Direct sampling and in situ observation of a persistent copepod aggregation in the mesopelagic zone of the Santa Barbara Basin

Observations from a one-person submersible (Wasp) in fall, 1982, revealed a persistent aggregation of non-migrating, Stage V copepodites of Calanus pacificus californicus Brodsky in a band 20±3 m thick at a depth of 450 m, about 100 m above the bottom of the Santa Barbara Basin, California. Copepod abundances, calculated from nearest-neighbor distances measured directly from the submersible, yielded maximum densities of 26×10⁶ copepodites m^-3. Quiescent behavior, low laminarinase activity, low protein content, high lipid content and evidence of low excretion rate all suggest that these copepodites were in a state of diapause. Diapausing C. pacificus californicus at other locations along the eastern Pacific coast were also captured in discrete depth plankton tows. Both the submersible observations and the net collections suggest that the dense aggregation of diapausing copepods we observed in the Santa Barbara Basin was a phenomenon associated with seasonal upwelling cycles, and that such aggregations occur during non-upwelling periods when food is scarce in surface waters. Numerous predators, especially the deep sea smelt Leuroglossus stilbius, were observed feeding upon the aggregated copepods; thus, in contrast to the conventional picture of surface-dominated food distribution, deep-water aggregations of C. pacificus californicus may support the mesopelagic community during periods of low food availability in surface waters.     

Feeding ecology of the loggerhead sea turtle Caretta caretta in the Northwestern Gulf of Mexico

Digestive tract contents collected from carcasses of 82 loggerhead sea turles (Caretta caretta) found on the south Texas coast from 1986 through 1988 were examined. Benthic invertebrates were the predominant prey. Sea pens (Virgularia presbytes), crabs, and mollusks accounted for 94% of the dry weight of the digestive tract samples. Temporal changes in the percent occurrence and percent dry weight of sea pens, crabs, and mollusks in the digestive tract samples were significant. Loggerheads fed primarily on sea pens during spring, then primarily on crabs during summer and fall. The increase of crabs in the loggerhead diet paralleled the annual increase in the abundance of crabs in the Gulf of Mexico. Sea pens were located nearshore at depths of 6 to 12 m and had a disjunct distribution. Present address: Texas A&M University, Department of Biology, College Station, Texas 77843-3258, USA     

Movements of bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in the northwestern Atlantic Ocean recorded by pop-up satellite archival tags

Pop-up satellite archival tags were implanted into 68 Atlantic bluefin tuna (Thunnus thynnus Linnaeus), ranging in size from 91 to 295 kg, in the southern Gulf of Maine (n=67) and off the coast of North Carolina (n=1) between July 2002 and January 2003. Individuals tagged in the Gulf of Maine left that area in late fall and overwintered in northern shelf waters, off the coasts of Virginia and North Carolina, or in offshore waters of the northwestern Atlantic Ocean. In spring, the fish moved either northwards towards the Gulf of Maine or offshore. None of the fish crossed the 45°W management line (separating eastern and western management units) and none traveled towards the Gulf of Mexico or the Straits of Florida (known western Atlantic spawning grounds). The greatest depth recorded was 672 m and the fish experienced temperatures ranging from 3.4 to 28.7°C. Swimming depth was significantly correlated with location, season, size class, time of day, and moon phase. There was also evidence of synchronous vertical behavior and changes in depth distribution in relation to oceanographic features.Communicated by J.P. Grassle, New Brunswick     

Interspecific variation in thermal denaturation of proteins in the congeneric musselsMytilus trossulus andM. galloprovincialis: evidence from the heat-shock response and protein ubiquitination

The genetic population structure of the precominant zooplankter, the copepod Calanus finmarchicus (Gunnerus), was examined to determine whether genetically distinct populations exist in the Gulf of Maine. C. finmarchicus was sampled in three regions of the Gulf of Maine (Great South Channel, spring 1989; northern Gulf of Maine, winter 1990; Great South Channel and Georges Bank, spring 1990). Copepods from seven locations in the Great South Channel, five in the northern Gulf of Maine and four on or near Georges Bank were assayed for allozyme variation and mitochondrial DNA variation of amplified 16S rRNA and cytochrome b genes. Restriction fragment length polymorphism (RFLP) analyses of both mitochondrial DNA genes revealed no variation among any of the individuals assayed. Analysis of five polymorphic allozyme loci revealed that genetic variation among the three geographic regions was low, and genetic identities were high between all locations (I>0.97). Most of the genetic variation was among locations regardless of region. Chi-square tests were used to examine genetic similarity between specific pairs of locations within and between regions. In the northern Gulf of Maine, genetic homogeneity occurred over larger spatial scales (hundreds of km) than in either the Great South Channel or Georges Bank (tens of km). Only copepods from the Bay of Fundy and Nova Scotian Shelf locations were genetically distinct from Wilkinson Basin copepods at two loci. Copepod populations from the northern locations may have been partially isolated or they may represent immigrant populations (e.g., from the Gulf of St. Lawrence). Several pairs of locations were genetically distinct at one or more loci in the two southern regions. Differences between locations in these regions may represent distinct populations advected into the areas at different times or from different sources (e.g., genetic variation may represent a mixture of genetically distinct northern and southern copepod populations). These results suggest extensive gene flow among populations of C. finmarchicus in the Gulf of Maine with some evidence of genetic population subdivision near the Gulf's northeastern and southern boundaries.     

Horizontal movement of ocean sunfish, <Emphasis Type="Italic">Mola mola</Emphasis>, in the northwest Atlantic

Data were retrieved from 25 ocean sunfish (Mola mola) that were tagged with pop-up satellite archival tags in the southern Gulf of Maine (n = 6), off Nantucket Island (n = 17), and off the coast of Georgia (n = 2) between September 2005 and March 2008. Tags remained attached from 7 to 242 days, with a mean attachment period of (X ± SD) 107.2 ± 80.6 days. Ocean sunfish tagged in the Gulf of Maine and southern New England left those areas in the late summer and early autumn and moved south along the continental shelf break. Fish traveled as far south as the Bahamas and the Gulf of Mexico. By moving south, sunfish experienced similar mean sea surface temperatures throughout the tagging period. The maximum straight-line distance traveled by a tagged Mola mola was 2,520 km in 130 days. Two tagged ocean sunfish entered the Gulf of Mexico, one in the December and one in July. Movements were associated with frontal features created by the Gulf Stream and fish moved farther offshore in 2007 when the Gulf Stream was deflected from the shelf break.     

Traditional occupancy–abundance models are inadequate for zero-inflated ecological count data

Traditional occupancy–abundance and abundance–variance–occupancy models do not take into account zero-inflation, which occurs when sampling rare species or in correlated counts arising from repeated measures. In this paper we propose a novel approach extending occupancy–abundance relationships to zero-inflated count data. This approach involves three steps: (1) selecting distributional assumptions and parsimonious models for the count data, (2) estimating abundance, occupancy and variance parameters as functions of site- and/or time-specific covariates, and (3) modelling the occupancy–abundance relationship using the parameters estimated in step 2. Five count datasets were used for comparing standard Poisson and negative binomial distribution (NBD) occupancy–abundance models. Zero-inflated Poisson (ZIP) and zero-inflated negative binomial (ZINB) occupancy–abundance models were introduced for the first time, and these were compared with the Poisson, NBD, He and Gaston's and Wilson and Room's abundance–variance–occupancy models. The percentage of zero counts ranged from 45 to 80% in the datasets analysed. For most of the datasets, the ZINB occupancy–abundance model performed better than the traditional Poisson, NBD and Wilson and Room's model. He and Gaston's model performed better than the ZINB in two out of the five datasets. However, the occupancy predicted by all models increased faster than the observed as density increased resulting in significant mismatch at the highest densities. Limitations of the various models are discussed, and the need for careful choice of count distributions and predictors in estimating abundance and occupancy parameter are indicated.     

Using the negative binomial distribution to model overdispersion in ecological count data

A Poisson process is a commonly used starting point for modeling stochastic variation of ecological count data around a theoretical expectation. However, data typically show more variation than implied by the Poisson distribution. Such overdispersion is often accounted for by using models with different assumptions about how the variance changes with the expectation. The choice of these assumptions can naturally have apparent consequences for statistical inference. We propose a parameterization of the negative binomial distribution, where two overdispersion parameters are introduced to allow for various quadratic mean-variance relationships, including the ones assumed in the most commonly used approaches. Using bird migration as an example, we present hypothetical scenarios on how overdispersion can arise due to sampling, flocking behavior or aggregation, environmental variability, or combinations of these factors. For all considered scenarios, mean-variance relationships can be appropriately described by the negative binomial distribution with two overdispersion parameters. To illustrate, we apply the model to empirical migration data with a high level of overdispersion, gaining clearly different model fits with different assumptions about mean-variance relationships. The proposed framework can be a useful approximation for modeling marginal distributions of independent count data in likelihood-based analyses.     

Quantitative distribution of herbivorous reef fishes in the Gulf of Aqaba (Red Sea)

The distribution of the main herbivorous fishes (Acanthuridae, Scaridae, Siganidae) was studied across a coral reef of the Jordanian coast in the Gulf of Aqaba (Red Sea). Visual counts were realized by diving along transects (200 m long and 5 m wide), parallel to the shore, at 10 stations located from the lagoon to 40 m deep on the outer reef slope. Herbivorous reef fishes reach their highest abundance on the reef front, where 234 fishes were counted per 1,000 m². Their density decreases on the reef flat, with an average of 150 fish 1,000 m^-2, and is lowest on the outer reef slope (69 fish 1,000 m^-2). Surgeonfishes form 63% of the herbivorous ichthyofauna, parrotfishes 35%, and rabbitfishes 2%. Families and species display different distributions according to biota. The Acanthuridae dominate on the reef flat, whereas the Scaridae are more numerous on the outer reef slope. The evolution of the social structure of the main species was observed: the adults generally school in the lagoon and on the reef flat, but are mainly solitary on the reef slope. The distribution of juvenile individuals is more restricted: they are concentrated on the reef front and on the upper part of the reef slope.This study is part of a cooperation programme between the University of Nice (France) and the University of Jordan, to study the ecology of the coral reefs and the surrounding waters of the Jordanian coast (Gulf of Aqaba, Red Sea)     

Impact of submersible pumps on Pb constituents in residential wells

Dissolved Pb in 51 domestic wells screened from 18 to 48 m in glacial tills and outwash deposits were examined in conjunction with the characteristics of their corresponding submersible pump. Pb concentrations, ranging from 0.8 to 24.9 μg l⁻¹, entering residential water supplies were measured during 2001–2004 in the Royal watershed, Maine. Principal component analyses assisted the weighting of pump age, well screen depth, and draw time variables. Preliminary Pb sequestration significance in the boreholes was predicted from geochemical speciation and synchrotron XAS analyses. Nascent ²⁰⁷Pb/²⁰⁶Pb and ²⁰⁸Pb/²⁰⁶Pb isotope analyses assisted the discrimination of possibly leached Pb from submersible pump materials among geogenic sources.     

Monitoring the movements of harbour porpoises (Phocoena phocoena) with satellite telemetry

The movements of nine harbour porpoises, Phocoena phocoena (L.), in the Bay of Fundy and Gulf of Maine were tracked using satellite telemetry. Transmitters were attached to the porpoises in August 1994 and 1995 after they were captured near Grand Manan Island at the mouth of the Bay of Fundy. Tracking periods ranged from 2 to 212 d (mean 50 ± 65 d). Porpoises exhibited a high degree of individual variation in movement patterns; five moved out of the Bay of Fundy into the Gulf of Maine. The porpoise with the longest tracking period moved extensively throughout the Gulf of Maine. These data suggest that seasonal movement patterns of individual harbour porpoises are discrete and are not temporally coordinated migrations. Porpoises that moved out of the Bay of Fundy into the Gulf of Maine did so following the 92 m isobath, which may represent an important movement corridor. The movement of porpoises from the Bay of Fundy into the Gulf of Maine supports the hypothesis that harbour porpoises from these two regions comprise a single population at risk of entanglement in both Canadian and US fisheries. Received: 4 February 1997 / Accepted: 25 August 1997     

一种用粪便样方法估计林麝种群密度的新方法

数量(密度)特征是动物种群的一个重要特征，是定点研究一个野生种群生态的一项非常重要的基础工作．其数量和年龄结构为进一步分析其种群的密度、移动、巢域和核域，尤其是为种群动态提供依据，也是珍稀动物保护和资源动物管理利用的先行工作和重要组成部份．关于野生动物种群数量的具体统计方法，有不少的文章报道，然而从动物的空间分布格局来研究大型脊椎动物却较为少见，这主要是受研究方法的限制．为此，作者对此进行了改进，提出了一种新的研究方法：临时空间样方法，并将其应用到冶勒自然保护区林麝的数量调查研究中去．结果表明：(1)集聚分布指数CA=12．09，说明冶勒自然保护区的林麝有较高的集聚度；(2)林麝粪堆在野外的分布属于聚集分布，并与负二项分布拟合较好；(3)在冶勒自然保护区，生活有62～98只左右的林麝，其密度为：1．0～1．6只／km^2．林麝在冶勒自然保护区呈现聚积分布的原因是由于环境和人类活动因素造成的．图1表2参24     

Sample based estimation of landscape metrics; accuracy of line intersect sampling for estimating edge density and Shannon’s diversity index

A recent trend is to estimate landscape metrics using sample data and cost-efficiency is one important reason for this development. In this study, line intersect sampling (LIS) was used as an alternative to wall-to-wall mapping for estimating Shannon’s diversity index and edge length and density. Monte Carlo simulation was applied to study the statistical performance of the estimators. All combinations of two sampling designs (random and systematic distribution of transects), four sample sizes, five transect configurations (straight line, L, Y, triangle, and quadrat), two transect orientations (fixed and random), and three configuration lengths were tested, each with a large number of simulations. Reference was 50 photos of size 1 km², already manually delineated in vector format by photo interpreters using GIS environment. The performance was compared by root mean square error (RMSE) and bias. The best combination for all three metrics was found to be the systematic design and as response design the straight line configuration with random orientation of transects, with little difference between the fixed and random orientation of transects. The rate of decrease of RMSE for increasing sample size and line length was studied with a mixed linear model. It was found that the RMSE decreased to a larger degree with the systematic design than the random one, especially with increasing sample size. Due to the nonlinearity in the definition of Shannon diversity estimator its estimator has a small and negative bias, decreasing with sample size and line length. Finally, a time study was conducted, measuring the time for registration of line intersections and their lengths on non-delineated aerial photos. The time study showed that long sampling lines were more cost-efficient than short ones for photo-interpretation.     

Small-scale spatial and temporal genetic structure of the Atlantic sea scallop (Placopecten magellanicus) in the inshore Gulf of Maine revealed using AFLPs

Despite long planktonic durations, many species of broadcast spawning invertebrates exhibit genetic structure at small spatial and temporal scales. Amplified fragment length polymorphisms were used to assess genetic variation in the sea scallop, Placopecten magellanicus, among four inshore and one offshore location in the Gulf of Maine and temporal genetic variation among age classes of sea scallops at one site. Our results indicated that genetic structure for P. magellanicus exists on smaller spatial scales (tens to hundreds of kilometers) than expected given the 40-day planktonic larval period. In addition, genetic differences among age classes may be influenced by inter-annual differences in larval supply or reproductive success. Future genetic studies should sample multiple age classes prior to comparison among locations.     

Age-structured modeling reveals long-term declines in the natality of western Steller sea lions.

Since the mid-1970s, the western Steller sea lion (Eumetopias jubatus), inhabiting Alaskan waters from Prince William Sound west through the Aleutian Islands, has declined by over 80%. Changing oceanographic conditions, competition from fishing operations, direct human-related mortality, and predators have been suggested as factors driving the decline, but the indirect and interactive nature of their effects on sea lions have made it difficult to attribute changes in abundance to specific factors. In part, this is because only changes in abundance, not changes in vital rates, are known. To determine how vital rates of the western Steller sea lion have changed during its 28-year decline, we first estimated the changes in Steller sea lion age structure using measurements of animals in aerial photographs taken during population surveys since 1985 in the central Gulf of Alaska (CGOA). We then fit an age-structured model with temporally varying vital rates to the age-structure data and to total population and pup counts. The model fits indicate that birth rate in the CGOA steadily declined from 1976 to 2004. Over the same period, survivorship first dropped severely in the early 1980s, when the population collapsed, and then survivorship steadily recovered. The best-fitting model indicates that in 2004, the birth rate in the central Gulf of Alaska was 36% lower than in the 1970s, while adult and juvenile survivorship were close to or slightly above 1970s levels. These predictions and other model predictions concerning population structure match independent field data from mark-recapture studies and photometric analyses. The dominant eigenvalue for the estimated 2004 Leslie matrix is 1.0014, indicating a stable population. The stability, however, depends on very high adult survival, and the shift in vital rates results in a population that is more sensitive to changes in adult survivorship. Although our modeling analysis focused exclusively on the central Gulf of Alaska, the western Gulf of Alaska and eastern Aleutians show a similar pattern of declining pup fraction with no increase in the juvenile, or pre-breeding, fraction. This suggests that declining birth rate may be a problem for western Steller sea lions across the Gulf of Alaska and into the Aleutian Islands.     

Density-related reproductive trade-offs in the green sea urchin, Strongylocentrotus droebachiensis

Green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), populations are being depleted rapidly in the Gulf of Maine and there is justified concern that the potential of this free-spawner to produce larvae may be severely inhibited. We evaluated the opposing effects of different population densities on gonad development and fertilization success, using population surveys and fertilization experiments. We determined gonad indices (gonad mass/body mass) over a range of population densities (0.1 to 250 ind. m⁻²) at seven sites in coastal Maine, USA, sampled at two depths (5 and 15 m). At shallow sites, we found that gonad indices declined by 50% over the 1500-fold range in adult population density. At 15 m deep locations, gonad mass was consistently low and did not vary significantly with density. Patterns of macroalgal abundance suggest urchins at high density and in deeper water were food limited. Because macroalgal cover co-varies inversely with sea urchin density, we designed field experiments to determine the interaction between sea urchin density and kelp canopy on fertilization success. On square arrays we manipulated the spacing of simulated urchins, but held their numbers constant (five sperm-filled syringes interspersed with four Nitex mesh egg containers permeable to sperm). These experiments, simulating the observed range of natural density, suggested that (1) fertilization rates decreased many times faster than individual gamete production increased over the same range in density, and (2) kelp increased fertilization success at high density when eggs were within 25 cm of a sperm source, but not when spaced 1 m apart. Additional laboratory fertilization experiments at ambient temperatures (3 to 5 °C) indicated that diluted sperm were viable for <1 h, but egg viability was virtually unchanged for >8 h. In short, to the individual the reproductive benefits of aggregating appear to outweigh the costs; and while sperm may be limiting at low population density, eggs may remain viable long enough to be fertilized by sperm from more distant males. Received: 10 February 1998 / Accepted: 22 January 1999     

Small-scale spatial and seasonal differences in the distribution of beach arthropods on the northwestern Tunisian coast. Are species evenly distributed along the shore?

Spatial distributions of sand beach arthropods were studied at Zouara (Nefza), a beach dune system along the northwestern coast of Tunisia. Two transects with pitfall traps, perpendicular to the shoreline and placed from sea towards land, captured spontaneously crawling arthropods. The pitfalls were used to obtain data on horizontal zonations for 2 days during different seasons (April, October). In April, in order to assess local, long-shore distributions of surface-active beach invertebrates, ten transects with pitfall traps were placed every 100 m, for 2 days. Core samples were also taken in correspondence with each of the ten transects to obtain the distribution of burrowing individuals. Sand samples were taken for successive laboratory analyses (salinity, grain size and organic contents). Local climatic conditions were also recorded. One isopod (Tylos europaeus, subdivided into three different size classes), two amphipods (Talitrus saltator, Talorchestia brito and indistinguishable juveniles) and four coleopteran species (Eurynebria complanata, Scarites laevigatus, Phytosus nigriventris, Phaleria acuminata) were considered. Mean zonations perpendicular to the shoreline showed that crustaceans were found closer to the sea-line limits than coleopterans and that between seasons some species changed their mean zonation more than others. Local, long-shore distributions of active and burrowing individuals showed that most species were unevenly distributed along the shore. These discontinuous distributions differed according to the species and age class and were mainly related to specific microclimatic conditions and to nutrition.     

Comparing species abundance models

Five regression models (Poisson, negative binomial, quasi-Poisson, the hurdle model and the zero-inflated Poisson) were used to assess the relationship between the abundance of a vulnerable plant species, Leionema ralstonii, and the environment. The methods differed in their capacity to deal with common properties of ecological data. They were assessed theoretically, and their predictive performance was evaluated with correlation, calibration and error statistics calculated within a bootstrap evaluation procedure that simulated performance for independent data.     

Spatial and temporal distributions of copepods to leeward and windward of Oahu, Hawaiian Archipelago

The spatial and temporal distributions of two island-associated copepod species, Undinula vulgaris Dana and Labidocera madurae Scott, were compared to the distributions of two open ocean species, Cosmocalanus darwinii Lubbock and Scolecithrix danae Lubbock, along 28-km windward and leeward transects off the island of Oahu, Hawaii. Samples were taken in September and December 1985 and April and June 1986. A warm, low salinity pool on the leeward side was a prominent feature during all transects except December. The abundances of the two oceanic species did not change significantly between leeward and windward stations, with distance from shore, or between September 1985 and April 1986 samples. As expected, very high abundances of U. vulgaris occurred at some nearshore stations, up to 3 g dry wt m⁻² for adults alone. Calculations of respiratory loss at these densities (0.7 g C m⁻² d⁻¹) suggest a high local productivity would be required to meet these demands. L. madurae, a surface-dwelling species normally restricted to within 1 km of shore, was an effective indicator species of nearshore water movement. It was more common in offshore samples on the leeward transects, rarely being found offshore on the windward side, consistent with prevailing currents and the presence of the leeward warm, low salinity pool. The occurrence of a strong mixing event in April 1986 resulted in L. madurae being distributed throughout the upper 100 m of the water column. The presence of oceanic species close to shore on the windward side also coincided with this wind-driven event. The primary environmental influence on vertical distributions was daytime cloud cover, with U. vulgaris tending to be found shallower on cloudy days. Of the two oceanic species, S. danae exhibited the most pronounced vertical migration, however, vertical distributions were not significantly correlated with environmental factors for either species. The abundant nearshore U. vulgaris population cannot be explained by differences in vertical distribution between it and the two oceanic species that might allow a physical mechanism to concentrate U. vulgaris. A high population growth rate is likely necessary to explain U. vulgaris' dominance. Received: 26 June 1998 / Accepted: 31 March 1999