Patch Occupancy and Potential Metapopulation Dynamics of Three Forest Mammals in Fragmented Afromontane Forest in South Africa

Abstract: We investigated the persistence of three medium-sized (2–9 kg), rare forest mammals in the fragmented mist-belt Podocarpus forests of the midlands of KwaZulu-Natal Province, South Africa. We recorded patch occupancy of blue duiker (   Philantomba monticola ), tree hyrax (   Dendrohyrax arboreus ), and samango monkey ( Cercopithecus mitis labiatus ) in 199 forest patches. Their rarity is ascribed to the fragmentation and destruction of their forest habitat. Incidence functions, derived from presence and absence data, were formulated as generalized linear models, and environmental effects were included in the fitted logistic models. The small and mostly solitary hyrax and duiker persisted in smaller patches than the large and social monkey. Although this result follows expectations based on relative home-range sizes of each species, the incidence probability of the samango monkey was invariant with increasing isolation, whereas a gradual decrease with increasing isolation was observed for the hyrax and duiker. Group dynamics may inhibit dispersal and increase the isolation effect in social species such as samango monkeys. A mainland-island metapopulation model adequately describes patterns of patch occupancy by the hyrax and duiker, but the monkeys' poor dispersal ability and obvious area-dependent extirpation suggest that they exist in transient, nonequilibrium (declining) metapopulations. Through identification of large forest patches for careful protection and management, the survival of all three species—especially the monkey—could be prolonged. Because no functional metapopulation may exist for the monkey, however, this is an emergency measure. For the duiker and hyrax, larger patches should form part of a network of smaller and closer patches in a natural matrix.     

Contrasts in Livelihoods and Protein Intake between Commercial and Subsistence Bushmeat Hunters in Two Villages on Bioko Island,Equatorial Guinea

Across West and Central Africa, wildlife provides a source of food and income. We investigated the relation between bushmeat hunting and household wealth and protein consumption in 2 rural communities in Bioko Island, Equatorial Guinea. One village was dedicated to commercial hunting, the other trapped game primarily for food. We tested whether commercial‐hunter households were nutritionally advantaged over subsistence‐hunter households due to their higher income from the bushmeat trade and greater access to wild‐animal protein. We conducted bushmeat‐offtake surveys in both villages (captures by hunters and carcasses arriving to each village). Mammals (including threatened primates: black colobus [Colobus satanas], Preussi's guenon [Allochrocebus preussi], and russet‐eared guenon [Cercopithecus erythrotis]), birds, and reptiles were hunted. The blue duiker (Philantomba monticola), giant pouched rat (Cricetomys emini), and brush‐tailed porcupine (Atherurus africanus) contributed almost all the animal biomass hunted, consumed, or sold in both villages. Monkeys and Ogilbyi's duikers (Cephalophus ogilbyi) were hunted only by commercial hunters. Commercial hunters generated a mean of US$2000/year from bushmeat sales. Households with commercial hunters were on average wealthier, generated more income, spent more money on nonessential goods, and bought more products they did not grow. By contrast, households with subsistence hunters spent less on market items, spent more on essential products, and grew more of their own food. Despite these differences, average consumption of vegetable protein and domestic meat and bushmeat protein did not differ between villages. Our results highlight the importance of understanding the socioeconomic and nutritional context of commercial and subsistence bushmeat hunting to correctly interpret ways of reducing their effects on threatened species and to enable the sustainable offtake of more productive taxa. Contrastes en el Sustento y la Ingesta de Proteínas entre Carne de Caza de Subsistencia y Comercial en Dos Aldeas en Isla Bioko, Guinea Ecuatorial     

Mass Differences among Male Mangabey Monkeys Inhabiting Logged and Unlogged Forest Compartments

Abstract: Remedies for reduction in primate numbers following rainforest habitat disturbance must incorporate understanding of how disturbance affects population biology. Between July 1996 and July 1998, I captured 31 adult male grey cheeked mangabeys (  Lophocebus albigena) from groups occupying logged and unlogged forest in Kibale National Park, Uganda. Captured males were weighed, measured, fitted with radiocollars, and radiotracked three to five times a week. Males from groups in the unlogged forest were significantly heavier than males from the logged forest. Males that immigrated into the study groups were significantly lighter than resident males in unlogged forest, but they did not differ in mass from males in logged forest. I tested hypotheses that these differences were due to differences in age, skeletal size, and tissue mass. Results support the hypothesis that differences are due to differences in tissue mass. Results of further tests showed that differences in tissue mass were explained more by differences in habitat type than by rates of male turnover. There was no evidence to suggest that differences in tissue mass were related to differences in costs of travel, crowding, or seasonal variation in habitat quality. My results suggest that selective logging may affect primate biomass, although these changes may or may not be reflected in estimates of primate density.     

Nestling provisioning in polygynous great reed warblers (Acrocephalus arundinaceus): do males bring larger prey to compensate for fewer nest visits?

Most studies of social polygyny in birds have examined male provisioning on the basis of the number of feeding visits. This may be misleading if males compensate for infrequent visits by bringing larger prey at each visit. We investigated nestling provisioning in the socially polygynous great reed warbler, Acrocephalus arundinaceus, in south Central Sweden in 1996–1997. We collected data on rate of feeding visits, prey size and the amount of biomass delivered by males and females. Males had lower rates of feeding visits and provided smaller prey to nestlings in secondary than in monogamous and primary nests. Secondary females had higher rates of feeding visits and brought larger prey than monogamous and primary females. These results confirm that secondary females face a potential cost of polygyny through a lower rate of male feeding, and that this cost was reinforced by the significantly lower male provisioning rate (biomass h^–1) at secondary nests. Secondary females compensated for the lack of male assistance by increasing their rate of feeding and bringing larger prey. As a result, offspring in nests of secondary females received as much food as did those in nests of primary females. Prey load size increased with the parent’s proportion of feeding visits, suggesting that parents use different feeding strategies depending on their amount of responsibility for nestling provisioning. We suggest that parents which take the main responsibility for nestling feeding have to forage further away from the nest, and based on optimal-foraging theory, they should then on average bring larger prey to their nest. Received: 4 April 1999 / Received in revised form: 12 October 1999 / Accepted: 23 October 1999     

Prey selectivity by crowned hawk-eagles on monkeys in the Kibale Forest,Uganda

Summary Results are presented from a 3.25-year study of a nesting pair of crowned hawk-eagles and an 18-year study of a primate community in the Kibale Forest, Uganda. The proportional composition of the living population of prey species was compared with that of eagle prey and animals dying from other causes. Monkeys were the predominant pry (83.7%). They also dominated the medium-to large-sized mammalian carcasses dying from other causes (88.9%). The eagles selected prey according to species, age, and sex. Selectivity by age and sex differed between prey species. Among red colobus monkeys, the eagles selected young juveniles and infants, but in four other monkey species they selected adult males. Eagle prey selectivity by species generally supports the hypothesis that polyspecific associations among the monkeys are effective deterrents against predation. The prey/predator ratio for the Kibale eagles was much higher, but the annual offtake of prey by the eagles was much lower than that of tropical felids. Mortality due to causes other than eagles was greater than expected in red colobus and less in redtails, but not significantly different from expected or equivocal in the other three monkey species. Other cause of mortality affected adult male and infant red colobus more than expected. Among the other four monkey species, significantly more adult males and fewer adult females died from these other causes than expected. Eagle predation had a major impact on the populations of adult males of both black and white colobus and blue monkeys and on both adult male and female mangabeys. The selectivity appeared to contribute significantly to the differential adult sex ratio in four monkey species, but not in red colobus. In contrast, mortality incurred during fighting among adult males probably accounted for the differential adult sex ratio in red colobus. Offprint requests to: T.T. Struhsaker at his current address     

Sea-bird aggregation at a deep North Pacific seamount

During June 1991, we studied sea birds at a mid-ocean seamount (Fieberling Guyot) in the eastern North Pacific Ocean. Avifaunal composition changed from small Procellariiformes [a storm-petrel; Oceanodroma leucorhoa (Vieillot)] away from the seamount to an assemblage dominated by larger tubenoses [mostly black-footed albatross Diomedea nigripes Audubon and Cook's petrel Pterodroma cookii (Gray)]. Compared to adjacent waters, sea-bird density and biomass within a 30-km radius centered on the seamount summit were 2.4 and 8 times higher, respectively. Individual sea-bird taxa were 2 to 40 times more abundant at the seamount relative to values reported previously from large-scale surveys of deep-ocean regions in the central North Pacific. In September 1991 we studied potential prey of sea birds in the upper water column using a neuston net and multiple opening-closing net system (MOCNESS) tows. Most potential prey types in the neuston exhibited no significant enhancement over the seamount. MOCNESS samples at 10 m depth, however, showed several prey types to be more abundant over the seamount, and the dominant size class of fish was slightly larger. We attribute the sea-bird aggregation observed at this seamount to changes in the abundance and/or behavior of pelagic organisms in the deep scattering layer (not adequately sampled in this study), perhaps augmented by migrations of seamount residents into the surface layers. Processes on and in the vicinity of seamounts may provide spatially-predictable prey to wide-ranging aerial sea birds foraging in this relatively austere environment.     

Predatory behavior of crowned hawk-eagles (Stephanoaetus coronatus) in Kibale National Park, Uganda

Evaluating the nature and significance of predation on populations of wild primates has been difficult given a paucity of data regarding the phenomenon. Here we addressed this problem in a 37-month study of the predatory behavior of crowned hawk-eagles living at the Ngogo study site in Kibale National Park, Uganda. We collected prey remains underneath the nests of two pairs of eagles and census data on potential prey species to investigate prey selection and the ecological impact of predation on the Ngogo primate population. Results indicate that primates form the vast majority of all prey items. Eagles prey selectively on monkeys according to sex and species. Male primates were taken more often than females, while two species, redtail monkeys and mangabeys, were captured significantly more and less, respectively, than chance expectation. In addition, there was no bias in the age of prey: adult and non-adults were killed in numbers roughly equal to their proportional representations in the forest. Further analyses indicate that a non-trivial fraction of the entire primate population at Ngogo succumbs to crowned hawk-eagle predation each year. These results reveal both parallels and contrasts with those reported previously. Some of the parallels are due to similarities in prey availability, while contrasts are likely related to methodological differences between studies, inter- individual variations in predator hunting styles, and differences in prey abundance, demography, and behavior. Received: 29 March 2000 / Revised: 6 June 2000 / Accepted: 15 October 2000     

Sea Otter Mortality Estimated from Carcasses Collected after the Exxon Valdez Oil Spill

Large numbers of sea otters (  Enhydra lutris) died following the 1989 Exxon Valdez oil spill. Two previous studies estimated spill-related mortality, one from the difference between counts of otters before and after the spill, and the other from the recovery rate of tagged carcasses that had been released at sea. I used a derivative of the second approach, but revised it to account for (1) moribund otters that had hauled out on shore; (2) carcasses that were collected at sea; and (3) differences in search effort in different areas of the spill—considerations that were not addressed previously. The mortality estimation procedure presented here, and applied to Prince William Sound, Alaska, had six input parameters, two with fixed values (the number of spill-related carcasses collected [391] and the number of otters taken alive that died in captivity [84]) and four that were assigned a range of values (the recovery rate for carcasses on the beach [60–90%], the proportion of recovered carcasses that were onshore [70–95%]), the proportion of offshore deaths recovered onshore (20–50%), and the proportion of moribund otters that hauled out (20–60%). Empirical estimates for these parameters were not available after the spill, but a range of plausible values for each was delimited using other available data. Randomly-selected combinations of values within these ranges produced mortality estimates for the sound ( x¯= 750, 5–95% quantiles ≈ 600–1000) that were lower than indicated by previous studies. These lower estimates do not diminish the tragic nature of the mortality, but instead highlight the significance of both the process and informational basis in producing estimates of catastrophic loss. Estimates of loss based on faulty or inadequate data may mislead investigations of population recovery and may jeopardize public trust in scientific assessments of future catastrophes.     

Diel variation and seasonal consistency in the fish assemblage of the non-estuarine Sulaibikhat Bay,Kuwait

In the period between September 1986 and August 1988, using an otter trawl, a total of 50 species from 30 families were captured in Sulaibikhat Bay, Kuwait. The dominant species were Liza carinata (Valenciennes), Pomadasys stridens (Forrskal) and Leiognathus brevirostris (Valenciennes). Although the total number of fishes captured in 1986–1987 and 1987–1988 differed by an order of magnitude, the seasonal changes in numbers, biomass and number of species in the intertidal and subtidal zones were consistent. Unbalanced three-way analysis of variance showed that significantly larger numbers of fishes and number of species, but not biomass, were captured in the intertidal at night compared to the intertidal during the day. This difference may be due to both increased net efficiency and movement of fishes into the intertidal zone at night to avoid piscivorous fishes that move from deep water into shallower water. During the day few fish are present in the intertidal zone and this may be to avoid predation by piscivorous birds. Several species of fishes were shown to have different patterns of behaviour during the diel period.     

Zooplankton capture by two scleractinian corals,Madracis mirabilis andMontastrea cavernosa,in a field enclosure

Capture of zooplankton by scleractinian corals has been noted for several species, yet quantitative information on rates of capture and differential capture by prey taxon has been lacking. We used field enclosures to examine prey capture for two coral species,Madracis mirabilis (Duchassaing and Michelotti) andMontastrea cavernosa (Linnaeus), on the north coast of Jamaica (Discovery Bay) in November 1989, February and March 1990, and January 1992.M. mirabilis has small polyps and a branching colony morphology (high surface/volume ratio), whereasM. cavernosa has large polyps and mounding colonies (low surface/volume ratio). Corals were isolated front potential prey, then were introduced into enclosures with enhanced zooplankton concentrations for 15- to 20-min feeding periods. Corals were fixed immediately after the experiment to prevent digestion, and coelenteron contents were examined for captured zooplankton. Plankton pumps were used to sample ambient zooplankton in the enclosures near the end of each run. Selectivity and capture rates were calculated for each prey taxon in each experiment; both indices were high for relatively uncommon large prey, and low for copepods, which were often the most common items in the plankton. Sizes of zooplankton captured by both species were generally larger than those available considering all prey taxa combined, but were almost the same for both coral species, even though the corals' polyp sizes are very different. This occurred primarily because small copepods, with low capture rates, dominated most plankton samples. For specific prey species, or group of species, there were few significant differences in size between the prey available and the prey captured.M. mirabilis, with small polyps, also captured far more prey per unit coral biomass than didM. cavernosa, with much larger polyps. We hypothesize that the large differences in capture rate of prey taxa are related to escape or avoidance behavior by those potential prey, and to the mechanics of capture, rather than to any selectivity by the corals.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

Habitat Mosaic, Wildlife Availability, and Hunting in the Tropical Forest of Calakmul, Mexico

Abstract: Habitat loss and subsistence hunting are two of the main activities that affect wildlife in frontier areas. We compared subsistence hunting patterns in four villages with different ethnic composition and degree of habitat disturbance in the vicinity of Calakmul Biosphere Reserve, Campeche, Mexico. We also compared differences between some of these villages in harvest composition and prey availability to determine hunting preferences. We used a Landsat TM satellite image to analyze the degree of disturbance around the villages. We conducted periodic surveys of subsistence hunting and prey availability. Wildlife availability was assessed monthly on nine transects (3000 m) established in the vicinity of three villages. The relative amount of disturbed habitat was smaller in an indigenous Maya village ( IV ) and larger in a mestizo village ( MV ). The two mixed-composition villages ( MCVs) had intermediate levels of disturbance. Ten species, four large and six small, of birds and mammals accounted for 97% of the hunting records. Hunting was more intense in IV and less intense in MCV1. The three village types had different hunting preferences. The habitat-mosaic composition in the vicinity of the villages influenced prey availability and subsistence-hunting preferences. Changes in the habitat mosaic were caused by the size of the holding and by ethnic composition. In spite of longer settlement time, the habitat mosaic in the vicinity of IV was less transformed than that of the other sites. Their larger holding size and greater diversity of economic activities may explain why the Mayas at IV have transformed the landscape less than the other groups and can hunt more and larger prey.     

Identifying important species that amplify or mitigate the interactive effects of human impacts on marine food webs

Some species may have a larger role than others in the transfer of complex effects of multiple human stressors, such as changes in biomass, through marine food webs. We devised a novel approach to identify such species. We constructed annual interaction-effect networks (IENs) of the simulated changes in biomass between species of the southeastern Australian marine system. Each annual IEN was composed of the species linked by either an additive (sum of the individual stressor response), synergistic (lower biomass compared with additive effects), or antagonistic (greater biomass compared with additive effects) response to the interaction effect of ocean warming, ocean acidification, and fisheries. Structurally, over the simulation period, the number of species and links in the synergistic IENs increased and the network structure became more stable. The stability of the antagonistic IENs decreased and became more vulnerable to the loss of species. In contrast, there was no change in the structural attributes of species linked by an additive response. Using indices common in food-web and network theory, we identified the species in each IEN for which a change in biomass from stressor effects would disproportionately affect the biomass of other species via direct and indirect local, intermediate, and global predator–prey feeding interactions. Knowing the species that transfer the most synergistic or antagonistic responses in a food-web may inform conservation under increasing multiple-stressor impacts.     

Prey selection by the scyphomedusan predator Aurelia aurita

We describe feeding behavior of Aurelia aurita (Linnaeus) using gut content analyses of field-collected specimens and a mesocosm experiment. The field studies were conducted in Narragansett Bay, Rhode Island, USA from March to April 1988, and the mesocosm studies were done at the Marine Ecosystems Research Laboratory at the University of Rhode Island. Patterns of prey selection changed with medusa diameter. Smaller medusae (12 mm diameter) consumed mostly hydromedusan prey whereas larger medusae (up to 30 mm diameter) ingested greater numbers of copepod prey. While larger medusae did feed on copepods, their diet also contained more barnacle nauplii and hydromedusae than expected from the relative abundances of these prey types in plankton samples. A marginal flow mechanism of feeding by A. aurita provided an explanation for the patterns of prey selection we observed in medusae of different sizes and among widely divergent prey types. Our data indicated that large prey, with escape speeds slower than the marginal flow velocities around the bell margins of A. aurita, made up a substantial fraction of the daily ration when they were available. Such prey species may be more important to nutrition than the more abundant copepods and microzooplankton. Successful development of young medusae may depend upon an adequate supply of slowly escaping prey.     

Fishery Stability, Local Extinctions, and Shifts in Community Structure in Skates

Abstract: Skates are arguably the most vulnerable of exploited marine fishes. Their vulnerability is often assessed by examining fisheries catch trends, but these data are not generally recorded on a species basis except in France. Aggregated skate catch statistics tend to exhibit more stable trends than those of other elasmobranch fisheries. We tested whether such apparent stability in aggregated catch trends could mask population declines of individual species. We examined two time series of species-specific surveys of a relatively stable skate fishery in the northeast Atlantic. These surveys revealed the disappearance of two skate species, long-nose skate (   Dipturus oxyrhinchus ) and white skate (   Rostroraja alba ) and confirmed a previously documented decline of the common skate (   D. batis ). Of the remaining five skate species, the three larger ones have declined, whereas two smaller species have increased in abundance. The increase in abundance and biomass of the smaller species has resulted in the stability of the aggregated catch trends. Because there is significant dietary overlap among species, we suggest the increase in abundance of the smaller species may be due to competitive release as the larger species declined. A consequence of this kind of stability is that declining species cannot be detected without species-specific data, especially in taxa exhibiting competitive interactions. This may explain why previously documented disappearances of two species of skates went unnoticed for so long. The conservation of skates and other elasmobranchs requires species-specific monitoring and special attention to larger species.     

Microbial decomposition of proteins and lipids in copepod versus rotifer carcasses

Zooplankton carcasses are common within aquatic systems and potentially serve as organic-rich substrates for bacteria. We compared the microbial decomposition of representative crustacean (copepod) and non-crustacean (rotifer) zooplankton carcasses and monitored changes in carcass protein and lipid contents. Our results showed that carcass decomposition was mainly driven by bacteria colonizing from the surrounding water. Carcass-associated bacteria displayed higher protease and lipase activities than free-living bacteria. Protein content of copepod carcasses decreased by 70% within the first 8 h and shifted from larger to smaller sized proteins, while protein loss in rotifer carcasses was insignificant. Carcass lipid content did not change significantly over 24 h in either zooplankton type, although polar branched fatty acids increased on copepod carcasses indicating an increase in viable microbial biomass. Our results suggest differential turnover of protein versus lipid within a zooplankton carcass and that carcasses from different zooplankton groups would affect water column microbial processes differently.     

Individual- and population-level responses of a keystone predator to geographic variation in prey

Investigating how food supply regulates the behavior and population structure of predators remains a central focus of population and community ecology. These responses will determine the strength of bottom-up processes through the food web, which can potentially lead to coupled top-down regulation of local communities. However, characterizing the bottom-up effects of prey is difficult in the case of generalist predators and particularly with predators that have large dispersal scales, attributes that characterize most marine top predators. Here we use long-term data on mussel, barnacle, limpet, and other adult prey abundance and recruitment at sites spread over 970 km to investigate individual- and population-level responses of the keystone intertidal sunstar Heliaster helianthus on the coast of Chile. Our results show that this generalist predator responds to changes in the supply of an apparently preferred prey, the competitively dominant mussel Perumytilus purpuratus. Individual-level parameters (diet composition, per capita prey consumption, predator size) positively responded to increased mussel abundance and recruitment, whereas population-level parameters (density, biomass, size structure) did not respond to bottom-up prey variation among sites separated by a few kilometers. No other intertidal prey elicited positive individual predator responses in this species, even though a large number of other prey species was always included in the diet. Moreover, examining predator-prey correlations at approximately 80, 160, and 200 km did not change this pattern, suggesting that positive prey feedback could occur over even larger spatial scales or as a geographically unstructured process. Thus individual-level responses were not transferred to population changes over the range of spatial scales examined here, highlighting the need to examine community regulation processes over multiple spatial scales.     

The effect of predation on artificial reef juvenile demersal fish species

There is a concern that artificial reefs (AR) may act purely as fishing aggregation devices. Predators attracted to ARs can influence the distribution and abundance of prey fish species. Determining the role of predators in AR is important in advancing the understanding of community interactions. This paper documents the effects of predation on fish assemblages of AR located near a coastal lagoon fish nursery. The Dicentrarchus labrax is a very opportunistic species preying on juveniles (0⁺ and 1⁺ age classes) of several demersal fish species on the ARs. Reef prey and sea bass abundance were negatively correlated. The mean numbers of prey per sea bass stomach increased with the increase of reef fish prey abundance, suggesting that predation has a significant influence, resulting in a decrease in prey abundance. Prey mortality (4–48%) of demersal reef fish associated species depends on bass density. Prey selection was related both with prey abundance and vulnerability. Results showed that D. labrax predation on AR-fish associated species can increase prey natural mortality. However, the role of bass predation on the ecological functioning of exploited ARs is not clear. There may be increases in local fishing yields due either to an increase in predator biomass through aggregation of sea bass attracted to ARs or to greater production. In contrast, predation on juveniles of economically important reef fish preys, especially the most frequent and abundant (Boops boops), can contribute to a decrease in recruitment to the fishery. Our results indicate that inter-specific interactions (predator–prey) are important in terms of conservation and management, as well as for the evaluation of the long-term effects of reef deployment. Thus, it is necessary to consider ecological interactions, such as predation, prior to the development and deployment of artificial habitats as a tool for rehabilitation.     

Occurrence of large numbers of carcasses of the large,grazing copepod Calanus cristatus from the Japan Sea

Carcasses of Calanus cristatus were discovered in plankton samples collected from the Japan Sea throughout the year from 1970 to 1985. Many carcasses of copepodite Stages IV and V occurred in the layer between 15 and 300 m below a distinct thermocline. The number of copepodite Stage V carcasses also peaked in the layer between 1 500 and 2 000 m. The highest density of copepodite Stages IV and V carcasses was 169 individuals per 1 000 m³ and 1 573 individuals per 1 000 m³, respectively. Carcasses of adults occurred at depths below 500 m and numbers of males and females per 1 000 m³ were 1 to 16 and 1 to 42, respectively. Living males were larger in catch number than living females, but the relationship for carcasses was the opposite. Weight of carcasses was 15 to 25% of living C. cristatus. Carcasses contained about 51% carbon and 8% nitrogen by weight. Carcasses may have been drifting for more than one year in the epipelagic layer under the thermocline because of their slow decomposition rate.