Growth of the Atlantic surfclam,Spisula solidissima,from Georges Bank to the Delmarva Peninsula,USA

Age/shell length data for offshore surfclam, Spisula solidissima (Dillwyn, 1817), populations were used to estimate the parameters of the von Bertalanffy growth model by time period and region, from Georges Bank in the north to the Delmarva Peninsula in the south. Randomization tests were used to compare curves. We tested the a priori hypothesis that growth curves would change over time in the south (i.e., New Jersey and Delmarva) but remain constant in the north (i.e., Long Island and South New England). This hypothesis was proposed because surfclam population structure in the south had been altered by the hypoxic event of 1976, and possibly by intense, long-term commercial harvesting. Northern regions, unaffected by these factors, served as natural controls. Based on a comparison of data collected in 1980 with pooled data from 1989 and 1992, the hypothesis was supported. Both the growth coefficient (k) and maximum shell length (L ) declined between two time periods in the two southern regions, while during the same time interval, no change occurred in the two northern regions. Differences in growth between regions were often statistically significant. For example, compared with the southern regions, the growth coefficient on Georges Bank was larger, and those clams attained a smaller maximum length. In a comparison of adjacent regions from Delmarva to S. New England, k increased from south to north. This could imply faster growth in cooler water, as well as no relationship between growth and primary productivity. Alternatively, size-selective mortality, imposed by the commercial fishery, was discussed as a mechanism that might account for this unexpected pattern.     

Density-dependent growth in the Atlantic surfclam, Spisula solidissima, off the coast of the Delmarva Peninsula, USA

Spisula solidissima (Dillwyn, 1817) is a large, suspension-feeding bivalve, whose range extends from Nova Scotia to South Carolina. This species is harvested commercially. Shell length and age data were collected for this species from 1980 to 1994 during surveys of population size and structure. These data were used to examine the relationship between the growth rate of S.␣solidissima and intraspecific density. The null hypothesis was that density (represented by number of individuals per tow) would have no effect on rate of growth. A negative relationship would support the alternative hypothesis, that intraspecific competition had taken place. This analysis focused on the surfclam population offshore from the Delmarva Peninsula, USA because: (1) a major recruitment event occurred in 1977, (2) clam fishermen had reported “stunted” surfclams in that area, (3) a wide range of local densities were available to examine, and (4) the existence of a closed area within the study area set up an interesting contrast with areas left open to harvesting. Maps of surfclam abundance across the Delmarva region demonstrate that areas of highest density have generally remained in the same location through time. The results suggested that intraspecific competition has been important in structuring this population. Based on data from 1980 to 1992, shell length was significantly reduced at high density, and a significant interaction between age and density was observed. Growth modeling indicated decreased asymptotic lengths and growth rates with increasing density. In nine out of ten pairwise randomization tests, fitted von Bertalanffy growth curves, representing different densities, were significantly different from each other. High densities of clams have persisted in the area that was closed to harvesting for 11 years (1980 to 1991). In 1994, length at age was significantly less in this closed area compared to that in the surrounding area. This effect was apparent in clams from 3 to 17 years of age, and most pronounced in the cohort that recruited to the Delmarva region in high numbers in 1977. Lower growth rates within the closed area have management implications for the optimal duration of closures. Received: 16 May 1997 / Accepted: 7 October 1997     

Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Growth,mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches

We studied some aspects of the population dynamics of the yellow clam Mesodesma mactroides (Deshayes, 1854) which inhabits the eastern sandy beaches of Uruguay. Observations were made from March 1983 through March 1985 (harvesting season), and additional data from January 1988 to December 1989 (closed season) were also included in the analyses. Parameters of growth and mortality were estimated by methods that make use of age or length information. An age/length relationship was constructed by counting growth rings. Results were consistent with those obtained with modal class progression analysis and other length/frequency methods. The growth curve obtained from the age/length relationship did not account for seasonality in growth rate, and a seasonal growth model explained growth oscillations in a better way. The instantaneous coefficient of natural mortality (M) estimated for the closed season was 1.64, whereas an increase in fishing mortality (F) was observed during 1984. Age composition did not change during the 2 yr of study, and the first 6 mm class accounted for over 50% of the population. The main contribution of recruits was observed between late summer and early fall (February to April), and the recruitment pattern suggested one main settlement period per year. Growth and recruitment success appeared to be regulated by density-dependence processes. Some implications for management are considered in this context. Methodological aspects concerning the reliability of empirical equations and programs for estimating growth and mortality are discussed.     

Genetic structure of deep-water redfish, Sebastes mentella, populations across the North Atlantic

Patterns of genetic diversity of the marine deep-water redfish, Sebastes mentella Travin, from 21 sampling locations (n=46 per sample, from 1995 to 1998) were examined throughout its range in the North Atlantic. Eight microsatellite loci were used in order to: (1) define the genetic structure and consider the possible influence of dispersal, geographic distance, oceanic currents and historical factors; and (2) relate the structure to present management units and practices. Three divergent population units (mean Š=0.012) associated with distinct geographic areas were defined: Eastern (Norway and Barents Sea), Panoceanic and Western (Gulf of St. Lawrence and offshore Newfoundland). The most important observation was the lack of genetic differences (mean Š=-0.0004) and lack of genetic isolation by geographic distance (r=0.318, P=0.112) among samples within the Panoceanic zone, from the Faroe Islands to the Grand Banks (6,000 km). A combination of vicariance, historical introgression with a closely related species, S. fasciatus, and contemporary patterns of oceanic circulation are likely to have shaped the observed population structure. These results only partially support current management units, and call for more integrated practices for regulating the exploitation of S. mentella throughout its extensive range.     

Comparative study of sympatric populations of two hyperiid amphipods,Primno johnsoni and P. evansi,from the eastern North Atlantic Ocean

Hyperiid amphipods are common components of oceanic plankton communities, and yet information is lacking on the ecology and biology of the majority of species. This study compares sympatric populations of two phrosinids, Primno johnsoni Bowman and P. evansi Sheader, using material collected on R.R.S. Discovery Cruises 121 and 140 in the eastern North Atlantic Ocean. The two species were essentially very similar in terms of their vertical distribution and diurnal migration, and in the timing and sequence of reproductive events (onset of maturity, ovary/oocyte development, marsupial development). Mature female and male size ranges were comparable for the two species, males maturing at a small body size, with little subsequent growth, and females producing a succession of 5 (P. evansi) or 6 (P. johnsoni) broods maximum. The species differed significantly in egg size and in total egg output. These dissimilarities were accentuated by differences in the patterns of female mortality in the two species, such that actual egg output per female was 1.8 times greater (2.4 times in terms of total egg volume) in P. johnsoni than in P. evansi. These differences were reflected in the relative size of populations in the field. Although both species are capable of consuming a wide range of planktonic prey, it is assumed that gelatinous species are important, especially as hosts for the parasitoid juvenile stages, and it is suggested that host-specificity might act to separate the niches.     

Reproductive cycle,development and recruitment of two geographically separated populations of the dendrochirote holothurian Aslia lefevrei

The boreo-mediterranean holothurian Aslia lefevrei (Barrois) occupies cryptic habitats from low-water mark to ca. 20 m. During the present work specimens were collected from the west coast of Ireland (February 1979 to August 1980), where A. lefevreiis a prominent element in two discrete epibenthic assemblages, and from the north-west of Spain (September 1981 to August 1982), where this holothurian is densely aggregated in association with raft cultivation of the mussel Mytilus edulis. Its reproductive cycle, spawning process, embryological development, settlement, and early growth rates have been documented by means of laboratory and field studies. Depending on geographic location, spawning in the wild occurs between January and May. Development to the gastrula takes some 3 to 4 d. Uniformly ciliated dipleurula/auricularia hatch 12 to 24 h after gastrulation. The pentactula stage is attained approximately 7 d after fertilization, following which the larva settles. Newly settled individuals utilize the five primary tentacles for both feeding and locomotion. With the advent of small numbers of podia in the ambulacral regions, two of the primary tentacles prolong their locomotory and anchorage capabilities while the remaining tentacles develop adult characteristics and function solely in feeding. Growth was followed for an estimated 30 mo period using length-frequency analyses. In the first year of life, A. lefevrei attain an average length of 3.4 cm; individuals on the sea floor can attain up to 18 cm, indicating a possible life expectancy of 10 yr or more.Contribution No. 301 from the School of Marine Sciences, University College Galway, Ireland     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

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.     

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

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

Activity and metabolism of larval Atlantic cod (Gadus morhua) from Scotian Shelf and Newfoundland source populations

Patterns of activity and metabolism were investigated in larval Atlantic cod (Gadus morhua L.) between December 1991 and July 1992: (1) throughout larval development; (2) between two genetically discrete populations (Scotian Shelf and Newfoundland) and (3) as a function of two different culture temperatures. During the yolk-sac stage (0 to 5 d post-hatch), changes in swimming speed were not related to mass-specific metabolic rates; no portion of the mass-specific oxygen consumption could be explained by changes in activity. In the mixed feeding stage (6 to 14 d posthatch), there was a tendency for oxygen consumption to be related to changes in swimming speed. In the exogenous feeding stage (>14 d post-hatch), oxygen consumption significantly increased with swimming speed. These ontogenetic patterns of activity and metabolism were the same for larvae from the Scotian Shelf and Newfoundland populations. However, over the entire larval life and among ontogenetic stages, the metabolic cost of activity (mass-specific O₂ consumption/swimming speed) of Scotian Shelf larvae was significantly higher than that of Newfoundland larvae. When cod larvae, that had developed at 5°C, were acutely exposed to 10°C, Scotian Shelf larvae had a higher intrinsic cost of activity than Newfoundland larvae, over the entire larval life. During the exogenous feeding stage, the mean metabolic cost of activity for Newfoundland larvae raised at 10°C and tested at 10°C was significantly higher and more variable than that of larvae raised at lower temperatures. However, the metabolic cost of activity of larvae raised and tested at 10°C was not significantly different between source populations. Together these findings suggest that differences in swimming energetics reflect changing energy requirements for activity among ontogenetic stages, and reflect adaptation to regional environments among genetically discrete populations.     

The role of sexual harassment in cave and surface dwelling populations of the Atlantic molly, Poecilia mexicana (Poeciliidae, Teleostei)

Sexual coercion is a common behaviour for males trying to compensate for being inferior in male competition and/or female choice. We measured the cost of male sexual harassment for females as reduced feeding time in three populations of the Atlantic molly (Poecilia mexicana). Of these populations, one originated from a typical river habitat, another from a lightless cave chamber and the third one from a small, milky creek outside the cave. We gave hungry females an opportunity to feed in the presence of either another female or a male. We found the following: (1) male sexual harassment represents a cost in the river dwelling population and both small and large males significantly reduce female feeding time; and (2) sexual harassment was not detected in the other two populations (cave and cave entrance). In the cave molly, small males are at a disadvantage in female choice, but predation by an aquatic heteropteran selects against large male body size.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

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.     

Genetic and taxonomic relationships among Northeastern Atlantic and Mediterranean populations of the soft coral Alcyonium coralloides

Species boundaries among taxa of colonial marine organisms are often obscured by intraspecific morphological and ecological variation; genetic comparisons of recognized “ecotypes” frequently reveal them to be reproductively isolated species. Based on morphological similarities, it has been proposed that the Mediterranean soft coral Alcyonium (=Parerythropodium) coralloides Pallas, 1766 and its Atlantic congener A. hibernicum belong to one highly variable and geographically widespread species, A. coralloides. I collected A. coralloides from ten Atlantic and three Mediterranean locations in 1990 and 1994, and used differences in colony form, substrate use and color to separate them into five distinct morphotypes. Two occur sympatrically in the Mediterranean (M1, M2) and three have overlapping distributions in the Atlantic (A1, A2, A3). I used allozyme electrophoresis to compare morphotypes genetically at 14 enzyme loci. Where two morphotypes occurred sympatrically, fixed allelic differences at 4 to 6 loci indicated reproductive isolation. In all but one pairwise comparison (M1 and A2), morphotypes whose ranges did not overlap were also separated by large genetic distances. From these results I suggest that the five morphotypes represent four distinct species. A. coralloides comprises two morphotypes (M1, A2) with relatively high genetic identity. A. hibernicum (=A1) is reproductively isolated from A. coralloides and should be retained as a valid species; levels of genetic diversity and heterozygosity within populations support the absence of outcrossing in this reportedly asexual species. Morphotypes M2 and A3 are also reproductively isolated from A. coralloides; they are taxonomically distinct from but belong to the same phylogenetic clade as A. hibernicum. Although preliminary observations suggest that differences in reproductive timing maintain species boundaries in sympatry, wider geographic sampling will be required to elucidate the events leading to speciation within this species complex. Received: 8 May 1998 / Accepted: 8 October 1998     

Parasitism in a social wasp: effect of gregarines on foraging behavior, colony productivity, and adult mortality

Behavior in eusocial insects likely reflects a long history of selection imposed by parasites and pathogens because the conditions of group living often favor the transmission of infection among nestmates. Yet, relatively few studies have quantified the effects of parasites on both the level of individual colony members and of colony success, making it difficult to assess the relative importance of different parasites to the behavioral ecology of their social insect hosts. Colonies of Polybia occidentalis, a Neotropical social wasp, are commonly infected by gregarines (Phylum Apicomplexa; Order Eugregarinida) during the wet season in Guanacaste, Costa Rica. To determine the effect of gregarine infection on individual workers in P. occidentalis, we measured foraging rates of marked wasps from colonies comprising both infected and uninfected individuals. To assess the effect of gregarines on colony success, we measured productivity and adult mortality rates in colonies with different levels of infection prevalence (proportion of adults infected). Foraging rates in marked individuals were negatively correlated with the intensity of gregarine infection. Infected colonies with high gregarine prevalence constructed nests with fewer brood cells per capita, produced less brood biomass per capita, and, surprisingly, experienced lower adult mortality rates than did uninfected or lightly infected colonies. These data strongly suggest that gregarine infection lowers foraging rates, thus reducing risk to foragers and, consequently, reducing adult mortality rates, while at the same time lowering per-capita input of materials and colony productivity. In infected colonies, queen populations were infected with a lower prevalence than were workers. Intra-colony infection prevalence decreased dramatically in the P. occidentalis population during the wet season.An erratum to this article can be found at     

Cadmium, zinc and copper accumulation in the squid Illex argentinus from the Southwest Atlantic Ocean

 The levels and burdens of Cd, Zn, Cu and Hg were measured in the dorsal mantle, digestive gland and gonads of the squid Illex argentinus, from the Southwest Atlantic Ocean. Mature and immature individuals of both sexes were analysed. Correlations of heavy metal concentrations and burdens with sex and food habits were studied. The highest metal levels were found, particularly for Cd, in the digestive gland, which attained a concentration of up to 270 μg g⁻¹ (wet weight). Dorsal mantle and gonads exhibited the same order of metal enrichment: Zn>Cu>Cd. Zinc and Cu levels were higher in ovaries than in testes and varied with the stage of maturation. The dorsal mantle exhibited the lowest heavy metal concentrations. Mercury levels were below the detection limit of the method in all the tissues analysed. Received: 7 September 1999 / Accepted: 9 March 2000     

Biomass of the invertebrate megabenthos from 500 to 4100 m in the northeast Atlantic Ocean

Data are presented on the biomass of the invertebrate megafauna at 22 stations on the continental slope in the Porcupine Seabight (PSB) (northeast Atlantic Ocean). Samples were collected between 1980 and 1982. Several units of biomass are used, all of which illustrate a decrease by a factor of about 30 from 500 to 4100 m. Lognormal curves were fitted to the data, the gradients of which were very similar for all biomass units and similar to the value for a transect down the continental slope in the western Atlantic. Biomass levels in the PSB are compared with those from other deep-sea environments. Some published values are more than ten times higher than the values reported here, while others are less than a tenth. The reasons for these differences and trends are discussed in terms of food supply. Sampling variability was examined at two stations, but by chance one (at 1300 m) appeared to encompass a sharp faunal discontinuity of the dominant fauna and the other (at 4000 m) contained very small numbers of large animals. For these reasons, sample variability was high at the repeat stations. Suspension-feeders and crustaceans dominated the biomass at upper-slope depths, while echinoderms were dominant on the middle and lower slope. As a result of this phyletic change, there was a small but insignificant decrease in mean body weight with increasing depth. Within phyla there was also a small but insignificant decrease with depth. If large species are excluded from the biomass/depth regression, the gradient changes considerably, demonstrating the increasing importance of small species at greater depths. The size distribution of megafaunal biomass was examined at several stations. This indicated that the megafauna form a functional group distinct from the macrofauna, just as the macrofauna are distinct from the meiofauna.