Underappreciated species in ecology: "ugly fish" in the northwest Atlantic Ocean.

Species shifts and replacements are common in ecological studies. Observations thereof serve as the impetus for many ecological endeavors. Many of the species now known to dominate ecosystem functioning were largely ignored until studies of those underappreciated species elucidated their critical roles. Recognizing the potential importance of underappreciated species has implications for functional redundancies in ecosystems and should alter our approach to long-term monitoring. One example of an applied ecological system containing species shifts, underappreciated species, and potential changes in functional redundancies is the topic of fisheries. The demersal component of many fish communities usually consists of high-profile and commercially valuable species that are targets of fisheries, plus a diverse group of lesser known species that have minimal commercial value and focus. Yet ecologically these traditionally nontargeted species are often a major biomass sink in marine ecosystems and can also be critical in the functioning of bentho-demersal food webs. I examined the biomass trajectories of several species of skates, cottids, lophiids, anarhichadids, zooarcids, and similar species in the northeast U.S. Atlantic ecosystem to determine whether their relative abundance has changed across the past four decades. Distribution and stomach contents of these species were also evaluated over time to further elucidate the relative importance of these species. Landings of these underappreciated bentho-demersal fish were also examined in comparison to those species that historically have been commercially targeted. Of particular emphasis was the evaluation of evidence for sequential stock depletion and the ramifications for functional redundancy for this ecosystem. Results indicate that some of these fish species are now the dominant piscivores, benthivores, and scavengers in this ecosystem. These formerly under-studied species generally have either maintained a consistent population size or have increased in abundance (and expanded in distribution) over the past several decades. Nontraditionally targeted fish species are an often overlooked but important component of bentho-demersal fish communities. Implications for the energy flow and resilience specifically for future fisheries and generally for harvesting biological resources are significant, remaining critical issues for the world's ecosystems.     

Species-specific differences in pure tonal whistle vocalizations of five western North Atlantic dolphin species

Summary Pure tonal whistle vocalizations from five species of dolphins found in the western North Atlantic had consistent, species-specific characteristics. The degree of differences between species, as based on the results of multivariate discriminant analysis (Fig. 2), correlated with the taxonomic and zoogeographic relations of the five dolphin species. Congeneric species had more similar vocalizations than species of different genera. Differences between sympatric species were greater than differences between allopatric species. Of the six whistle parameters measured, maximum frequency had the lowest coefficient of variation for all five species, and duration and number of inflection points had the highest coefficients of variation for all five species.     

Diets of four deep-water scaphopod species (Mollusca) in the North Atlantic and the Nordic Seas

The buccal pouch contents of four deep-water scaphopod (Mollusca) species, Pulsellum affine, Pulsellum teres, Siphonodentalium lobatum and Polyschides olivi, collected in the North Atlantic and the Nordic Seas, were examined. The buccal pouches of the examined scaphopod species contained almost exclusively foraminifers, which agrees with previous studies on shallow and deep-water scaphopods. The scaphopods seemed to be engulfing at least some of the most common foraminifer species, which are distributed in the cold waters of the Nordic Seas or the temperate waters in the northernmost part of the North Atlantic. About half of the food species (30 foraminifer species) occurred in only one scaphopod species, nearly always in very low numbers, and seemed to be an unimportant food source. The remaining and more frequent food species occurred in two or more scaphopod species. A similarity tree was calculated with parsimony and presence–absence coding of the food species in the scaphopods. This resulted in the similarity tree: 100%{P. teres 79%[P. affine 70%(P. olivi and P. lobatum)]}, when branch support was measured with percentage jackknife stability. A general trend was found towards larger food items being consumed by larger scaphopod species, indicated by the positive correlation between the size of the scaphopods and the food species. It is suggested that microdistribution of the foraminifers and the large individual size of the foraminifers may be important factors in avoiding predation by scaphopods.     

Venoms from six species of marine fish: lethal and hemolytic activities and their neutralization by commercial stonefish antivenom

A specimen of the stonefish Synanceja verrucosa was captured in Okinawa in March 1988. Live specimens of the scorpionfish Inimicus japonicus were purchased from the Tokyo Central Wholesale Market in November 1988 and those of four species of zebrafish Pterois lumulata, P. volitans, P. antennata and Dendrochirus zebra from an aquarium in December 1988. Crude venoms were extracted from dorsal spines of the six species. All venoms exhibited lethal activity against mice and hemolytic activity specific for rabbit erythrocytes. The lethal activity (or hemolytic activity) of each venom was very unstable to freezing, lyophilization and heating. Both lethal and hemolytic activities of S. verrucosa venom were remarkably neutralized by the commerical stonefish (Synanceja trachynis) antivenom. This antivenom was also effective, to some extent, in counteracting the other venom activities. The neutralizing capacities of the antivenom were calculated to be 7310 LD₅₀ ml^-1 for S. verrucosa venom and 1 220 to 2 990 LD₅₀ ml^-1 for the other venoms. Results of neutralization tests suggest that venoms from the six species were comparable in terms of antigenecity.     

Levels of selected heavy metals in commercial fish from five freshwater lakes,Pakistan

The levels of As, Cd, Cr, Cu, Fe, Mn, Ni, Pb, Zn and Hg are estimated by atomic absorption in the muscle of six commercial species of fish belonging to five freshwater lakes of Pakistan. The species included Mastacembelus armatus, Tor putitura, Mystus seenghala, Wallago attu, Catla catla and Labeo rohita. The fish were procured in comparable weight ranges so that a viable comparison of trace metal content could be effected. As, Fe, Pb, Zn and Hg showed elevated levels of 0.006–6.967, 0.933–6.133, 0.060–4.108, 0.978–5.363 and 0.030–3.211 μg/g, wet weight. The relevant statistical parameters, such as standard deviation, standard error, skewness and its t‐value are also reported for establishing the randomness of the distribution in relation to the corresponding data of freshwater lakes of the world and examined in view of species‐specificity and origin.     

Seasonal distribution of modern coccolithophores in the western North Atlantic Ocean

The seasonal successions of standing crop and floral assemblage were studied for oceanic coccolithophores at 5 weather stations in the western North Atlantic Ocean. The coccolithophores were less populous during winter at the northern stations, whereas they became scarce during summer at the southern stations. The average standing crop in the surface water was quite consistent throughout all climatic regions, being a few tens of thousand cells per liter. The average standing crop at the 100 m level showed a clear trend of increasing southwards. The population at this depth was only one-tenth as large as that of the surface level at the northernmost station, whereas it was almost two-thirds as large as the surface levels at the southernmost station. Many of the major species exhibited seasonally biased occurrences. The trend was mostly repeated in consecutive years, although the magnitude of their abundances fluctuated from year to year in some case. The average species composition was obtained for the 5 weather stations at the surfacewater (surface and 20 m levels combined) and 100 m levels; Emiliania huxleyi dominated the flora at all stations. The relationship between occurrences of major species and ambient water temperature was also studied. Although some species were stenothermal, the majority exhibited eurythermal characteristics; this accounts for the high specific diversity of the coccolithophore community throughout the wide range of waters studied.Contribution No. 2899 of the Lamont-Doherty Geological Observatory of Columbia University.     

Concentrations of V,Cr, Mn,Fe, Ni,Co, Cu,Zn, As and Cd in mesopelagic crustaceans from the North East Atlantic Ocean

Specimens of oceanic decapods, mysids and euphausiids, collected from the North East Atlantic Ocean during July 1985, were analysed for V, Cr, Mn, Fe, Ni, Co, Cu, Zn, As and Cd. Measurement of the metals was carried out using two techniques: inductively-coupled plasma-source mass spectrometry (ICP-MS) and atomic absorption (AAS). With the exception of the iron data, good agreement was observed between the data from the two techniques. Baseline-metal data are presented for the species measured, along with any effects of animal size on metal concentration. The data presented for oceanic animals from different trophic levels are important in studies of metal fluxes and for the assessment of markers within the food web in the oceans.     

Evolutionary genetics of the Mytilus edulis complex in the North Atlantic region

Genetic relationships among Mytilus populations throughout the North Atlantic region, including the Mediterranean and the Baltic Sea, were studied using enzyme electrophoresis. Three distinct groups of populations, each of a remarkably wide distribution, can be recognised on the basis of their multilocus allelic composition: (1) M. galloprovincialis L. of the Mediterranean and western Europe; (2) a genetically distinct form of M. edulis Lmk. from both the Baltic Sea and some localities in the Canadian Maritime Provinces (here provisionally termed the trossulus type mussel); and (3) the traditional Atlantic M. edulis populations of northwestern European coasts and most of eastern North America. These groups are regarded as representing three relatively old evolutionary lineages, which all deserve separate and equal systematic status. The main part of the differentiation at most of the loci studied is accounted for by this major systematic pattern, but considerable geographical differentiation within each of the three principal groups was also detected. At single loci, different electromorphs were found to prevail in disjunct populations of M. galloprovincialis (Mediterranean/Britain) and of the trossulus-type mussel (Baltic/eastern Canada). Within the Atlantic M. edulis, a major part of the differentiation is transoceanic. At one locus (Ap), geographic differentiation appeared to be relatively independent of the systematic boundaries; the possible role of interlineage hybridisation in contact areas in regulating the pattern of geographical variation is discussed.     

Benzo[a]pyrene oxidation and microsomal enzyme activity in the mussel (Mytilus edulis) and other bivalve mollusc species from the Western North Atlantic

Analysis of subcellular fractions revealed a complement of microsomal electron transport components including reductases and heme proteins in several organs of the three bivalve species Mytilus edulis, Macrocallista maculata and Area zebra. Dithionite difference spectroscopy of CO-treated microsomes yielded spectra typical of cytochrome P-450 in digestive gland and gill, with absorption maxima at 450 nm. A time-dependent reduction of cytochrome P-450 was also observed. The levels of these components and rates of microsomal benzo[a]pyrene (BP) metabolism were highest in the digestive gland, and were very similar between species. In M. edulis there was a suggested seasonal variation in BP metabolism but no population differences in this activity or in levels of other components. Digestive gland microsomal metabolites of BP identified by HPLC retention and UV spectroscopy included BP-1.6-quinone, BP-3,6-quinone and BP-6,12-quinone, which comprised 65% of the total metabolites, and dihydrodiols and phenols, the latter products consistent with cytochrome P-450 monooxygenation and expoxide hydrolase function. However, the inconsistent dependence of BP metabolism on NADPH, and inconsistent inhibition by CO suggest that catalyst(s) additional to cytochrome P-450 may be acting in BP metabolism. Based on these results and the prominent quinone formation, we speculate that peroxidative mechanism(s) may be involved. The role of peroxidative as well as well as monooxygenase reactions in the in-vivo disposition and effects of foreign chemicals in bivalves, and also the major function of cytochrome P-450 in these bivalves, remain to be established.Some of these results have appeared in preliminary form; J. J. Stegeman, Sea Grant Annual Report, Words Hole Oceanographic Institution, p 15, 1981     

A process-convolution approach to modelling temperatures in the North Atlantic Ocean

This paper develops a process-convolution approach for space-time modelling. With this approach, a dependent process is constructed by convolving a simple, perhaps independent, process. Since the convolution kernel may evolve over space and time, this approach lends itself to specifying models with non-stationary dependence structure. The model is motivated by an application from oceanography: estimation of the mean temperature field in the North Atlantic Ocean as a function of spatial location and time. The large amount of this data poses some difficulties; hence computational considerations weigh heavily in some modelling aspects. A Bayesian approach is taken here which relies on Markov chain Monte Carlo for exploring the posterior distribution.     

Tracking adult North Atlantic bluefin tuna (Thunnus thynnus) in the northwestern Atlantic using ultrasonic telemetry

Ultrasonic, depth-sensitive transmitters were used to track the horizontal and vertical movements, for up to 48 h, of 11 adult (136 to 340 kg estimated body mass) North Atlantic bluefin tuna (Thunnus thynnus Linnaeus). Fish were tracked in October 1995, September and October 1996, and August and September 1997 in the Gulf of Maine, northwestern Atlantic. The objective was to document the behavior of these fish and their schools in order to provide the spatial, temporal, and environmental information required for direct (i.e. fishery-independent) assessment of adult bluefin tuna abundance using aerial surveys. Transmitters were attached to free-swimming fish using a harpoon attachment technique, and all fish remained within the Gulf of Maine while being followed. Most of the bluefin tuna tagged on Stellwagen Bank or in Cape Cod Bay (and followed for at least 30 h) held a predominately easterly course with net horizontal displacements of up to 76 km d⁻¹. Mean (±SD) swimming depth for all fish was 14 ± 4.7 m and maximum depth for individuals ranged from 22 to 215 m. All but one fish made their deepest excursions, often single descents, at dawn and dusk. In general, adult bluefin tuna spent <8% of their time at the surface (0 to 1 m), <19% in the top 4 m, but >90% in the uppermost 30 m. Mean (±SD) speed over ground was 5.9 km h⁻¹, but for brief periods surpassed 20 to 31 km h⁻¹. Sea surface temperatures during tracking were 11.5 to 22.0 °C, and minimum temperatures encountered by the fish ranged from 6.0 to 9.0 °C. Tagged bluefin tuna and their schools frequented ocean fronts marked by mixed vertebrate feeding assemblages, which included sea birds, baleen whales, basking sharks, and other bluefin schools. Received: 19 July 1999 / Accepted: 25 March 2000     

Patterns of temperature adaptation in North American Atlantic coastal actinians

Although acutely measured rate processes in coelenterates show close correlation with the Van't Hoff Q₁₀ rule and the distribution of many coelenterate groups is correlated with temperature, little is known about coelenterate temperature adaptation. Analysis of lethal temperatures shows that the southern distribution of 3 species of North Atlantic actinians is correlated with their upper lethal temperature. Oxygen consumption data from acute measurements indicate conformity to the Q₁₀ rule. Oxygen consumption data from animals pre-exposed to various temperatures indicate that oxygen consumption is capable of acclimation to temperature. Metridium senile from Massachusetts shows positive acclimation, typical of a poikilotherm partially regulating its metabolic rate in response to temperature change. Two species from Virginia, Haliplanella luciae and Diadumene leucolena, appear to show a reverse pattern. Their response includes encystment and negative adjustment of metabolic rate, showing evasion rather than regulation in response to environmental change. Temperature has a marked effect on parameters of the activity pattern of Diadumene leucolena, but the temperature sensitivity of activity is not correlated with that of oxygen consumption.     

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.     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic

Fishes and zooplankton were obtained (March–April 1979 and partly in August 1974) from 45 hauls taken during the day and at night in the central equatorial Atlantic between Latitude 3°N and 2°S from the surface to 1250-m depth, using the RMT 1+8, a combined opening-closing plankton and micronekton trawl. The vertical distribution of 30 myctophid species is described. All species migrate in a diel pattern, Ceratoscopelus warmingii and Lampanyctus photonotus down to at least 1250 m. During daytime most species aggregated at 400-to 700-m depth, therefore only partly occupying the depth of the Deep Scattering Layer (400 to 500 m at 15 kHz). The feeding patterns of seven of the most abundant species were compared, with a total of 1 905 stomach contents being analysed. All seven species are regarded as opportunistic predators, which feed predominantly during the night on calanoid copepods. A total of 66 species of calanoid copepods were identified among the prey items, with smaller species definitely being in the minority. Stomachs of C. warmingii (700 to 1 250 m depth) and Lepidophanes guentheri (500 to 900 m depth) from daytime samples contained copepod species restricted to the upper 150 m of the water column, including Undinula vulgaris, Nannocalanus minor, and Euchaeta marina, thereby confirming an extended vertical migration of predators. Differences in diet and preferences between species in their total food spectrum are described.     

Contrasting feeding patterns among species of fish larvae from the tropical Andaman Sea

Feeding habits of tropical fish larvae were analysed in a comparative study of four species (Scorpaenodes sp., Carangoides sp., Acanthocepola sp. and Cynoglossus sp.) from the Andaman Sea. We investigated morphological characteristics and their potential influence on larval feeding, and looked for common patterns in larval prey preference. Gut contents of a total of 300 larvae were examined and compared with local zooplankton composition. The feeding habits of the investigated larvae shared a number of characteristics. During ontogeny both the preferred prey size and the number of prey in the gut increased, and across all larval size classes the relative prey size spectrum stayed constant, of approximately the same magnitude for all four species. On the other hand, larval feeding also differed in a number of aspects, especially differences in the taxonomic composition of preferred prey were apparent. Scorpaenodes sp. preferred abundant and large prey taxa, Acanthocepola sp. and Carangoides sp. preferred large, but less common prey taxa, while Cynoglossus sp., which had the relatively smallest mouth size, preferred smaller sized prey groups. Hence, the findings indicate that from an offset of common characteristics, especially related to prey size preference, larvae have their individual feeding patterns related to specific morphology and patterns of distribution.Communicated by M. Kühl, Helsingør     

Phylogeography of the rock-pool copepod Tigriopus brevicornis (Harpacticoida) in the northern North Atlantic, and its relationship to other species of the genus

We investigated relationships among North Atlantic Tigriopus brevicornis populations and their relationships to Mediterranean T. fulvus and North American T. californicus, using crossing experiments and mitochondrial DNA sequencing. All T. brevicornis populations tested were interfertile, while interspecific crosses produced either no offspring or offspring that did not survive past the larval stage, with the exception of a few T. brevicornis × T. californicus crosses that produced mature adults. DNA sequencing of a fragment of mitochondrial cytochrome oxidase I (COI) showed that samples of T. brevicornis from Iceland, the Faroes, Ireland, Scotland and Nova Scotia formed a single shallow clade. In contrast, T. brevicornis from more southern populations in France and Portugal formed a clade with substantially greater branch lengths. Tigriopus brevicornis was monophyletic, and T. brevicornis plus Mediterranean T. fulvus were together also monophyletic. The phylogeography of T. brevicornis closely mirrored that found in T. californicus, with substantially reduced interpopulation divergence at northern latitudes. The known distribution of T. brevicornis in Iceland and the Faroes is shown and dispersal mechanisms and habitat selection briefly discussed.     

Breeding and larval distribution of the pteropod Clione limacina in the North Atlantic,Subarctic and North Pacific Oceans

The pteropod Clione limacina (Phipps, 1774) is an arcticboreal, circumpolar species, which is widely distributed in the North Atlantic and Subarctic Oceans; it also occurs in the North Pacific Ocean (in the Oyashio and neighbouring waters) and along the Atlantic coast of North America in the waters of the cold Labrador current to the Cape Hatteras region (35° N). The distribution of C. limacina larvae in the plankton of the Norwegian, Barents and White Seas, the Bear Island-Spitsbergen region of the Greenland Sea, the Newfoundland Grand Bank and the Flemish-Cap Bank region of the North-western Atlantic Ocean, and the Kurile-Kamchatka region of the North-western Pacific Ocean has been studied, and information from literature concerning the reproduction and larval occurrence of the species is summarized. Throughout its distributional are, spawning of C. limacina is characterized by the same general ecological pattern. This species breeds and spawns in all types of water masses occurring within the vertical range which it commonly inhabits — from surface layers to 500 m water depth. In all local populations of the species, the most intensive spawning is correlated with the spring/summer period of annual heating of the local waters, and the highest abundance parallels maximum growth of phytoplankton which serves as food for veligers and early polytrochous larvae. After the end of this period, spawning intensity in all local C. limacina populations declines sharply, but spawning continues at low intensity during the autumn/winter season, being practically continuous throughout the year. Distribution patterns of C. limacina larvae are determined by those of their parental forms (the parental forms spawn in the zones permanently inhabited). The earliest larval stages of C. limacina (veligers) are present predominantly in the upper 100 or 200 m water layer, i.e. in the zone of high phytoplankton abundance. Polytrochous larvae, after becoming predaceous feeders, are distributed throughout the whole water column from the surface to 500 m depth, similar to adult C. limacina. As with the adults, larvae are present (within the species' distribution area) in all types of water masses. Since the beginning of the twentieth century, in the course of the warming of the Arctic Ocean, the southern race of C. limacina (formerly a summer/autumn seasonal invader in the Norwegian Sea) has become a permanent component of the plankton fauna of the Norwegian and Barents Seas in regions influenced by the Norwegian-Northcape Current System.     

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     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin