Distribution of secondary metabolites in the sponge Oceanapia sp. and its ecological implications

The Micronesian sponge Oceanapia sp. has an unusual growth form that consists of an irregular turnip-shaped base, which is buried in the substrate. One to several fistules, which protrude through the sand, are attached to the base of the sponge. On top of each fistule is a small fragile capitum. We examined whether this conspicuous red-colored sponge was chemically defended and if intraspecimen variation existed in the distribution of secondary metabolites between different parts of the sponge. Furthermore we assessed the deterrent properties of the secondary metabolites to generalist and more specialized fish predators. We also wanted to see if the optimal defense theory holds in the case of a marine invertebrate. According to the theory, organisms evolve and allocate defenses in a way that maximizes individual fitness, assuming that defenses are costly to the fitness of the organisms. We were able to evaluate this hypothesis, since the different sponge parts in Oceanapia sp. were at different risk to damage by predators and had a different value in terms of fitness loss to the sponge (the capitum probably plays a role in asexual propagation). Concentrations of crude organic extract increased from the base to the capitum of the sponge. The major secondary metabolites kuanoniamine C and D also showed a sharp increase from the basal root to the capitum. There was no difference in structural material or ash content between the base and the fistule of the sponge, but fiber and protein content were significantly higher in the fistule. The methanol fraction was highly deterrent in field feeding assays towards generalist reef fish at base concentration. It also deterred feeding by the spongivorous angelfish Pomacanthus imperator in laboratory feeding experiments at the same concentration. The field feeding assays with pure compounds showed that kuanoniamine C and D deterred feeding by natural assemblages of reef fishes at fistule concentrations, confirming their role as defensive agents. The intraspecimen variation of secondary metabolites in Oceanapia sp. supports the optimal defense theory by showing the highest concentrations in those parts of the sponge that are most visible to predators and are likely to be most important for inclusive fitness. Received: 5 May 1999 / Accepted: 16 September 1999     

Chemical studies on Indopacific Ceratosoma nudibranchs illuminate the protective role of their dorsal horn

Summary. Two Ceratosoma species, C. trilobatum and C. gracillimum (Mollusca: Gastropoda: Opistobranchia), collected along the South China Sea coast, have been studied in order to investigate their defensive strategies. Both species contain four typical sponge furanosesquiterpenoids, probably with a dietary origin. The tissue distribution of these compounds has been studied by ¹H-NMR analysis and preliminary bioassays have been carried out in order to evaluate the ecological role of the isolated metabolites. The main sesquiterpene component in both species was (–)-furodysinin (2), which showed significant feeding-deterrent and ichthyotoxic properties in bioassays with freshwater organisms. The work has been performed studying a very limited number of molluscs that prevented the realization of ecologically relevant experiments. However, the almost exclusive presence of compound 2 in the crude extract of the mantle glands concentrated in the dorsal horn of C. gracillimum strongly supports the defensive role of that dorsal protuberance, which probably acts as a defensive lure attracting potential predators to the part of the animal that contains the distasteful chemicals.     

Chemoecological studies of the exocrine glandular larval secretions of two chrysomelid species (Coleoptera):Phaedon cochleariae andChrysomela lapponica

Summary The exocrine glandular secretions of larvae of the subfamily Chrysomelinae are known to repel conspecific adults, other competitive phytophagous insects and natural enemies. InPhaedon cochleariae, the intraspecific activity of tlc fractions of the larval secretion was tested in order to examine the ecological significance of two fractions containing minor components and a fraction containing the major compound, the cyclopentanoid monoterpene (epi)chrysomelidial. InChrysomela lapponica, the defensive activity of the larval secretion against ants is known from specimens feeding upon willow or birch. The feeding preferences of larvae and adults ofC. lapponica from a Finnish and a Czech population were tested. The Finnish individuals significantly preferred feeding uponSalix borealis, whereas they hardly fed upon birch. The Czech specimens clearly preferred birch (Betula pubescens) to willow species. Application of salicin onto leaves of a willow species free of this phenolglycoside revealed that the Finnish individuals preferred feeding upon leaves with salicin. On the other hand, the Czech individuals avoided feeding upon leaves ofB. pubescens treated with salicin. The chemical composition of the glandular secretion of the Finnish larvae differed from the one of the Czech larvae. GC-MS-analyses of the secretions revealed that salicylaldehyde was the only major component of the secretion of Finnish larvae feeding upon the salicin-containing willowS. borealis. The glandular secretion of the Czech larvae feeding upon birch contained numerous esters of isobutyric acid and 2-methylbutyric acid. When Czech larvae had fed upon a salicin-containing willow (S. fragilis), the major compounds of their secretion were benzoic acid, salicylalcohol and benzoic acid esters; salicylaldehyde was only detected in traces. Thus,C. lapponica individuals from the Finland population adapted so closely to a salicincontaining willow that they clearly prefer this plant for food and that they obviously derive their main larval defensive compound (salicylaldehyde) from their host-plant.     

Sponge-feeding fishes of the West Indies

In an analysis of the stomach contents of 212 species of West Indian reef and inshore fishes, sponge remains were found in 21 species. In eleven of these, sponges comprised 6% or more of the stomach contents; it is assumed that these fishes feed intentionally on sponges. Sponges comprise over 95% of the food of angelfishes of the genus Holacanthus, over 70% of the food of species of the related genus Pomacanthus, and more than 85% of the food of the filefish, Cantherhines macrocerus. Lesser quantities of sponges are ingested by the remaining fish species. Fishes that feed on sponges belong to highly specialized teleost families, suggesting that this habit has evolved in geologically late time. The small number of fish species that concentrate on sponges as food suggests that the defensive characters of sponges—mineralized sclerites, noxious chemical substances, and tough fibrous components—are highly effective in discouraging predation. The two sponges most frequently eaten by fishes have a low percentage of siliceous spicules relative to organic matter, but among the 20 next most frequently consumed species no striking correlation occurs with respect to spicule content. Color and form of the sponge show no special correlation with frequency of occurrence in fish stomachs. Three species of fishes appear to concentrate on one species of sponge, but in these cases over 60% of the food taken consists of a variety of other organisms. Those fishes, more than half of whose diet consists of sponges, tend to sample a wide variety of species. No strong evidence is provided by our data that fish predation is a significant factor in limiting sponge distribution in the West Indian region.     

Surface sequestration of chemical feeding deterrents in the Antarctic sponge<Emphasis Type="Italic"> Latrunculia apicalis</Emphasis> as an optimal defense against sea star spongivory

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.Communicated by P.W. Sammarco, Chauvin     

Dietary preferences of the opisthobranch mollusc Stylocheilus longicauda for secondary metabolites produced by the tropical cyanobacterium Lyngbya majuscula

Pure compounds isolated from the cyanobacterium Lyngbya majuscula Gomont were evaluated in an artificial diet for their influence on the feeding preferences of the sea hare Stylocheilus longicauda (Quoy and Gaimard, 1824), which lives in and feeds on this filamentous cyanobacterium (blue-green alga). Microcolin B, ypaoamide, malyngolide and other natural products acted as feeding deterrents at natural concentrations. At lower concentrations, sea hares were indifferent to ypaoamide and malyngolide in their diets. In contrast, barbamide stimulated sea hare feeding at the concentrations normally found in L. majuscula. Malyngamides and majusculamides, the most common natural products found in samples of L. majuscula from Guam, increased sea hare feeding at low concentrations and inhibited feeding at the higher concentrations that occurred in some collections of L. majuscula. Dietary selection of cyanobacteria by S. longicauda may be regulated by the concentration of specific chemical cues produced by L. majuscula. Received: 28 January 1997 / Accepted: 24 April 1998     

Ecological role for pteroenone,a novel antifeedant from the conspicuous antarctic pteropod Clione antarctica (Gymnosomata: Gastropoda)

Dense populations of the antarctic pteropod Clione antarctica (Smith) offer a rich source of potential nutrients and energy to planktivorous predators. Nonetheless, antarctic fish do not prey on C. antarctica. Employing flash and high-pressure liquid chromatographic techniques, a linear -hydroxyketone, pteroenone (C₁₄H₂₄O₂) was isolated from whole tissues of C. antarctica. When embedded in alginate food pellets at ecologically relevant concentrations, pteroenone caused significant feeding deterrence in Pagothenia borchgrevinki and Pseudotrematomas bernacchii, two antarctic fish known to feed on planktonic organisms. Concentrations of pteroenone were variable between pteropods (0.056 to 4.5 mg ml^-1 tissue), but even those individuals with the lowest natural concentration contained levels five-fold greater than the lowest effective feeding-deterrent concentration (0.012 mg ml^-1 alginate). Chemical analysis indicated that the primary dietary item of the carnivorous C. antarctica, the shelled pteropod Limacina helicina, does not contain pteroenone. This suggests that C. antarctica does not derive this defensive compound from its diet. This is the first example of a defensive secondary metabolite in a pelagic gastropod.     

Cytotoxic haplosclerid sponges preferred: a field study on the diet of the dotted sea slug<Emphasis Type="Italic"> Peltodoris atromaculata</Emphasis> (Doridoidea: Nudibranchia)

Trophic specializations are widespread among opisthobranch molluscs. One purported example from the Mediterranean Sea is the dotted sea slug Peltodoris atromaculata. It has been hypothesized that this species is strongly monophagous on the sponge Petrosia ficiformis. However, the small amount of evidence that has been found for this hypothesis is based just on laboratory tests. Here we study the feeding habits and the diet of Peltodoris atromaculata in its natural habitat. We observed and videotaped 161 individuals together with the organisms on which they were found (their living substrata). Feeding scars were identified and videotaped as well. Individuals and their living substrata were sampled for further analysis in the laboratory. The composition of faeces of Peltodoris, especially undigested sponge spicules, was analyzed by light and scanning electron microscopy and compared to the composition of the living substrata. Most of the faecal samples consisted of undigested sponge spicules. Although Peltodoris was found on 11 species of sponges, only 2 of them, Petrosia spp. and Haliclona fulva, form its diet (76% out of n=121 samples). In accordance with this, feeding scars in the habitats were exclusively observed on these two sponges. Estimation of electivity indices suggests that Haliclona is preferred over Petrosia. One remarkable feature of the exclusive feeding of Peltodoris on Petrosia and Haliclona is that both sponges share specific fulvinol-like polyacetylenes that show cytotoxic activity in bioassays. Potential benefits and evolutionary aspects of this trophic specialization are discussed. Besides sponge-containing faeces, we found spicule-free faeces (24%, n= 29). These were very small in volume compared to sponge-containing faeces, and only few distinct structures were present. However, the use of food other than sponges is not necessarily indicated by this, because the spicule-free faeces might also represent left-overs from the stomach and digestive gland after sponge spicules have been released.Communicated by O. Kinne, Oldendorf/Luhe     

Ecological roles of natural products from the marine sponge Geodia corticostylifera

In the Brazilian coast, high numbers of the small brittle star Ophiactis savignyi usually live associated with the sponge Geodia corticostylifera (Demospongiae, Geodidae), but not with other sympatric sponge species. In order to check whether this association was related only with the physical shelter provided by the sponge body or was chemically mediated, the crude organic extract of G. corticostylifera was added to sponge mimics made of phytagel and spongin skeleton. Control and treated mimics were simultaneously offered to previously sponge-associated O. savignyi in both static seawater and flow-through laboratory experiments. Ophiuroids were allowed to move towards the preferred mimic. The defensive properties of the sponge extract against fish predation and fouling were also evaluated. Chemotaxis assays showed that symbiotic ophiuroids were able to chemically recognize its host sponge, moving significantly more towards mimics containing G. corticostylifera extract. Chemical deterrence assays showed that the natural concentration of the extract of this sponge was also able to inhibit generalist fish predation on field experiments and the attachment of the common mussel Perna perna in laboratory assays. These results indicate that the crude extract of G. corticostylifera plays multiple functions in the marine environment, presumably being responsible for a closer association of this sponge with O. savignyi, providing protection for this ophiuroid and inhibition of epibionts on itself.     

Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Inter-individual variation in sequestration (as measured by energy dispersive spectroscopy) predicts efficacy of defensive secretion in lubber grasshoppers

Summary. We tested whether inter-individual variation in concentration of sequestered compounds predicts defensive efficacy of the defensive secretion of onion-fed Eastern lubber grasshoppers. When fed onion, lubbers have the ability to sequester sulfur compounds into their defensive secretion, and the secretion from onion-fed lubbers is more deterrent of ants than the secretion from lubbers fed other diets (Jones et al. 1989). To test three hypotheses, we developed a method for measuring total sequestered sulfur in the secretion using energy dispersive spectroscopy (EDS). First, we found that lubbers fed an ad libitum, monophagous diet of onion produce more secretion and have higher concentrations of total sulfur in their secretions than lubbers fed other diets. These sulfur concentrations varied three-fold; we believe that this variation in the composition of defensive secretions stems from physiological variation. Second, our method was sensitive enough to detect sulfur in the secretion of some lubbers that had been fed onion only once. Third, the sulfur concentration of a secretion sample significantly predicted its ability to deter ants from foraging. Hence, our results suggest that inter-individual variation in defensive chemistry in insects offered the same diet may be ecologically important. Received 25 November 1997; accepted 5 February 1998.     

Turning the game around: toxicity in a nudibranch-sponge predator–prey association

Escalation theory proposes enemy-related selection as the most relevant factor of natural selection among individual organisms. When hazardous to predators, prey might be considered enemies that influence predator evolution. Opisthobranch molluscs that prey on chemically defended prey are an interesting study case on this subject. Predation on chemically defended species paved the way for opisthobranchs to enter in an arms race, developing means to detoxify and/or excrete harmful compounds, which led to the sequestration of those compounds and their self-defensive use, an escalation of defenses. Here we aim to understand whether the opisthobranch predator is better protected than its chemically defended prey, using as predator–prey model, a nudibranch (Hypselodoris cantabrica) and the sponge it preys upon (Dysidea fragilis), and from which it obtains deterrent chemical compounds. Specimens of both species were collected on the Portuguese coast, and their crude extracts were analyzed and used in palatability tests. Nudibranchs revealed a higher natural concentration of crude extract, probably due to a progressive accumulation of the compounds. Both predator and prey extracts revealed similar mixtures of deterrent metabolites (furanosesquiterpenes). Palatability tests revealed a more effective deterrence in the nudibranch extracts because significant rejection rates were observed at lower concentrations than those necessary for the sponge extracts to have the same effect. We concluded that the predator is chemically better protected than its prey, which suggests that its acquisition of chemical defenses reveals a defensive escalation.     

Chemical ecology and origin of defensive compounds in the Antarctic nudibranch Austrodoris kerguelenensis (Opisthobranchia: Gastropoda)

The common Antarctic nudibranch Austrodoris kerguelenensis (Bergh) contains diterpene diacylglycerides only present in its external body parts. These compounds provide a chemical defense against sympatric predators, such as the seastar Odontaster validus Koehler. Bioassays conducted with O. validus revealed that live nudibranchs, mantle tissue and Et₂O extract of the A. kerguelenensis mantle deterred feeding by the seastar. Further bioassays testing organic fractions of the Et₂O mantle extract showed that the diterpene diacylglycerides, as well as corresponding monoacylglycerides and monoacylglycerides of regular fatty acids, were responsible for the feeding deterrence in O. validus. We suggest that A. kerguelenensis derives the bioactive diacylglycerides by de novo biosynthesis rather than by sequestration from its sponge diet, since the mollusk does not contain active metabolites in the viscera, and neither the active compounds nor precursors were detected in the sponge diet. Furthermore, A. kerguelenensis did not show a strong chemodetection or feeding preference for its main diet, hexactinellid sponges, in Y-maze and food choice experiments, respectively.     

Mimicry: imitative depiction of discharged defensive secretion on carapace of an opilionid

Summary. The opilionid, Parampheres ronae, has a pair of orange markings on its carapace at a location where some other opilionids discharge yellowish, quinone-containing defensive fluid. P. ronae itself produces a defensive secretion, but the fluid is quinone-free and nearly translucent. We suggest that the orange markings in P. ronae are aposematic in the sense that they are imitative of the glandular emissions of quinone-producing opilionids.     

Spatio-temporal variations in the diet and stable isotope composition of the Argentine hake <Emphasis Type="Italic">Merluccius hubbsi</Emphasis> Marini, 1933 of the continental shelf of southeastern Brazil

The feeding ecology of Merluccius hubbsi was investigated in 2 regions of SE Brazil. The major food sources for the hakes were fish, crustaceans, and squid. In the upwelling system of Cabo Frio, the diet was very similar in the summers of 2001/2002 and spring 2002; fish were the most important prey followed by crustaceans. In Ubatuba, euphausiids were an important prey during the winter 2001 (100 m), while in the summer 2002, fish and amphipods predominated in the diet in the shallower site (40 m) and squid in the deeper site (100 m). The hakes showed temporal differences in stable isotope signatures in both regions, while C:N ratios varied only in Cabo Frio. δ¹⁵N and δ¹³C (bulk and corrected for lipid content) increased with fish length, which seems to be related to the increasing importance of fish and decreasing importance of euphausiids and amphipods in the diet of larger hakes. The mean trophic level of 3.7 for M. hubbsi was estimated using δ¹⁵N of bivalves as baseline and the fractionation of 3.4‰ between trophic levels.     

Video-monitored predation by Caribbean reef fishes on an array of mangrove and reef sponges

Although predation by fishes is thought to structure benthic invertebrate communities on coral reefs, evidence to support this claim has been difficult to obtain. We deployed an array of eight sponge species on Conch Reef (16 m depth) off Key Largo, Florida, USA, and used a remote video-camera to record fish activity near the array continuously during five daylight periods (6 h for 1 d, at least 11.5 h for 4 d) and one night period (11 h). Of the eight sponge species, four were from adjacent reefs (Agelas wiedenmayeri, Geodia neptuni, Aplysina fistularis, and Pseudaxinella lunaecharta), and four were from a nearby mangrove habitat (Chondrosia collectrix, Geodia gibberosa, Halichondria sp., andTedania ignis). Each species of reef sponge was chosen to match the corresponding mangrove species in form and color (black, brown, yellow, and red, respectively). Predation events only occurred during daylight hours. Tallies of the number of times fishes bit sponges revealed intense feeding by the expected species of sponge-eating fishes, such as the angelfishHolacanthus bermudensis, H. tricolor, andPomacanthus arcuatus, the cowfishLactophrys quadricornis, and the filefishCantherhines pullus, but surprisingly also by the parrotfishSparisoma aurofrenatum andS. chrysopterum. Of 35 301 bites recorded, 50.8% were taken by angelfish, 34.8% by parrotfish, and 13.7% by trunkfish and filefish. Mangrove sponges were preferred by all reef fishes; 96% of bites were taken from mangrove species, with angelfish preferringChondrosia collectrix and parrotfish preferringGeodia gibberosa. Fishes often bit the same sponge repetitively, and frequently consumed entire samples within 30 min of their deployment. Sponge color did not influence fish feeding. Two of the four mangrove sponge-species deployed on the array were also found living in cryptic habitats on adjacent reefs and were rapidly consumed by fishes when exposed. Our results demonstrate the importance of fish predation in controlling the distribution of sponges on Caribbean reefs.     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Growth and grazing responses of tintinnid ciliates feeding on the toxic dinoflagellate Heterocapsacircularisquama

Growth and feeding rates of two tintinnid species, Favellaazorica and Favellataraikaensis, were determined under various concentrations of the dinoflagellate Heterocapsacircularisquama which has been reported as highly toxic to shellfish. Mean growth rates of F. azorica and F. taraikaensis on a diet of H.circularisquama (ca. 10² cells ml⁻¹) were 2.15 and 1.97 doublings d⁻¹, respectively. These values are similar to those on a diet of Heterocapsatriquetra which is suitable food for various zooplankton. However, growth rates of both tintinnid species decrease with increasing concentrations of >10³ cells ml⁻¹ of H. circularisquama. In particular, H. circularisquama under conditions of >10³ cells ml⁻¹ caused mortality in F.taraikaensis, probably due to toxins. Clearance and ingestion rates of F. azorica on H. circularisquama were 4.1 to 27.5 μl ind⁻¹ h⁻¹ and 1.5 to 28.7 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10⁴ cells ml⁻¹ and those of F. taraikaensis were 0.9 to 22.1 μl ind⁻¹ h⁻¹ and 0.1 to 13.0 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10³ cells ml⁻¹. Both clearance and ingestion rates on H.circularisquama were higher for replicates fed on H.triquetra. Daily grazing impact of the two species of Favella on the initial stage of a bloom of H.circularisquama were estimated to reach 6 to 50% of H. circularisquama at a concentration of 540 cells ml⁻¹, indicating that grazing by tintinnids such as Favella spp. may significantly regulate the initial stages of blooms of H. circularisquama. Received: 3 January 1997 / Accepted: 27 January 1997     

Hydroquinones from defensive secretion of a giant Pacific millipede, <Emphasis Type="Italic">Acladocricus setigerus</Emphasis> (Diplopoda: Spirobolida)

Summary. The presence of considerable quantities of hydroquinones including hydroquinone, 2-methylhydroquinone, 2,3- dimethoxyhydroquinone, 2-methoxy-3-methylhydroquinone, and 2,3-dimethoxy-5-methylhydroquinone renders the defensive secretion of Acladocricus setigerus (Silvestri, 1897) significantly different from that of other quinone-producing millipedes. In addition, two hitherto undescribed compounds, namely, 2,3-dimethoxy-5-methylhydroquinone and 2-methyl-3,4-methylenedioxyphenol, were characterized from the defensive secretion. However, it is uncertain if the latter compound is formed after the release of the secretion. The two major compounds, 2-methyl-1,4-benzoquinone and 2-methoxy-3-methyl-1,4-benzoquinone, constitute about 75% of the defensive fluid. Furthermore, hydrocarbons, which are typically present in the secretions of most other arthropods that use benzoquinones as repellents, are notably absent in the secretion of A. setigerus.     

Predation on Euphausia pacifica by demersal fishes: predation impact and influence of physical variability

The importance of Euphausia pacifica Hansen to the demersal fish community off Sendai Bay, northern Japan, was studied based on a total of 256 trawl samples collected during May, October and November, from 1989 to 1992. The samples were classified into 16 assemblages according to year, month and depth. The importance of E. pacifica to the total diet of each of the assemblages was measured using an index considering both fish diet and species composition. A total of 24 fish species out of 87 were found to ingest E. pacifica. Of these, Gadus macrocephaus and Theragra chalcogramma had the highest predation impact due to their dominance in the fish assemblages and the high proportion of E. pacifica in their diets. The contribution of E. pacifica to the total diet of the fish assemblage was highest in the shallow (≤300 m) regions during May, accounting for an average of 38.5% for the 4 years. However, during May 1990, when the warm Kuroshio Extension prevailed, the contribution was higher in the deep (>300 m; 22.6%) region, reflecting active and/or passive movements of E. pacifica. The average predation impact was maximal in the May/shallow assemblage (mean ± SE; 4.6 ± 1.4 kg wet wt km² d⁻¹) and was minimal in the November/shallow assemblage (0.4 ± 0.3). Annual estimates of consumption by demersal fish fishes ranged from 43 to 128 metric tons, representing 15 to 64% of the annual commercial catch of E. pacifica by local fisheries. It is therefore suggested that the fisheries on E. pacifica have the potential to considerably impact the demersal fish assemblages. Received: 3 December 1997 / Accepted: 10 June 1998