition, the terpane distributions of the mussel fractions differed markedly from that of Forties crude oil, particularly in the relative abundance of diterpanes to triterpanes, which was higher in the mussel fractions than in the oil, and in the absence of 17α(H), 18α(H), 21ß(H)‐18, 30‐bisnorhopane, a known component of Forties crude. Furthermore, the ratio of C₂₉ to C₃₀ triterpanes was greater than unity in the mussels but much less than unity in the Forties oil. These data preclude Forties or other similar North Sea crudes as the major source of pollutant aliphatic hydrocarbons in the mussels. The relative concentrations and distributions of steranes and triterpanes suggest that the most likely source of pollutants is a Middle Eastern based oil derived either from rig activities, or from background pollution in the North Sea. The low concentration of ≥ C₂₇ steranes in the more polluted mussels (e.g. Forties B; UCM > 200 ppm dry wt.) suggests that gas oils used on the platforms may be the major source of petrogenic hydrocarbons in these samples.     

Determination of the framework conditions and research–development needs for the dissemination of cleaner (sustainable) production applications in Turkey

SUMMARY

A cross-section survey of five states in Nigeria was derived through a stratified random sampling technique in 1999–2000 to assess the contribution of traditional knowledge in environmental conservation. Data were collected through a combination of questionnaire survey and discussions with stakeholders, with field assessment of herb gardens. Interviews were held with 1953 respondents composed of women, men and youths drawn from 20 settlements in 10 local government areas in the savanna and rainforest ecological zones.

The results derived from data analysis indicated that: (i) the rural communities were willing to accept integrated rural land-use practices that would conserve species and habitats, sustain the rural economy and the socio-cultural lifestyles of the people; (ii) the people at the local level had some basic knowledge about species and habitats, and have been conserving the forests with traditional land tenure rights; and (iii) environmental projects acceptable to the rural communities must be beneficial to the people, participatory, integrated with traditional knowledge and based on local needs.     

After the flames then what? exploring the linkages between wildfires and household food security in the northern Savannah of Ghana

ABSTRACT

This study deploys the livelihoods theory in political ecology together with integrated methods including remote sensing, participatory geographic information systems, interviews and focus group discussions to explore the intricate and dynamic linkages between perennial wildfires and household food insecurity among smallholder farmers in the northern savannah zone of Ghana. Our results suggest that wildfires are a major pathway to household food insecurity in two ways, as perceived by the farmers. First, in instances where wildfires destroy the major crops of smallholders, they are unable to find enough food for the lean season which results in transitory food insecurity. Secondly, in the long term, the environmental impacts of wildfires on the soil lead to lower yields and inadequate food to meet household nutritional needs. Farmers then deploy certain strategies such as early-harvesting and switching to early-maturing crops to avoid with the perennial fires and these strategies further reinforce other dimensions of food insecurity such as nutrition insecurity. Because livelihoods strategies such as crop farming, hunting, charcoal production and herding intersect in the northern savannah zone, we recommend policy options that will engage community members in finding common ways of using shared spaces sustainably. Government should also adopt policies that will encourage domestication of certain livelihood strategies such as cattle herding.     

Size,spatial, and bathymetrical distribution of the ascidian Halocynthia papillosa in Mediterranean coastal bottoms: benthic–pelagic coupling implications

Benthic suspension feeders are abundant in Mediterranean coastal environments, though most of them are threatened due to overexploitation, climate change impacts, and unregulated diving or fishing practices. Little is known about most of the coastal communities in terms of large-scale distribution and realistic benthic–pelagic coupling implications, which are keys to understand and manage those threatened ecosystems. The active suspension feeder Halocynthia papillosa (one of the most common ascidian species of the Mediterranean Sea) was selected as a model organism to help to understand the ecological role in benthic–pelagic coupling processes and its importance as a carbon sink (an essential ecosystem service). The spatial and bathymetrical distribution of this organism has been studied using remotely operated vehicle video transects. The species was distributed throughout the study area, with a maximum density of 4 specimens m^?2. The highest abundances and the biggest sizes were observed on the range of 20–50 m depth. The role as carbon and nitrogen sink of this suspension feeder has been quantified coupling distribution data with existing in situ studies of feeding and respiration. Along the 1.24 ha of the study area, H. papillosa yearly ingested 519.4 g C and 31.4 g N and retained 20.2 g C. As long as the physiological data are known, this new methodology could be very useful in assessing bentho–pelagic links and the capacity of being C and N sinks of a wide range of species. This new approach may be essential for the future management of benthic communities.     

The flow structure in the wake of a fractal fence and the absence of an “inertial regime”

Recent theoretical work has highlighted the importance of multi-scale forcing of the flow for altering the nature of turbulence energy transfer and dissipation. In particular, fractal types of forcing have been studied. This is potentially of real significance in environmental fluid mechanics where multi-scale forcing is perhaps more common than the excitation of a specific mode. In this paper we report the first results studying the detail of the wake structure behind fences in a boundary layer where, for a constant porosity, we vary the average spacing of the struts and also introduce fractal fences. As expected, to first order, and in the far-wake region, in particular, the response of the fences is governed by their porosity. However, we show that there are some significant differences in the detail of the turbulent structure between the fractal and non-fractal fences and that these override differences in porosity. In the near wake, the structure of the fence dominates porosity effects and a modified wake interaction length seems to have potential for collapsing the data. With regards to the intermittency of the velocities, the fractal fences behave more similarly to homogeneous, isotropic turbulence. In addition, there is a high amount of dissipation for the fractal fences over scales that, based on the energy spectrum, should be dominated by inter-scale transfers. This latter result is consistent with numerical simulations of flow forced at multiple scales and shows that what appears to be an “inertial regime” cannot be as production and dissipation are both high.     

Gametogenesis, spawning behavior, and early development in the “iceworm”Hesiocaeca methanicola (Polychaeta: Hesionidae) from methane hydrates in the Gulf of Mexico

Gametogenesis, spawning behavior, and early development have been described in the methane-seep polychaete Hesiocaecamethanicola Desbruyères and Toulmond, 1998, the first documentation of the reproductive biology of any cold-seep polychaete species. The worms were collected at the Green Canyon site in the Gulf of Mexico (27°44.77N; 91°13.33W) in August 1997. The gonads are situated in the neuropodia in association with an intraepithelial capillary system. Oogenesis is intraovarian, with oocytes remaining in the ovary until late vitellogenesis when they are released into the coelom for final growth and differentiation. Vitellogenesis involves both autosynthetic and heterosynthetic processes and, along with ovarian structure, suggests a relatively slow process of egg formation. Sperm differentiation is intertesticular until early spermiogenesis, when spermatid tetrads complete development in the coelom. The mature spermatozoon is an ect-aquasperm consisting of a tapering, digitate acrosome, a spherical nucleus, a midpiece containing from six to eight mitochondria and glycogen stores, and two centrioles associated with the flagellum. Both sexes undergo broadcast spawning, but the males spawn exclusively through the anus, a behavior previously unknown in polychaetes. Artificial fertilization resulted in spiral cleavage and development to the trochophore stage. Oocyte size-frequency analysis of adult females indicates a wide range of oocyte sizes and vitellogenic stages, suggesting that asynchronous gametogenesis occurs. Received: 13 July 2000 / Accepted: 29 November 2000