Latitudinal variation in local interactions and regional enrichment shape patterns of marine community diversity

While communities are shaped by both local interactions and enrichment from the regional species pool, we propose a hypothesis that the balance of these forces shifts with latitude, with regional enrichment dominating at high latitudes and local interactions dominating at low latitudes. To test this hypothesis, we conducted a latitudinal-scale experiment with marine epifaunal communities. In four regions of the North Atlantic Ocean and Caribbean Sea, we used mimics of ecosystem engineers to manipulate biogenic structural complexity. We iteratively evaluated diversity patterns of experimental communities up to one year after deployment. Additional data were also collected from one of our tropical sites 2.5 years after initial deployment. As hypothesized, we found a reciprocal latitudinal gradient in the effects of the structurally complex mimics and regional enrichment. In the tropics, local diversity was always higher in association with the mimics than in exposed areas that were more open to predation. This effect was consistent across two spatial scales and beyond the one-year timescale of the experiment. In temperate communities, no consistent effects of the mimics on diversity were observed. However, the proportion of species from the regional species pool that were present at the local scale increased from the tropics to the temperate zone, consistent with the hypothesis that higher-latitude communities may experience greater influence from the regional species pool than communities at low latitudes. This study represents the first large-scale experimental demonstration that suggests that the relative impact of local interactions and regional enrichment on community diversity may depend on latitude.     

Reproduction on the edge: large-scale patterns of individual performance in a marine invertebrate

Reproductive output is a central attribute of life history, providing a measure of individual and population performance. The fields of ecology, biogeography, and evolutionary biology take disparate approaches in addressing spatial variation in reproduction, and thus we lack clear predictions for how reproductive output should vary geographically. We empirically investigate these contrasting theoretical approaches by determining geographic patterns in reproductive output for intertidal populations of the purple sea urchin, Strongylocentrotus purpuratus, at 15 sites spanning a large geographic distance (9 degrees span of latitude) from central California, USA, to Baja California, Mexico. Contrary to predictions from biogeography, some of the highest values of reproductive output are at sites near the species' southern range boundary. Additionally, sea urchins do not exhibit a latitudinal gradient in reproduction, but rather show considerable mesoscale variation in reproductive output. Spatial analyses reveal that this variation is correlated with coastal topographic features that are known to influence the pattern of nearshore nutrient flux and circulation. We hypothesize that urchins' reproductive output may be driven by the spatial distribution of their food supply, drift macroalgae, the abundance of which is influenced by both nutrient supply and alongshore transport processes that are coupled to topographic features. Large-scale studies such as this provide valuable insight into the causes of species' range limits, population connectivity, habitat reserve design, and forecasting the effects of climate change on species' distributions.     

Partitioning the factors of spatial variation in regeneration density of shade-tolerant tree species

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.     

Climate vs. soil factors in local adaptation of two common plant species

Evolutionary theory suggests that divergent natural selection in heterogeneous environments can result in locally adapted plant genotypes. To understand local adaptation it is important to study the ecological factors responsible for divergent selection. At a continental scale, variation in climate can be important while at a local scale soil properties could also play a role. We designed an experiment aimed to disentangle the role of climate and (abiotic and biotic) soil properties in local adaptation of two common plant species. A grass (Holcus lanatus) and a legume (Lotus corniculatus), as well as their local soils, were reciprocally transplanted between three sites across an Atlantic-Continental gradient in Europe and grown in common gardens in either their home soil or foreign soils. Growth and reproductive traits were measured over two growing seasons. In both species, we found significant environmental and genetic effects on most of the growth and reproductive traits and a significant interaction between the two environmental effects of soil and climate. The grass species showed significant home site advantage in most of the fitness components, which indicated adaptation to climate. We found no indication that the grass was adapted to local soil conditions. The legume showed a significant home soil advantage for number of fruits only and thus a weak indication of adaptation to soil and no adaptation to climate. Our results show that the importance of climate and soil factors as drivers of local adaptation is species-dependent. This could be related to differences in interactions between plant species and soil biota.     

Geographical variation in the behaviour of a central place forager: Antarctic fur seals foraging in contrasting environments

Foragers show adaptive responses to changes within their environment, and such behavioural plasticity can be a significant driving force in speciation. We investigated how lactating Antarctic fur seals, Arctocephalus gazella, adapt their foraging within two contrasting ecosystems. Location and diving data were collected concurrently, between December 2003 and February 2004, from 43 seals at Bird Island, where krill, Euphausia superba, are the main prey, and 39 at Heard Island, where mostly fish are consumed. Seals at Heard Island were shorter and lighter than those at Bird Island and they spent longer at sea, dived more frequently and spent more time in the bottom phase of dives. Generalized additive mixed effects models showed that diving behaviours differed between the islands. Both populations exploited diel vertically migrating prey species but, on average, Heard Island seals dived deeper and exceeded their estimated aerobic dive limits. We propose that the recovery of the Heard Island population may be limited by the relative inaccessibility and scarcity of food, whereas at Bird Island, the presence of abundant krill resources helps sustain extremely high numbers of seals, even with increased intra- and inter-specific competition. Both populations of fur seals appear to be constrained by their physiological limits, in terms of their optimal diving behaviour. However, there does appear to be some flexibility in strategy at the level of trip with animals adjusting their time at sea and foraging effort, in order to maximize the rate of delivery of energy to their pups.     

DMSP-lyase activity in five marine phytoplankton species: its potential importance in DMS production

Activity of DMSP-lyase, which cleaves dissolved DMSP (henceforth DMSP_d-lyase), was examined in five axenically cultured phytoplankton species, including both DMSP-producing and non-DMSP-producing species. High DMSP_d-lyase activity was found in two DMSP producers, Heterocapsa triquetra strain NIES-7 and Scrippsiella trochoidea strain NIES-369 (Dinophyceae). The DMS production rates at 100 nM DMSP_d were 0.5 fmol cell⁻¹ min⁻¹ for H. triquetra and 0.3 fmol cell⁻¹ min⁻¹ for S. trochoidea. In a non-DMSP producer, Heterosigma akashiwo strain NIES-6 (Raphidophyceae), the DMSP_d-lyase activity was not found. Two DMSP-producing Prymnesiophyceae species, Isochrysis galbana strain CCMP-1323 and Gephyrocapsa oceanica strain NIES-353, did not show any obvious activity either, in contrary to other authors' findings on Phaeocystis sp., another DMSP-producing Prymnesiophyceae species. The comparison of the DMSP_d-lyase activity of the two Dinophyceae species with bacterial DMSP consumption and DMS production activity in Tokyo Bay showed that the DMSP_d-lyase activity of H. triquetra and S. trochoidea could be an important mechanism for DMS production during their blooms. Received: 9 April 1999 / Accepted: 10 December 1999     

Male choice and competition in Tetraopes tetraophthalmus: effects of local sex ratio variation

Summary Male milkweed beetles (Tetraopes tetraophthalmus) are shown to exhibit choice between potential mates. Males, however, also exhibit direct competition for access to potential mates. These differences in male behavior are dependent on the local sex ratio, with males becoming increasingly choosy when the sex ratio is female-biased, and more competitive when the sex ratio is malebiased. Females show neither choice nor competition. In both the laboratory and in the field, body size is important in determining patterns of competition and choice. Larger males win most aggressive encounters, and males generally prefer large females as mates. The combination in a single sex, of competition for mates and mate choice is not predicted by classical sexual selection theory. Male choice and competition may have evolved in T. tetraophthalmus due to the combination of (1) the mating system (2) the pattern of variation in female fecundity, and (3) the spatially and temporally variable operational sex ratios. This explanation is not dependent on high heritability of body size.     

Genetic variation in marine teleosts: High variability in habitat specialists and low variability in habitat generalists

Genetic variation was reviewed in 106 species of marine teleosts. Two heterozygosity estimates were used, one including all protein and enzyme loci and a second excluding the non-enzymatic protein loci. Mean heterozygosities are 0.055±0.036 based on all loci in 106 species and 0.060+0.038 based on enzymatic loci in 89 species. A significant negative correlation was noted between heterozygosity and the proportion of general protein loci included in the estimate. A comparison was made of heterozygosities among taxonomic orders and families, life zones, reproductive mode, geographical range and size. High levels of genetic variation are found in Clupeiformes, Atheriniformes, Pleuronectiformes, temperate pelagic, tropical, intertidal-sublittoral and wide-range species. Low levels of genetic variation are found in Gadiformes, Scorpaeniformes, temperate demersal, polar, and narrowrange species. The most striking differences in heterozygosities are between temperate demersal flatfishes and temperate demersal round fishes. It is suggested that much of the data can be explained by a habitat specialist-generalist model, with high heterozygosities in specialists and low heterozygosities in generalists, but that this is only one of a mosaic of factors which influence genetic variation.     

Nutritional geometry and macronutrient variation in the diets of gannets: the challenges in marine field studies

Foraging theory proposes that the nutritional driver of food choice and foraging in carnivores is energy gain. In contrast, recent laboratory experiments have shown that several species of carnivore select prey that provides a diet with a specific balance of macronutrients, rather than the highest energy content. It remains, however, to be determined how nutritionally variable the foods of predators in the wild are, and whether they feed selectively from available prey to balance their diet. Here, we used a geometric method named the right-angled mixture triangle (RMT) for examining nutritional variability in the prey and selected diets of a group of wild carnivores and marine top predators, the gannets (Morus spp.). A prey-level diet analysis was performed on Australasian gannets (M. serrator) from two New Zealand locations, and the macronutrient composition of their chosen prey species was measured. We use RMT to extend the comparison in the compositions of foods and diets from Australasian gannets from Australia as well as Northern gannets (M. bassanus) and Cape gannets (M. capensis). We found nutritional variability at multiple scales: intra- and interspecific variability in the pelagic fish and squid prey themselves; and intra- and interspecific variability in the diets consumed by geographically disparate populations of gannets. This nutritional variability potentially presents these predatory seabirds with both opportunity to select an optimal diet, and constraint if prevented from securing an optimal diet.     

Alkaline phosphatase activity in marine macrophytes: histochemical localization in some widespread species in southern Spain

Alkaline phosphatase activity (APA) was determined in 44 species of marine macrophytes collected throughout 1991 and 1992 along the southern coast of Spain. Activity varied between 0.83 mol paranitrophenol (pNP) released g^-1 dry wt h^-1 in Ulva rigida var. gigantea and 238.8 mol pNP g^-1 dry wt h^-1 in Bangia fuscopurpurea. Using a histochemical method, APA sites were located in five of these species: Corallina elongata Ellis et Soland, Gelidium latifolium (Grev.) Thur. et Born., G. sesquipedale (Clem.) Born. et Thur., Porphyra umbilicalis (L.) Kützing and Zostera noltii Hornem. Enzymatic activity was found in the outer part of the thallus, either on the cell wall or in the cortical cells. In the marine phanerogam Z. noltii activity was also located in the vascular bundle. The results suggest that APA is present in many, if not all, marine macrophytes from southern Spain, and plays a relevant role in the utilization of dissolved organic phosphorus compounds.     

Seasonal variation in some population parameters of Euchaeta species (copepoda: Calanoida)

Studies of the biology of oceanic copepods are few relative to those of coastal species. Females of the genus Euchaeta have spermatophores attached to the genital somite by the male and carry their broods of eggs attached to this somite, so defining the breeding season. Populations of E. norvegica in the fjordic environment of Loch Etive (collected between 1971 and 1974 and from 1978 to 1979), Scotland and in the marginal oceanic region of the Rockall Trough, northeastern Atlantic Ocean (collected between 1973 and 1976) and of a further eight species of Euchaeta in the Trough are examined. Seasonal changes in the incidence of egg masses and spermatophores attached to the females, sex ratio and population numbers are determined. Sampling errors in the estimation of these parameters are larger in the oceanic Rockall Trough. The Loch Etive populations of E. norvegica produce two generations per year with a proportion, variable between years, of the population producing a third generation. This species produces a single annual generation in the Rockall Trough as do E. acuta and E. pseudotonsa. Two generations per year are probably passed through by E. gracilis in the Trough while the large bathypelagic E. sarsi may produce a single generation in the autumn of every second year. The other bathypelagic species, E. barbata, E. scotti, E. abbreviata and E. longissima, breed continuously throughout the year and no estimate of their generation times is possible. Consequently, this approach to population analysis is useful where breeding is seasonal, but resolving generation time in bathypelagic crustaceans remains a problem.     

Regional coexistence and local dominance in Chaoborus: species sorting along a predation gradient

Variation in the intensity of predation across the well-known environmental gradient of freshwater habitats from small, ephemeral ponds to large, permanent lakes is a key factor in the development and maintenance of aquatic community structure. Here, we present data on the distribution and abundance of four species of Chaoborus (Diptera: Chaoboridae) across this environmental gradient. Chaoborus show a distinct pattern of species sorting when aquatic systems are divided into fish and fishless environments, and this pattern is consistent with species traits known to influence their vulnerability to fish predation (i.e., pigmentation, diel vertical migration [DVM] behavior, and body size). To test whether fish are the drivers of this pattern, we created a gradient in fish density by stocking bluegill sunfish (Lepomis macrochirus) into 15 experimental ponds in southwestern Michigan, USA, and then allowed Chaoborus species to colonize. There was clear evidence of species sorting along the predation gradient; Chaoborus americanus was most abundant in the fishless ponds, C. flavicans was neutral in response to fish, and C. punctipennis and C. albatus were most abundant at high fish biomass, a response consistent with their field pattern. Furthermore, prey preference experiments confirm that size selective predation and differences in Chaoborus species traits contribute to the pattern of Chaoborus abundance and distribution.     

Identifying trophic variation in a marine suspension feeder: DNA- and stable isotope-based dietary analysis in Mytilus spp.

Accurate field data on trophic interactions for suspension feeders are lacking, and new approaches to dietary analysis are necessary. Polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) was integrated with stable isotope analysis to examine dietary patterns in suspension-feeding Mytilus spp. from seven spatially discrete locations within a semi-enclosed marine bay (Strangford Lough, Northern Ireland) during June 2009. Results of the two methods were highly correlated, reflecting dietary variation in a similar manner. Variation in PCR-DGGE data was more strongly correlated with the principal environmental gradient (distance from the opening to the Irish Sea), while values of δ ¹³C and δ ¹⁵N became progressively enriched, suggesting a greater dependence on animal tissue and benthic microalgae. Diatoms and crustaceans were the most frequently observed phylotypes identified by sequencing, but specific DNA results provided little support for the trophic trends observed in the stable isotope data. This combined approach offers an increased level of trophic insight for suspension feeders and could be applied to other organisms.     

Genetic variation in seven goby species (Perciformes: Gobiidae) assessed by electrophoresis and taxonomic inference

Data on the electrophoretic patterns of 20 enzyme loci were used to investigate relationships among seven Mediterranean goby species. The study evidenced different levels of genetic variability among the species. On the whole, the degrees of genetic differentiation were consistent with the current taxonomic recognition of species and genera. Phenetic and phylogenetic analyses agreed in clustering the samples in three groups, i.e. the Gobius (including Zosterisessor ophiocephalus), the Lesueurigobius species and Pomatoschiarua minutus. The taxa analysed probably evolved following the complex geological and climatic events that have been affecting the Mediterranean basin ever since the Miocene. Received: 20 June 1998 / Accepted: 5 February 1999     

Heritable pheromone blend variation in a local population of the butterbur borer moth Ostrinia zaguliaevi (Lepidoptera: Crambidae)

Summary. In a local population of Ostrinia zaguliaevi Mutuura & Munroe (Lepidoptera: Crambidae), extensive variation was found in the blend ratio of three sex pheromone components, (Z)-9-tetradecenyl acetate (Z9-14:OAc, 10.2-63.8%), (Z)-11-tetradecenyl acetate (Z11-14:OAc, 32.2-86.8%), and (E)-11-tetradecenyl acetate (E11-14:OAc, 2.1-11.9%). The variation was observed over a three-year period (2002-2004). Mother-daughter regression analyses have shown that although the heritability of the minor component E11-14:OAc was not significant, the heritability of the proportions of Z9-14:OAc and Z11-14:OAc were substantial (0.5–0.6). In artificial selection experiments, the mean % Z9-14:OAc in the sex pheromone changed significantly within three generations (37% in the control line, 48 and 52% for two lines selected for increase, and 23 and 30% for two lines selected for decrease). Despite these changes, the amounts of fatty acyl pheromone precursors, (Z)-9-, (Z)-11- and (E)-11-tetradecenoate, in the pheromone gland were not significantly affected by the selection. Taken together, variation in the pheromone blend of O. zaguliaevi is likely to be attributable to a few genes involved in the reduction or acetylation of fatty acyl pheromone precursors, the last two steps in pheromone biosynthesis.     

Predicted impact of ocean acidification on a marine invertebrate: elevated CO<Subscript>2</Subscript> alters response to thermal stress in sea urchin larvae

Ocean acidification (OA) and the biological consequences of altered seawater chemistry have emerged as a significant environmental threat to healthy marine ecosystems. Because a more acidic ocean interferes with fixation of calcium carbonate to form shells or calcified skeletons, future ocean chemistry may significantly alter the physiology of calcifying marine organisms. These alterations may manifest themselves directly in the calcification process, or have synergistic effects with other environmental factors such as elevated temperatures. New tools permit us to explore subtle changes in gene expression patterns in response to environmental conditions. We raised sea urchins (Strongylocentrotus franciscanus) under conditions simulating future atmospheric CO₂ levels of 540 and 970 ppm. When larvae raised under elevated CO₂ conditions were subjected to 1-h acute temperature stress, their ability to mount a physiological response (as measured by expression of the molecular chaperone hsp70) was reduced relative to those raised under ambient CO₂ conditions. These results represent the first use of gene expression assays to study the effects of OA on sea urchin development. They highlight the importance of looking at multiple environmental factors simultaneously as this approach may reveal previously unsuspected biological impacts of atmospheric changes.     

Providing a common diet to different marine decapods does not standardize the fatty acid profiles of their larvae: a warning sign for experimentation using invertebrate larvae produced in captivity

Larval decapods are commonly produced in captivity and employed in experiments to evaluate interspecific physiological and biochemical differences. Currently, it is still unknown if different decapod species provided a common diet and exposed to identical abiotic conditions produce newly hatched larvae (NHL) with similar fatty acid (FA) profiles. This study analyzed the FA composition of NHL from five marine shrimp species (Lysmata amboinensis, L. boggessi, L. debelius, L. seticaudata and Rhynchocinetes durbanensis) fed a common diet and stocked at constant temperature. FA profiles of NHL differed significantly within and among genera. NHL from species unable to molt from zoea I to zoea II in the absence of food (L. amboinensis, L. debelius and R. durbanensis) displayed the lowest FA contents. Researchers must be aware that providing a common diet to different species, even if closely related, may not standardize the FA profile of NHL produced in captivity.     

Marine calanoid resting eggs in Norway: Abundance and distribution of two copepod species in the sediment of an enclosed marine basin

Abundance and distribution of calanoid resting eggs in sediment were examined in 1988–1989 in an enclosed marine basin (Svartatjønn; western Norway). The basin undergoes rotenone treatment and draining every winter. Egg densities were highest in autumn, when up to 2 × 10⁶ eggs m^–2 were found, and lowest in summer. Egg densities were also highest in the deeper parts of the basin and in the upper 1 cm of sediment. Hatching was initiated in late January by pumping warmer deep water from the outside area into the basin. Evidence is given for resting-egg formation inEurytemora affinis Poppe, 1880 andAcartia clausi Gurney, 1931 (present phenotype reclassified asA. teclae by Bradford 1976: N. Z. Jl mar. Freshwat. Res. 10: 159–202). Seasonal differences in inhibition of hatching, together with the longevity and tolerance of these eggs, strongly suggest that they are diapause eggs.