Food resource partitioning in a Mediterranean demersal fish assemblage: the effect of body size and niche width

We investigated the effects of body size, feeding strategy and depth distribution on the trophic resource partitioning among the 26 dominant fish consumers in a fish assemblage on the central Mediterranean shelf-break. The fish assemblage was structured in two major trophic guilds: epibenthic and zooplanktonic feeders, according to the position of each predator along the benthos–plankton gradient. Within each main guild, the species were segregated along a prey-size or fish-size gradient into five further guilds. Fish size and prey size were strongly correlated, indicating that the prey-size niche can be well explained by predator size. Fish consumers showed a significant negative correlation between the similarity in prey type and the similarity in depth distribution; most species with similar trophic preferences segregated along the depth dimension. The only predators overlapping in both food and depth preferences were those with a more specialist trophic behavior. These results suggest that fish body size and depth preferences are the two main niche dimensions, explaining a large part of the coexistence between the Mediterranean shelf-break fish consumers.     

Day/night vertical distribution of euphausiids in the eastern tropical Pacific

Between March 23 and April 4, 1981, samples were taken in the eastern tropical Pacific. The day/night vertical distribution of euphausiid species and biomass are described and contrasted in detail on two eastern tropical Pacific stations, the DOME station, in a region of continuous upwelling and the BIOSTAT station, in a nonupwelling area. The effects of various biological parameters, such as temperature, salinity and oxygen concentrations on the distributions of the species are examined. The numbers of euphausiids m^-2 on both stations were highest during the day, indicating that avoidance of the sampler was not a problem. During the day the largest concentration of adult euphausiids was between 300 and 350 m whereas the juveniles were concentrated between 170 and 80 m on both stations. Very few individuals were found within the oxygen minimum layer, but low concentrations of some species were found below the oxygen minimum down to 1 000 m. At night the euphausiid concentration migrated upward into the mixed layer (20 to 30 m) at BIOSTAT and to the base of the mixed layer at the DOME. Significant differences in the night depths of the species were found on both stations. The oxygen minimum layer appeared to act as a barrier to the vertical distribution of all species. Only two species were found in water with an oxygen concentration of <0.1 ml O₂ l^-1. Twentyone species of euphausiids were found on the two stations but the adult population was dominated by only two or three species on both stations. The reproductive state of the species suggested that some species reproduced earlier on the DOME than on BIOSTAT. Analysis of the depth distribution by cluster analysis showed that the most abundant species occupied different depths during the night and day at BIOSTAT but the two most abundant species were concentrated at the same depth at the DOME station although portions of each species population occupied different pelagic zones.     

Feeding ecology of benthopelagic zooplankton on an eastern tropical Pacific seamount

The gut contents of dominant deep-sea benthic boundary layer zooplankton (primarily copepods and mysids) and the vacuole contents of phaeodarians collected and preserved in situ at four depths (from 724 to 3112 m) on an eastern tropical Pacific seamount (Volcano 7; 13°23N, 102°27W) between 23 November and 4 December 1988 were analyzed using transmission electron microscopy (TEM). Suspended, and sinking plus resuspended particles, were quantitatively sampled to characterize potential food sources. The broad oxygen minimum characteristic of this region intersects the summit of the seamount and affects the feeding ecology of these organisms. Several copepods and mysids and an amphipod contained guts packed with what appeared to be gram-positive bacteria, an unusual finding. We hypothesize that the source of these bacteria-like bodies was a mat or aggregate that originated in the oxygen minimum or at its upper or lower interface. The presence of the bacteria-like bodies in 43 to 100% of the particlefeeding zooplankton that were sectioned and that had gut contents, suggests that the bacteria-like bodies are an important food source. The diverse gut and vacuole contents of other detritivores were similar among depths. Particles and microorganisms from the depths were also similar. This finding can be explained by the rapid sinking of particles and aggregates from surface waters and their relatively intact transit through the broad oxygen minimum with its reduced populations of zooplankton. The presence of algal cells in guts and vacuoles of benthic boundary-layer zooplankton suggests that these zooplankton select particles of recent surface origin for consumption. The presence of the guts filled with bacteria-like bodies shows that some deep-sea copepods and mysids that are normally generalist feeders can specialize opportunistically. The similarity of gut contents of crustaceans and vacuole contents of phaeodarians suggests that these two very different groups of particle feeders utilize similar food sources in the deep sea.     

Molecular phylogeny and DNA barcoding of tropical eastern Pacific shallow-water gorgonian octocorals

The octocoral fauna inhabiting the shallow waters (<50 m) of the eastern Pacific has been the subject of renewed interest, and the taxonomy of the most important genera in the region has been reviewed and clarified. Many new species have been described, significantly increasing the known biological diversity of the region. Despite their importance as potential sister-groups of Caribbean octocorals, the phylogenetic relationships of eastern Pacific octocorals remain poorly studied. Here, using partial mitochondrial MutS and igr1-COI sequences, we provide a phylogenetic assessment of a broad sample of eastern Pacific shallow-water octocorals and investigate their phylogenetic relationships with Caribbean gorgonians. We corroborate the monophyly of Pacifigorgia, Leptogorgia and Eugorgia and provide evidence of a close relationship between Swiftia and Psammogorgia, currently placed in Plexauridae. In addition, the phylogenies obtained here provide insights into the historical biogeography and phylogenetic diversity of the eastern Pacific octocoral assemblages and on character evolution among this diverse faunal assemblage. Finally, we evaluate the classification power of DNA barcoding for identifying species of shallow-water eastern Pacific octocorals and assess the use of a nuclear intron (SRP54) to supplement traditional mitochondrial barcodes in this group of organisms.     

Short-term temporal patterns of early recruitment of coral reef fishes in the tropical eastern Pacific

Short-term temporal patterns of recruitment have been described in a variety of coral reef fishes and have often been related with lunar and tidal cycles. While the relative importance of lunar and tidal factors in determining recruitment patterns has been difficult to assess, most studies have been done in the Caribbean and Indo-Pacific, where tidal amplitudes are small. We studied the short-term temporal dynamics of fish recruitment at Gorgona Island (tropical eastern Pacific), where there is a large tidal amplitude (~4.4 m). Every other day during three consecutive months in 1998, we directly measured the magnitude of reef fish recruitment to standardized coral units (SCUs) isolated from natural reefs. A total of 40 species from 21 families settled on the SCUs. Of 11 species with sufficient numbers for meaningful statistical analyses, two (Lutjanus guttatus and Pomacanthus zonipectus) had lunar recruitment with peaks near the new moon; three combined species of antennariids showed semilunar recruitment with peaks near moon quarters; and eight other species showed sporadic and aperiodic recruitment pulses. The contribution of lunar (moonlight intensity) and tidal factors (tidal amplitude and net tidal flow) to recruitment dynamics varied among species, although it was generally low (<18%) even among species with periodic patterns, except perhaps in L. guttatus. In this species, recruitment magnitude correlated negatively with moonlight intensity, accounting for 34.5% of the variance. Post-settlement predation by roving predators may be one cause of this relationship. In the remaining species, particularly those with sporadic and aperiodic recruitment pulses, stochastically varying weather and oceanographic events may be more important in determining temporal variation in recruitment.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.     

Food availability as a selective factor on the chemical compositions of midwater fishes in the eastern North Pacific

In September and October 1980 we examined the relationships between food availability, depth, and chemical composition among 12 midwater fish species, from three adjacent areas of the eastern North Pacific: the eastern gyre, the California Current, and the transitional region between them. By comparing trends in chemical composition across a geographical productivity gradient, the influence of food availability could be examined both with and without depth as a dependent variable. In general, caloric density, lipids, and water content showed consistent trends along both vertical and horizontal gradients of food availability. Lipids and caloric contents were lowest among bathypelagic species and among fishes from the gyre. Water content was highest in the gyre and among deeper-living species. While protein content as a percentage of the total wet weight declined with depth, there was no clear trend among mesopelagic species between stations. Protein as percent of the total organic content showed no decrease with depth, but was significantly higher in the gyre among mesopelagic species. These findings suggest that compositional trends in water, lipids, and caloric content are correlated with food availability, regardless of any depth-related factors. In contrast, protein did not vary consistently with food availability and thus may be linked to other factors.     

Metabolic suppression in thecosomatous pteropods as an effect of low temperature and hypoxia in the eastern tropical North Pacific

Many pteropod species in the eastern tropical North Pacific Ocean migrate vertically each day, transporting organic matter and respiratory carbon below the thermocline. These migrations take species into cold (15–10°?C) hypoxic water (<20?μmol O₂ kg^?1) at depth. We measured the vertical distribution, oxygen consumption and ammonia excretion for seven species of pteropod, some of which migrate and some which remain in oxygenated surface waters throughout the day. Within the upper 200?m of the water column, changes in water temperature result in a?~60–75?% reduction in respiration for most species. All three species tested under hypoxic conditions responded to low O₂ with an additional?~35–50?% reduction in respiratory rate. Combined, low temperature and hypoxia suppress the metabolic rate of pteropods by?~80–90?%. These results shed light on the ways in which expanding regions of hypoxia and surface ocean warming may impact pelagic ecology.     

All-offspring dispersal in a tropical mammal with resource defense polygyny

In polygynous mammals, males are usually responsible for gene flow while females are predominantly philopatric. However, there is evidence that in a few mammalian species female offspring may disperse to avoid breeding with their father when male tenure exceeds female age at maturity. We investigated offspring dispersal and local population structure in the Neotropical bat Lophostoma silvicolum. The mating system of this species is resource defense polygyny, with the resource being active termite nests, excavated by single males, which are then joined by females. We combined field observations of 14 harems during 3 years and data about the genetic structure within and between these groups, calculated with one mitochondrial locus and nine nuclear microsatellite loci. The results show that both male and female offspring disperse before maturity. In addition, we estimated life span of excavated termite nests and the duration they were occupied by the same male. Our findings suggest that long male tenure of up to 30 months is indeed a likely cause for the observed dispersal by female offspring that can reach maturity at a low age of 6 months. We suggest that dispersal by offspring of both sexes may occur quite frequently in polygynous tropical bats and thus generally may be more common in mammals than previously assumed.     

Ecological metrics of biomass removed by three methods of purse-seine fishing for tunas in the eastern tropical Pacific Ocean

An ecosystem approach to fisheries management is a widely recognized goal, but describing and measuring the effects of a fishery on an ecosystem is difficult. Ecological information on the entire catch (all animals removed, whether retained or discarded) of both species targeted by the fishery and nontarget species (i.e., bycatch) is required. We used data from the well-documented purse-seine fishery for tunas (Thunnus albacares, T. obesus, and Katsuwonus pelamis) in the eastern tropical Pacific Ocean to examine the fishery's ecological effects. Purse-seine fishing in the eastern tropical Pacific is conducted in 3 ways that differ in the amount and composition of target species and bycatch. The choice of method depends on whether the tunas are swimming alone (unassociated sets), associated with dolphins (dolphin sets), or associated with floating objects (floating-object sets). Among the fishing methods, we compared catch on the basis of weight, number of individuals, trophic level, replacement time, and diversity. Floating-object sets removed 2-3 times as much biomass as the other 2 methods, depending on how removal was measured. Results of previous studies suggest the ecological effects of floating-object sets are thousands of times greater than the effects of other methods, but these results were derived from only numbers of discarded animals. Management of the fishery has been driven to a substantial extent by a focus on reducing bycatch, although discards are currently 4.8% of total catch by weight, compared with global averages of 7.5% for tuna longline fishing and 30.0% for midwater trawling. An ecosystem approach to fisheries management requires that ecological effects of fishing on all animals removed by a fishery, not just bycatch or discarded catch, be measured with a variety of metrics.     

Inter-island movements of scalloped hammerhead sharks (Sphyrna lewini) and seasonal connectivity in a marine protected area of the eastern tropical Pacific

Marine top predators are common at offshore bathymetric features such as islands, atolls, and seamounts, where most pelagic reef fish reside, while certain sharks perform inter-island movements between these formations. Scalloped hammerhead sharks are known to school in great numbers at small islands and seamounts in the eastern tropical Pacific (ETP) and are very susceptible to fisheries while moving into the open sea. It is, therefore, essential to understand hammerhead inter-island movements and environmental effects to provide baseline information for their conservation and management within and beyond an insular marine protected area. Movements of scalloped hammerheads were analyzed in the Galapagos Marine Reserve (GMR) and ETP, and environmental factors were linked to their movements. Hammerheads were tagged (N = 134) with V16 coded pingers (July 2006 to July 2010) in the northern Galapagos and detected at listening stations around four islands in the GMR and two isolated islands in the ETP, 700 and 1,200 km away. Hammerheads formed daytime schools at specific locations, but dispersed at night. Overall, more daytime than nighttime detections were recorded at all receivers in the northern Galapagos Islands, and more detections in the up-current sides of these islands. Hammerheads remained more days at the northern islands during part of the warm season (December–February) compared to the cool; however, fewer individuals were present in March–June. Movement modes were diel island excursions (24-h cycles) in the northern Galapagos and inter-island in the GMR and ETP at different scales: (1) short back-and-forth (<50 km, SBF), <5 days cycles, (2) medium distance (50–300 km, MDT), 5–20 days, and (3) long distance (>300 km, LDT), 15–52 days. The high degree of inter-island connectivity of hammerheads within the northern GMR is striking compared to the almost nil movement to the central GMR. A seasonal migratory pattern to locations offshore is indicated by (1) fewer hammerheads observed in the northern GMR during part of the warm season (March–June) and (2) evidence of LDT movements from the northern GMR to other islands in the ETP. LDT movements of mature female hammerheads are possibly associated with pupping areas. Our results indicate that currents, season, and individual behavior mainly drive inter-island movements of hammerheads at small (SBF) and medium (MDT) scales. These findings have important implications for the management of a highly mobile and endangered top predator within a marine protected area and beyond.     

Seasonal and decadal changes in distribution patterns of Halobates (Hemiptera: Gerridae) populations in the eastern tropical Pacific

Five species of the marine insect Halobates share similar ecology but have distinct biogeographic ranges in the eastern tropical Pacific, a region from approximately 75°W–160°W and 10°S–35°N. Between 2001 and 2010, the Sea Education Association collected Halobates from 682 neuston tows (surface net 1 m × 0.5 m, 335-μm mesh) during fifteen cruises between San Diego, USA, Mexico and Tahiti. Total Halobates spp. densities varied substantially from year to year, but our data do not show a sustained change from a data set collected 40 years earlier from 1967 to 1968 (Cheng and Shulenberger in Fish Bull 78(3):579–591, 1980). Halobates are sensitive to sea surface temperature and we observed significant differences in species distributions over time, but these were not due to differences in water temperature or climate change. Our analyses show that the patterns observed are attributable to substantial but previously undescribed seasonal shifts that occur each year in the ranges for both Halobates sobrinus and Halobates micans. There is substantial overlap in ranges during seasonal shifts, but very little co-occurrence of H. sobrinus and H. micans in individual net tows, suggesting biological mechanisms rather than physical factors are restricting distribution and co-occurrence of these two species.     

Spatial and temporal variability of fish assemblage in Kuwait Bay

Spatial and temporal variability of fish assemblages in Kuwait Bay was studied using the catch data acquired through monthly otter trawl sampling from June 2002 through July 2004. We compared species composition using relative species density and cluster analysis among different seasons (summer’02, summer’03, winter’03, and spring’04) and areas (center, south 1, south 2, west, north and east). Significant differences in total species, species richness (SR), diversity (H′), and fish abundance were tested among (between) different seasons and areas using analysis of variance (ANOVA) and Duncan’s test. The results showed that the catches consisted of 80 species representing 41 families with Leiognathus bindus and Plicofollis tenuispinis dominating numerically. Spatially, species composition fell into three main areas: (1) east; (2) south 1 and south 2; and (3) north, center, and west. Temporally, winter’03 and spring’04 were closely related, but we found low similarity between the two summers and even lower similarity between the summers and other seasons, indicating high intra-annual variation. L. bindus was abundant in most of the areas except south 1, south 2 and the east area. Spatial changes of abundance of the two dominant species showed a complementary trend. Where L. bindus was numerically high, P. tenuispinis was low, and vice versa. Significant lower total species number and SR were observed in south 2 than all the other areas except the east, and significant higher fish abundance occurred in the summers than during the other seasons. Overall, it was concluded that variation in fish assemblages was area dependent, while fish abundance was mainly influenced by seasonal changes.     

Patterns of spatial and species abundance within the larval fish assemblage of the North Pacific central gyre during late summer

The larval fish assemblage in the epipelagic zone of the North Pacific central gyre near Lat. 28°N, Long. 155°W during late summer was sampled with stratified opening/closing bongo nets. This assemblage exhibited recurring patterns of spatial and species distributions. Spatial patterns included species vertical distributions, co-occurrence and pathchiness; species patterns included species composition and abundance relations. Both types of patterns changed with depth, the most dramatic change occurring in relation to the bottom of the summer mixed layer. An examination of the physical and biological environments of the epipelagic zone with relation to the summertime species and spatial patterns and to seasonal changes in larval fish species structure indicates that the thermal structure of the upper water column may exert a major influence on overall ichtyoplanton species and spatial structures within the gyre.     

Influence of the biological and physical environment on the vertical distribution of mesozooplankton and micronekton in the eastern tropical Pacific

The day/night vertical distributions of mesozooplankton and micronekton biomass and that of a large number of copepod species to a depth of 1 000 m are described and contrasted in detail from two eastern tropical Pacific stations, the DOME station, in a region of continuous upwelling and the BIOSTAT station, in a non-upwelling area. The effects of various biological parameters such as the zones of primary production and chlorophyll maxima plus physical parameters, such as temperature, salinity and oxygen concentrations, on the distributions of the species and mesozooplankton and micronekton biomass are examined. The thermocline depth appeared to have the greatest influence of all the physical factors on the vertical distribution of the copepod species. The vertical distribution of many species was truncated in the region of the oxygen minimum layer with very few species found below this layer. The vertical distribution of copepod molts in the top 1 000 m showed that the molts probably originated at the depths at which they were found and that the molts from the euphotic zone did not reach the deep water. Copepod species association at different depths showed that few species had common depth distributions during both day and night. The highest degree of association occurred at night in the region of the thermocline. At depths below the thermocline, the degree of similarity in vertical distributions was greatly reduced. The biomass of mesozooplankton at these two stations is compared to other regions of the tropical oceans described in the literature and shown to be among the highest values recorded. The oxygen minimum layer and the thermocline had the greatest impact on species vertical distribution. The biomass of mesoplankton was about twice as high and the number of copepods between three and four times as high at the DOME site as compared to the BIOSTAT site, and evidence suggested that this was influenced by the higher primary production in this region. The difference in micronekton biomass between sites was less, with DOME having about 1.2 to 2.2 the biomass. This suggests that the influence of the higher level of primary productivity at the DOME was evidence of the higher trophic levels.     

Feeding in a larval fish assemblage: the nutritional significance of an estuarine plume front

Gut fullness of larval fishes was used to determine the nutritional significance of an estuarine plume front off Botany Bay, SE Australia, on three days in March/April 1990. Fishes were captured in three different water masses (estuarine plume, front and shelf water), each separated by 200 m, using a 260 m mesh purse-seine net. Overall, the gut-fullness index (GFI) of 260 fish larvae combined from eight families (Gerreidae, Mugilidae, Mullidae, Sparidae, Blenniidae, Kyphosidae, Monodactylidae, Pomacentridae), was significantly greater in the plume and front water on two of the sampling occasions, but no difference was detected on the third occasion. Trends in GFI among families were inconsistent with respect to the front. The mugilid (Liza argentea) fed equally and abundantly in all water masses (gut fullness>90%), while the kyphosid (Kyphosus spp.) had a significantly greater GFI in the plume compared to the shelf water. In general, the response of GFI to the front varied between dates and amongst taxa. Diet analysis showed that mugilids selected for copepod nauplii within the plume and shelf water [alpha selectivity index (ASI) of (0.22 to 0.98)] and for harpacticoid copepods within the front (ASI=0.92), representing characteristic diets identified by canonical discriminant analysis. Kyphosids selected positively for copepod nauplii in all three water masses on both occasions (ASI>0.96). In general, diets of larval fish were taxon-specific, and responded variably to the three habitats. Comparison of diets across water masses was complicated by the large number of families, which rarely occurred in all water masses. The nutritional significance of an estuarine plume front varies between species of larval fish, and there are no obvious trends that can be applied to the larval fish community in general.     

Spawning migrations and local movements of a tropical sparid fish

Between June 1994 and September 1996, a mark–recapture study was conducted in the Hinchinbrook Channel area of tropical northeastern Australia to investigate the movements of the sparid fish Acanthopagrus berda. A total of 962 A. berda were tagged, 922 within a single estuary system, Deluge Inlet. No movements among estuaries were detected. Of the fish tagged in Deluge Inlet, 12.9% were recaptured. Two movement regimes were detected: local movements (mostly <500 m) during non-spawning times, and longer migrations (up to 3.12 km) during spawning seasons, to and from a spawning site at the estuary mouth. Although there appeared to be a major spawning aggregation in the mouth of Deluge Inlet, ripe female A. berda were present in upstream sites, indicating that spawning probably occurs there also. This contrasts to the situation in South Africa, where A. berda is reported to spawn only close to estuary mouths. Differences such as this highlight our limited knowledge of variations in the reproductive biology of widespread species such as A. berda. Received: 25 November 1997 / Accepted: 10 September 1998     

Mesopelagic fish zoogeography in the central tropical and subtropical Pacific Ocean: species composition and structure at representative locations in three ecosystems

Mesopelagic fish species are not uniformly distributed throughout the tropical and subtropical Pacific Ocean. There are reasonably clear-cut faunal assemblages with characteristics species. The geographic ranges of the assemblages conform to those patterns shown by zooplankton. As part of a 5 yr (August 1969-March 1974), seven-cruise, ecological study, replicate sampling in central gyral waters included additional quantitative sampling at a point along the equator at 155°W to compare species composition and structure at this location with points in the central gyres. These comparisons show that the distribution of individuals among species displays similar structure in all three areas, but species composition and the relative abundances of shared species differ. There are species restricted to the low-productivity central gyres, and subgroups of these are restricted to specific gyres. The gyral faunas differ markedly from the high-productivity species assemblage found along the equator. With the exception of the eastern tropical Pacific, where low oxygen concentrations probably limit species distributions, the primary productivity regime is the most likely factor defining the distributions of most mesopelagic fishes in the tropical and subtropical Pacific. The mechanisms for the maintenance of these geographic patterns are not clear, but the correlation of the patterns with the productivity regime leads one to hypothesize that the ultimate factor is food.