Genetic differentiation and reproductive incompatibility among Baja California populations of the copepod Tigriopus californicus

Populations of the harpacticoid copepod Tigriopus californicus were sampled from five sites from San Diego, California, to Playa Altamira, Baja California, Mexico. Allozyme analyses revealed that all the study populations are sharply differentiated genetically. At the extreme, two populations, Punta Baja and Playa Altamira, have no alleles in common at the seven allozyme loci studied. All pairwise interpopulation crosses successfully produced F₂ hybrids except those involving the Playa Altamira population. All crosses using Playa Altamira females failed to produce F₁ hybrids, while Playa Altamira males successfully produced F₁ progeny with females from all other sites. These F₁ offspring, however, were completely sterile (with San Diego and Punta Banda females) or only occasionally produced F₂ offspring (with Punta Morro and Punta Baja females). These results suggest that allopatric differentiation among Baja populations has resulted in exceptionally high levels of genetic divergence and nearly complete reproductive isolation of the Playa Altamira population, which should now be recognized at the semispecies (or perhaps sibling species) level.     

Cannibalistic behaviour in the harpacticoid copepod Tigriopus fulvus

The results reported here demonstrate the existence of cannibalistic behaviour in laboratory-reared Tigriopus fulvus (Fischer, 1860) females towards first-stage non-related nauplii; mothers, sparing their own offspring, do not share this behaviour. The phenomenon is related to the probability of nauplius/female encounters. It is suggested that mother/offspring recognition is mediated by a chemical compound.     

Identification of sex, age and species-specific proteins on the surface of the harpacticoid copepod Tigriopus japonicus

 Behavioral experiments have shown that male copepods of the species Tigriopus japonicus (Nori) can distinguish species, sex, and developmental stage of potential mates using contact chemoreception. Lectin-binding patterns on the body surface of females have indicated that surface-bound glycoproteins may be important signals in mate choice. In the present study, the proteolytic enzyme trypsin was used to cleave surface proteins from females, reducing their attractiveness to males. The protein fragments released were used to make monoclonal antibodies. Three levels of screening were used to identify monoclonal antibodies that recognized proteins involved in mate recognition. One monoclonal antibody bound to the terminal urosome and lateral prosome of CV females, and its binding significantly decreased female attractiveness to males. Western blotting showed that this antibody bound the trypsin-cleaved fragment and proteins of homogenized CV females and virgin adult females, but did not bind proteins of homogenized males, CIII females, or females of T. californicus or T. fulvus. This antibody recognized proteins on the surface of females that may enable males to discriminate conspecifics, sex, and age. It is likely that this molecule has a central role in the evolution of reproductive isolation in this group. Received: 22 July 1999 / Accepted: 21 March 2000     

Assessment of individual and combined toxicities of three heavy metals (Cu,Cd and Hg) by using Tigriopus fulvus

The toxicities of copper, cadmium, and mercury ions and their binary and ternary mixtures were studied using the copepods Tigriopus fulvus. The LC50 values measured after 48 h exposure to single metal solutions revealed a toxicities rank as follows: CdT. fulvus. The combined effect of the metals was found to be antagonistic for Cu?Cd, Hg?Cd, and Cu?Cd?Hg, additive for Hg?Cu.     

Survival and sex ratios of the intertidal copepod,Tigriopus californicus,following ultraviolet-B (290–320 nm) radiation exposure

The marine harpacticoid copepod, Tigriopus californicus (Baker), is a successful colonizer of supralittoral splash pools from Torch Bay, Alaska, to Baja California, Mexico. As these pools are subject to abundant amounts of direct solar radiation, it was of interest to determine the sensitivity of T. californicus to ultraviolet-B (UV-B, 290 to 320 nm) radiation. During 1980, copepods were raised under diurnal conditions in the laboratory and fed a mixture of unicellular algae and bacteria. Larval and adult stages were irradiated for 0 to 26 h on a rotating turntable under enhanced ultraviolet radiation. Following irradiation, the copepods were maintained in the culture area and checked daily for survival. All life stages of T. californicus tolerated enhanced UV-B radiation irrespective of exposure period; in contrast, survival of an irradiated planktonic copepod, Acartia clausii (Giesbrecht), was significantly lower. Additionally, there appeared to be asex ratio shift in adults that developed from the youngest naupliar stages (N₁/N₂) of T. californicus that were irradiated. Extraction and tentative identification of the pigment(s) responsible for the bright orange color of the carapace of T. californicus were performed. These pigments may function to absorb UV radiation and protect internal structures from photochemically induced damage.     

Mate choice in the face of both inbreeding and outbreeding depression in the intertidal copepod Tigriopus californicus

 In species vulnerable to both inbreeding and outbreeding depression, individuals might be expected to choose mates at intermediate levels of genetic relatedness. Previous work on the intertidal copepod Tigriopus californicus has repeatedly shown that crosses between populations result in either no effect or hybrid vigor in the first generation, and hybrid breakdown in the second generation. Previous work also shows that mating between full siblings results in inbreeding depression. The present study again found inbreeding depression, with full sibling mating causing significant fitness declines in two of the three populations assayed. In the mate choice assays, a single female was combined with two males. Despite the costs of both inbreeding and outbreeding, mate choice showed clear inbreeding avoidance but no clear pattern of outbreeding avoidance. This lack of outbreeding avoidance may be attributed either to the temporary increase in fitness in the F₁ generation or to the absence of selection for premating isolation in wholly allopatric populations with infrequent migration. If this inability to avoid unwise matings is common to other taxa, it may contribute to the problem of outbreeding depression when allopatric populations are mixed together. Received: 18 May 1999 / Accepted: 25 January 2000     

Mitochondrial DNA lineages in the European populations of mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.)

Marine mussels (Mytilus spp.) belong to a group of benthic species crucial to coastal ecosystems in Europe and are important for the cultivation industry. In the present study, the nuclear adhesive protein marker (Me15/16) was used for identification of Mytilus species in coastal areas, on a large geographic scale in Europe. Pure M. edulis populations were found in the White Sea and Iceland. M. edulis, M. trossulus and their hybrids were found in the Baltic Sea and the North Sea (Oosterschelde, The Netherlands). M. galloprovincialis, M. edulis and their hybrids occurred in Ireland. M. galloprovincialis populations were observed in the Sea of Azov (Black Sea), the Mediterranean and Portugal. The mitochondrial (mt) DNA coding-region ND2-COIII was studied by PCR (polymerase chain reaction) and RFLP (restriction fragment length polymorphism) assay methods. The mtDNA control region was studied by PCR. Substantial differentiation in the frequency of female haplotypes among the studied populations in Europe was observed. Despite isolation between the Mytilus taxa on a macro-geographic scale, considerable mitochondrial gene flow occurred between populations, with introgression in hybrid zones on a more local geographic scale in Europe. MtDNA of the Atlantic Iberian (Portugal) population of M. galloprovincialis was more similar to mtDNA in populations of M. galloprovincialis and M. edulis from the Atlantic coasts of the Ireland and M. edulis from the North Sea, than to M. galloprovincialis from the Mediterranean. Lower polymorphism of mtDNA in populations of the Baltic and Azov Sea mussels in comparison with other European populations was observed and can be explained by the recent history of both seas after the Pleistocene glaciation. In the M. galloprovincialis population from the Azov Sea, the presence of the male-inherited (M) genome was demonstrated for the first time by sequencing the control region and was observed at high frequency. Possible influence of mussel culture on geographic distribution of the Mytilus taxa in Europe is discussed.     

Phylogeographic patterns of the mysid Mesopodopsis slabberi (Crustacea, Mysida) in Western Europe: evidence for high molecular diversity and cryptic speciation

The phylogeographic patterns among populations of Mesopodopsis slabberi (Crustacea, Mysida), an ecological important mysid species of marine and estuarine habitats, were analysed by means of DNA sequencing of a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) and the 16S ribosomal RNA genes. Samples of M. slabberi collected from five Atlantic and two Western Mediterranean populations were investigated. Very high levels of within-population molecular diversity were observed in all samples (mean h=0.807 and π=0.0083), with exception of the Mediterranean Ebro population which contained only one haplotype. Differentiation among populations was high, and a clear phylogeographic break was observed between the Atlantic and Mediterranean populations. Moreover, a strong differentiation was detected between both populations in the Western Mediterranean Sea (Alicante and Ebro delta), while two divergent lineages occurred in sympatry within the Atlantic Mondego estuary. The high congruence between both the COI and 16S rRNA sequence data, the reciprocal monophyly of the different mitochondrial clades and the levels of nucleotide divergence between them suggest the presence of a complex of cryptic species within M. slabberi. Estimations of divergence time between the different mitochondrial lineages indicate that a split occurred during the late Miocene/early Pliocene. Such a divergence could be concordant with vicariant events during sea-level drops within the Mediterranean region at that time. However, within the Mediterranean Sea, the potential of divergence through ecological diversification cannot be ruled out.     

Mitochondrial cytochrome b variation in sleeper sharks (Squaliformes: Somniosidae)

Sleeper sharks are a poorly studied group of deep-sea sharks. The subgenus, Somniosus, contains three morphologically similar species: S. microcephalus found in the Arctic and North Atlantic; S. pacificus in the North Pacific; and S. antarcticus in the Southern Ocean. These sharks have been reported mainly in temperate to polar waters and occasionally in subtropical locations. They have not been recorded in tropical waters. This study investigates the relationships among the accepted species of Somniosus through analysis of mitochondrial cytochrome b sequence variation. Seventy-five samples were examined from four sampling locations in the North Pacific, Southern Ocean and North Atlantic. Twenty-one haplotypes were found. A minimum spanning parsimony network separated these haplotypes into two divergent clades, an S. microcephalus and an S. pacificus/antarcticus clade, strongly supporting the distinction of S. microcephalus as a separate species from the Pacific sleeper shark species. Analysis of genetic structure within the S. pacificus/antarcticus clade (analysis of molecular variance, allele frequency comparisons, and a nested clade analysis) showed limited or no differences amongst three populations. Further examination of genetic variation at more variable mtDNA and nuclear markers is needed to examine the species status of S. pacificus and S. antarcticus.     

Geographic and reproductive isolation in the marine Harpacticoid Copepod Tisbe

Within the genus Tisbe (Copepoda: Harpacticoida), there is a wide range of reproductive isolation. In the European area, in addition to species where (between geographic populations) the reproductive barrier is practically absent, there are others which show a varying degree of incompatibility. The present paper deals with the results of cross-breeding experiments carried out between American and European populations. The respective intrapopulation crosses served as controls. Transatlantic crosses have been attempted with the following species: T. lagunaris, T. bulbisetosa, T. holothuriae, T. battagliai, and T. clodiensis. The American and European populations of the first 4 species are interfertile, and the F₁ hybrids produce viable offspring. Moreover, the interpopulation crosses, compared to the controls, exhibit a slight superiority whose nature is, however, still unclear. The fifth species, T. clodiensis, behaves differently. The absence of viable offspring from interpopulation crosses, or the death of the hybrids before reaching the adult stage, indicate that, in this species, the barrier to gene flow is complete. Populations of T. clodiensis show, in addition, signs of relative intraspecific incompatibility. The problem of the different degree of reproductive isolation in the genus Tisbe, with special regard to the role played by ecological factors in determining the isolation patterns, is discussed.     

Population structure of the planktonic copepod Calanus pacificus in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Feeding deterrence properties of apo-fucoxanthinoids from marine diatoms. II. Physiology of production of apo-fucoxanthinoids by the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana,and their feeding deterrent effects on the copepod Tigriopus californicus

The marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana have been shown to produce apo-fucoxanthinoid compounds which act as feeding deterrents against the harpacticoid copepod Tigriopus californicus. The amounts and types of apo-fucoxanthinoids produced were species specific. Th. pseudonana produced small quantities of apo-12-fucoxanthinal and apo-13-fucoxanthinone only during senescence, while P. tricornutum produced much greater quantities of these two compounds during both log and senescence phases, in addition to producing a third compound, apo-10-fucoxanthinal, only during senescence. For both species, production of apo-fucoxanthinoids increased as the cells entered senescence phase due to phosphate limitation. The amounts of apo-fucoxanthinoids necessary to reduce feeding in T. californicus by 50% ranged from 2.22 to 20.2 ppm. This range was approximately 1000 times lower than the total apo-fucoxanthinoid concentration in P. tricornutum. The amounts of apo-fucoxanthinoids necessary to cause a 50% mortality in a population of T. californicus ranged from 36.8 to 76.7 ppm. Thus, these compounds are present in concentrations which may have ecological significance in the control of bloom formation and grazing. The production of apo-fucoxanthinoids may be a phenomenon common to many diatoms, particularly as they enter senescence due to nutrient limitation.     

Amphi-Atlantic phylogeography of direct-developing lineages of Lasaea, a genus of brooding bivalves

Direct-developing intertidal Lasaea spp. occur in the North Atlantic as both continental margin and oceanic island populations. We conducted a molecular phylogenetic analysis of representative populations in order to test colonization hypotheses for North Atlantic oceanic islands. Thirty individuals each were collected in 1995 and 1996 from two continental putative source populations (Florida, Iberia) and two oceanic island populations (Bermuda, Azores). They were sequenced for a 462 nucleotide portion of the mitochondrial large ribosomal subunit (16S) gene. No amphi-Atlantic genotypes were detected: Bermudan lineages co-clustered exclusively with Floridian congeners, and Azorean samples formed an exclusive clade with Iberian haplotypes. Our data indicate that geographical proximity to continental source populations is a better predictor of phylogenetic relationships in North Atlantic Lasaea spp. than present-day oceanic surface circulation patterns. The phylogenetic trees generated are not consistent with colonization of oceanic islands by indirect-developing ancestral lineages or by truly trans-oceanic rafting events. However, they are consistent with predicted topologies resulting from limited (≤ 2000 km), long-distance colonization by rafting (against present-day circulation patterns in the case of the Azores) and from anthropogenic introductions. Received: 17 December 1998 / Accepted: 7 June 1999     

Patterns of mtDNA diversity in North Atlantic populations of the mussel Mytilus edulis

Patterns of (female) mitochondrial DNA diversity were investigated in the blue mussel Mytilus edulis. Mytilus edulis is a ubiquitous member of contemporary North Atlantic hard-substrate communities and well represented in studies of this region. Mytilus edulis was surveyed in North America and Europe, as well as mid-Atlantic sites in Greenland, Iceland, and the Faroe Islands. Mitochondrial DNA sequences revealed considerable population structure but no monophyly of haplotypes between any major regions. Coalescent analyses suggest that migration across the Atlantic Ocean has prominently been from North American source populations and that Greenland was colonized recently and exclusively from North America. In North America, there was support for two regional groups along the North American coastline. Surprisingly, we also found evidence of recombination between some Mytilus edulis and Mytilus galloprovincialis female mtDNA sequences, particularly in northern Europe. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Tropical fishes in a temperate sea: evolution of the wrasse <Emphasis Type="Italic">Thalassoma pavo</Emphasis> and the parrotfish <Emphasis Type="Italic">Sparisoma cretense</Emphasis> in the Mediterranean and the adjacent Macaronesian and Cape Verde Archipelagos

The northeastern Atlantic and the Mediterranean Sea share geological histories and display great faunal affinities. The majority of the Mediterranean species have Atlantic origins, with a few species with tropical affinities. These include the parrotfish Sparisoma cretense and the wrasse Thalassoma pavo that are restricted to the subtropical northeastern Atlantic, the Macaronesian archipelagos (Azores, Madeira, and Canaries) and the southern Mediterranean. The Pleistocene glaciations have been described as having different effects on the fauna of the two regions. During glacial peaks, Mediterranean waters remained warmer than those of the adjacent Atlantic. Within the eastern Atlantic, the effects of Pleistocene glaciations were differentiated. Here, we perform a comparative analysis focusing on T. pavo and S. cretense populations from the northeastern Atlantic and the Mediterranean to assess the effects of Pleistocene glaciations in these two species. Sequences from the mitochondrial control region were obtained and analyzed combining phylogeographic and demographic approaches. Gene flow between Atlantic and Mediterranean populations was shown to be very high. The Mediterranean populations of T. pavo and S. cretense showed high levels of genetic diversity, even in the eastern basin, pointing to an ancient colonization event. This suggests that both species must have been able to persist in the Mediterranean during the cold Pleistocene periods. Historical migration estimates revealed a Mediterranean towards Atlantic trend in the case of T. pavo, which may reflect the re-colonization of areas in the Atlantic by fish that survived the cold phases in relatively warmer Mediterranean refugia. Our data also showed that within the Macaronesian Archipelagos, migrations occurred from Madeira towards the Azores, for both T. pavo and S. cretense, thus supporting a post-glacial colonization of the Azores by fish that persisted in the warmer region of Madeira. Similar geographic distributions, thermal affinities, and means of dispersion for T. pavo and S. cretense resulted in a similar response to the effects of Pleistocene glaciations, as evidenced by identical phylogeographic patterns.     

Feeding deterrence properties of apo-fucoxanthinoids from marine diatoms. I. Chemical structures of apo-fucoxanthinoids produced by Phaeodactylum tricornutum

Methanol extracts of freshly harvested cells of the marine diatom Phaeodactylum tricornutum were found to produce a feeding deterrent effect in the copepod Tigriopus californicus. Bioassay guided fractionation of the methanol extracts led to the isolation of four compounds possessing feeding deterrent activity. The compounds were identified as apo-10-fucoxanthinal (1), apo-12-fucoxanthinal (2), apo-12-fucoxanthinal (3), and apo-13-fucoxanthinone (4) by detailed spectroscopic analysis and comparison with authentic compounds produced semi-synthetically from fucoxanthin. Compounds 1 to 4 exhibited feeding deterrent responses in T. californicus at concentrations of less than 20 ppm.     

Genetic structure of the flounders Platichthys flesus and P. stellatus at different geographic scales

The genetic structure of the flounders Platichthys flesus L. and P. stellatus Pallas was investigated on different spatial scales through analysis of allozyme variation at 7 to 24 polymorphic loci in samples collected from different regions (Baltic Sea, North Sea, Brittany, Portugal, western Mediterranean, Adriatic Sea, Aegean Sea and Japan) in 1984 to 1987. No geographic variation was evident within a region. Some pattern of differentiation by distance was inferred within the Atlantic, while the Mediterranean comprised three geographically isolated populations and was itself geographically isolated from the Atlantic (fixed allele differences at up to three loci were found among P. flesus populations from the Atlantic, the western Mediterranean, the Adriatic Sea, the Aegean Sea and also P. stellatus from the coast of Japan). Sea temperature during the reproductive period probably acts as a barrier to gene flow between populations. Genetic distances among European flounder populations (P. flesus) were higher than, or of the same magnitude as, the genetic distance between Pacific (P. stellatus) and European flounder populations, suggesting that P. flesus is paraphyletic and/or there is no phylogenetic basis to recognising P. stellatus as a different species. The divergence between P. flesus and P. stellatus was thus inferred to be more recent than the divergence between the present P. flesus populations from the NE Atlantic and eastern Mediterranean. The eastern Mediterranean populations are thought to originate from the colonisation of the Mediterranean by a proto-P. flesus/P. stellatus ancestor, whereas the present western Mediterranean population has undergone a more recent colonisation event by P. flesus. Patterns of mitochondrial DNA variation, established on a smaller array of P. flesus samples, were in accordance with the geographic patterns inferred from the allozyme survey. In addition, they supported the hypothesis of a two-step colonisation of the western Mediterranean. These results contribute to our understanding of the biogeography of the Mediterranean marine fauna, especially the group of boreal remnants to which P. flesus belongs. Received: 7 February 1997 / Accepted: 26 March 1997     

Experimental interbreeding between geographically separated populations of the marine wood-boring isopod Limnoria tripunctata with preliminary indications of hybrid vigor

Populations of the marine wood-boring isopod Limnoria tripunctata Menzies were cultivated and interbred in the laboratory in order to determine whether geographically separated populations would interbreed. Two series of interbreeding studies were conducted. In the first, field populations from 11 Atlantic and Caribbean sites and 3 Pacific sites were interbred with a North Western Atlantic population from Beaufort, North Carolina (USA). Viable crossings, to at least F ₁, occurred in all but one case—that of a BeaufortxChatham (Massachusetts, USA) cross. In the second series of experiments, field populations from 9 Atlantic and Caribbean sites were crossed with populations from St. Teresa, Florida, Gulf of Mexico. In this case also, all crosses except one produced viable offspring, not only to the F ₁, but to the F ₂ generations as well. The St. Teresa x Chatham cross was not successful. Geographic distance was not a factor in the success of crossbreeding between disjunct populations. Chatham populations (mainly) are now believed to represent a valid but previously obscure species, Limnoria tuberculata Sowinski, and not a variant of L. tripunctata Menzies. Enhanced viality or vigor, as determined from rate of wood destruction, population growth rate and survival, was indicated in several crossings, but this was not necessarily correlated with distance between populations. Observations were made on the manner in which the species has achieved a worldwide distribution, and a hypothesis is put forward to explain the evolution of species related to L. tripunctata.Contribution from Zoology Department, Duke University, Durham, North Carolina, and Duke University Marine Laboratory, Beaufort, North Carolina, and the Department of Oceanography, No. 289, Florida State University, Tallahassee, Florida. Support in part from NSF Grant No. GB-6898.     

Feeding deterrent and toxicity effects of apo-fucoxanthinoids and phycotoxins on a marine copepod (Tigriopus californicus)

Using the marine harpacticoid copepod Tigriopus californicus, the effects of phytoplankton feeding deterrents and toxins were differentiated and measured. Eight compounds were tested for feeding deterrence and toxicity responses: four apo-fucoxanthinoids (apo-10′-fucoxanthinal, apo-12′-fucoxanthinal, apo-12-fucoxanthinal, and apo-13′-fucoxanthinone) and four well-known phycotoxins (domoic acid, okadaic acid, microcystin-LR, and a mixture of PSP-1 toxins). Since several of these compounds exhibited both feeding deterrence and toxicity, a model was developed to deconvolute the observed toxicity response from the observed feeding deterrence response, and to classify these compounds based on the degree of toxicity and/or feeding deterrence they exhibited towards T. californicus. Microcystin-LR, the PSP-1 toxins, and the four apo-fucoxanthinoids behaved only as feeding deterrents at low concentrations. Okadaic acid exhibited both toxicity and feeding deterrence at low concentrations, with the threshold concentration for feeding deterrence at a lower level than the threshold concentration for toxicity. Domoic acid acted only as a toxin at low concentrations, with all decreases in feeding resulting from the death of the copepod. Received: 4 November 1996 / Accepted: 3 December 1996