Latitudinal trends in size-dependence of bioluminescence in the midshipman fish Porichthys notatus

The bioluminescent fish Porichthys notatus (plainfin midshipman), has a discontinuous distribution along the Pacific coast of North America. The fish is present from Cape Mendocino southward to Baja California, Mexico, absent off the coast of Oregon, USA, and abundant, northward, in Puget Sound, Washington. Interestingly, the population in Puget Sound lacks the substrate (luciferin) necessary for the luminescence reaction and, despite possessing an otherwise fully functional photophore system, is nonluminescent. The California population of P. notatus is uniformly luminescent south of Monterey Bay, but 15% of the speciments tested from San Francisco Bay and the Gulf of the Farallons have been reported to be nonluminescent. Explanations for nonluminescent midshipman in both Puget Sound and the San Francisco Bay area have focussed on a dietary requirement for luciferin. To gain further insight into reasons for nonluminescence in the San Francisco Bay region, the distribution of bioluminescence in P. notatus was studied from Monterey Bay to Cape Mendocino during 1985. A complex pattern of bioluminescence was found, in which nonluminescent individuals reflected neither a local anomaly in the San Francisco Bay region nor a simple gradient of decreasing luminescence towards the northern end of the range of the California population. Instead, a distinct size-dependent component in luminescence capability of the fish was observed. Aspects of the life history of P. notatus and related factors which might influence the bioluminescence characteristics of this population are discussed.Please address all correspondence to Dr. F.I. Tsuji at the Osaka Bioscience Institute     

Disruptive counterillumination and its anti-predatory value in the plainfish midshipman Porichthys notatus

The plainfin midshipman Porichthys notatus Girard occurs in nearshore waters off the western coast of North America. We found that the ventrally directed photophores of juvenile laboratory-grown P. notatus function in counterillumination. Their bioluminescence matched downwelling light to obscure the silhouette as viewed from below. P. notatus did not obligatorily counterilluminate at all times in appropriate light. A hydrodynamic stimulus, in addition to appropriate downwelling light, was required to induce counterillumination. This is perhaps an adaptation to conserve its Vargula-type luciferin, which is obtained from its diet. The angular distribution of light produced by P. notatus approximated that typical of downwelling light in the ocean. There was a direct relationship between downwelling irradiance and the maximum intensity of P. notatus luminescence. Insufficient light was produced to replace the light blocked by the fish under lighting conditions typical of the natural environment. The luminescence was, however, effective in eliminating the silhouette of P. notatus as observed with an image-intensifying camera and by dark-adapted human observers. Consequently, the effectiveness of counterillumination probably originates in part from the disruptive pattern of the luminous photophores. P. notatus from the Puget Sound population, which lacks a luciferin source and is non-luminous, displayed counterillumination behavior when its luminescent system was activated by force-feeding the fish with dried Vargula hilgendorfi. Experiments using adult P. notatus as predators on a mixed population of non-luminous (Puget Sound) and luminous (Santa Barbara Channel) juvenile P. notatus revealed that under conditions of dim overhead illumination non-luminous P. notatus were preyed upon at twice the rate of luminous individuals. This is the first experimental evidence suggesting that counterillumination is effective as an anti-predatory strategy. Received: 25 August 1998 / Accepted: 25 March 1999     

Photopigment and spectral sensitivity in the bioluminescent fish Porichthys notatus

Digitonin extracts from retinas of the marine fish Porichthys notatus, from California and Puget Sound (USA), contain a single photopigment with maximum absorbance at 498 nm. In the presence of hydroxylamine the final product of bleaching absorbs maximally at 367 nm, indicating that the photopigment is based on retinal. Spectral sensitivity measurements performed on intact eyes and eye-cup preparations in darkadapted and selectively light-adapted states yield curves which suggest that the retina of P. notatus contains at least 3 different types of receptors. In the darkadapted state, the spectral sensitivity curve has a broad peak, with a maximum between 480 and 500 nm. In the blue and red light-adapted states, in addition to depressing the sensitivity, the shape of the sensitivity curve also changes. Under these conditions new peaks appear which suggest the presence of at least two different types of cones, with sensitivity maxima in the blue and the green regions of the spectrum, respectively. Comparison of the spectral sensitivity curves with the in vivo bioluminescence emission spectrum of the fish gives an especially good correspondence and suggests that the fish is able to perceive the bioluminescence of its own species.     

Population interconnectivity and implications for recovery of a species of concern, the Pacific hake of Georgia Basin

To aid the recovery of a species, understanding the extent to which populations are connected is useful for targeting conservation efforts. Pacific hake within waters of Puget Sound, Washington State, USA, and Georgia Strait, British Columbia, Canada are listed as a species of concern under the U.S. Endangered Species Act due to dramatic declines in the Puget Sound population. To assess the role of dispersal in the recovery of Pacific hake, we sought to quantify patterns of connectivity between populations in Puget Sound and Georgia Strait. Using natural chemical markers from otoliths of fish sampled from these two populations, we linked natal signatures of fish to signatures of individuals from known spawning grounds. Results indicated that 82 % of individuals collected from Puget Sound (n = 78) were estimated to have originated there, while 40 and 92 % of the individuals collected from two cohorts within Georgia Strait (n = 9 and 24, respectively) had originated from Puget Sound. A trend of “population abandonment” of fish from Puget Sound suggests that recovery of this Pacific hake population will depend on local management practices.     

Zinc and cadmium in the Pacific hake Merluccius productus off the western U.S. coast

Concentrations of zinc and cadmium in muscle tissue or whole Pacific hake, Merluccius productus (Ayres), increase with increasing total wet weight, especially for smaller fish. These trends appear to be related to the euphausiid diet of hake. Except for the isolated population in Puget Sound, no relationship between zinc or cadmium concentration and collection site is apparent.     

Larval development of the crabCancer magister in temperature regimes simulating outer-coast and inland-water habitats

Postlarval megalopae of the Dungeness crabCancer magister inhabiting offshore coastal waters along the west toast of North America are larger and settle earlier than do those occupying the inland waters of the Puget Sound basin (Washington, USA, and British Columbia, Canada). The Puget Sound habitat is characterized by low initial temperatures that steadily increase during the course of zoeal development, while offshore temperatures are more moderate and stable. Larvae were raised in the laboratory from hatching to megalopa in three temperature treatments: (1) constant 10 °C (CO), (2) a regime that temporally simulated temperatures found off the central California coast during the larval season (CA); and (3) a regime simulating temperatures found within the Puget Sound basin (PS). Zoeal duration was 44% longer in the PS treatment than in the other two, at least partially accounting for observed differentes between outer-toast and inland-water settlement times. Although differentes were measured in megalopal weights and carapace lengths among treatments, results do not explain differences in megalopal size observed between outer-toast and inland-water individuals. Survival to megalopa was highest in the PS treatment, with the difference due to significantly lower mortality than in CO and CA treatments during the terminal zoeal stage. Daily instantaneous mortality rates were lower in the PS than in CO or CA treatments, indicating Chat extended larval duration will not necessarily result in reduced settlement success.     

Colonization of the Southern Patagonia Ocean by Exotic Chinook Salmon

Abstract:  Anadromous salmonids have been particularly successful at establishing wild populations in southern Patagonia, in contrast to their limited success elsewhere outside their native ranges. The most recent such discovery is a spawning population of Chinook salmon in the Santa Cruz River, which flows into the Atlantic Ocean from Argentina. We used mitochondrial DNA analysis to discriminate between alternative potential sources of this population and were able to discard in situ introductions of fish imported directly from California in the early twentieth century. Our results showed that the fish most likely came from Puget Sound, Washington, imported into southern Chile for salmon-ranching experiments in the 1980s. This finding provides concrete evidence of colonization of Atlantic rivers from Pacific locations. The southern Pacific and Atlantic oceans provide a favorable marine environment for the success of invading salmon. In particular, the waters associated with fjords, southern channels, and the inshore portion of the Patagonian shelf provide a rather bounded, continuous waterway for exotic anadromous salmonids, rich in diverse forage species.     

Differences in morphology and habitat use among the native mussel <Emphasis Type="Italic">Mytilus trossulus</Emphasis>, the non-native <Emphasis Type="Italic">M. galloprovincialis,</Emphasis> and their hybrids in Puget Sound,Washington

Mytilus galloprovincialis (Mg), the Mediterranean blue mussel, is sympatric with the native M. trossulus (Mt) throughout much of the north Pacific, likely as the result of human introduction. We investigated the distribution of the two species and their hybrids (Mgt) in Puget Sound, Washington, to determine whether differences occur in habitat preference between the two species and hybrids. In addition, we investigated whether there were consistent size and shape differences between the native and introduced mussels and hybrids. Measurements of over 6,000 mussels from 30 sites, of which 1,460 were genotyped for a species-specific genetic marker, revealed that Mg and Mgt can be found throughout Puget Sound. Mg and Mgt were larger and exhibited a greater height:length ratio than Mt. Frequencies of Mg and Mgt were higher in subtidal habitats, such as docks, than on intertidal rocks, walls or pilings. Within intertidal habitats, Mg and Mgt were more frequent than Mt in the lower reaches of the intertidal. At slightly more than half the sites the frequency of the three genotypes accorded with random mating expectations suggesting no consistent barriers to gene flow between species. The standardized random sampling methods and simple morphometric identification techniques described here can be used to test whether the frequency of invasive mussels changes over time and space in Puget Sound.     

Dinoflagellate luminescence increases susceptibility of zooplankton to teleost predation

The burglar alarm theory of bioluminescence was investigated by determining predation rates of a nocturnal teleost predator,Porichthys notatus, on nonluminescent kelp mysids illuminated by dinoflagellate flashes, between the fall and spring of 1989/1990. Mysids (Holmesimysis costata) were placed in aquaria containing varying concentrations (0 to 40 cells/ml) of the dinoflagellatePyrocystis fusiformis and a single midshipman fish. Controls usedP. fusiformis during their luminescence-inhibited day phase. Mysid swimming movements readily stimulated dinoflagellate luminescence. Flashes and prey strikes were observed simultaneously by image-intensifying and infrared video cameras on a splitscreen monitor. Predation rates increased at dinoflagellate concentrations of 3 to 15 cells/ml and decreased below controls at levels>20 cells/ml. Videotape analysis showed that at low concentrations (2 to 5 cells/ml), strike success rates exceeded 75% if prey were previously illuminated by a flash, but dropped below 50% at higher cell densities. Increased predation was attributed to luminescence revealing prey position. The decrease at higher concentrations was considered to be due to greater flash frequency providing a more diffuse and confusing target. The study demonstrates the effects of secondary luminescence on zooplankton predation at normally encountered dinoflagellate concentrations.     

Evidence for multiple recruitment-cohorts of Puget Sound Dungeness crab,Cancer magister

While sampling intertidally in Puget Sound, Washington, USA, for juvenile Dungeness crab (Cancer magister) in 1984, we found evidence of two distinct cohorts of the same year-class based on sizes of first-instar juveniles (J1) and the spatial/temporal patterns of settlement. In 1988, three distinct cohorts were observed to settle in Puget Sound and its approaches. Settlement of one cohort occurred during May in the Strait of Juan de Fuca and in those areas of Puget Sound closest to the Strait. J1 individuals of this cohort were large (x=7.4 mm carapace width, CW) and comparable in both size and timing of settlement to populations along the Washington coast (e.g. Grays Harbor and Willapa Bay). Initial settlement density of the May cohort was as high as 215 crabs/m² in intertidal eelgrass beds along the Strait of Juan de Fuca and decreased to <2 crabs/m² within Puget Sound and the Strait of Georgia. A second cohort apparently originated in Hood Canal (a deep inland fjord), its size upon settlement in June was significantly smaller (J1 x= 5.3 mm CW) than the May cohort, and it was limited to Hood Canal and areas of Puget Sound close to the mouth of Hood Canal. A third cohort, which settled in late July and August, was the smallest of the three cohorts (J1 x= 4.8 mm CW), and was widely distributed around Puget Sound from Seattle in the south to the USA/Canadian border in the north. We hypothesize that most juvenile recruitment in Hood Canal and Puget Sound originates from parental stocks endemic to their respective basins (Hood Canal and Puget Sound cohorts), but that, on occasion, oceanographic conditions allow substantial influx of Pacific Ocean Dungeness crab larvae (oceanic cohort) through the Strait of Juan de Fuca into Puget Sound. Tracking of spatial/temporal settlement patterns and comparison of J1 sizes proved useful for estimating the probable sources and dispersion of Dungeness crab larvae. Differences in size and time of settlement between various larval cohorts of C. magister may prove useful as biomarkers for tracing circulation patterns within and between inland waters of Washington and the Pacific Ocean. Causes of smaller size and later settlement of the Puget Sound cohort relative to oceanic conspecifics of the same year-class are discussed.Contribution No. 856, School of Fisheries, University of Washington, Seattle     

Disjunct Sea of Cortez–Pacific Ocean Gillichthys mirabilis populations and the evolutionary origin of their Sea of Cortez endemic relative, Gillichthys seta

The shortjaw mudsucker, Gillichthys seta, an intertidal goby endemic to the Sea of Cortez, has been proposed to be the paedomorphic derivative of the longjaw mudsucker, Gillichthys mirabilis. G. mirabilis is a disjunct species, with populations found along the Pacific coast of central California to central Baja California, and with isolated populations found in the northern Sea of Cortez. Previous studies have suggested that the endemic paedomorph form speciated in sympatry with the Sea of Cortez population of G. mirabilis. Alternatively, this speciation event could have occurred before the separation of G. mirabilis populations into two disjunct entities. To test these alternative hypotheses, we collected adult individuals from both species throughout their ranges from December 1997 to November 1998. We amplified and sequenced 142 partial [527 base pairs (bp)] mitochondrial cytochrome b regions and 18 nuclear creatine kinase introns (140 bp). We found that Pacific populations of G. mirabilis separated into two distinct clades, possibly reflecting a phylogeographic break found in other fish species along the Baja California coast at Punta Eugenia. These two Pacific populations were well separated from Sea of Cortez populations. Furthermore, our results indicate that the split between Sea of Cortez and Pacific populations of G. mirabilis occurred well after the speciation event that separated G. mirabilis from its paedomorphic counterpart, G. seta. Received: 30 March 2000 / Accepted: 4 September 2000     

Apparent predation on ichthyoplankton by zooplankton and fishes in nearshore waters of southern California

Plankton collected at discrete depths in Santa Monica Bay, California, USA, during January 1982 were examined for fish eggs and larvae that had been attacked or consumed by zooplankton. The bongo net remained open for only 3 min and samples were preserved within 5 min of capture. Juvenile and adult fishes that had been captured by otter trawl and preserved within 20 min of capture were examined for ingested fish eggs and larvae. Three copepods (Corycaeus anglicus, Labidocera trispinosa, and Tortanus discaudatus), one euphausid larva (Nyctiphanes simplex), one amphipod (Monoculoides sp.), and an unidentified decapod larva were found attached to fish larvae in the preserved plankton samples (attachment to 23% of the fish larvae was observed in one sample). Overall, about 5% of the white croaker (Genyonemus lineatus) larvae and 2% of the northern anchovy (Engraulis mordax) larvae had attached zooplankton predators. Most fish larvae with attached zooplankton predators were small. We found no indication of zooplankton predation on fish eggs. Few fish eggs and larvae were found in the digestive tracts of juvenile or adult fishes, and the ingested fish larvae were relatively large. The discussion considers apparent preyspecificity of the zooplankton predators as well as potential biases that may be associated with preserved samples collected by nets.     

Correlation of bioluminescence emissions with ventral photophores in the mesopelagic squidAbralia veranyi (Cephalopoda: Enoploteuthidae)

Specimens of the squidAbralia veranyi were collected off the Bahamas in 1989. The bioluminescence of the ventral photophores was recorded on videotape at 11 to 12°C and 24°C, in conjunction with measurements of the spectral emission at the same temperatures. At 11 °C, the spectrum was unimodal, peaking at 490 nm with a narrow bandwidth. At the higher temperature a shoulder at about 440 nm appeared in the emission spectrum. The short, bright flashes from the subocular photophores had a broader bandwidth and a shorter wavelength emission maximum (475 nm) than that of the ventral photophores. Video recordings at the two temperatures showed no changes in either the identities of the luminescing photophores or their relative intensities. One structurally distinct type of photophore was not illuminated. We conclude that the observed spectral changes were produced within one group of photophores, and that recruitment of the unilluminated photophores would produce an additional spectral component, previously reported from another species ofAbralia (A. trigonura).     

Recent introduction of an Asian estuarine copepod,Pseudodiaptomus marinus (Copepoda: Calanoida), into southern California embayments

Past studies made between 1938 and the mid-1970's of coastal embayments between northern Baja California and Oregon (USA) established the presence of only one species of the calanoid copepod genus Pseudodiaptomus, P. euryhalinus. This species is endemic to warm temperate latitudes of western North America. We now report the presence of breeding populations of the Asian calanoid copepod P. marinus in two Southern California embayments, Mission Bay and Agua Hedionda Lagoon. The Mission Bay population was monitored monthly from December 1986 to June 1987 and showed evidence of continuous breeding and growth. The regional, endemic species, P. euryhalinus, was not observed in either embayment. Mechanisms that might account for the apparent recent introductions of P. marinus into Southern California include dispersion by ocean circulations and transport by transoceanic ships. In the present case, the explanation is that the introduction is most likely a direct consequence of aquaculture projects. Remaining to be confirmed by systematic sampling is the apparent disappearance of P. euryhalinus from embayments dominated by P. marinus.     

The dynamics of phytoplankton blooms in puget sound a fjord in the Northwestern United States

This paper describes a quantitative investigation of relationships between the growth of phytoplankton, and climatic and hydrodynamci conditions in temperate fjords with marked tides, as exemplified by Puget Sound, Washington (USA). Algal growth in the open waters of the central basin of the Sound is dominated by a number of intense blooms beginning in late April or May and recurring throughout the summer. Rarely, and only briefly, does nitrate become exhausted. The phytoplankton production rate in the central basin of Puget Sound is about 465 g C m^-2 year^-1. During the springs of 1966 and 1967, oceanographic measurements were carried out at a mid-channel station with sufficient frequency to allow investigation of physical and biological processes with time scales of the order of a day. The principal investigative tool is a numerical model in which the hydrodynamical conditions are represented by an approximate analysis of the gravitational convection mode of circulation. Algal concentration is represented as a continous function of space and time in the model which ascribes changes in phytoplankton density to variations in photosynthetic and respiratory activity, algal sinking, grazing by herbivores, and to mixing and advection. Computations adequately reproduce the principal features of phytoplankton concentrations observed during 75 days and 35 days in the springs of 1966 and 1967, respectively. Numerical experiments assess the relative importance of various processes which govern the level of primary production in Puget Sound. It is concluded that phytoplankton growth is limited by a combination of factors, including vertical advection and turbulence, modulation of underwater light intensity by self-shading and inorganic particulates, sinking of algal cells, and occasional rapid horizontal advection of the population from the area by sustained winds. The high primary productivity of the Sound is due to intensive upward transport of nitrate by the estuarine mechanism. These results should be generally applicable to other temperate fjords because of the largely conventional choice of the biological functions.     

Distribution of dimorphic males of three species of Nematoscelis (Euphausiacea)

Samples collected from the eastern Atlantic Ocean between 40° and 4° N with rectangular and ring nets contained male specimens of Nematoscelis atlantica, N. microps, and N. tenella (Euphausiacea) with saddle-shaped thickenings of chitin on the dorsal surface of some abdominal segments, and with enlarged photophores on the abdominal segments immediately posterior to those with thickenings. Two forms of each species are distinguished by the number and the position of chitinous thickenings and enlarged photophores. Except in a zone off W. Africa (approximately between 22° and 16° N) each form of N. atlantica and N. microps occurs as a distinct population. These populations appear to be characteristic of either the North or South Atlantic Central water, and the overlap zone corresponds approximately with the physical boundary between these water masses and with a faunal transition zone found in this region by other workers. For N. tenella it appears that only one form occurs as a separate population south of 18° N in South Atlantic Central water. Between 28° and 18° N the two forms co-exist.     

Assessing Changes in Amphibian Population Dynamics Following Experimental Manipulations of Introduced Fish

Abstract: Sport‐fish introductions are now recognized as an important cause of amphibian decline, but few researchers have quantified the demographic responses of amphibians to current options in fisheries management designed to minimize effects on sensitive amphibians. Demographic analyses with mark–recapture data allow researchers to assess the relative importance of survival, local recruitment, and migration to changes in population densities. I conducted a 4‐year, replicated whole‐lake experiment in the Klamath Mountains of northern California (U.S.A.) to quantify changes in population density, survival, population growth rate, and recruitment of the Cascades frog (Rana cascadae) in response to manipulations of non‐native fish populations. I compared responses of the frogs in lakes where fish were removed, in lakes in their naturally fish‐free state, and in lakes where fish remained that were either stocked annually or no longer being stocked. Within 3 years of fish removals from 3 lakes, frog densities increased by a factor of 13.6. The survival of young adult frogs increased from 59% to 94%, and realized population growth and recruitment rates at the fish‐removal lakes were more than twice as high as the rates for fish‐free reference lakes and lakes that contained fish. Population growth in the fish‐removal lakes was likely due to better on‐site recruitment of frogs to later life stages rather than increased immigration. The effects on R. cascadae of suspending stocking were ambiguous and suggested no direct benefit to amphibians. With amphibians declining worldwide, these results show that active restoration can slow or reverse the decline of species affected by fish stocking within a short time frame.     

Red bioluminescence in fishes: on the suborbital photophores of Malacosteus,Pachystomias and Aristostomias

Many deep-sea animals produce blue bioluminescence, but species of three genera of midwater dragonfishes also produce red light. In addition to numerous small body photophores, species of these genera (Malacosteus, Pachystomias and Aristostomias) have large suborbital photophores that emit red light and postorbital ones that emit blue light. Pachystomias microdon additionally has a red-emitting preorbital photophore. Fluorescence measurements from the intact suborbital photophores, and from their exposed cores, confirm the previous hypothesis that the red light emitted by Malacosteus is spectrally altered by a superficial shortwave cutoff brown filter. No such filter is present in the other two genera. Studies of the anatomy of the photophores show that the suborbital photophore of Malacosteus is similar in general organisation to other photophores, including the postorbital photophore. The red-emitting photophores of Pachystomias and Aristostomias, however, have a unique organisation, in which the bulk of the photophore comprises a gland that lies outside the thick reflector. Strands of tissue run from the gland into the photogenic core of the photophore through numerous pores in the reflector. The production and use of red light by these fishes is discussed in the context of these results.Electronic Supplementary Material Supplementary material is available for this article at     

Public Willingness to Pay for Recovering and Downlisting Threatened and Endangered Marine Species

Abstract: Nonmarket valuation research has produced economic value estimates for a variety of threatened, endangered, and rare species around the world. Although over 40 value estimates exist, it is often difficult to compare values from different studies due to variations in study design, implementation, and modeling specifications. We conducted a stated‐preference choice experiment to estimate the value of recovering or downlisting 8 threatened and endangered marine species in the United States: loggerhead sea turtle (Caretta caretta), leatherback sea turtle (Dermochelys coriacea), North Atlantic right whale (Eubalaena glacialis), North Pacific right whale (Eubalaena japonica), upper Willamette River Chinook salmon (Oncorhynchus tshawytscha), Puget Sound Chinook salmon (Oncorhynchus tshawytscha), Hawaiian monk seals (Monachus schauinslandi), and smalltooth sawfish (Pristis pectinata). In May 2009, we surveyed a random sample of U.S. households. We collected data from 8476 households and estimated willingness to pay for recovering and downlisting the 8 species from these data. Respondents were willing to pay for recovering and downlisting threatened and endangered marine taxa. Willingness‐to‐pay values ranged from $40/household for recovering Puget Sound Chinook salmon to $73/household for recovering the North Pacific right whale. Statistical comparisons among willingness‐to‐pay values suggest that some taxa are more economically valuable than others, which suggests that the U.S. public's willingness to pay for recovery may vary by species.     

Population genetics and systematics of theLeptasterias hexactis (Echinodermata: Asteroidea) species complex

Morphological characters and 14 enzyme loci were examined for 1 040 sea stars, currently recognized as forms ofLeptasterias hexactis, from Lynn Canal, Alaska, and Puget Sound, Washington, USA, between March 1988 and April 1989. Three morphologically and two genetically distinctLeptasterias forms were identified. The most common form found at both localities wasL. epichlora (Brandt)sensu Verrill.L. hexactis (Stimpson)sensu Verrill co-occurred withL. epichlora at all study sites and apparently hybridizes extensively withL. epichlora in the Puget Sound region, but rarely, if at all, in Alaska. The presumptive product of this hybridization morphologically resembledL. aequalis (Stimpson)sensu Fisher, and was conspicuously absent from Alaskan samples. Considerable genetic distance existed betweenL. epichlora andL. hexactis (Nei'sD=0.19±0.01) and moderate genetic differentiation occurred between populations of each species from Alaska and Washington (Weir and Cockerham'sF _RT =0.29±0.04 forL. epichlora and 0.21±0.15 forL. hexactis). A significant (p<0.05) deficiency in the proportion of heterozygous individuals was found compared to Hardy-Weinberg expectations (Wright's fixation index,F _ID=0.12±0.04 and 0.31±0.08 forL. epichlora andL. hexactis, respectively). However, mean observed heterozygosity for each species (0.09±0.03, 0.14±0.04 and 0.14±0.04 forL. epichlora, L. hexactis andL. aequalis, respectively) fell within the range of reported values for other asteroid species (ca. 0.04 to 0.37). The results of this study indicate that considerable genetic integrity is maintained betweenL. epichlora andL. hexactis, which warrants their recognition as distinct species despite their apparent hybridization in the Puget Sound region.