Wax-ester metabolism in the orange roughyHoplostethus atlanticus (Beryciformes: Trachichthyidae)

Freshly caught orange roughy (Hoplostethus atlanticus) obtained at the edge of the Chatham Rise, east of the South Island of New Zealand, in June 1988, were used to investigate tissue-specific lipid synthesis and to obtain blood for plasma lipoprotein analysis. Tissue slices prepared from an intraneurocranial fat deposit, pieces of skull (neurocranium), swimbladder and liver were incubated with radioactive acetate, palmitate and oleate, and the labelled lipids extracted and analysed. All four tissues could incorporate acetate label into fatty acids which were themselves incorporated into triacylglycerols and phospholipids although not into wax esters, the most abundant lipid class in these tissues. Exogenous palmitate and oleate were also incorporated into more complex lipids, with the label from oleate (but not palmitate) being found in wax esters as well as triacylglycerols and phospholipids. The distribution of the label in the wax esters showed that some reduction of the fatty acid to fatty alcohol had occurred. The only exception to this pattern was the swimbladder, which incorporated small amounts of palmitate-label into both the fatty acid and fatty alcohol portions of the wax esters. Lipoproteins were isolated from serum by centrifugation. All lipoprotein classes contained phospholipid, cholesterol, triacylglycerol, cholesteryl esters and wax esters. A very low-density lipoprotein class also contained large amounts of unesterified fatty acids, which are possibly artefacts resulting from storage of the samples. Significant quantities of-3 fatty acids were also found in the serum lipids.     

Chemical recognition of queen cells by honey bee workersApis mellifera (Hymenoptera: Apidae)

Summary Honey bee workers are able to nurse or to destroy and thus to recognize the capped queen cells containing a pupa. Fatty acid esters, especially methyl oleate, methyl palmitate and ethyl oleate were found in significant amounts on the queen pupal cuticle. Methyl oleate, the major component, along with smaller amounts of methyl linoleate and methyl linolenate, were involved in the recognition of queen cells by workers. In natural conditions of the colony, queen cells containing a paraffin pupal lure with methyl oleate were accepted 5.9 days by workers, releasing about 1.8 queen pupa equivalents during that period, when control cells (without ester) were kept only 2.1 days. Although these esters are non specific to honey bees, they are of great importance in social regulation of the honey bee colony.     

Phylogeography of surfclams,<Emphasis Type="Italic"> Spisula solidissima</Emphasis>, in the western North Atlantic based on mitochondrial and nuclear DNA sequences

The Atlantic surfclam, Spisula solidissima (Dillwyn), is broadly distributed in sandy sediments of the western North Atlantic between the Gulf of St. Lawrence and the Gulf of Mexico. In the United States, a substantial commercial fishery between Long Island and Cape Hatteras harvests offshore populations of one subspecies, S. s. solidissima. A smaller coastal form, S. s. similis Say (also known as S. s. raveneli Conrad), has a partially sympatric geographic distribution, but differs in several life-history characteristics. DNA sequence variation in mitochondrial cytochrome oxidase I (COI) and in introns at two nuclear calmodulin loci was examined to measure genetic divergence between the two subspecies and to test for population structure among populations of S. s. solidissima. Surfclams were collected from seven localities between 1994 and 2001. Based on both mitochondrial and nuclear DNA variation, the two subspecies of S. solidissima are reciprocally monophyletic, with a net COI divergence of 13.9%, indicating long-term reproductive isolation. The only significant differentiation among populations of S. s. solidissima (based on an AMOVA analysis of COI sequences) was between the Gulf of St. Lawrence and more southerly populations. A long internal branch in the S. s. solidissima genealogy coupled with low haplotype diversity in the northern-most population suggests that populations north and south of Nova Scotia have been isolated from each other in the past, with gene exchange more recently. Populations of S. s. similis from Atlantic and Gulf of Mexico coasts had a net COI divergence of 9.2%. Thus, diversification of Spisula spp. clams in the western North Atlantic involved an early adaptive divergence between coastal and offshore forms, with later barriers to dispersal emerging in the offshore form from north to south and in the coastal form between Atlantic and Gulf of Mexico populations.     

Mating expenditures reduced via female sex pheromone modulation in the primitively eusocial halictine bee, Lasioglossum (Evylaeus) malachurum (Hymenoptera: Halictidae)

The present paper reports on behavioral experiments and gas chromatographic analysis of chemical communication in the mating biology of the primitively eusocial sweat bee Lasioglossum (Evylaeus) malachurum. In a dual-choice experiment, a female made odorless was significantly less attractive than an untreated one. Attraction in L. (Evylaeus) malachurum is therefore mediated by a female-produced sex pheromone. Further bioassays showed that unmated gynes are more attractive to males than mated ones. Males are able to differentiate between the two groups of females as little as 3 h after mating. Biotests with different samples obtained from attractive gynes showed surface extracts to be most attractive. Behavioral tests with synthetic copies of the compounds identified as cuticular constituents of virgin gynes were highly attractive to males; the volatile bouquets consisting of n-alkanes, n-alkenes and iso- pentenyl esters of unsaturated fatty acids were the most attractive samples. Isopentenyl esters of unsaturated fatty acids were the key compounds in inducing male inspections as well as stimulating pounces and copulatory attempts. Virgin and nesting gynes differed clearly in the relative and absolute amounts of the volatiles on the cuticle. The total amount of volatiles was significantly higher in virgin gynes and decreased in breeding queens. Hydrocarbons were the dominant group of compounds in both groups of females. The relative amounts of the wax-type ester, hexadecyl oleate, iso- pentenyl esters, and a hitherto unidentified steroid were higher in attractive virgin gynes, while the relative proportions of hydrocarbons and lactones dominated in nesting queens. The site of sex pheromone production in attractive young L. (Evylaeus) malachurum gynes remains unknown. Head glands or Dufour's gland secretions may be involved. Another possible source of the `active principle' found among the cuticular lipids could be glandular cells of the epidermis. The significance of modulation of female sex pheromone composition is discussed in terms of a reduction in mating expenditures. Received: 30 April 1998 / Accepted after revision: 24 July 1998     

Biosynthesis of wax esters in oceanic crustaceans

Slices from the hepatopancreas of various oceanic curstaceans incorporated radioactivity into wax esters from ¹⁴C glucose and ¹⁴C aspartic acid to a lesser extent and from ¹⁴C palmitic acid to a much greater extent. Radioactivity was incorporated from ¹⁴C palmitic acid into both fatty acid and fatty alcohol moieties of wax esters, the percentage of total radioactivity present in alcohol moieties being greater in deep-living than in shallow-living species. Cell-free preparations from the hepatopancreas but not from muscle, supplemented with ATP and reduced pyridine nucleotides, incorporated radioactivity from 1-¹⁴C palmitoyl Coenzyme A into both fatty alcohol and fatty acid moieties of wax esters. Incorporation into fatty alcohol was NADPH- rather than NADH-specific. Preparations from deep-living species had a greater percentage of total radioactivity in the fatty alcohol moieties of wax esters than preparations from shallow-living species. We conclude that the level of wax esters in a given species is correlated with the rate at which the species biosynthesises these lipids de novo; deep-living species have higher rates of wax ester biosynthesis and higher levels of wax esters than shallow-living species. The results support the thesis that wax esters in oceanic crustaceans are derived largely from the animals' internal biosynthetic activities, presumably in response to particular biochemical and/or physiological requirements, rather than from their diets.     

A functional interpretation of cilia and mucocyte distributions on the abfrontal surface of bivalve gills

The lack of fundamental data on the abfrontal surface of bivalve gills has prompted a comparative study of cilia and mucocytes on this surface, using scanning electron microscopy and histology on eight species of bivalves, representing seven families and the four major gill types: Mytilus edulis, Modiolus modiolus, Arca zebra, Placopecten magellanicus, Crassostrea virginica, Spisula solidissima, Mercenaria mercenaria, and Mya arenaria. Abfrontal cilia and mucocytes were found in all species studied, with types and densities differing within and between gill types. The three species of homorhabdic filibranchs presented different densities of abfrontal cilia and mucocytes, from very dense in M. edulis to sparse in A. zebra. The heterorhabdic gills had intermediate cilia and mucocyte densities, with highest concentrations of both abfrontal cilia and mucocytes on the principal filaments. The eulamellibranchs showed low ciliary densities together with high mucocyte densities, especially in S. solidissima, where the abfrontal mucocytes were glandular. These results indicate that: (1) the abfrontal surface is a vestigial mucociliary surface; (2) the abfrontal surface cannot participate in water pumping in most species, due to low ciliary densities; and (3) species with high densities of abfrontal mucocytes could utilize abfrontal mucus to reduce drag, especially in the highly fused gills, such as those of the eulamellibranchs. The differing distributions of abfrontal cilia and mucocytes may reflect different selective pressures acting on the gills within the various taxa. Received: 12 February 2000 / Accepted: 10 September 2000     

The fate of dietary lipids in the Arctic ctenophore Mertensia ovum (Fabricius 1780)

Lipids of the Arctic ctenophore Mertensia ovum, collected from Kongsfjorden (Svalbard) in 2001, were analysed to investigate seasonal variability and fate of dietary lipids. Total lipids, lipid classes and fatty acid and alcohol compositions were determined in animals, which were selected according to age-group and season. Changes in lipids of age-group 0 animals were followed during growth from spring to autumn. Total lipids increased from May to September. Lipids as percentage of dry mass were lowest in August indicating their use for reproduction. Higher values occurred in September, which may be due to lipid storage for overwintering. Wax esters were the major lipid class accounting for about 50% of total lipids in age-group 0 animals from July and August. Phospholipids were the second largest lipid fraction with up to 46% in this age-group. The principal fatty acids of M. ovum from all age-groups were 22:6(n-3), 20:5(n-3) and 16:0. Wax ester fatty alcohols were dominated by 22:1(n-11) and 20:1(n-9) followed by moderate proportions of 16:0. The unique feature of M. ovum lipids was the high amount of free fatty alcohols originating probably from the dietary wax esters. In May, free alcohols exhibited the highest mean proportion with 14.6% in age-group 0 animals. We present the first data describing a detailed free fatty alcohol composition in zooplankton. This composition was very different from the alcohol composition of M. ovum wax esters because of the predominance of the long-chain monounsaturated 22:1(n-11) alcohol accounting for almost 100% of total free alcohols in some samples. The detailed lipid composition clearly reflected feeding of M. ovum on the herbivorous calanoid species, Calanus glacialis and C. finmarchicus, the abundant members of the zooplankton community in Kongsfjorden. Other copepod species or prey items seem to be less important for M. ovum.     

Studies on the mechanism of biosynthesis of wax esters in Euchaeta norvegica

Preparations of Euchaeta norvegica catalyse the incorporation of (1-¹⁴C) hexadecanol, (9,10-³H₂) oleic acid, (U-¹⁴C) glucose, (U-¹⁴C) alanine and (U-¹⁴C) aspartic acid into wax esters. Both the fatty alcohol and fatty acid moieties of the wax esters are labelled, indicating de novo biosynthesis. A limited interconversion of fatty acid and fatty alcohol has been demonstrated. Incorporation of glucose into wax esters is diminished by prior starving of E. norvegica; it is also reduced in the presence of ecdysterone. Formation of wax esters from glucose is inhibited by increased oxygen tensions, rotenone and dinitrophenol, and is relatively insensitive to decreased oxygen tensions and cyanide. The data are discussed with particular reference to the reasons why wax esters are biosynthesised by calanoids.     

Ecological investigations on the zooplankton community in balsfjorden,northern Norway: Lipids and fatty acids in Meganyctiphanes norvegica,Thysanoessa raschi and T. inermis during mid-winter

Total lipid of Meganyctiphanes norvegica (M. Sars) contained 53% triacylglycerols and traces of wax esters, that of Thysanoessa raschi (M. Sars) contained 44% triacylglycerols and 10% wax esters and that of T. inermis (Krøyer) contained 28% triacylglycerols and 40% wax esters. The triacylglycerols of M. norvegica were relatively rich in 20:1 and 22:1 fatty acids and its traces of wax esters resembled those of calanoid copepods. The triacylglycerols of both Thysanoessa species were deficient in 20:1 and 22:1 fatty acids but were richer in 16:1(n-7) and 18:1 (n-7) acids than those of M. norvegica. The wax esters of T. raschi contained phytol as almost the only fatty alcohol and were rich in 16:0 and 18:1 (n-9) fatty acids. The wax esters of T. inermis contained mainly 16:0 and 14:0 fatty alcohols with lesser amounts of phytol and their dominant fatty acid was 18:1, especially the (n-9) isomer. The triacylglycerols of T. inermis had 18:4 (n-3) as the major polyunsaturated fatty acid. From these and other aspects of fatty acid and fatty alcohol analyses it is concluded that a major foodstuff of M. norvegica in Balsfjorden is wax ester-rich calanoid copepods. T. raschi and especially T. inermis are concluded to have much more preference for phytoplanktonic food. Results are discussed in terms of current knowledge of the lipid chemistry of krill in the northern and southern hemispheres.     

Metabolic relationships between fatty alcohol and fatty acid in the liver of Squalus acanthias

The high speed supernatant fraction of a homogenate of Squalus acanthias L. liver catalyses the oxidation of oleyl alcohol to oleic acid by an NAD-dependent process. Reduction of fatty acid to fatty alcohol could not be demonstrated. Liver lipids of S. acanthias contain about 60 and 40% of triglycerides and diacyl glyceryl ethers, respectively, with only traces of wax esters. Serum lipids contain 28, 15 and 26% of triglycerides, diacyl glyceryl ethers and wax esters, respectively. Cell-free fractions of S. acanthias liver catalyse the formation of wax esters by an ATP¹-dependent process. The bulk of wax ester synthesis occurs in the high-speed supernatant fraction of liver. The rate of formation of the ester bond in wax esters is comparable to the rate of synthesis of ester bonds in triglycerides, and greatly exceeds the rate of formation of the ether bond in diacyl glyceryl ethers. Results are discussed in terms of possible factors controlling the levels of neutral lipids in S. acanthias.     

Hydrolysis of a model surfactant as measured using ACYL coenzyme a synthetase

Hydrolysis of a model nonionic surfactant, [1‐¹⁴C]methyl palmitate, was compared between porcine esterase and lipase using a new hydrolase assay. The assay incorporates acyl coenzyme A (CoA) synthetase to convert the hydrolytic product of methyl palmitate, palmitic acid, to its acyl CoA derivative; palmitoyl CoA is separated from unreacted substrate for quantitation by a highly efficient extraction. The assay achieves quantitative separation between product and substrate due to the high water solubility of the acyl CoA derivative, eliminating the need for time‐consuming chromatographic separations. After 60 min under optimal conditions, only 20 U/mL porcine hepatic esterase hydrolyzed 93.6+0.9% of 20μM methyl palmitate, while 100U/mL porcine pancreatic lipase was required to hydrolyze only 82.3 ±0.7% of the same substrate. While both enzymes detoxified the surfactant, esterase was more efficient, possibly indicating preferential specificity for simple monoesters; generally selective for endogenous triacylglycerols, lipase may be less specific for surfactants. However, together both enzymes may enable mammals to hydrolyze ingested nonionic surfactants from oil spill dispersants, reducing their toxicity.     

Lipogenesis in the arctic euphausiid Thysanoessa inermis

Biosynthesis of lipids by Thysanoessa inermis collected from Balsfjorden, northern Norway, in May 1980, was examined in vitro. The highest concentration of lipid within the krill was in the hepatopancreas, and this organ was the most active in esterifying free fatty acids into wax esters. The hepatopancreas (i.e., thoracic contents) incorporated (¹⁴C) glucose, (¹⁴C) alanine and ³H₂O into wax esters, with the fatty alcohol moieties being labelled more than the fatty acids. (¹⁴C) fatty acid was incorporated preferentially into the fatty acid moieties of wax esters, this incorporation being markedly stimulated by free fatty alcohol. It is concluded that the fatty alcohols of wax esters are preferentially biosynthesized de novo from dietary protein and carbohydrates, whereas the fatty acids derive preferentially from dietary lipid. On the basis of ³H incorporated from ³H₂O, the hepatopancreas in a 50 mg II-group (2 yr old) individual of T. inermis is capable of biosynthesizing de novo, approximately 0.1 mg of lipid (as fatty acids) per day at 5°C.     

Naticid snail predation on early post-settlement surfclams (<Emphasis Type="Italic">Spisula solidissima</Emphasis>) on the inner continental shelf of New Jersey,USA

Naticid gastropod predators leave characteristic boreholes in the shells of their bivalve prey that allow the evaluation of spatial differences in occurrence and intensity of predation. This approach, extensively used in paleobiological research, was used to compare spatial variation in predation by moonsnails (Euspira heros Say) on initial recruits of the Atlantic surfclam (Spisula solidissima Dillwyn). A four-year sampling at two 12-m deep stations on either flank of Beach Haven Ridge on the inner continental shelf detected temporal and spatial differences in surfclam densities following a large settlement pulse in early July. Between-station density differences were large during the first peak in surfclam density, but decreased in August-September, when densities at both stations also decreased. Surfclam seasonal peaks were followed by peaks in density of articulated shells (recent mortality due to non-crustacean predators and/or intrinsic mortality) and surfclams with boreholes (mortality due to naticids). The proportion of surfclams with boreholes was consistently higher (more than twice, on average) at the station where the highest densities of surfclams were also detected. These spatial differences are interpreted as differences in predation intensity resulting from a corresponding higher abundance of moonsnails at the same station. Although naticid predation is not the primary source of surfclam mortality, it consistently contributes to the reduction of spatial differences in density initially created by dissimilar levels of larval surfclam settlement.     

Ontogenetic and seasonal changes in lipid and fatty acid/alcohol compositions of the dominant Antarctic copepods Calanus propinquus,Calanoides acutus and Rhincalanus gigas

Lipid compositions of the dominant Antarctic copepods Calanoides acutus, Rhincalanus gigas and Calanus propinquus from the Weddell Sea have been investigated in great detail. Copepods were collected during summer in 1985 and late spring/early winter in 1986. The analyses revealed specific adaptations in the lipid biochemistry of these species which result in very different lipid components. The various copepodite stages of C. acutus synthesize wax esters with long-chain monounsaturated moieties and especially the alcohols consisted mainly of 20:1(n-9) and 22:1(n-11). R. gigas also generates wax esters, but with moieties of shorter chain length. The fatty alcohols consisted mainly of 14:0 and 16:0 components, while the major fatty acids were 20:5, 18:4 and 22:6, of which 18:4 probably originated from dietary input. In contrast, C. propinquus accumulates triacylglycerols, a very unusual depot lipid in polar calanoid copepods. Major fatty acids in C. propinquus were the long-chain monounsaturates 22:1(n-9) and 22:1(n-11), which may comprise up to 50% of total fatty acids. In C. acutus and C. propinquus there was a clear increase of long-chain fatty acids with increasing developmental stage. In contrast, the fatty acid and alcohol composition of the R. gigas copepodite stages were characterized by the dominance of the polyunsaturated fatty acids as well as high amounts of the monounsaturates 18:1(n-9) and 16:1(n-7). There was a considerable decrease of the dietary fatty acid 18:4(n-3) towards the older stages during summer; in late winter/early spring 18:4 was only detected in very low amounts. This tendency was also found in the other two species, but was less pronounced. In all three species dry weight and lipid content increased exponentially from younger to older stages. The highest portion of wax esters, or of triacylglycerols in C. propinquus, was found in the adults. Dry weight and lipid content were generally higher during summer. In late winter/early spring the variability was more pronounced and lipid-rich specimens showed a selective retention of long-chain monounsaturated fatty acids, whereas in lipid-poor specimens these fatty acids were very much depleted.     

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

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

Diurnal lipid and mucus production in the staghorn coral Acropora acuminata

Net ¹⁴C-accumulation into lipids of Acropora acuminata was rapid and increased with light intensity. Dark ¹⁴C-incorporation was less than 1% noon maximum. Structural lipids were the first radioactively labelled lipid types showing linear ¹⁴C-uptake kinetics. Storage lipids showed non-linear, power-curve kinetics for ¹⁴C-uptake. The rate of ¹⁴C-incorporation into triglycerides and wax esters was maximal during early afternoon and at midday, respectively. Electron microscopic evidence is given for zooxanthellae being primary sites for synthesis of lipids which are exuded from chloroplasts and transferred to animal tissues. Free lipid droplets and crystalline inclusions (wax ester) were common in animal tissues, the inclusions being often associated with mucus-producing cells. The diurnal rate of mucus production was constant. However, ¹⁴C-mucus-lipid production showed a light-dependent diurnal pattern and accounted for 60 to 90% total ¹⁴C of mucus during periods of photosynthetically-saturating light. Here, ¹⁴C was primarily associated with wax esters which were always present in the mucus-lipid. ¹⁴C-triglycerides occur in mucus released only during the day. Lipid and mucus synthesis is discussed in relation to the carbon budget of A. acuminata, in which mucus represented a loss of 40% net C fixation.     

Distribution of a Community of Mammals in Relation to Roads and Other Human Disturbances in Gabon,Central Africa

We present the first community‐level study of the associations of both roads and other human disturbances with the distribution of mammals in Gabon (central Africa). Our study site was in an oil concession within a littoral mosaic landscape. We conducted surveys along 199 line transects and installed camera traps on 99 of these transects to document mammal presence and abundance. We used generalized linear mixed‐effect models to document associations between variables related to the ecosystem (land cover, topography, and hydrology), roads (coating, width of rights of way, condition, type of vehicle used on the road, traffic level, affiliation of users, and general type of road), and other human disturbances (urbanization, agriculture, hunting, logging, gathering, and industrial activities) and the abundance or presence of 17 species or groups of mammals including elephant (Loxodonta cyclotis), buffalo (Syncerus caffer), sitatunga (Tragelaphus spekei), red river hog (Potamochoerus porcus), smaller ungulates, gorilla (Gorilla gorilla), chimpanzee (Pan troglodytes), side‐striped jackal (Canis adustus), carnivores, monkeys, and large rodents. Some types of roads and other human disturbances were negatively associated with the abundance or presence of elephants, buffalos, gorillas, sitatungas, some monkeys, and duikers. The pattern of associations of mammals with roads and other human disturbances was diverse and included positive associations with road presence (red river hog, some monkeys, and duikers), agriculture (sitatunga, small carnivores, and large rodents) and industrial activities (sitatunga, red river hog, red duikers, and side‐striped jackal). Our results suggest that the community of mammals we studied was mostly affected by hunting, agriculture, and urbanization, which are facilitated by road presence. We recommend increased regulation of agriculture, hunting, and road building in the area. Distribución de una Comunidad de Mamíferos en Relación a Carreteras y Otras Perturbaciones Humanas en Gabón, Africa Central     

Importance of wax esters and other lipids in the marine food chain: Phytoplankton and copepods

Wax esters, which function as reserve fuels, account for 25 to 40% of the lipid of the pelagic copepod Calanus helgolandicus (Copepoda, Calanoida). In laboratory experiments with these crustaceans, diatoms (Lauderia borealis, Chaetoceros curvisetus, and Skeletonema costatum) and dinoflagellates (Gymnodinium splendens), which contained no wax esters, were used as food. Changes in the food concentration affected both the amount of lipid and the composition of the wax esters. Since the fatty acids of the triglycerides and wax esters of C. helgolandicus resembled the dietary fatty acid composition, it appeared that copepods incorporated their dietary fatty acids largely unchanged into their wax esters. The polyunsaturated alcohols of the wax esters did not correspond in carbon numbers or degrees of unsaturation to the dietary fatty acids. We postulate two different metabolic pools to explain the origin of these long chain alcohols. The phospholipid fatty acids were not affected by changes in the amount or type of food, probably because of their structural function.     

Age-structure, recruitment, and adult mortality in populations of the Atlantic surfclam, Spisula solidissima, from 1978 to 1997

The Atlantic surfclam, Spisula solidissima (Dillwyn, 1817), is a dominant member of the benthic community on the continental shelf from Georges Bank to North Carolina, USA. This bivalve has supported a major fishery, primarily off New Jersey and the Delmarva Peninsula, since the 1960s. Early papers documented that these populations were at historical lows in the mid-1970s owing to commercial harvesting and a hypoxic event off New Jersey. It was also shown that major recruitment took place off New Jersey in 1976 and off the Delmarva Peninsula in 1977. Because the size frequencies of surfclams from federal surveys do not show distinct year classes, there has been uncertainty about the number of year classes in these populations throughout the 1980s and 1990s. The present study describes changes in population age- and size-structure from 1978 to 1997 in federal waters (≥5.5 km from shore) of the USA. Given the 30 to 35 year life span of S. solidissima, these populations could be composed of many year classes. Yet, these populations were composed of only two to three year classes in 1978. Through annual recruitment, the number of year classes increased over time, and populations off New Jersey and the Delmarva Peninsula contained at least 19 year classes in 1997. This major change in population structure over time was not evident from examination of available size-frequency data, and could only be inferred from data on age-composition. Throughout the 1980s and 1990s, the surfclam fishery was supported by multiple year classes. The mean and variance of recruitment to the New Jersey region, as indicated by the abundance of 4-year-olds over time, was greater than that off Delmarva, particularly between 1980 and 1986. The instantaneous rate of adult mortality, which includes the effect of harvesting, was approximately 0.26 yr⁻¹ in each region. Received: 11 September 1998 / Accepted: 1 February 1999     

Release of sex pheromone and its precursors in the pine sawfly Diprion pini (Hym., Diprionidae)

Summary. The first identification of a sex pheromone of a pine sawfly (Hymenoptera, Diprionidae) dates back almost thirty years. Since then, female-produced pheromones of over twenty diprionid species have been investigated by solvent extraction followed by separation and identification. However, no study has shown what the females actually release. Collection of airborne compounds using absorbtion on charcoal filter as well as solid phase microextraction (SPME) followed by analysis employing gas chromatography combined with mass spectrometry (GC-MS), revealed an unusual system in Diprion pini, in which the pheromone precursor alcohol, 3,7-dimethyl-2-tridecanol, is released together with acetic, propionic, butyric and isobutyric acids. The corresponding acetate, propionate and butyrate esters of 3,7-dimethyl-2-tridecanol were also found in the samples. All esters were electrophysiologically active, and the propionate and isobutyrate were attractive in trapping experiments. Based on these and earlier reported results, it seems that at least in part of its range, the pheromone response of D. pini is not very specific with regard to the functional group, as long as this is an ester.