Relationship between zooplankton distribution,geographic characteristics and hydrographic patterns off the Catalan coast (Western Mediterranean)

Astract Northern anchovy larvae, Engraulis mordax, were collected on five cruises covering the Southern California Bight, one cruise in April 1984, four cruises in January, February and May 1986. The amounts of triacylglycerol, cholesterol and polar lipid were measured in individual anchovy larvae as indicators of their nutritional condition. There was a significant difference in each of the size-specific lipid components between the stations for four of the five cruises. This indicates that all the ocean habitats within a cruise were not equal in promoting growth (and presumably survival) in the anchovy larvae. There were also differences between cruises, where the percentage of the larvae judged to be in poor condition varied from 8 to 27%. Canonical correlation analyses revealed a significant relationship in most cases between the larval fish parameters and the physical and biological data collected from each occupied station. Cholesterol and triacylglycerol were the most important larval components, and the short-term energy storage component in the copepod Calanus pacificus was the most important station component. The data, however, are not sufficient to identify and characterize stations in the field which produce anchovy larvae in good condition.     

Nutritional condition and growth rate of anchovy larvae (Engraulis mordax) in the California Current: two contrasting years

A large number of northern anchovy larvae, Engraulis mordax, were collected in the Southern California Bight on two CalCOFI cruises during spring 1987, and the amounts of triacylglycerol, cholesterol and polar lipids were measured in each individual larva using the latroscan TLC/FID (thin-layer chromatography/flame ionization detection) system. The 1987 larvae were compared to larvae taken in the Bight during the previous year. Canonical correlation analyses revealed a significant relationship between the larval lipid components and physical/biological data collected at each occupied station during the late 1987 spring cruise, similar to that recorded in 1986. The short-term nutritional condition of the copepod Calanus pacificus was the most important station variable in both cruises. Their triacylglycerol:cholesterol ratio indicated that few of the 1987 larvae had been starving, but their very low contents of length-specific cholesterol and polar lipid indicated that they had been growing very slowly. Otolith readings on defatted larvae confirmed that the 1987 larvae had been growing significantly more slowly than the 1986 larvae. Furthermore, the data suggested that in the 1986 population the faster growing larvae were also those with the highest lipid content. This situation was reversed for the 1987 larvae, when the faster growing larvae in the population seemed to achieve their faster growth only at the expense of a lowered lipid content. The abundance of anchovy larvae was much higher in spring 1987 than in spring 1986 but, despite this, the apparent recruitment (number of Age 0 fish in July of each year) was not higher in 1987.     

Developmental changes in the composition of myofibrillar proteins in the swimming muscles of Atlantic herring,Clupea harengus

Changes in myofibrillar protein composition during development have been investigated in the swimming muscles of the Atlantic herring Clupea harengus L. using a range of electrophoretic techniques. The main muscle-fibre type of larvae, and the fast- and slow-muscle fibres of adult fish were found to contain distinct isoforms of myosin heavy chain (MHC) and myosin light chain 2 (LC2). Larval LC2 was present as a minor component of adult fast-muscle myosin. In contrast, larval and adult fast-muscle myosin appeared to contain identical alkali light chains. Tropomyosin and troponin C were also identical in larval and in adult fast-muscle. All three muscle-fibre types contained unique isoforms of troponin T (TNT) and troponin I (TNI). Larval muscle had multiple isoforms of TNT, some of which may correspond to embryonic forms. It was concluded that although the main muscle-fibre type in larvae shares some myofibrillar proteins with adult fast muscle, it also contains characteristic isoforms of MHC, TNI, TNT and LC2 and therefore represents a distinct fibre type. The particular combination of myofibrillar proteins present at any developmental stage was found to be dependent on the rearing temperature. For example, a higher proportion of embryonic TNT isoforms were present at hatching in larvae reared at 5°C than at either 10 or 15°C. Over a period of 7 d, there was a gradual reduction in the number of TNT isoforms, but the pattern in 5°C larvae after 7 d still did not resemble that in 1 d-old larvae reared at 15°C.     

A correction to: biological and oceanographic insights from larval labrid (Pisces: Labridae) identification using mtDNA sequences

Previously published mtDNA cytochrome b sequences for Xyrichtys novacula (Linnaeus) and two morphologically distinct types of Xyrichtys larvae (Pisces: Labridae) included human cytochrome b sequence, presumably due to sample contamination and/or poor preservation of sample DNA. Those sequences had been used to identify the two types of Xyrichtys larvae as X. novacula, but owing to the contamination, the identifications are invalid. Fresh specimens were collected: X.␣martinicensis (Cuvier and Valenciennes), X. novacula, X. splendens (Castelnau), and Xyrichtys sp. larvae of each of the two morphotypes (dorsal-forward eye and ventral-forward eye). The cytochrome b fragment was amplified from each specimen using the polymerase chain reaction. Comparison of the sequences with human cytochrome b sequence confirmed that the new sequences were not contaminated. The interspecific differences in Xyrichtys sequences were less than previously reported, but still greater than observed in many fish genera. Distance and parsimony analyses indicated that X. novacula and X. martinicensis were more closely related to each other than to X. splendens. This conclusion differs from our previous conclusion that X. martinicensis and X. splendens were more closely related to each other than to X. novacula. Distance and parsimony analyses also demonstrated that both larval morphotypes were X. novacula and, thus, our previous conclusions regarding larval ecology and oceanographic transport remain unchanged. Received: 12 March 1997 / Accepted: 24 October 1997     

Distribution and seasonal abundance of Polygordius spp. (Class: Polychaeta; Family: Polygordiidae) exo- and endolarvae in the southern Mid-Atlantic Bight,USA

The importance life history plays in understanding population dynamics and the functional roles of species for predicting climate change scenarios are well established. Yet, in the marine environment, the complete life history is unknown for many species, especially the link between morphologically and ecologically distinct planktonic larvae, and their corresponding benthic adult forms. Integration of meroplankton abundance, benthic adult species, larval morphology, and molecular data was employed to unravel the complete life history of Polygordius, a dominant polychaete in sandy shelf sediments of the Mid-Atlantic Bight. Polygordius species are unusual, having two distinct planktonic larval forms: an exolarva and an endolarva. Extensive sampling in the southern Mid-Atlantic Bight with careful preservation of meroplankton (2006–2007) revealed the identity of multiple stages of exo- and endolarvae, and their spatial, seasonal, and vertical distribution. Molecular and morphological evidence indicated exolarvae are Polygordius jouinae and endolarva are an undescribed species. Structure and development of these larvae differed greatly. P. jouinae exolarvae were found off Delaware Bay to North Carolina. At some stations, they were abundant, with densities up to 4,013 m^?3, comprising >90 % of the total meroplankton. Exolarvae spent up to a month in the plankton starting in March/May depending on year, settlement began in July when larvae were at least 2 mm in length, and by October were no longer observed in the plankton. These findings are consistent with the distribution patterns and life cycle known for adults. This is the first report of endolarvae north of Cape Hatteras.     

Variation in pelagic larval growth of Atlantic billfishes: the role of prey composition and selective mortality

Atlantic blue marlin (Makaira nigricans) and sailfish (Istiophorus platypterus) larvae were collected from 10 monthly cruises (June–October 2003 and 2004) across the Straits of Florida to test (1) whether growth differed between the more productive western region near the Florida shelf, and the less productive eastern region toward the Bahamas, and (2) whether growth was related to prey consumption. Examination of larval sagittal otoliths revealed that instantaneous growth and daily growth during the first 2–3 weeks of life did not vary significantly between the two regions for either species. However, recent growth during the last two full days prior to collection was greater in the west for blue marlin larvae. Recent growth of blue marlin larvae <9 mm SL (primarily zooplanktivorous) was significantly related to prey composition (faster growth when higher proportions of Farranula copepods were consumed). Western larvae grew faster and had higher proportions of Farranula in their guts. Trends for sailfish larvae were not significant. In both species, comparison of early growth between <9 and ≥9 mm SL size groups indicated that growth trajectories diverged around 5–8 mm SL, the time when billfish larvae become capable of piscivory. Significantly faster growth of larger (older) larvae suggests that mortality was selective for fast growers and that the transition to piscivory may be a critical point in the early life of billfish.     

Does cannibalism in Spodoptera frugiperda (Lepidoptera: Noctuidae) reduce the risk of predation?

The incidence of cannibalism of larval Spodoptera frugiperda (Lepidoptera: Noctuidae) on maize under field conditions was investigated using field cages. Cannibalism was found to account for approximately 40% mortality when maize plants were infested with two or four fourth-instar larvae over a 3-day period. Field trials examined the effect of larval density on the prevalence of natural enemies of S. frugiperda. The abundance of predators (earwigs, staphylinids, other predatory beetles, and Chrysoperla spp.) was significantly greater on maize plants with higher levels of larval feeding damage, while the relationship between predator abundance and number of S. frugiperda larvae per plant was less clear. As larval damage is probably a more reliable indicator of previous larval density than numbers collected at an evaluation, this indicates that predation risk will be greater for larvae living in large groups. Parasitism accounted for 7.1% mortality of larvae in sorghum, and involved six species of Hymenoptera and Tachinidae. There was no effect of larval density or within-plant distribution on the probability of larval attack by parasitoids. The selective benefits of cannibalism, in relation to the risk of predation and parasitism, are discussed. Received: 23 March 2000 / Accepted: 24 June 2000     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Feeding by larvae of two different developmental modes in Streblospio benedicti (Polychaeta: Spionidae)

Females of the spionid polychaete Streblospio benedicti (Webster) produce either small eggs (60–70 μm diameter) and planktotrophic larvae, or large eggs (100–200 μm) and lecithotrophic larvae that reportedly do not feed. This intraspecific polymorphism, a form of poecilogony, is potentially useful in studies of larval ecology and evolution, but necessary data on larval form and function are lacking. This study describes the morphology and nutritional biology of larvae obtained from Atlantic (South Carolina) and Pacific (California and Washington) populations from 2003 to 2005. The two types of larvae produced by Atlantic S. benedicti differed greatly in length (229±22 μm SD for planktotrophs vs. 638±40 μm for lecithotrophs) and chaetiger number (2–5 vs. 10–11) at release from the female’s brood pouch. Planktotrophic larvae bore long provisional chaetae on their first chaetiger; provisional chaetae were absent in lecithotrophic larvae. Larvae from Pacific populations were all of the lecithotrophic form, and were similar to their Atlantic counterparts in all respects. High-speed video microscopy revealed that both types of larvae used opposed bands of cilia to capture suspended particles and transport them to the mouth, where they were often ingested. Lecithotrophic larvae reared with suspended phytoplankton (Rhodomonas sp., 10⁴ cells ml⁻¹) for 2 days grew significantly faster than sibling larvae reared without added food, indicating that these larvae can digest and assimilate ingested food. Larvae of S. benedicti that develop from large eggs are thus facultative planktotrophs instead of obligately non-feeding lecithotrophs, a result that affects the interpretation of comparative studies of the ecology and evolution of larvae in S. benedicti and certain other marine invertebrates.     

Advanced age for sexual maturity in the ocean quahog Arctica islandica (Mollusca: Bivalvia)

A study of gonadal state in 39 small specimens of Arctica islandica Linné collected in April 1976 from the Middle Atlantic shelf of North America suggests that sexual maturity is reached at a later age than has been reported for other shelf bivalves. Eight of the specimens had gonads in the undifferentiated state. On the basis of annual internal growth banding in the shells of these immature specimens, age ranged between 4 and 14 years. Average age was 9.38 years (standard deviation, s,= 3.54). Shell length ranged from 24 to 47 mm (average 38.85 mm, s=9.25). Differentiated gonads were found in specimens as young as 6 years. Age of maturity shows a wide range and may be dependent upon growth rate and locality.     

Inheritance of a nuclear DNA polymorphism assayed in single bivalve larvae

We examined the inheritance of theMytilus edulis CaM-1 Intron 3 locus, a non-coding DNA locus with two potentially neutral length-variants. The polymerase chain reaction (PCR) was used to determine theCaM-1 genotype for 799 larvae obtained from 11 laboratory crosses. Larvae were typed singly only 8 to 24 h after fertilization. We find evidence that each allele can be inherited from either sex, that there are no barriers to fertilization between gametes of different genotypes, and that most larvae have genotypes compatible with Mendelian inheritance of the locus. Deviations from expected genotype frequencies were found in some crosses; we suggest that a contributing factor is aneuploidy in early larvae, but a major cause is more likely to be selection at a locus linked toCaM-1, occurring under the artificial laboratory culture conditions. This study demonstrates the feasibility of applying molecular genetic techniques to early larval stages of marine bivalves, and presents a new non-destructive biopsy method for DNA analysis from living adult mussels.     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

Anguilliform larvae collected off North Carolina

The distinctive larval stage of eels (leptocephalus) facilitates dispersal through prolonged life in the open ocean. Leptocephali are abundant and diverse off North Carolina, yet data on distributions and biology are lacking. The water column (from surface to 1,293 m) was sampled in or near the Gulf Stream off Cape Hatteras, Cape Lookout, and Cape Fear, North Carolina during summer through fall of 1999–2005, and leptocephali were collected by neuston net, plankton net, Tucker trawl, and dip net. Additional samples were collected nearly monthly from a transect across southern Onslow Bay, North Carolina (from surface to 91 m) from April 2000 to December 2001 by bongo and neuston nets, Methot frame trawl, and Tucker trawl. Overall, 584 tows were completed, and 224 of these yielded larval eels. The 1,295 eel leptocephali collected (combining all methods and areas) represented at least 63 species (nine families). Thirteen species were not known previously from the area. Dominant families for all areas were Congridae (44% of individuals, 11 species), Ophichthidae (30% of individuals, 27 species), and Muraenidae (22% of individuals, ten species). Nine taxa accounted for 70% of the overall leptocephalus catches (in order of decreasing abundance): Paraconger caudilimbatus (Poey), Gymnothorax ocellatus Agassiz complex, Ariosoma balearicum (Delaroche), Ophichthus gomesii (Castelnau), Callechelys muraena Jordan and Evermann, Letharchus aliculatus McCosker, Rhynchoconger flavus (Goode and Bean), Ophichthus cruentifer (Goode and Bean), Rhynchoconger gracilior (Ginsburg). The top three species represented 52% of the total eel larvae collected. Most leptocephali were collected at night (79%) and at depths > 45 m. Eighty percent of the eels collected in discrete depth Tucker trawls at night ranged from mean depths of 59–353 m. A substantial number (38% of discrete depth sample total) of larval eels were also collected at the surface (neuston net) at night. Daytime leptocephalus distributions were less clear partly due to low catches and lower Tucker trawl sampling effort. While net avoidance may account for some of the low daytime catches, an alternative explanation is that many species of larval eels occur during the day at depths > 350 m. Larvae of 21 taxa of typically shallow water eels were collected at depths > 350 m, but additional discrete depth diel sampling is needed to resolve leptocephalus vertical distributions. The North Carolina adult eel fauna (estuary to at least 2,000 m) consists of 51 species, 41% of which were represented in these collections. Many species of leptocephali collected are not yet known to have juveniles or adults established in the South Atlantic Bight or north of Cape Hatteras. Despite Gulf Stream transport and a prolonged larval stage, many of these eel leptocephali may not contribute to their respective populations.     

Population genetics of the blue crabCallinectes sapidus: modest population structuring in a background of high gene flow

To determine the genetic population structure of blue crabs (Callinectes sapidus Rathbun), electrophoretic allozyme analysis was performed on 750 individuals collected from 16 nearshore locations ranging from New York to Texas, USA. Twenty enzymes and non-enzymatic proteins coded by 31 presumptive loci were examined. Twenty-two loci were either monomorphic or polymorphic at less than theP ₉₅ level; alleles for these polymorphic loci were geographically dispersed. Allele frequencies for three of the remaining polymorphic loci were homogeneous over all populations, as were levels of polymorphism and heterozygosity. Phenograms generated by the UPGMA (unweighted pair-group method using arithmetic averages) and distance Wagner methods exhibited no geographic pattern in the clustering of populations. Estimates ofN _em (effective number of migrants per generation between populations) indicated substantial gene flow, with aalues sufficiently high to infer panmixia between all blue crab populations from New York to Texas. However, despite this high level of gene flow, two striking patterns of geographic differentiation occurred: genetic patchiness and clinal variation. Allele frequencies atEST-2, GP-1, IDHP-2, DPEP-1, DPEP-2, andTPEP exhibited genetic patchiness on local and range-wide geographic scales, and allele frequencies atEST-2 varied temporally. Genetic patchiness in blue crabs is likely to be the result of the pre-settlement formation and subsequent settlement of genetically heterogeneous patches of larvae; allele frequencies of those larval patches may then be further modified through ontogeny by localized selection. In the Atlantic Ocean, a regional latitudinal cline ofEST-2 allele frequencies was superimposed on the range-wide genetic patchiness exhibited by that locus. This pattern against a background of high gene flow is highly likely to be maintained by selection. In estuaries along the Atlantic Ocean coast, a combination of low adult long-distance migration and a high retention rate of locally spawned larvae could serve to segregate populations and allow for the development of the geographic cline inEST-2. The Gulf of Mexico showed no apparent cline, perhaps due to long-distance migration of females in some regions of the Gulf, or to masking by genetic patchiness. These results emphasize the importance of both ecological and evolutionary time scales and structuring mechanisms in determining genetic population structure.     

Feeding ecology of Greenland halibut and sandeel larvae off West Greenland

Feeding ecology of Greenland halibut (Gr. halibut) (Reinhardtius hippoglossoides) and sandeel (Ammodytes sp.) larvae on the West Greenland shelf was studied during the main part of the productive season (May, June and July). Copepods were the main prey item for larval Gr. halibut and sandeel, constituting between 88 and 99% of the ingested prey biomass. For both species, absolute size of preferred prey increased during ontogeny. However, preferred copepod size in relation to larval length differed markedly. In Gr. halibut, the relative size of the prey declined during growth of the larvae, while it remained constant for sandeel at a level of 2.7% of larval length. This led to a reduction in prey niche overlap between the two species. The available prey copepod biomass differed distinctly across the shelf area. In May, the prey density of Gr. halibut was the highest in the off-shelf area in Davis Strait. In June and July, the prey-rich areas for both species were mainly located at the slopes of the banks and at the shelf break area. Gut fullness was higher in these areas than in neighbouring areas, suggesting that the larval food resource could be scarce. The feeding ecology of Gr. halibut and sandeel could explain why larval abundance indices of the two species have historically shown opposite responses to yearly environmental conditions and total zooplankton occurrence.     

Potential impact of climate warming on the recruitment of an economically and ecologically important species, the European lobster (Homarus gammarus) at Helgoland, North Sea

A laboratory-based study was performed to assess the impact of climate warming on the recruitment of the endangered population of the European lobster (Homarus gammarus) at Helgoland (North Sea, German Bight). Egg-bearing females collected in situ just after spawning in late summer were subjected to various seasonal temperature regimes. Regimes with elevated temperatures (mild winters) resulted in a strong seasonal forward shift of larval hatching. Hatching took place at significantly lower temperatures than under regimes with normal winters. Experiments on larval development across a range of constant temperatures showed that no successful larval development occurred at temperatures below 14°C. Larval survival increased from 9% at 14°C to 80% at 22°C, while duration of larval development decreased correspondingly from 26 to 13 days. We hypothesize that an ongoing warming of the North Sea will strongly affect the recruitment success of the Helgoland lobster, mainly resulting from a decoupling of the seasonal peak appearance of larvae from optimal external conditions (temperature, food availability) for larval development.     

Early life histories in the bryozoan<Emphasis Type="Italic"> Schizobrachiella sanguinea</Emphasis>: a case study

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (15 days from settlement) and late (4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed.Communicated by S.A. Poulet, Roscoff