Alongshore transport of a toxic phytoplankton bloom in a buoyancy current: Alexandrium tamarense in the Gulf of Maine

We examined the mechanisms controlling blooms of the toxic dinoflagellate Alexandrium tamarense Lebour and the concomintant patterns of shellfish toxicity in the southwestern Gulf of Maine, USA. During a series of cruises from 1987 to 1989, hydrographic parameters were measured to elucidate the physical factors affecting the distribution and abundance of dinoflagellates along this coast. In 1988 and 1989 when toxicity was detected in the southern part of this region, A. tamarense cells were apparently transported into the area between Portsmouth, New Hampshire, and Cape Ann, Massachusetts, in a coastally trapped buoyant plume. This plume appears to have been formed by the outflow from the Androscoggin and Kennebec Rivers. Flow rates of these rivers, hydrographic sections, and satellite images led us to conclude that the plume persisted for about a month, and extended alongshore for several hundred kilometers. The distribution of cells followed the position of the plume as it was influenced by wind and topography. When winds were downwelling-favorable (to the southwest), cells were moved alongshore to the south, and were held to the coast; when winds were upwelling-favorable (to the northeast),the plume sometimes separated from the coast, advecting the cells offshore. In 1987 when no plume was present, A. tamarense cells were scarce, and no toxicity was recorded at the southern stations. The alongshore advection of toxic cells within a coastally trapped buoyant plume can explain the details of the temporal and spatial patterns of shellfish toxicity along the coast. We hypothesize that (1) the source of the A. tamarense populations is in the north, possibly associated with the Androscoggin and Kennebec estuaries, that (2) toxicity patterns follow a predictable relationship with river flow volume and timing of flow peaks and that (3) wind stresses directly influence the distribution of low salinity water and the dinoflagellate cells. Local, in situ growth of dinoflagellates can be an important factor initiating toxic dinoflagellate blooms. However, these data demonstrate the significant role of alongshore transport of established populations of A. tamarense in controlling the location and timing of paralytic shellfish poisoning (PSP) outbreaks in May and June along the southwestern coast of the Gulf of Maine.     

Acoustically detected year-round presence of right whales in an urbanized migration corridor

Species' conservation relies on understanding their seasonal habitats and migration routes. North Atlantic right whales (Eubalaena glacialis), listed as endangered under the U.S. Endangered Species Act, migrate from the southeastern U.S. coast to Cape Cod Bay, Massachusetts, a federally designated critical habitat, from February through May to feed. The whales then continue north across the Gulf of Maine to northern waters (e.g., Bay of Fundy). To enter Cape Cod Bay, right whales must traverse an area of dense shipping and fishing activity in Massachusetts Bay, where there are no mandatory regulations for the protection of right whales or management of their habitat. We used passive acoustic recordings of right whales collected in Massachusetts Bay from May 2007 through October 2010 to determine the annual spatial and temporal distribution of the whales and their calling activity. We detected right whales in the bay throughout the year, in contrast to results from visual surveys. Right whales were detected on at least 24% of days in each month, with the exception of June 2007, in which there were no detections. Averaged over all years, right whale calls were most abundant from February through May. During this period, calls were most frequent between 17:00 and 20:00 local time; no diel pattern was apparent in other months. The spatial distribution of the approximate locations of calling whales suggests they may use Massachusetts Bay as a conduit to Cape Cod Bay in the spring and as they move between the Gulf of Maine and waters to the south in September through December. Although it is unclear how dependent right whales are on the bay, the discovery of their widespread presence in Massachusetts Bay throughout the year suggests this region may need to be managed to reduce the probability of collisions with ships and entanglement in fishing gear.     

Biogeography of toxic dinoflagellates in the genusAlexandrium from the northeastern United States and Canada

Twenty-eight strains of toxic dinoflagellates in the genusAlexandrium from the northeastern United States and Canada were characterized on the basis of morphology, bioluminescence capacity, mating compatibility, and toxin composition. The distributions of these characters were evaluated in the context of regional patterns of paralytic shellfish poisoning (PSP) and coastal hydrography. Two morphospecies were identified-A. tamarense Lebour andA. fundyense Balech. The two are interspersed geographically though there are areas, such as the Gulf of Maine, where apparently onlyA. fundyense occurs. Southern waters (Cape Cod, Connecticut, and Long Island) have especially diverse populations. The two species are sexually compatible. Virtually all northern isolates are bioluminescent, whereas southern isolates include bioluminescent and non-bioluminescent strains. Cluster analyses, based on high performance liquid chromatography (HPLC) determinations of the suite of toxins produced by each isolate, revealed two and perhaps three distinct groups. One is comprised almost exclusively of northern strains, and the other of southern strains. A Cape Cod cluster may be separable from the southern group. These analyses explain a previously reported north-to-south trend of decreasing toxicity, as the northern isolates produce greater proportions of the more potent toxins than do southern forms. The overall perspective is that the biogeography of toxicAlexandrium spp. in the study region is not that of a single, widespread, homogeneous population, but rather is comprised of several sub-populations, each with its own physiological characteristics and history. Two scenarios are considered with respect to this regional biogeography. The first invokes recent and continuing dispersal of isolates to the south from a center of origin in the north, followed by recombination and strong selection. The second holds that the northern and southern populations diverged from a common ancestor (vicariance), but now represent localized populations with little mixing of genotypes. Neither hypothesis can be completely refuted by the data presented here, though the weight of the evidence favors the latter. The correct scenario may be a combination of both, with recent and continuous speading occuring within the Gulf of Maine and perphaps the Gulf of St. Laerence, but with endemic localized populations persisting without genetic exchange in most southern locations. These data also indicate that although morphological criteria separate toxicAlexandrim isolates from the study region into two morphospecies, these assignments do not coincide with clusterings based on toxin composition or allozyme electrophoresis, and they are further violated by mating results. A revision of taxon designations to the varietal level could be justified.     

Monitoring the movements of harbour porpoises (Phocoena phocoena) with satellite telemetry

The movements of nine harbour porpoises, Phocoena phocoena (L.), in the Bay of Fundy and Gulf of Maine were tracked using satellite telemetry. Transmitters were attached to the porpoises in August 1994 and 1995 after they were captured near Grand Manan Island at the mouth of the Bay of Fundy. Tracking periods ranged from 2 to 212 d (mean 50 ± 65 d). Porpoises exhibited a high degree of individual variation in movement patterns; five moved out of the Bay of Fundy into the Gulf of Maine. The porpoise with the longest tracking period moved extensively throughout the Gulf of Maine. These data suggest that seasonal movement patterns of individual harbour porpoises are discrete and are not temporally coordinated migrations. Porpoises that moved out of the Bay of Fundy into the Gulf of Maine did so following the 92 m isobath, which may represent an important movement corridor. The movement of porpoises from the Bay of Fundy into the Gulf of Maine supports the hypothesis that harbour porpoises from these two regions comprise a single population at risk of entanglement in both Canadian and US fisheries. Received: 4 February 1997 / Accepted: 25 August 1997     

Movements of lobsters (Homarus americanus) tagged in the Bay of Fundy,Canada

Of the 18 359 American lobsters (Homarus americanus) tagged and released at three locations in the Bay of Fundy during 1977–1980, 5 375 lobsters (29.3%) were recaptured within 6 yr of release. Mature lobsters (95-mm carapace length, CL) on average moved significantly greater distances than immature lobsters (<95-mm CL). Many (16.4%) of the mature lobsters were recaptured >92.6 km (50 nautical miles) from the release sites; the farthest distance moved was 798 km for a male at liberty for 3.5 yr. Examination of the direction of movements indicates some intermixing of lobsters within the Bay of Fundy, and throughout the Gulf of Maine and the adjoining Continental Shelf and slopes. Mature lobsters were recaptured in deeper waters during winter than during summer. The seasonal shallow-deep migration of mature lobsters in the Bay of Fundy is possibly associated with maximizing degree-days for molting, growth, gonadal development, and egg development. In some areas, mature females on average moved farther and seasonally earlier into deeper waters than mature males. Although the seasonal migration resulted in many lobsters returning to the original release area year after year, some lobsters made extensive long-distance movements away from the Bay of Fundy. Temperature-dependent, seasonal deep-shallow migration can explain both the local returns and the long distance migrations of mature lobsters in the Bay of Fundy and the Gulf of Maine. To achieve appropriate seasonal temperature regimes, mature lobsters have to move different distances and directions, depending on local topography.     

Toxic effects of a bloom of the diatom Rhizosolenia chunii on shellfish in Port Phillip Bay,Southeastern Australia

A bloom of the diatom Rhizosolenia chunii Karsten sensu Sundström occurred in Port Phillip Bay (38°S; 145°E), southeastern Australia, between late August and mid-October 1987. Coincident with this bloom, mussels (Mytilus edulis planulatus), scallops (Pecten alba) and flat oysters (Ostrea angasi) throughout the Bay developed a bitter taste that was so unpleasant and persistent that mussels became unmarketable for 7 mo. The bitter taste was concentrated in the digestive gland, which also showed extensive inflammation and degeneration. Digestive gland lesions were evident in mussels during September 1987, but became progressively more severe and, in the chronic end-stage were associated with high shellfish mortality 3 to 8 mo after the bloom had ceased. This study provides strong indirect evidence that the bitterness and the toxicity were caused by a bloom of R. chunii. R. chunii is the first species of diatom to be associated with shellfish mortality and, as the mortality occurred many months after the bloom had ceased, we suggest that the toxicity of certain algal blooms may be difficult to detect rather than uncommon.     

Horizontal movement of ocean sunfish, <Emphasis Type="Italic">Mola mola</Emphasis>, in the northwest Atlantic

Data were retrieved from 25 ocean sunfish (Mola mola) that were tagged with pop-up satellite archival tags in the southern Gulf of Maine (n = 6), off Nantucket Island (n = 17), and off the coast of Georgia (n = 2) between September 2005 and March 2008. Tags remained attached from 7 to 242 days, with a mean attachment period of (X ± SD) 107.2 ± 80.6 days. Ocean sunfish tagged in the Gulf of Maine and southern New England left those areas in the late summer and early autumn and moved south along the continental shelf break. Fish traveled as far south as the Bahamas and the Gulf of Mexico. By moving south, sunfish experienced similar mean sea surface temperatures throughout the tagging period. The maximum straight-line distance traveled by a tagged Mola mola was 2,520 km in 130 days. Two tagged ocean sunfish entered the Gulf of Mexico, one in the December and one in July. Movements were associated with frontal features created by the Gulf Stream and fish moved farther offshore in 2007 when the Gulf Stream was deflected from the shelf break.     

Interspecific variation in thermal denaturation of proteins in the congeneric musselsMytilus trossulus andM. galloprovincialis: evidence from the heat-shock response and protein ubiquitination

The genetic population structure of the precominant zooplankter, the copepod Calanus finmarchicus (Gunnerus), was examined to determine whether genetically distinct populations exist in the Gulf of Maine. C. finmarchicus was sampled in three regions of the Gulf of Maine (Great South Channel, spring 1989; northern Gulf of Maine, winter 1990; Great South Channel and Georges Bank, spring 1990). Copepods from seven locations in the Great South Channel, five in the northern Gulf of Maine and four on or near Georges Bank were assayed for allozyme variation and mitochondrial DNA variation of amplified 16S rRNA and cytochrome b genes. Restriction fragment length polymorphism (RFLP) analyses of both mitochondrial DNA genes revealed no variation among any of the individuals assayed. Analysis of five polymorphic allozyme loci revealed that genetic variation among the three geographic regions was low, and genetic identities were high between all locations (I>0.97). Most of the genetic variation was among locations regardless of region. Chi-square tests were used to examine genetic similarity between specific pairs of locations within and between regions. In the northern Gulf of Maine, genetic homogeneity occurred over larger spatial scales (hundreds of km) than in either the Great South Channel or Georges Bank (tens of km). Only copepods from the Bay of Fundy and Nova Scotian Shelf locations were genetically distinct from Wilkinson Basin copepods at two loci. Copepod populations from the northern locations may have been partially isolated or they may represent immigrant populations (e.g., from the Gulf of St. Lawrence). Several pairs of locations were genetically distinct at one or more loci in the two southern regions. Differences between locations in these regions may represent distinct populations advected into the areas at different times or from different sources (e.g., genetic variation may represent a mixture of genetically distinct northern and southern copepod populations). These results suggest extensive gene flow among populations of C. finmarchicus in the Gulf of Maine with some evidence of genetic population subdivision near the Gulf's northeastern and southern boundaries.     

Genetic differentiation ofMytilus edulis in eastern North America

There is significant differentiation at five polymorphic loci ofMytilus edulis among certain geographical areas of the Atlantic coast of North America. Non-metric multidimensional numerical methods distinguished three population groups: (I) populations south of Cape Cod, (II) populations throughout the Gulf of Maine, Gulf of St. Lawrence, areas of both southern and northern Newfoundland, and southern Hudson Bay, and (III) populations in southeastern Nova Scotia, northern Newfoundland and Hudson Strait, Quebec. Each subset consists of populations that exhibit characteristic multilocus, multiple allele genotypes. Populations in Groups II and III are spatially interdigitated among each other. At least one geographical area of mixing between genetically distinct populations occurs in northeastern Newfoundland. There is no evidence for interbreeding among genetically distinct individuals in mixed populations, suggesting the possibility that populations in the Atlantic Canadian Provinces and areas of northern Canada may consist of two distinct species.     

Succession and growth limitation of phytoplankton in the Gulf of Bothnia (Baltic Sea)

A one year field study of four stations in the Gulf of Bothnia during 1991 showed that the biomass was ca. two times, and primary productivity ca. four times, lower in the north (Bothnian Bay) than in the south (Bothnian Sea) during the summer. Nutrient addition experiments indicated phosphorus limitation of phytoplankton in the Bothanian Bay and the coastal areas in the northern Bothnian Sea, but nitrogen limitation in the open Bothanian Sea. A positive correlation between the phosphate concentration and the production/biomass ratio of phytoplankton was demonstrated, which partly explained the differences in the specific growth rate of the phytoplankton during the summer. Differences in photosynthetic active radiation between the stations also showed a covariation with the primary productivity. The relative importance of nutrient or light limitation for photosynthetic carbon fixation could not, however, the conclusively determined from this study. Marked differences in phytoplankton species composition from north to south were also observed. The number of dominating species was higher in the Bothnian Sea than in the Bothnian Bay. The distribution of some species could be explained as due to nutrient availability (e.g. Nodularia spumigena, Aphanizomenon sp.), while salinity probably limits the distribution of some limnic as well as marine species. The potentially toxic phytoplankton N. spumigena, Dinophysis acuminata and Chrysochromulina spp. were common in the Bothnian Sea but not in the Bothnian Bay. The pico- and nanoplankton biomass during late summer was higher than previously reported due to a revised carbon/volume ratio.     

Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

Recent Invasions of the Gulf of Maine: Three Contrasting Ecological Histories

Introduced species are common members of estuarine communities where their role as competitors and predators is of concern, This paper examines the invasion of Gulf of Maine benthic habitats by the ecologically similar alien invertebrates Styela clava, Botrylloides diegensis , and Membranipora membranacea .
Styela clava increased slowly in abundance at study sites in Beverly, Massachusetts and Portsmouth, New Hampshire. We found no evidence of competitive dominance by S. clava , even though it is the competitive dominant in similar habitats elsewhere. Botrylloides diegensis rapidly became a dominant species after its arrival in the Great Bay Estuary, but this dominance was short-lived. B. diegensis persists in the estuary as an early colonist of primary space and as an epibiont on secondary substrates in established communities. Membranipora membranacea became the dominant epiphyte on laminarian kelps within two years. Although M. membranacea overgrew the native epiphytes Obelia geniculata and Electra pilosa in the overwhelming majority of encounters these native species are more common on other algal hosts. Therefore, competitive dominance is not likely a factor in the successful invasion of the Gulf of Maine by M. membranacea .
These species provide evidence for opposing views of the role of competition in mediating community invasion. We show that ecological similarity among species is not an accurate criterion to predict either the mechanism of invasion or the means of persistence. In addition, these data indicate that biological invasions must be examined on broad spatial and temporal scales; short-term or narrowly focused studies can lead to incorrect conclusions.     

Paralytic shellfish poison in Spisula solidissima: Anatomical location and ozone detoxification

The surf clam Spisula solidissima, when exposed to a northern bloom of the toxic dinoflagellate Gonyaulax tamarensis, concentrates paralytic shellfish poison (PSP) and retains it for periods of over 1 year. The purpose of this investigation was to identify those tissues in which S. solidissima concentrates PSP and to examine the efficacy of ozone gas in PSP detoxification. Various levels of the toxin were found in every untreated tissue examined: the mantle and gill containing high concentrations (>1600 g/100 g tissue); the visceral mass, siphon, and foot showing less toxicity (1100 to 200 g/100 g tissue); and the adductor muscle yielding a level of toxin considered safe for human consumption (<60 g/100 g tissue). Toxic clams exposed to ozonized seawater for 2 weeks exhibited rapid detoxification in all tissues examined.This work was supported, in part, by a grant from the Massachusetts Science and Technology Foundation, Wakefield, Massachusetts 01880, USA.     

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     

Horizontal movements of Atlantic blue marlin (<Emphasis Type="Italic">Makaira nigricans</Emphasis>) in the Gulf of Mexico

We examined movements of Atlantic blue marlin (Makaira nigricans) from the Gulf of Mexico based upon 42 pop-up archival transmitting (PAT) tags. Long deployments (including one 334-day track) revealed diverse movement patterns within the Gulf of Mexico. North–south seasonal changes in blue marlin distribution showed strong correspondence with established seasonal patterns of sea surface temperature and primary production. During the summer spawning season, blue marlin utilized outer shelf and shelf edge waters in the northern Gulf of Mexico, and longer duration tracks indicated overwintering habitats in the Bay of Campeche. Egress occurred throughout the year and was difficult to determine because some tracks ended in the Straits of Florida (n = 3) while other tracks recorded movement through it or the Yucatan Channel (n = 4). Our results indicate that Atlantic blue marlin have a more restricted geographic range of habitats than previously recognized and that the Gulf of Mexico provides spatially dynamic suitable habitat that is utilized year-round through seasonal movements.     

Trace Metals in New England Marine Sediments: Casco Bay, Maine, in relation to Other Sites

The concentrations in surface sediments of the trace metals Cd, Cr, Cu, Ni, Pb and Zn have been determined at 32 stations in the Casco Bay region of the Gulf of Maine. The metals are not distributed homogeneously but exhibit elevated levels around the Portland waterfront and generally low levels at offshore and tidally scoured stations. Comparison of these results to those from both industrialized and non-industrialized sites throughout New England indicates that the sediments in parts of the Casco Bay region are affected by trace metals.     

Primary productivity in the Gulf of California

Primary productivity data from 6 cruises in the Gulf of California are summarized. A total of 31 stations with 133 samples was available. The Gulf of California represents a subtropical area with exceptionally high rates of primary productivity. This fact may be accounted for by temporary seasonal upwelling, a wind-mixed water column down to the shallow thermocline, especially in the northern part of the Gulf, and by tidal mixing, especially in the Ballenas Channel between Baja California and the island Angel de la Guardia. Rates of primary productivity in the Gulf of California are comparable to those in areas such as the Bay of Bengal, the upwelling areas off the west coast of Baja California, or North Africa. They are about 2 to 3 times greater than that in the open Atlantic or the open Pacific at similar latitudes.Contribution from the Scripps Institution of Oceanography, University of California, San Diego. This work was part of the Scripps Tuna Oceanography Research Program. It was supported by the Bureau of Commercial Fisheries under contract number 14-17-0007-963. Shiptime was provided from funds of the National Science Foundation.     

Effects of Trichodesmium spp. blooms on penaeid prawn larvae

The effects of blooms of the cyanobacterium Trichodesmium spp. on penaeid prawn larvae were examined using in situ and laboratory rearing experiments and plankton surveys in Albatross Bay, Gulf of Carpentaria, Australia. The in situ experiments demonstrated that, during a bloom of Trichodesmium spp., larvae of the prawn Penaeus merguiensis did not develop beyond the first protozoea stage, and survival was low compared with times when diatoms were dominant in the same study area. Laboratory experiments confirmed the in situ results. None of the prawn larvae fed Trichodesmium sp. in laboratory experiments developed beyond the first protozoeal stage. In contrast, 94% of prawn larvae fed the green flagellate Tetraselmis suecica successfully developed to the second protozoea stage. Electron microscopy of larvae gut-contents revealed that Trichodesmium spp. were ingested by larvae but were of no nutritional value, resulting in starvation. A 7 yr plankton survey, from 1985 to 1992, showed that minimum abundance of prawn larvae occurs during the annual summer blooms of Trichodesmium spp. and that maximum abundance of prawn larvae generally occurs just after the bloom. There was a negative correlation between the abundance of larvae and the abundance of Trichodesmium at individual sites, one offshore and one inshore, indicating that the blooms affect the survival of larvae. We conclude that variations in both timing and magnitude of Trichodesmium blooms are important determinants of prawn larvae abundance in Albatross Bay. Received: 28 April 1997 / Accepted: 2 April 1998     

Lethal effects of Northwest Atlantic Ocean isolates of the dinoflagellate, Scrippsiella trochoidea, on Eastern oyster (Crassostrea virginica) and Northern quahog (Mercenaria mercenaria) larvae

The thecate dinoflagellate Scrippsiella trochoidea is a cosmopolitan, bloom-forming alga that has been generally considered non-toxic. Here, we report that environmentally relevant cell densities (10⁴ cells mL⁻¹) of Scrippsiella trochoidea strains isolated from the Northwest Atlantic Ocean caused 100% mortality in Eastern oyster (Crassostrea virginica) larvae during 3-day exposures while parallel control larvae exhibited 100% survival. S. trochoidea also exhibited lethal effects on Northern quahog (Mercenaria mercenaria) larvae (70% mortality during 3-day exposure) but were non-toxic to juvenile fish (Cyprinodon variegates). The cultures of S. trochoidea were more lethal to Northern quahog larvae than ten other species of harmful algae, including the highly toxic species Cochlodinium polykrikoides. Scrippsiella trochoidea cultures within later stages of growth were more toxic than exponential growth stages to bivalve larvae, and the toxicity was dose dependent. Furthermore, toxicity was maintained in the cultures that were sonicated, boiled, and frozen as well as in resuspended residues of the culture but was significantly lower in cell-free culture media. Collectively, these results suggest that S. trochoidea causes mortality in bivalve larvae through a physicochemical rather than strictly chemical mechanism, such as clogging of larval feeding apparatuses by materials produced by S. trochoidea (e.g., lipids, extracellular polysaccharides, and/or cell debris) which accumulate as cells in culture or blooms age. This is the first report of the lethal effects of Scrippsiella trochoidea on shellfish larvae.     

Paralytic shellfish toxins sequestered by bivalves as a defense against siphon-nipping fish

Saxidomus giganteus (butter clams), are known to sequester diet-derived paralytic shellfish toxins (PST), highly potent neurotoxins, in their siphons. Captive staghorn sculpins (Leptocotus armatus), a marine fish species known to crop bivalve siphons, developed a significant aversion to siphons from toxic but not non-toxicS. giganteus following a single conditioning feeding of toxic siphon tissues. Control fish showed no aversive response to siphons from non-toxicS. giganteus during 11 feeding sessions over 56 d. Aversive and non-aversive behavior varied with the toxicity of the siphons, but not with the geographic origin of the clams. Both experimental and control fish ate freely and showed no aversion to siphons from toxic littleneck clams (Protothaca staminea). Littleneck clams, unlikeS. giganteus, retain PST in their visceral mass but not in their siphons. Both toxic and non-toxicS. giganteus extended their siphons significantly more often and higher above the sediment surface during dark hours, but toxicS. giganteus extended their siphons higher than non-toxic individuals. These results support the hypothesis that siphon-nipping by fish may have selected for the retention of PST in butter clam siphons as a chemical defense.