Trophodynamics and seasonal cycle of the copepod Pseudocalanus acuspes in the Central Baltic Sea (Bornholm Basin): evidence from lipid composition

Seasonal lipid dynamics of the copepod Pseudocalanus acuspes were studied in the Bornholm Basin (Central Baltic Sea) on a monthly basis from March 2002 until March 2003 and were interpreted in light of life cycle strategies and diet selection. The individual total lipid content of females ranged from 0.9 to 1.8 μg, with relative wax ester content reaching a significant maximum in May (44% of total lipids) and minimum (17% of total lipids) in April and November. Significant changes in size, lipid content, lipid classes and fatty acid composition of structural as well as storage lipids suggested five characteristic seasonal phases that were induced by different feeding histories and environmental conditions. Storage lipids were characterized by 18:1(n−9) as major component, which ranged between 44% of total fatty acids in June and 23% in February. The strong coherence between 18:1(n−9) in the seston lipids and the occurrence of ciliates emphasized the importance of ciliates in the diet of P. acuspes. As indicated by changes in the amounts of fatty acid markers, other food sources varied over the year, suggesting an opportunistic feeding behavior. The spring period was characterized by an increase in typical diatom and dinoflagellate markers, whereas other sources, potentially cyanobacteria, became more important during summer. The life cycle strategy is discussed with respect to extant adaptations to high latitudinal habitats.     

Discrimination among and variation within species of Pseudocalanus based on the GPI locus

Patterns of allozyme variation at the GPI (glucose phosphate isomerase) locus showed that the recently described or clarified Pseudocalanus newmani, P. acuspes, P. moultoni and P. minutus are genetically isolated from each other even though morphologically they are very similar, and in places sympatric. This genetic isolation was inferred from the presence of private alleles in some species, and from differences in their degrees of observed heterozygosity and in the observed frequencies of shared alleles in all the species examined. Temporal variations were not detected in P. newmani, P. acuspes and P. minutus. Populations of P. newmani and P. moultoni from the Atlantic and Pacific coasts of North America have apparently diverged. No such divergence was detected among samples of P. acuspes from Nova Scotia, the Canadian Arctic and the Baltic sea, and of P. minutus from Nova Scotia and Hudson Bay. P. elongatus collected in the English Channel differed from P. acuspes from the Baltic Sea.     

Comparative ultrastructure of lipid storage sites in femaleEuchaeta marina andPleuromamma xiphias (Copepoda: Calanoida)

FemaleEuchaeta marina (Prestandrea, 1833) have one large, thin-walled lipid sac, whereas femalePleuromamma xiphias (Giesbrecht, 1889) have two separate and morphologically distinct lipid storage sites. One lipid site inP. xiphias corresponds to the mesenteric tissue that surrounds the anterior region of the midgut. The morphology of these cells resembles that of mammalian brown adipocytes. The cytoplasm is filled with extensive smooth endoplasmic reticulum, numerous mitochondria and several deposits of intracellular lipid. The second lipid site ofP. xiphias lies in the posterior region of the metasome and resembles the thin-walled lipid sac ofE. marina. Both lie adjacent to, but are not contiguous with, the narrow mesenteric tissue surrounding the last region of the midgut. Both sacs contain a single, large deposit of intracellular lipid enclosed by a very thin rim of cytoplasm and resemble mammalian white adipocytes. The different habitats and reproductive processes of these two copepod species may relate to the observed variations in lipid cell morphology. The reserve lipid inE. marina plays a primary role in reproduction and is linked closely with the continuous cycle of oocytic maturation. The lipids synthesized and stored byP. xiphias, a strong vertical migrator, may be influenced by food availability, a function of their mesopelagic habitat. The primary role of the reserve lipids in this copepod may be to provide energy during migrations and between feeding periods, with relatively less lipid being allocated to reproduction.     

Proximate composition of Antarctic mesopelagic fishes

Eleven mesopelagic fish species from the Weddell/Scotia Sea region of the Antarctic captured during the austral spring 1983, austral fall 1986, and austral winter 1988, were analyzed for proximate composition. Water, ash level, protein, lipid and carbohydrate were examined in relation to depth of occurrence and season. No depth-related trends were evident, primarily due to a low species diversity and minimal differences in those species' vertical distributions. The Antarctic speciesElectrona antarctica showed a significant increase in lipid level (% wet wt and % ash-free dry wt) between spring, fall and winter. The increase may signify an accumulation over the productive season, possibly as a reserve for the winter months. Lipid levels (% wet wt and % ash-free dry wt) were significantly lower in the Weddell Sea specimens examined in this study than in previously examined identical and congeneric species taken during the same season from a more productive near-shore Antarctic region. Comparisons with congeners and confamilials from tropical-subtropical and temperate systems revealed variable trends. The Antarctic speciesE. antarctica andCyclothone microdon had lower water and protein (% wet wt) levels than similar species from tropical-subtropical or temperate regions. Lipid levels of the two species are similar to temperate individuals, while energy levels are slightly higher. In contrast, species of the genusBathylagus show no trends in composition as a function of latitude. Differences in productivity, water-column temperature-structure, and seasonality are important considerations when examining trends among mesopelagic species.     

Lipid composition of twenty-two species of Antarctic midwater zooplankton and fish

Total lipids, hydrocarbons, wax esters, triacylglycerols, and phospholipids were determined for 22 major biomass species of zooplankton and fish in an Antarctic mesopelagic community that were collected in 1982 and 1983. Lipid levels were similar to levels in more temperate mesopelagic species. Total lipid concentration was depth related, with all lipid-rich species being collected at depths greater than 230 m. Wax ester content in copepods (60 to 70% of the total lipid) was generally higher than in subtropical species. Lipids indicated three predator-prey relationships (Parandania boecki-Atolla wyvillei, Thysanoessa macrura-Calanoides acutus andEurythenes gryllus-Atolla wyvillei). These were confirmed by gut content analyses. The mesopelagic fishBathylagus antarcticus, Pleuragramma antarcticum, andProtomyctophum bolini stored most lipid intramuscularly, whereasElectrona antarctica andGymnoscopelus nicholsi contained extensive stores in subcutaneous lipid sacs. The intramuscular lipids inP. antarcticum and the subcutaneous lipid sacs ofE. antarctica were primarily wax esters, possibly used for increased buoyancy or long-term energy storage. Unlike the odd-carbon preference of aliphatic hydrocarbons which typifies terrestrial plants and temperate marine organisms, even-carbon chain-length paraffins predominate in 80% of the Antarctic species analyzed. Although the source of these even-carbon n-alkanes cannot be determined from our data, their dominance in the species analyzed suggests that an unusual biochemical pathway may be responsible for their synthesis in this ecosystem.     

Fluids and jellies of the acusticolateralis system in relation to body fluids in Coryphaenoides rupestris and other fishes

The marine deep-water fish Coryphaenoides rupestris has a lymph-like fluid in its large cranial lateral-line canals. A similar canal endolymph is also found in a few other fishes possessing closed lateral-line canals. We have analysed, chemically, the canal endolymph from 3 species of teleosts. In C. rupestris, the ionic composition resembles that of blood plasma, but the content of potassium is slightly lower. In Lota lota, the canal endolymph has a high potassium concentration, as does the inner-ear endolymph. In Glyptocephalus cynoglossus, the ionic composition of the canal endolymph resembles sea water. In all species, the concentrations of glucose and protein are much lower than in the blood plasma. Analyses were further made on endolymph and perilymph of the inner ear, on jelly-like material from the acusticolateralis system, and on body fluids from a small number of fish species, even elasmobranchs and a holocephalan. The water phase of the cupulae of the lateral-line system of C. rupestris had an ionic composition similar to that of the endolymph surrounding the cupulae. High potassium and low values for glucose and protein were found in the inner-ear endolymph of all species investigated. No carbonic anhydrase activity was found in the lateral-line system of C. rupestris, whereas a low activity of this enzyme was found in the inner ear of Gadus morrhua. The investigation shows that the composition of the fluids in contact with the mechano-receptors of the lateral-line system differs between different species of fishes. A few observations were made on the structural organization of the cranial lateral-line system in C. rupestris.     

Annual population development and production by small copepods in Disko Bay,western Greenland

The population of small copepod species (approximately <1 mm) were investigated during an annual cycle in Disko Bay, western Greenland. The small species considered were Acartia longiremis, Pseudocalanus spp., Oithona spp., Oncaea spp., Microsetella spp., and Microcalanus spp. Most of the small species were present in the surface waters year round and numerically dominated the community, and in biomass from late summer and throughout winter. Oithona spp. was numerically the main contributor, while Pseudocalanus spp. dominated in terms of biomass. In the uppermost 50 m, maximum abundance, biomass and secondary production were observed in late September after the phytoplankton production practically had terminated and the winter initiated. The free spawning Acartia longiremis showed a strong seasonal fluctuation in biomass and egg production, in contrast to the egg carrying species Pseudocalanus spp. and Oithona spp. These had a long spawning season and maintained a more stable biomass year round. Secondary production was estimated by three different ways: (1) based on the obtained specific egg production rates, (2) a temperature dependent equation, and (3) a multilinear regression taking temperature, body weight and chlorophyll into consideration. The contribution of the small species was insignificant when compared to the large Calanus species during the spring- and post-bloom. However, during late summer and winter, where Calanus had left the upper water strata for hibernation, the small species played a crucial role in the pelagic carbon cycling.     

Effects of water-soluble fraction of the Mexican crude oil “Isthmus Cactus” on growth,cellular content of chlorophylla,and lipid composition of planktonic microalgae

Phytoplankton species were grown in batch cultures in the presence of the water-soluble fraction (WSF; 50 and 100%) of a Mexican crude oil (Isthmus Cactus). The algae exhibited various responses ranging from retarded growth to stimulation of growth. The cellular content of chlorophylla and the lipid composition of the algae were examined. Four algae, the bacillariophytesNitzschia closterium andAsterionella glacialis, the cryptophytesRhodomonas lens, and the chlorophyteDunaliella tertiolecta, exhibited retarded growth. In most of these algae, cellular chlorophylla, lipid pigments, glycolipids and triglycerides decreased whereas sterols and hydrocarbons accumulated. Phospholipids did not exhibit any specific pattern of change during the experiments. The cyanophyteAgmenellum quadruplicatum and the bacillariophyteSkeletonema costatum were less sensitive to the WSF. The cell yield of the dinophyteProrocentrum minimum was stimulated by the WSF. In these three latter species, lipid pigments were enhanced or remained at control levels. We concluded that the toxic effect of the WSF disrupts the biosynthesis mechanisms required for a functional photosynthetic apparatus (biosynthesis of chlorophylla, glycolipids and lipid pigments) in sensitive algae, a phenomenon coupled to sterol accumulation in these algae.     

Reproduction of asteroids from Balsfjorden,Northern Norway: Analyses of lipids in the gonads of Ctenodiscus crispatus,Asterias lincki and Pteraster militaris

Seasonal lipid analyses were performed on the gonads and residual bodies of Ctenodiscus crispatus (Retzius) and Asterias lincki (Müller & Troschel), collected from Balsfjorden, northern Norway during March 1978–February 1980. The ovaries of both species had high percentages of total lipid (in the region of 60% of the dry weight) that were relatively constant throughout the season. Testes from both species had in the region of 20% of their dry weight as lipid, again with little seasonal variation. The majority of the total lipid in the ovaries of C. crispatus was accounted for by alkyldiacylglycerols and there was little seasonal variation in lipid class composition. Phospholipid accounted for the bulk of the lipid in the testes of C. crispatus, where alkyldiacylglycerols were present but in minor amounts. A slight degree of seasonal variation occurred in the composition of testis lipid in that maximal percentages of triacylglycerols and alkyldiacylglycerols occurred in January–April. The ovaries of A. lincki contained both triacylglycerols and alkyldiacylglycerols as major neutral lipids. Significant seasonal variation occurred in lipid class composition in the ovaries in that neutral lipids showed transient minima in February–April. The testes of A. lincki had substantial amounts of both triacylglycerols and alkyldiacylglycerols: maximal percentages of these lipids occurred in January–March. Analyses of gonads of near-mature Pteraster militaris (O. F. Müller) revealed that the ovaries were rich in lipid that was predominantly alkyldiacylglycerols, whereas the testes had much lower levels of lipid accounted for mainly by phospholipids. The residual bodies of all three asteroids had significant but small amounts of alkyldiacylglycerols in their lipids. A. lincki had triacylglycerols as a major lipid class in its body lipids, whereas in C. crispatus the major body lipid was phospholipid. Little seasonal variation was seen in the composition of the residual body lipid of either C. crispatus or A. lincki. The results are consistent with an aseasonal reproductive pattern in C. crispatus, although a main breeding activity occurred in mid-winter. A. lincki is a more seasonal breeder, but also had maximal activity in winter. Irrespective of their breeding habits, all three asteroids produced eggs rich in alkyldiacylglycerols. A major role for this lipid in the development of the large lecithotrophic eggs of asteroids is indicated.     

Seasonal and annual variability in an epifaunal community in the German Bight

While the benthic infauna of the North Sea has been studied intensively over the past decades, few studies have focused on the larger mobile epifauna. Studies carried out to date have described the distribution of epifaunal communities over the whole of the North Sea, but variability within the identified communities, which occurs on a much smaller scale, has so far remained unstudied. This is the first study to describe seasonal and annual variability of an epifaunal assemblage in the German Bight area, where environmental conditions are highly variable. The benthic community was sampled with a 2 m beam trawl from 1998 to 2001. The echinoderms Ophiura albida and Asterias rubens and the crustacean Pagurus bernhardus were the dominant species caught throughout the study period. Overall the species composition of the catches was relatively consistent, while abundances of dominant species fluctuated considerably between sampling periods. Differences between sampling periods were not only influenced by the abundances of dominant species, but also by less dominant species such as Ophiura ophiura, Astropecten irregularis, Corystes cassivelaunus, Crangon crangon and Aphorrahis pespelicani. The abundances of these species varied annually and seasonally in the assemblage. Clear differences between summer and winter in the species composition, abundance and biomass were identified. Annual and seasonal changes were most likely linked to migratory movements of epifauna into and out of the area under investigation. Temporal changes in species composition and abundance correlated best with water temperatures, while the spatial distribution of the total biomass over the whole sampling period was correlated with sediment characteristics. Anthropogenic influences such as fishing activity and chronic large-scale eutrophication are thought to have influenced the community on a long-term basis, but have been considered unlikely causes for the short-term variability described by this study.Communicated by J.P. Thorpe, Port Erin     

Characteristic features of body composition and metabolism in some interzonal copepods

Protein, lipid, phosphorus, and organic carbon contents, as well as electron transport system (ETS) activity, lactatedehydrogenase activity, and gut evacuation rate, were measured in four interzonal species of Pacific copepods:Calanus australis, C. pacificus, Eucalanus inermis, andE. elongatus f.hyalinus, collected at the upwelling areas off Peru (8°S) and California (27°N), and in the middle of the North Pacific (30°N), from February to April 1987. The two Eucalanidae species —E. inermis andE. elongatus — have distinctive biochemical and elemental body composition and rates of main physiological processes. Relative protein, lipid, phosphorus, and organic carbon contents (µg mg^–1 wet weight) in these species were, respectively, ca. 1/7 to 1/10, 1/5 to 1/20, 1/5 to 1/10, and 1/5 those inCalanus spp. Likewise, oxygen uptake rate per unit of wet weight (based on ETS activity) inE. inermis andE. elongatus was 5 to 10% of that in calanids; a similar difference was found in phosphorus excretion rate. In addition, gut evacuation rates inE. inermis andE. elongatus were ca. one-fifth of those inCalanus spp. Based on these data, we considered the eucalanids as belonging to a distinctive physiological group, figuratively named jelly-body copepods. In contrast with calanids, active lactatedehydrogenase has been found in the bodies ofE. inermis andE. elongatus, apparently allowing them to survive for a long time in layers of extremely low oxygen content (<0.2 ml l^–1). The adaptive value of physiological features in these eucalanids and typical calanids is compared.     

Biochemical composition of cephalopods with different life strategies,with special reference to a giant squid,<Emphasis Type="Italic"> Architeuthis</Emphasis> sp.

The present study is a comprehensive comparison of the biochemical composition (protein, lipid, glycogen and cholesterol contents, and amino acid and fatty acid profiles) of 14 species of cephalopods with different life strategies (benthic, nektobenthic, benthopelagic and pelagic) in distinct habitats (neritic, demersal, oceanic and deep sea), with special emphasis placed on a male Architeuthis sp. The giant squid showed a significantly lower protein and total amino acid content in the gonad and digestive gland. The major essential amino acids were leucine, lysine and arginine. The major non-essential amino acids were glutamic acid, aspartic acid and proline. The benthopelagic cirrate octopus Opisthoteuthis sp. showed a lower content of these nitrogen compounds in the muscle. In respect to lipid and fatty acid contents, the giant squid showed the highest values in the gonad and digestive gland and the lowest in the muscle. Most of saturated fatty acid content was presented as 16:0; monounsaturated fatty acid content, as 18:1 and 20:1; and polyunsaturated fatty acid content, as 20:4n-6, 20:5n-3 and 22:6n-3. The highest cholesterol content in the gonad was detected in Opisthoteuthis sp. and Architeuthis sp.; in the digestive gland. in Todarodes sagittatus; and in the muscle, in Sepia elegans. The highest glycogen value in the gonad was detected in Octopus vulgaris; in the digestive gland and muscle, the highest values were attained in Opisthoteuthis sp. The relationships between life strategies and biochemical composition were investigated and principal component analysis (PCA) was performed. The different cephalopod life strategies could be well separated on the basis of the first two principal components. The nektobenthic and pelagic strategies were clearly separated from the benthic, suggesting that these groups of species are characterized by lower lipid content and higher protein content in the gonad. A rather similar life-strategy distinction was obtained for the digestive gland. The benthopelagic strategy was also well separated from benthic and pelagic strategies and from Architeuthis sp. In the muscle, the results indicated lower nitrogen and carbon compound contents in Architeuthis sp. and Opisthoteuthis sp. The environmental conditions that possibly explain the Architeuthis sp. stranding, the relationships between biochemical compositions and the life strategies of the different cephalopod species studied are discussed.Communicated by S.A. Poulet, Roscoff     

Floristic composition and vegetation ecology of the Malindi bay coastal dune field, Kenya

A short outline is given of the floristic composition, structure and distribution of coastal dune vegetation found at Malindi Bay, Kenya. The area was studied by air photo interpretation and field sampling to determine the relationship of plants to aeolian features. TWINSPAN classification was used to distinguish geomorphological units on the basis of their species composition. In this paper, an inventory and first quantitative analysis of vegetation distribution is presented. We identified 174 plant species from 62 families in the sand dunes and several plant communities are distinguished based on the species content and the connection with morphological units.Papilionaceae with 18 species andPoaceae with 17 species were the most represented families. A distinct zonal distribution of the plant communities was found. The most important plant species are the pioneer vegetation consisting ofHalopyrum mucronatum, Ipomoea pescaprae andScaevola plumieri. The woody shrub species which have colonized the established primary dunes and hummock dunes areCordia somaliensis, Pluchea discoridis, Tephrosia purpurea (dunensis). Succulent herbs were identified in the dune slacks and salt marsh that are moist and damp environments.     

Neutral lipids in macroalgal spores and their role in swimming

We followed changes in the neutral lipid content of actively swimming zoospores of the palm kelp Pterygophora californica in a laboratory experiment to investigate the degree to which spore swimming is fueled by endogenous lipid reserves. The neutral lipid content of individual zoospores during the experiment was measured by flow cytometry using Nile Red, a fluorescent stain that is specific for neutral lipid. Results showed that photosynthesis greatly influenced lipid consumption during zoospore swimming. We found no detectable change in the neutral lipid content of zoospores after 30 h of swimming under conditions where light was near the optimum for photosynthesis. By contrast, neutral lipid declined by ≃43% over 30 h in zoospores kept in the dark. To evaluate whether lipid reserves are generally related to spore motility in macroalgae, we surveyed spore lipid-content and composition in species with motile spores and non-motile spores using thin-layer chromatography (TLC), and flame-ionization detection (FID). We observed substantial differences in lipid content and composition among the 20 species examined. Spores high in total lipid (as estimated by the ratio of lipid:carbon) generally had a large amount of neutral lipid; motile spores had significantly more lipid and a significantly larger neutral lipid fraction than non-motile spores. The kelps as a group had the highest total lipid content and the largest neutral lipid fraction, while non-motile spores of red algae were generally low in total lipid and in the proportional abundance of neutral lipid. Phospholipids accounted for more than half of all lipid in 14 of the 20 species examined, while neutral lipid accounted for the majority of lipid in all five species of kelp examined. Triacylglycerols, which function primarily in energy storage, were the primary form of neutral lipid in all but one species of kelp (Agarum fimbriatum), whereas free fatty acids were the dominant form of neutral lipid in most red algae and in brown algae that had a small neutral lipid fraction. Our results are largely consistent with the hypothesis that macroalgae use endogenous lipid reserves to fuel spore-swimming. The small amounts of triacylglycerols observed in the motile spores of several species of brown and green algae, however, indicate that the amount of lipid reserves needed to fuel spore-swimming may be influenced by a variety of factors including swimming behavior, photosynthetic efficiency, and the light environment inhabited by spores. Received: 8 September 1998 / Accepted: 8 January 1999     

Lipids of some Caribbean and Red Sea corals: total lipid,wax esters,triglycerides and fatty acids

The Caribbean reef-building corals Porites porites (Pallas) and Montastrea annularis (Ellis and Solander) and the Red Sea corals Pocillopora verrucosa (Ellis and Solander), Stylophora pistillata (Esper) and Goniastrea retiformis (Lamark) were analysed for total lipid, wax ester and triglyceride content, and fatty acid composition. M. annularis contained about 32% of dry weight as total lipid, whereas much lower values of between 11 and 17% were recorded for the other four species. It is concluded that there is greater variation in coral total lipid than hitherto thought. The total lipid contained a substantial proportion of wax ester (22 to 49%) and triglyceride (18 to 37%). The storage lipids (wax esters and triglycerides) accounted for between 6 and 20% of the dry weight and between 46 and 73% of the total lipid. Variation in lipid content between species could not be attributed to geographical location, but the low values for total lipid in Red Sea corals may in part be due to environmental factors as these samples were collected in winter. All corals analysed contained high levels of saturated fatty acids, the most abundant fatty acids being 16:0, 18:0 and 18:1n-9. Marked differences were observed in polyunsaturated fatty acid (PUFA) content between species, with comparatively low levels of 10 and 11% of fatty acids being recorded in M. annularis and G. retiformis, respectively. The values for the other species ranged between 21 and 37%. Fatty acid composition may vary according to the proportions of fatty acids obtained from diet, algal photosynthesis and synthesis by the animal tissues.     

Habitat partitioning among four elasmobranch species in nearshore,shallow waters of a subtropical embayment in Western Australia

There is a need to explore, in an integrated and statistical manner, how the number of species, relative abundance, species composition and life-cycle stages of elasmobranchs in nearshore waters vary among habitat types and during the year. Therefore, four sites in a large marine embayment, each representing a different habitat type, were sampled at regular intervals. These sites were: (1) unvegetated, with no vegetation within at least 200 m; (2) unvegetated, immediately adjacent to sparse mangroves; (3) unvegetated, immediately adjacent to dense mangroves; and (4) vegetated, with seagrass (Posidonia australis) throughout and in the vicinity. Gill netting caught 10 shark species (5 families), 5 ray species (4 families) and 12 teleost species (10 families). Carcharhinus cautus, which contributed approximately 60% to the numbers of elasmobranchs caught, completed its life cycle in nearshore, shallow waters. Negaprion acutidens, Carcharhinus brevipinna, Carcharhinus limbatus and Rhizoprionodon acutus used these waters as a nursery area. C. cautus was caught mainly in the unvegetated sites, particularly in those near mangroves. N. acutidens was caught entirely in unvegetated sites, while R. acutus, C. brevipinna and Chiloscyllium punctatum were caught predominantly or exclusively in seagrass. The mean number of species and mean catch rate of elasmobranchs were greatest for the seagrass site and least for the unvegetated site with no vegetation within at least 200 m and were significantly less for the latter site than for the unvegetated site immediately adjacent to dense mangroves (P<0.05). The numbers of species and catch rates of elasmobranchs were significantly greater in summer and autumn than in winter (P<0.05) and, in the case of number of species, also than in spring (P<0.05). We conclude that the spatial and food resources in the nearshore, shallow waters of Shark Bay are partitioned among elasmobranch species, thus reducing the potential for competition among these species for the resources in those waters.Communicated by G.F. Humphrey, Sydney     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Nutritional quality of macrophytes eaten and not eaten by two temperatezone herbivorous fishes: A multivariate comparison

The mean annual chemical compositions (ash, lipid, carbohydrate, protein, nitrogen and carbon) of 23 species of macrophytes (22 seaweeds and 1 seagrass) from a rocky intertidal habitat on the central California coast were determined from December 1981 through December 1982. These data were used to test the hypothesis that the 13 red and green seaweeds eaten by the two principal herbivorous fishes (Cebidichthys violaceus andXiphister mucosus) at the site are higher in nutritional quality than the 9 red and brown seaweeds and the seagrass not consumed by these fishes. A MANOVA using the ash, lipid, carbohydrate and nitrogen data showed that the centroids of the dietary and nondietary species groups were significantly different. In a two-group discriminant analysis that followed, only two species were misclassified as members of the opposite group. Multigroup discriminant analysis of the 23 macrophytes resulted in some overlap among dietary and nondietary species. Species were discriminated on the first canonical axis by ash content and on the second and third axes by lipid and carbohydrate contents. Nitrogen contributed little to the overall discrimination of species in the analysis. The nondietary red algaCorallina vancouveriensis, with its high ash content and therefore relatively low nutritional quality, was clearly separated from all other species in the analysis. Brown algae were of higher nutritional quality, but are not eaten by the two fishes, possibly because these seaweeds produce indigestible carbohydrates and secondary compounds. However, the exclusion from the diets of several red algal species that were virtually indistinguishable from the dietary red algae remains unexplained.Contribution No. 64 from the Ocean Studies InstitutePlease address all correspondence and requests for reprints to Dr. Horn at Fullerton     

Biochemical and energetic composition of bathyal echinoids and an asteroid,holothuroid and crinoid from the Bahamas

The biochemical and energetic composition of body components of ten species of bathyal echinoids, and an asteroid, a holothuroid and a stalked crinoid were determined from individuals sampled from a variety of deep-water sites near the Bahamas (north Caribbean Sea) in October 1988. When compared with other studies of echinoderms, no geographic- or depth-related differences in biochemical or energetic composition were found. Body-wall tissues were composed primarily of skeletal material (mineral ash), but were comparatively high in organic material in the echinothuriid echinoids, and the asteroid and holothuroid. Gut tissues and pyloric cecae had high levels of lipid and protein, indicating their potential role in nutrient storage. Body-wall tissues were generally low in energy, but were highest in the echinoidsAraeosoma belli (7.7 kJ g^–1 dry wt) andSperosoma antillense (8.0 kJ g^–1 dry wt), the asteroidOphidiaster alexandri (8.9 kJ g^–1 dry wt), and the holothuroidEostichopus regalis (13.1 kJ g^–1 dry wt). Energy levels of gut and pyloric cecal tissues were two to three times higher than those of body-wall tissues. Total somatic tissue energy values varied greatly among species, ranging from 1.5 kJ in the echinoidAspidodiadema jacobyi to 142.1 kJ inE. regalis. As the bathyal echinoderms examined in this study occur in great abundance, they represent a significant reservoir of organic and inorganic materials and energy in deep-water benthic systems.     

Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae)

Summary Sex pheromone communication in the nine European species of small ermine moths (Yponomeuta) is reviewed in regard to the potential role of pheromones in the speciation process. Six of the nine species studied (viz.,Y. evonymellus, Y. cagnagellus, Y. padellus, Y. irrorellus, Y. plumbellus, andY. vigintipunctatus) use a mixture of (E)-11-and (Z)-11-tetradecenyl acetate in different ratios as primary pheromone components, with combinations of tetradecyl acetate, (Z)-9-tetradecenyl acetate, (Z)-11-hexadecenyl acetate and the corresponding alcohols of the acetates as additional pheromone components. Analysis of (Z)- to (E)-11-tetradecenyl acetate ratios produced by individual females of these species demonstrated significant variation among females of all species. However, the ranges of ratios produced byY. cagnagellus, Y. irrorellus, andY. plumbellus, sharing the same host-plant species, spindle tree, did not overlap. Niche separation of all six species mentioned required consideration of at least one additional pheromone component or of temporal aspects. The remaining three species,i.e. Y. malinellus, Y. mahalebellus andY. rorellus, have pheromones that differ qualitatively.Biosynthetic routes to the pheromone components identified are proposed on the basis of fatty acid pheromone precursors found in the pheromone glands. A phylogenetic tree for the genus is constructed based on allozyme frequency data and changes in pheromone composition are superimposed on this tree. We suggest that the ancestral ermine moth pheromone is a mixture of (Z)-11- and (E)-11-tetradecenyl acetate and the corresponding alcohols, and a scenario of how present-day patterns evolved is outlined. The pheromone differences among the three species using spindle tree as their host-plant might have evolved throughreproductive character displacement upon secondary contact between populations that had already diverged genetically in allopatry. Pheromone differences within the so-calledpadellus-complex (includingY. cagnagellus, Y. mahalebellus, Y. malinellus, Y. padellus, andY. rorellus) in which species might have originated sympatrically, may have evolved byreinforcing selection as these species still hybridise and produce viable offspring when confined in cages. The role of pheromones in reproductive isolation amongYponomeuta species is emphasised by (1) the function of pheromone components of some of the species as behavioural antagonists to other species, (2) the cross-attraction under experimental conditions between allochronic species with similar pheromones, and (3) the formation of hybrids in the laboratory between species that are isolated in nature by pheromone differences.