Chemical composition of midwater fishes as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

The variation with depth in water, lipid, protein, carbon and nitrogen contents (% wet weight) of 42 species of midwater fishes, collected in November 1976 off the west coast of Oahu in the Hawaiian Archipelago, was measured. The Hawaiian fishes show significant relationships between these components and depth of occurrence. The slopes of these relationships are not significantly different from those reported for midwater fishes from off California, USA. However, the fishes from Hawaii have significantly lower lipid levels and higher protein levels than do the species from off California. The deep-living Hawaiian species (500 m and deeper) have significantly lower lipid (% wet weight), but there is no significant difference in protein (% wet weight). The difference in lipid contents at all depths appears to be an evolved characteristic, with the greater lipid levels off California being selected for by greater spatial and temporal variation in the food supply for these fishes off the California coast than off Hawaii. The higher protein contents in the shallow-living Hawaiian fishes appear to reflect greater muscle power selected for in these fishes by the greater water clarity, and therefore greater reactive distances, in the surface layers off Hawaii. These conclusions support the general hypothesis that the lower protein contents of bathypelagic fishes are not directly selected by food limitation at depth, but rather result from the relaxation of selection for rapid-swimming abilities at greater depths due to the great reduction at greater depths in the distance over which visual predator-prey interactions can take place. The lower lipid levels in the deeper-living species are apparently made possible by the reduced metabolic rates of these species which reduces their need for energy stores.     

Metabolic rates of midwater crustaceans as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

During July of 1983, 1986, and 1987, we measured rates of oxygen consumption of 234 individuals of 17 species of midwater crustaceans (orders Decapoda, Mysidacea, and Euphausiacea) off the Hawaiian islands at depths from the surface to greater than 1200 m. The routine metabolic rates declined with increasing depths of the species' occurrence to an extent greater than could be accounted for by depth-related changes in body size or water temperature. Most species appeared able to regulate their oxygen consumption down to the lowest oxygen partial pressures found in their depth range (20 mm Hg O₂), but did not regulate to such low oxygen partial pressures as did similar midwater crustaceans off California, where oxygen levels reach as low as 6 mm Hg. Metabolic rates of the shallower-living, but not the deepest-living Hawaiian crustaceans were significantly higher than those of Californian crustaceans. This is interpreted as indicating that the metabolic rates of midwater crustaceans are not adapted specifically to differing levels of primary production and that the decline with depth of metabolic rates in these species is not the result of food limitation at depth. The data are, however, consistent with the hypothesis that lower metabolic rates at depth are due to the relaxation of selection pressures relating to visual predation near the surface.     

Triacylglycerols and wax esters in the lipids of deep midwater teleost fishes of the Southern California Bight

Total lipid contents and the proportions of triacylglycerols and wax esters were examined in 23 species of deep-sea fishes collected between 1977 and 1980 in the Santa Catalina, San Clemente and San Pedro Basins off Southern California, USA. Mean total lipid content ranged from 1 to 22% of wet weight. Triacylglycerols accounted for 1 to 91% of total lipids present and were more abundant than wax esters in the lipids of all but a single species. Wax ester contents ranged from 0 to 24% of total lipids but in 20 species were 10% of the lipids. These results do not substantiate the previously suggested idea that wax esters are important constituents of the lipids of deep-sea fishes. Moreover, the hypothesis that wax ester content increases with increasing depth of occurrence is not supported. To assess the importance of the lipids present in bouyancy regulation, the water content and swimbladder inflation of these species were also examined. Neither total lipids nor wax esters appear to play an important role in buoyancy regulation in the majority of these fishes. Eight species have gas-filled swimbladders. Of the species lacking inflated swimbladders, 7 have watery bodies (87 to 95% H₂O) and 6 with low lipid (8.3%) and water (85%) contents may need to swim continuously to maintain their position in the water column. Lipids, largely in the form of triacylglycerols, may be important in the buoyancy of three species with lipid contents>10% of wet weight. Two of these species also possess inflated swimbladders. The reasons for the production of large quantitites of was esters by certain deep-sea teleosts, especially gonostomatids and myctophids, remain unknown.     

Chemical composition,metabolic rates and feeding behavior of the midwater ctenophore Bathocyroe fosteri

Individuals of the midwater ctenophore Bathocyroe fosteri (0.01 to 1.6 g dry weight, DW) were collected from Bahamian waters by the submersible Johnson-Sea-Link during May and September/October 1983 and October/November 1984 from 530 to 700 m depth. Metabolic rates were measured and showed oxygen consumption to be in the range of 0.01 to 0.18 mg O₂ g^-1 DW h^-1 at temperatures ranging from 9° to 12°C. Ammonium excretion (0.01 to 0.14 g-at N g^-1 DW h^-1) was typically low. Energy expenditures estimated from respiration data (ca. 7% body C d^-1) indicated that one to three midwater crustacean prey (ca. 150 g C d^-1) could provide the daily maintenance ration required by a 40 mm ctenophore. These metabolic characteristics complemented in situ observations of poor locomotor ability and passive feeding behavior.     

Oxygen consumption of midwater fishes and crustaceans from the eastern Gulf of Mexico

Oxygen consumption was measured as a function of temperature, oxygen partial-pressure (P_O2)and species depth of occurrence for twenty-three species of midwater fishes and crustaceans collected from the eastern Gulf of Mexico from June 1981 to July 1985. Q₁₀s (7° to 20°C) of 3.90 and 3.24 were recorded for myctophid and non-myctophid fish groups, respectively, while values of 2.22, 2.19, 2.19 and 2.54 were calculated for sergestid, penaeid, carid and euphausiid crustacean groups, respectively. Q₁₀s were consistent for species within each group. All of the species tested regulated their oxygen consumption to P_O2levels normally encountered within the eastern Gulf. Values of critical partial pressure (P_c) ranged from 20 to 40mm Hg and increased slightly with increasing temperature and respiration rate. Declining respiration with increasing minimum depth of occurrence was primarily a function of temperature alone. Changes in size, dry weight and water content contributed only a small fraction to the observed decrease. This finding contrasts with studies from the eastern Pacific Ocean, where temperature is a minor contributor to changes in respiration rate with depth.     

Distribution and abundance of the epipelagic mollusk Carinaria japonica in waters off Southern California

The distribution and abundance of the heteropod mollusk Carinaria japonica Okutani is described from midwater trawl records off Southern California (USA) and northern Baja California (Mexico). Maximal numbers of individuals were recorded during August in oceanic waters off Southern California. In oceanic waters near Guadalupe Island, Baja California, low population levels were recorded irrespective of season. In neritic waters over the continental shelf off Southern California, C. japonica were present in low numbers, mainly between July and December when conditions seemed most favorable for their entry into the Southern California Eddy.     

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

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

Patterns of growth,energy utilization and reproduction in some meso- and bathypelagic fishes off Southern California

We have studied growth, energy use and reproduction in 4 mesopelagic fishes and 5 bathypelagic fishes living off Southern California (USA). All of the mesopelagic species underwent diurnal vertical migrations, while none of the bathypelagic species did so. The life histories of these pelagic fishes were compared among themselves and with epipelagic sardines and anchovies studied by others. The epipelagic species had the highest growth rates (estimated from otoliths, expressed in standard length or kilocalories), the mesopelagic species had the lowest growth rates and the bathypelagic species had intermediate growth rates. The relatively rapid growth rates of the bathypelagic fishes were achieved by high relative growth efficiencies made possible by low metabolic rates. Of the species studied, the lifespans of the epipelagic and bathypelagic species ranged from 4 to 8 yr and the lifespans of mesopelagic species from 5 to 8 yr. Data on egg diameters suggest that the mesopelagic species first reproduce in their 3rd yr, while the bathypelagic species do so in their last year. Epipelagic fishes generally have a large size, rapid growth, long life and early, repeated reproduction. Mesopelagic fishes are characterized by small size, slow growth, long life and early, repeated reproduction. Bathypelagic fishes generally have large size, rapid growth, somewhat shorter lives and late reproduction, which is possible a single event. The latter pattern is evidently feasible only in a rather stable environment where juvenile survivorship would always display relatively low variability. Many unusual characteristics of deep-living animals have possibly been selected by factors peculiar to the environment; however, such characteristics are just as likely to have been selected by factors equally present in many other environments, but not expressed there due to masking selective forces. In particular, we have in mind the darkness, stability and homogeneity of the bathypelagic realm as phenomena which represent the effective absence of many selective forces.The first two authors contributed equally and their names are listed in alphabetical order     

Variability in the gut contents of individual A cartia tonsa from waters off Southern California

The gut contents of 169 individual Acartia tonsa from Los Angeles Harbor, California, USA, were measured during a 24 h period (16–17 June 1986) by gut pigment (fluorescence) and microscopic analyses. Individual gut-pigment levels varied 10-fold or more within sampling intervals. Some copepods with moderate (0.2 to 0.5 ng) to high (>0.5 ng) gut-pigment levels were present in samples from both day and night collections. While the percentage of copepods containing>0.5 ng pigment was about the same during the day (8%) as at night (10%), the percentage of copepods with 0.2 to 0.5 ng pigment rose from 17% during the day to 55% at night. Significant differences between pigment levels in copepods collected before and after evening twilight were suggestive of a nocturnal feeding habit regardless of intense individual variability in gut-pigment content. Food in the gut was distributed in parcels, indicative of intermittent feeding that potentially contributes to individual variability. Feeding was not synchronized during most of the day and night, but synchrony increased at evening and morning twilights. Although synchrony declined after evening twilight, individual gut-pigment contents were relatively elevated in most of the nighttime samples. Thus, active feeding seems neither to require nor to imply synchrony.     

Nitrogen supply,tissue composition and frond growth rates for Macrocystis pyrifera off the coast of southern California

Seasonal changes in ambient NO ₃ ^– and NH ₄ ⁺ , tissue composition (N, C, and C/N ratio), and frond growth rates for Macrocystis pyrifera (L.) Agardh were examined. Ambient NO ₃ ^– showed distinct seasonal variations. Frond growth rates were variable, but showed no clear correlation to ambient NO ₃ ^– . The average N content of plant tissue did, however, show the same seasonal variations as ambient NO ₃ ^– . The longitudinal distribution of total tissue N and various components of tissue N along fronds were also analyzed. Several distinct patterns were found: high levels of protein N at growing tips and elevated levels of soluble N in lower parts of the frond. Free amino acids accounted for a major portion of the soluble N, but neither NO ₃ ^– nor NH ₄ ⁺ accumulated in the plant tissue. The longitudinal distribution of N along the fronds is compared to reported variations in C metabolism, and it is concluded that C and N sourcesink relations do not always coincide and bidirectional translocation may occur.     

Ants as a Measure of Effectiveness of Habitat Conservation Planning in Southern California

Abstract: In the United States multispecies habitat conservation plans were meant to be the solution to conflicts between economic development and protection of biological diversity. Although now widely applied, questions exist concerning the scientific credibility of the conservation planning process and effectiveness of the plans. We used ants to assess performance of one of the first regional conservation plans developed in the United States, the Orange County Central‐Coastal Natural Community Conservation Plan (NCCP), in meeting its broader conservation objectives of biodiversity and ecosystem‐level protection. We collected pitfall data on ants for over 3 years on 172 sites established across a network of conservation lands in coastal southern California. Although recovered native ant diversity for the study area was high, site‐occupancy models indicated the invasive and ecologically disruptive Argentine ant (Linepithema humile) was present at 29% of sites, and sites located within 200 m of urban and agricultural areas were more likely to have been invaded. Within invaded sites, native ants were largely displaced, and their median species richness declined by more than 60% compared with uninvaded sites. At the time of planning, 24% of the 15,133‐ha reserve system established by Orange County NCCP fell within 200 m of an urban or agricultural edge. With complete build out of lands surrounding the reserve, the proportion of the reserve system vulnerable to invasion will grow to 44%. Our data indicate that simply protecting designated areas from development is not enough. If habitat conservation plans are to fulfill their conservation promise of ecosystem‐level protection, a more‐integrated and systematic approach to the process of habitat conservation planning is needed.     

Distribution and community structure of midwater fishes in relation to the subtropical convergence off eastern Tasmania,Australia

Midwater fishes were sampled at night to a depth of 400 m in oceanic waters east of Tasmania, Australia, in May/June of 1992, 1993 and 1994. We examined whether there were differences in the distribution and density of these fishes in relation to the subtropical convergence (STC) separating East Australia Current (EAC) water from subantarctic water (SAW) to the south. A total of 23999 fishes from 107 taxa and 43 families were identified. Four new records were identified from the study area. Myctophids and stomiatoids were the main fish taxa captured. We found no evidence of increased density of the combined fish catch in any particular area. However, differences between areas in the density of individual species were noted, although no species was confined to any one area. Ceratoscopelus warmingi, Lobianchia dofleini and Vinciguerria spp. were most abundant in the EAC, whereas Lampichthys procerus was most abundant in the SAW. Only Diaphus danae was found in significantly higher numbers in the STC. Multivariate analysis revealed that community structure in the EAC was significantly different from that of the SAW but not from the STC. Further, when separated by depth, the shallow EAC group was significantly different in all group comparisons bar that with the shallow STC. Our data indicated that the thin EAC layer above 200 m could be distinguished by its fauna from the SAW. However, we could not detect a separate community in the STC. We conclude, therefore, that the STC is not an area of increased micronekton abundance, nor does it contain a distinct community, during the Australian autumn/winter.     

Rainwater composition as a function of the cloud type

The influence of the type of clouds that produce precipitation in the rainwater composition was analyzed. Logroño, a remote station in the North of Spain was chosen for the analyses. Results prove that the rainwater composition from Cumulonimbus is different to the composition of the rainwater from the other clouds. In addition, the source of NO₃ ^? and SO₄ ^2? is studied in the different types of clouds. It is shown that the source is the soil except Stratocumulus formed from Cumulus in which the source is the gas‐particle conversion.     

Correlation of primary production as measured aboard ship in Southern California Coastal waters and as estimated from satellite chlorophyll images

Chlorophyll images suitable for the regional analysis of the Southern California Bight were obtained by satellite (Nimbus-7 Coastal Zone Color Scanner) on 27 February, 6 March and 11 March 1979. An algorithm to convert satellite chlorophyll data to primary production has been developed. In addition, primary production was measured along with other biological and physical factors from ships cruising in the Southern California Bight during late February and early March 1979. There was fair agreement between shipboard measurements and satellite estimations; for example, primary production (mg carbon m^-2 d^-1) averaged 554 mg from shipboard measurement and 403 mg from satellite estimation when averaged for the entire Bight for the course of the study. The high standard error associated with the variance in productivity index used in our algorithm prevented more accurate conversion of chlorophyll concentrations into primary productivity. However, the synoptic data provided by concurrent ship and satellite measurements can be used to quantify, and perhaps reduce, sampling errors associated with ship sampling alone and to provide estimates of the accuracy of primary productivity on a regional basis.     

Enzyme activities of fish skeletal muscle and brain as influenced by depth of occurrence and habits of feeding and locomotion

Activities of lactate dehydrogenase (LDH), pyruvate kinase (PK), malate dehydrogenase (MDH) and citrate synthase (CS) were measured in the white skeletal muscle of marine fishes having different depths of occurrence and different feeding and locomotory strategies. There were significant depth-related differences in the two glycolytic enzymes, LDH and PK. LDH activity was most variable, and differed by 3 orders of magnitude between the most active shallow-living species and certain deep-sea fishes likely to have only minimal capacities for active locomotion. Superimposed on the depth-related patterns was a high degree of interspecific variation (up to 20-fold) in enzymic activity among species from any given range of depth of occurrence. Variation of both LDH and PK activities, noted for shallow- and deep-living fishes, seems to be largely accounted for by differences in feeding habits and locomotory performance. Active pelagic swimmers have much higher activities of LDH and PK than, for example, deep-living sit-and-wait predators. Benthopelagic fishes like rattails and the sablefish have the highest activities found among deep-living fishes, suggesting that these species engage in relatively active food-searching behavior compared to most other deep-sea fishes. The activity of CS, an enzyme of the citric acid cycle and an indicator of aerobic metabolism, varied little among species. Thus, the large interspecific variation in glycolytic potential (LDH and PK) among species is not associated with a similar variation in aerobic metabolism of white muscle. The much higher and more variable activity of MDH relative to CS suggests that, in addition to its function in the citric acid cycle, MDH may play an important role in redox balance in fish white muscle. In a comparison of white muscle composition between the shallow- and deep-living species, water content did not differ significantly, but protein content was significantly higher in shallow- than in deep-living fishes (211 and 130 mg g^-1 wet wt of muscle, respectively). The differences in muscle protein content are small relative to the differences between shallow- and deep-living species in LDH, PK and MDH activities. Thus, depthrelated differences in muscle enzymic activity are caused by factors other than enzyme dilution. Enzyme activities (LDH, PK and CS) in brain tissue were relatively constant among species regardless of depth of occurrence or feeding and locomotory habits. Habitat and lifestyle do not seem to influence the demands for overall metabolic function in brain. The utility of muscle enzymic activity data for making predictions about the ecological characteristics of difficult-to-observe, deep-living, fishes is discussed.     

Changes in photosynthetic pigment concentration in seaweeds as a function of water depth

We conducted a study of the relationship between changes in photosynthetic pigment content and water depth in Great Harbor near Woods Hole, Massachusetts, USA, on the green algae Ulva lactuca and Codium fragile and the red algae Porphyra umbilicalis and Chondrus crispus. A calibrated underwater photometer equipped with spectral band filters measured light attenuation by the water column. The depth required for a 10-fold diminution of photon flux was 3.6, 5.3, 6.0 and 6.0 m for red, blue, yellow and green light, respectively. Seaweeds were attached to vertically buoyed lines and left to adapt for 7 days; then, with their positions reversed, they were allowed to readapt for 7 days. All species showed greater photosynthetic pigment content with increased depth. Further, the ratio of phycobiliproteins and chlorophyll b to chlorophyll a increased with depth. Changes in pigment content were reversible and occurred in the absence of cell division. There was a net loss of pigments near the surface (high irradiance), and subsequent synthesis when seaweeds were transferred to a position deep in the water column (low irradiance). In contrast, seaweeds which were found in intertidal habitats changed only their pigment concentration, and not pigment ratio, a phenomena analogous to higher plant sun and shade adaptation. Therefore, seaweeds modify their photon-gathering photosynthetic antennae to ambient light fields in the water column by both intensity adaptation and complementary chromatic adaptation.     

Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panamá

The community structure of the decapod crustacean fauna of 7 tropical, shallowwater, marine habitats (sandy beaches, mangrove swamps and rocky intertidal habitats on both the Pacific and Caribbean coasts of Panama, and Pocillopora damicornis coral habitat of the Bay of Panama) were examined and analyzed for species composition and relative abundances. Collections from the 7 habitats yielded 4361 individuals, representing 236 species. The number of species per habitat was (Pacific, Caribbean): sandy beach (16, 7); mangrove (20, 17); P. damicornis (53); rocky intertidal (78, 67). There were more species represented by more individuals in the Pacific habitats. An index of faunal similarity was calculated for each pair (Pacific-Caribbean) of habitats. This index is the number of ecologically similar congeneric species which occurred in both habitats expressed as a percentage of the total number of species present in the pair of habitats. For the sandy beach communities there were three Pacific species which were similar to three Caribbean species, a similarity of 6/23 or 26%. The index of similarity for the mangrove communities is 54% and for the rocky intertidal communities it is 37%. The P. damicornis community has affinities with the Pacific rocky intertidal community (18%), with that of the Caribbean rocky intertidal (16%) and with that of Indo-West Pacific pocilloporid corals (20%). A few specialized species dominated each of the communities. The habitats and the number of species accounting for over half of the individuals present are (Pacific, Caribbean): sandy beach (1, 1); mangrove (4, 4); P. damicornis (5); and rocky intertidal (3, 6). Most of the species in each community were represented by one or a few individuals.     

Proximate and elemental composition and energy content of mesopelagic crustaceans from the Eastern Gulf of Mexico

The proximate and elemental chemical compositions of 25 species of pelagic decapod and mysid crustaceans collected from the eastern Gulf of Mexico (27°N; 86°W, 1984 to 1989) was examined. Water level ranged from 63 to 95% and increased slightly with species' increased depth of occurrence. Protein levels were generally high (1.5 to 18.3% wet wt, WW; 27.6 to 62.4% ash-free dry wt, AFDW) and comprised the primary organic component in the majority of species. Protein, both as % WW and % AFDW, decreased with increased depth of occurrence. In contrast to protein, lipid levels were low (0.5 to 8.9% WW; 5.7 to 60.9% AFDW), and increased with increased depth of occurrence. Carbon and nitrogen best mirrored measured lipid and protein levels when considered as non-protein carbon and non-chitin nitrogen, respectively. C:N ratios increased with increased depth, consistent with changes in protein and lipid with depth. Because of their compositional attributes, resident Gulf of Mexico species have a low total wet weight energy content relative to species from more productive regions. Energy content showed no significant trend with depth. Vertical migration patterns were distinctly different between shallow-and deep-living gulf species and these differences were largely responsible for the relationships observed between composition and depth. In migrating species, the protein and nitrogen content were higher, the lipid and carbon contents and C:N ratio lower, than in non-migrating species. Three deep-living species of the genus Acanthephyra were found to be compositionally atypical, resembling shallow, migrating types rather than other deep-living, non-migratory species.