Feeding rate of slimy sculpin and burbot on young lake charr in laboratory reefs

Synopsis Predation and contaminants are two possible factors in the poor recruitment of young lake charr Salvelinus namaycush in the Great Lakes. We measured the feeding rate of slimy sculpins Cottus cognatus and burbot Lota lota on young lake charr (uncontaminated young from eggs of a hatchery brood stock and contaminated young from eggs of Lake Michigan lake charr) in laboratory test chambers with a cobble substrate. The median daily consumption rate of sculpins for all tests was 2 lake charr eggs (N = 22 tests; 95% confidence interval, 0–13) and 2 lake charr free embryos (N = 31 tests; 95% confidence interval, 0–10). Feeding rate did not differ between hatchery and contaminated prey. Slimy sculpins continued to feed on lake charr when another prey organism, the deepwater amphipod Pontoporeia hoyi, was present. Feeding by burbot on free embryos (4–36 d^–1) increased as the mobility of young increased, but burbot consumed about 10% of their body weight weekly in free-swimming young (140–380 d^–1). Predation on lake charr eggs by sculpins could be considerable over the 100 to 140 d incubation period, and burbot could eat large numbers of free-swimming lake charr as the young fish left the reef. Predation pressure on young lake charr may inhibit rehabilitation of self-sustaining populations of lake charr on some reefs unless a critical egg density has been reached.     

Shifts in abundance and growth of slimy sculpin in response to changes in the predator population in an arctic Alaskan lake

Lake trout (Salvelinus namaycush) in Toolik Lake are tightly coupled to the benthos, since they have no pelagic forage fishes. Slimy sculpins (Cottus cognatus) are a prey of lake trout and the soft sediment chironomids are an important prey for the sculpin. Our previous work showed that the median size of lake trout in Toolik Lake had decreased significantly between 1977 and 1986, and smaller lake trout are likely to be less effective as sculpin predators. Using our historic data on the slimy sculpin population from 1978, we took advantage of the recent change in the predator community to examine for subsequent changes in the sculpin community. Between 1978 and 1987, the percentage of slimy sculpin caught in the soft sediments has increased (25% to 39.5%). In 1987 there was a significant difference in the mean weight of sculpin caught on different substrates. The mean individual weight of sculpins increased from the nearshore rock area to the rock/soft-sediment interface to the soft sediments. There was no difference in mean individual weight with habitat in 1978. The mean total length at age for slimy sculpins during this time has also increased significantly. We suggest that the risk of predation while foraging in the soft sediments has declined. The increased use of the soft sediment area appears to have contributed to their increased growth, likely due to greater food abundance.     

The influence of riparian vegetation on the functional organization of four Hong Kong stream communities

Pontoporeia affinis Lindström predominated profundal macroinvertebrate communities in eutrophic Lake Mälaren. Amphipod densities, at both stations studied, averaged > 50% of overall total macroinvertebrate densities. The life history of P. affinis was two years, with two cohorts distinguished, both exhibiting similar patterns in annual growth. During the sixteen years of this study, amphipod growth seemed to exhibit long-term cyclic fluctuations, with periods of 9–10 yrs. at Görväln and 5–6 yrs. at Grönsöfjärden. Amphipod growth was significantly (negatively) correlated with conspecific density and biomass. The mechanism(s) governing Pontoporeia population oscillations is (are) not clear. It is postulated that changes in food availability, as the result of overgrazing, may adversely affect Pontoporeia recruitment and subsequently be observed in the form of long-term population oscillations as noted in Lake Mälaren.     

Seasonal feeding ecology of co-existing native and invasive benthic fish along a nearshore to offshore gradient in Lake Michigan

Relative abundance, diet composition and feeding strategy were determined for three benthic fish, the native deepwater sculpin Myoxocephalus thompsonii (Girard, 1851) and slimy sculpin Cottus cognatus (Richardson, 1836), and the invasive round goby Neogobius melanostomus (Pallas, 1814), along a nearshore to offshore gradient in southeastern Lake Michigan during March–December 2010, 2015, and 2016. Round goby were most abundant in the nearshore (<25 m), slimy sculpin were most abundant in the transitional zone (35–65 m), and deepwater sculpin were most abundant in the offshore zone (>75 m). Despite a large degree of spatial separation, some species did overlap, with slimy and deepwater sculpin occurring in sympatry throughout the year in the offshore and transitional zones, and round goby overlapping with both sculpin species seasonally in the transitional zone. Deepwater sculpin exhibited specialization on Mysis diluviana in all depth regions. Slimy sculpin in the offshore reduced diet overlap with deepwater sculpin by specializing on fish eggs during spring and fall, whereas in the transitional depth zone, there was considerable overlap between sculpin species due to the high importance of Mysis in diets. The invasive round goby had a mixed diet, with some diet overlap with native sculpin, especially slimy sculpin, in the transitional zone. In the nearshore zone, round goby displayed a generalized diet with many prey contributing to the diet, but the average contribution of any prey was generally low. Spatial separation and variable feeding strategies help reduce, but not eliminate shared resource use amongst these benthic fish in Lake Michigan.     

Basin-scale Patterns in the Drift of Embryonic and Larval Fishes and Lamprey Ammocoetes in two Coastal Rivers

We studied the distribution and abundance of drifting embryonic and larval fishes and lampreys in the Smith and Van Duzen rivers of northern California, U.S.A. We collected seven fish species in four families and at least one lamprey species in the drift. All taxa drifted almost exclusively at night. Sculpins, Cottus aleuticus and C. asper, outnumbered all other taxa, comprising 63% of the catch in the Van Duzen River and 90% of the catch in the Smith River. We estimated that sculpin drift reached 3×10⁷ individualsh^–1 during the relatively high flow period from late winter through early summer. Most sculpin in these two rivers appeared to drift to the estuaries; we estimated 2.5×10⁹ sculpin embryos and larvae reached the Smith River estuary in 1995. In contrast to the sculpins, the patterns in the drift of other taxa suggest limited transport to the estuaries. Suckers, Catostomus occidentalis in the Van Duzen River, C. rimiculus in the Smith River, threespine stickleback, Gasterosteus aculeatus, and lamprey, Lampetra tridentata and possibly L. richardsoni, drifted at much lower rates and later in the year than sculpins. In the Van Duzen River, drift appeared to serve as a dispersal mechanism for only one of three introduced cyprinids. California roach, Lavinia symmetricus, drifted at low rates throughout the summer while we captured only seven Sacramento pikeminnow, Ptychocheilus grandis, and no speckled dace, Rhynichthys osculus. The information gathered on the drift of early life history phases is germane to both the conservation of native fishes and management of non-indigenous species in coastal rivers.     

The life history,production and food habits of Pontoporeia affinis Lindström (Crustacea: Amphipoda) in mesotrophic Lake Erken

The life history and feeding habits of Pontoporeia affinis Lindström were studied in mesotrophic Lake Erken during 1982 to 1983. The greatest densities and biomasses of P. affinis occurred at 12 m (662 ind./m² and 118.2 mg dw/m², respectively) with values decreasing at the shallower and deeper regions. The life cycle of Pontoporeia was one year, with recruitment occurring in April, at which time the majority of the population consisted of individuals <2 mm. Overall the gut content of Pontoporeia was predominantly composed of detritus (97.6%) with algae accounting for 2.3%. Annual production was highest at 12 m (386.2 mg dw/m²) and lowest at 16 m (8.1). The highest production occurred during September, coinciding with the autumn Bacillariophyceae bloom.     

Comparative ecology of prickly sculpin,Cottus asper,and coastrange sculpin,C. aleuticus,in the Eel River,California

Synopsis We documented species' distributions, size structure of populations, abundance in mainstem and tributary streams, habitat use, and diets of prickly sculpin, Cottus asper, and coastrange sculpin, C. aleuticus, in the Eel River drainage of California, to determine the processes allowing coexistence of these very similar fishes. We observed prickly sculpins at 43 sites and coastrange sculpins at 34. The species co-occurred at 26 sites. Young-of-year coastrange sculpins were only observed within 42 km of the ocean, but young-of-year prickly sculpins were present throughout the species range. Mean, maximum, and minimum lengths of coastrange sculpins were positively correlated with distance from the ocean but no significant relationships were found for prickly sculpins. Absolute abundance of both species was highest in mainstem habitat (prickly sculpins = 0.6 sculpins m^–2 and coastrange sculpins = 0.4 sculpins m^–2) . Tributary densities of both species tended to be less than 0.1 sculpins m^–2. The species inhabited very similar habitats and had very similar diets. Coastrange sculpin populations in upstream areas were maintained by immigration from downstream areas in contrast with prickly sculpin populations that produced young-of-year fish throughout their range. Densities were probably not high enough for interspecific interactions to be important. The factors limiting the upstream distribution of the species may include high water temperatures, stability of the stream bed, and behavior of the fish. In the past, the range of sculpins within the Eel River drainage probably fluctuated with changing physical conditions. Recent introductions of exotic species that compete with and prey upon sculpins, and ongoing human activities in the drainage could result in major reductions in the distribution and abundance of one or both species.     

Prédation exercée par le chabot (Cottus Gobio L.) sur l'alevin de truite commune (Salmo trutta L.): taille maximale de capture des alevins par les chabots

In laboratory conditions, where predation by sculpins (Cottus gobio L.) upon brown trout fry (Salmo trutta L.) was strong, predation was shown to be closely linked to total length of prey and predator. The limit of this predation can be defined by the equation: Y = 0.484 X + 5.8 (Y = total length of the trout fry in mm and X = total length of the sculpin in mm).

     

Size-selective predation by fourhorn sculpin,Myoxocephalus quadricornis (L.) (Pisces) on Mesidotea entomon (L.) (Crustacea,Isopoda)

During three different seasons Mesidotea entomon specimens from fourhorn sculpin stomachs were analyzed, and compared with the M. entomon population in the field. The field samplings were carried out in the northern Bothnian Sea. Feeding experiments revealed fourhorn sculpins to be highly selective when feeding on M. entomon.The fourhorn sculpin most preferably selected large M. entomon in the field as well as in the laboratory experiments. The preference for large M. entomon remained after correction for availability of differently sized M. entomon. In summer the actual sizes of M. entomon eaten by fourhorn sculpin were smaller than in autumn and winter. During all three seasons the oldest M. entomon were the ones most preferred. Ultimate effects of predation by fourhorn sculpin on the life-history of M. entomon are discussed.     

Growth and age of staghorn sculpins of the genus <Emphasis Type="Italic">Gymnocanthus</Emphasis> (Cottidae) in Peter the Great Bay and adjacent waters of Primorye (Sea of Japan)

Age and growth have been studied in four species of staghorn sculpins of the genus Gymnocanthus inhabiting Primorye (the Russian sector of the Sea of Japan). Large black edged and purplegray sculpins, G. herzensteini and G. detrisus, inhabited deeper waters compared with threaded sculpin G. pistilliger and (especially) whip sculpin G. intermedius. The size differences between different sculpin species could already be revealed in the diameter of mature oocytes and the length of hatching larvae and fries passing to the benthic phase of development. The maximum size in these species correlated with their maximum age. G. herzensteini, G. detrisus, G. pistilliger, and G. intermedius could attain the age of no less than 16+, 15+, 9+, and 7+, respectively. The gains in fish of the same age were also greater in species attaining greater size values. In all the species, females lived longer than males and were larger than males of the same age. Throughout the life cycle, the growth rate gradually came down; the decrease was especially pronounced during the period of sexual maturation. In the southern part of the geographic range, the specimens of purplegray sculpin attained smaller size and showed lower growth rate compared with the fish from the northern part of the range. The threaded sculpin demonstrated the inverse relationships. In general, females were dominating in samples of staghorn sculpins; however in small and medium size groups, the proportion of males was greater, which was due to their smaller, compared with females, absolute gains. In elder groups, the proportion of males gradually went down, as they were dying earlier than females.     

Factors influencing densities and distributions of Pontoporeia hoyi in Lake Ontario

The physical and chemical characteristics of the sediment-water interface greatly influence distributions of the bottom-feeding amphipod Pontoporeia hoyi which is a major component of the Great Lakes aquatic food-webs. Sediment-water interface samples from the Great Lakes indicate that Pontoporeia densities are positively correlated with sediment concentrations of organic carbon, particularly in the upper depth range of the species. Pontoporeia is not found in anoxic sediments. Growth rates and production of Pontoporeia are influenced by the quality of food supplies and there appears to be a seasonal response to the down-flux of plankton detritus. Comparisons of numerations and distribution patterns also revealed an apparent positive relationship between Pontoporeia and sediment-associated bacteria. Interlake comparisons of population densities provide no clear evidence of any large scale impact by contaminated sediment on this species. Depending upon the extent to which organic carbon (OC) is assimilated in the water column, bottom sediments may store and buffer the quantity of OC available to Pontoporeia. Thus, if the annual down-flux of OC remains generally constant, populations of Pontoporeia should also remain constant unless influenced by changes in predation intensity.Numerations of Pontoporeia revealed that populations of this amphipod are much larger in Lake Michigan than in Lake Ontario. We suggest that much of this difference can be accounted for by different forms of predator-prey interaction near the base of the benthic food-web. Commercial fisheries data tend to support this idea. We also speculate that differences in the concentration of dissolved silica may figure prominantly among factors that affect Pontoporeia population densities.Under proposed new taxonomy (Bousfield, 1989), Pontoporeia hoyi is now classified as Diporeia hoyi. However, for convenience, we have retained the commonly used genus Pontoporeia throughout this text.     

Comparative life histories and microhabitat use in three sympatric sculpins (Cottidae:Cottus) in northeastern California

Synopsis Life histories of three sculpins endemic to the Pit River system, northeastern California, are detailed.Cottus pitensis is widely distributed throughout the drainage. This species is typically found in rapidly flowing, shallow water on coarse substrate. It grows to a large size, >100 mm, and lives to 5 years. Spawning occurs annually, in early spring. Fecundity is relatively low, <320 oocytes.Cottus asperrimus is restricted to a large population in Fall River and smaller concentrations in Hat Creek and Pit River. It is a relatively small fish, <80 mm, and lives to 5 years. It occurs in deep, slow-moving, often spring-fed streams, typically on fine substrates. Spawning lasts from autumn to early spring. Males establish nests on hard substrates and guard several clutches of eggs. Fecundity ranges from 140–580 oocytes.Cottus klamathensis macrops is relatively rare in the drainage. The largest concentration is in the Hat Creek system, smaller populations exist in Fall River and Pit River. This sculpin is found on coarse substrate, often where aquatic vegetation is abundant. It is typically found in slow, relatively shallow water. It grows to >100mm and lives to 5 years. Males guard eggs during the short spawning season beginning in late winter. Fecundity in this species reaches a maximum of 650 oocytes. Life history strategies vary within the genus, yet the life histories of these sculpins are more similar to each other than they are to many of their congeners. This within-group similarity may have resulted from exposure of all three species to similar environmental pressures. This is predicted by life-history theory.     

Feeding interactions and diet of carnivorous fishes in the Shelikhov Bay of the Sea of Okhotsk

The food spectra, trophic statuses, and feeding interrelations of three most abundant benthic carnivorous fish species inhabiting the Shelikhov Bay—the Pacific cod Gadus macrocephalus, the great sculpin Myoxocephalus polyacanthocephalus, and the Okhotsk sculpin M. ochotensis—are considered based on materials collected during the complex survey of the RV Professor Kaganovsky of the TINRO-Center, in September 2004. It was found that these species were facultative predators with wide food spectra. The significance of prey objects in the diet of the Okhotsk sculpin was as follows: crustaceans, fish, and mollusks. Great sculpin and Pacific cod preyed mostly on fish, then on crustaceans, and mollusks. Pacific cod ate equal proportions of fish and decapods. All the species had age-related variability of diet. The potential competition of great sculpin and Okhotsk sculpin for food was mitigated by the difference in the depths of their ranges, as well as by morphological (body size) and behavioral peculiarities in the areas where their habitats overlapped, and in microecosystems. The most probable competition was among Pacific cod 30–60 cm long and Okhotsk sculpin 20–50 cm in length, as well as among cod and great sculpins of all sizes.     

Multiple density dependence in two sub-populations of the amphipod Monoporeia affinis: a potential for alternative equilibria

Possible mechanisms for differences in population densities and dynamics were investigated in the amphipod Monoporeia affinis at two deep sites in the northern Bothnian Sea. The two sites were sampled yearly for 10 years. Average sizes, growth and mortality of the different age-classes were estimated from the cohort structure of the two populations. Laboratory experiments also investigated the ability of the common predatory isopod Saduria entomon to cause densitydependent (DD) mortality of the prey M. affinis. At site A, 43 m depth, the average density of M. affinis was twice as high as at site B, 81 m depth. The fluctuations in density were asynchronous between the two sites. Recruitment and subadult sizes of Monoporeia affinis were density dependent at both sites. The main functional difference between the two populations seemed to be the DD mortality of the 1 + cohort that occurred only at the low-density site B. A corresponding DD mortality was found in the predation experiments at densities of 1 + m. affinis corresponding to those found at site B. The potential importance of the predator was also indicated by a significant negative correlation between the biomass of S. entomon and the rate of change in M. affinis density in the field. The similarities in the abiotic factors between the two sites suggested that differences in carrying capacity should be small. The results could be explained by the predation regulation hypothesis for the low-density population at site B, while at site A M. affinis seemed to be regulated by intra-specific competition and limited by predation. It is suggested that in this simple predator-prey system there is potential for the existence of alternative equilibria.     

Why do juvenile fourhorn sculpin, Triglopsis quadricornis, appear in the pelagic habitat at night?

Synopsis Challenging the generally accepted view of the fourhorn sculpin Triglopsis quadricornis as a cold-stenothermal and true benthic species, novel findings of juveniles in the pelagic habitat of Lake Vättern, Sweden, have been examined and critically evaluated. Young-of-the-year and small-sized, sexually mature individuals, ranging 27–45 and 82–110 mm in total length, were recorded in nocturnal trawl catches from pelagic midwater in August. The sculpin had fed extensively on both benthic and semi-pelagic crustaceans, including large cyclopoid copepods, Mysis relicta, Pallasea quadrispinosa, Monoporeia affinis, Gammaracanthus lacustris, and Saduria entomon. Based on an extensive literature review a series of ecological and evolutionary reasons for a pelagic behavior of this profundal coldwater fish is suggested. Nocturnal, semi-pelagic feeding, and consecutive, thermotactic, vertical migrations up in midwater as shown by Wurtsbaugh & Neverman (1988) in order to maximize energy intake after feeding, are possible adaptations of these juvenile sculpin for minimizing size-dependent mortality due to intra- and interspecific predation and competition from other deepwater fishes.     

Geology and environmental significance of sediment distributions in an area of the submerged Niagara escarpment,Georgian Bay

The study site covers a spur of the submerged Niagara escarpment and part of the Flowerpot basin of Georgian Bay. Bedrock outcrops occur on islands and shoals but much of the area is covered by glacial till (10–20 m thick). Glaciolacustrine clay occurs in thin patches on the spur; in the deepest parts of the area, however, it is nearly 70 m thick. The distributions of Recent sediments which comprise lag-gravels, sands and silty clays, reflect major postglacial lake level changes in the Huron — Georgian Bay basin.Sediment geochemistry in Georgian Bay is similar to many other parts of the Great Lakes but calcium charbonate levels are lower than expected, except in some deposits of gaciolacustrine clay. There is little evidence of anthropogenic impact in the content of trace metals, although Pb likely includes a portion derived from atmospheric loading; Hg is usually present at very low levels but there are anomalously high values in some deepwater modern muds.In the bottom sediment habitat of modern benthos, the distribution of chironomids seems to reflect silt content. Oligochaetes are least in areas of thin Recent mud or exposed glaciolacustrine clay. Pontoporeia and sculpins are most numerous at depths between 50–110 m. Bottom tracks and other bed marks cover extensive areas of the deepwater lake sediments.     

Notes onDiphucephala affinis (Coleoptera:Scarabaeidae) associated with flowers ofHibbertia andAcacia in Western Australia

Notes are provided on the association of the beetleDiphucephala affinis (Scarabaeidae) with yellow flowers ofHibbertia hypericoides, H. huegelii (Dilleniaceae),Acacia pulchella, andA. stenoptera (Mimosaceae). Observations were undertaken during September 9–19, 1979 at S. Perth, Western Australia. They indicated thatD. affinis is not a pollinator ofHibbertia as suggested in the literature, but may play a small role in the pollination of someAcacia species.     

Dynamics of Fish Colonization of an Experimental Artificial Reef in Peter the Great Bay,Sea of Japan

An experimental artificial reef (AR) was built in Peter the Great Bay (Sea of Japan) to compensate for the biotope of Zostera destroyed by sea urchins. After eight years, the number of fish species on the AR increased from 5 to 18 and the fish biomass increased from 3.07 up to 37.1 g/m². Nonmigrating species (Opisthocentrus, young-of-the-year rockfishes, elegant sculpin) formed the bulk of the population; and migrant species (flounders, frog and great sculpins, sea raven) made up the greater portion of the biomass (up to 34 g/m²). Cage reefs are recommended to compensate for destroyed habitats, particularly Zostera beds.Original Russian Text Copyright ¢ 2005 by Biologiya Morya, Markevich.     

Spatial and temporal variation of benthic Cladocera (Crustacea) studied with activity traps in Lake Myvatn, Iceland

Benthic Cladocera were studied with a modified type of an activity trap in Lake Myvatn in 1990–1992. After feasibility experiments, the operation time and the distance of the traps from the benthic substrate were adjusted in order to minimize the effects of diurnal variation and the trapping of planktonic organisms. The trap catches of Eurycercus lamellatus were positively correlated with their abundance at the bottom as estimated by grab sampling (r = 0.910, P < 0.001). The usefulness of the activity trap was demonstrated by: (1) a lake-wide survey of the benthic Cladocera; (2) a study of the seasonal variation in the size distribution, abundance and sex ratio of E. lamellatus; and (3) a study of the seasonal succession of Chydorus sphaericus and Alona affinis. The variation of benthic Cladocera among 21 trap sites distributed on a 1–2-km scale across the lake exceeded the within-site variation. The sampling sites could be divided into five main groups based on cluster analysis. Eurycercus lamellatus was the most common species in the mat of filamentous green algae (Cladophorales). Alonella nana dominated the area of spring water inflow in the north basin and Macrothrix hirsuticornis the area of spring water inflow in the southeastern part of the lake. In other parts of the lake either Chydorus sphaericus or Alona quadrangularis tended to dominate. The size distribution and sex ratio of E. lamellatus was followed at two sites through one summer. In early summer most individuals were females less than 1.45 mm long. Around mid-summer they had grown to 0.69–3.1 mm. By the end of August the size distribution had become bimodal, with a large number of small males and a smaller number of much larger females. The seasonal succession in the abundance of E. lamellatus, A. affinis and C. sphaericus was followed at four sites over two seasons. With some exceptions the abundance of a species followed a similar seasonal trajectory on the different stations in any one year. There was, however, a marked difference between the two years (1991 and 1992), probably related to different temperatures.     

The Distribution of Nearshore Fishes in Kelp and Eelgrass Communities in Prince William Sound, Alaska: Associations with Vegetation and Physical Habitat Characteristics

The nearshore (less than 20m depth) demersal fish community in Prince William Sound, Alaska, is dominated by Pacific cod, Gadus macrocephalus, pricklebacks (mostly Arctic shanny Stichaeus punctatus), gunnels (mostly crescent gunnels Pholis laeta), a variety of greenlings (Hexagrammidae) and sculpins (Cottidae). During summer, the spatial distribution of fishes, over scales of 100's of m to 10's of km, varied by habitats characterized by different vegetation types. Juvenile Pacific cod and greenlings were numerically dominant in eelgrass, Zostera marina, beds. Pricklebacks and sculpins were dominant in areas with an understory of the kelps Agarum cribrosum and Laminaria saccharina. Greenlings and sculpins were the most abundant demersal fishes in more exposed sites with a canopy of Nereocystis luetkeana and an understory of L. bongardiana. Measured habitat variables, including vegetation type, slope, vegetation biomass, and substratum type, explained a significant proportion of the variation in the presence or absence of most fishes. The relative importance of different habitat characteristics varied between taxonomic groups of fishes. Vegetation type explained a significant proportion of variation for cod, rockfishes, and ronquils. Juvenile cod were closely associated with eelgrass, while rockfish and ronquils were associated with kelps. Pricklebacks and rockfishes were more frequently observed on steeply sloped shorelines, while ronquils were more often found at sites with higher biomass of vegetation. Within A. cribrosum habitats, more greenlings and sculpins were present at sites where algal biomass was higher. Also, sculpins were more abundant in deeper water and gunnels were more abundant in shallow water within this habitat. These associations may not have been causative. However, evidence suggests that some differences between fish communities in eelgrass and Agarum beds may have been causally related to vegetation characteristics. The possible roles of different vegetation types as refugia from predators or as sources of prey are discussed.