Predator-prey relations between age-1+ summer flounder (Paralichthys dentatus, Linnaeus) and age-0 winter flounder (Pseudopleuronectes americanus, Walbaum): predator diets, prey selection, and effects of sediments and macrophytes

Laboratory experiments and weekly trammel net surveys in the Navesink River, New Jersey (USA) were used to examine the predator-prey interaction between age-1+ summer flounder (Paralichthys dentatus) and age-0 winter flounder (Pseudopleuronectes americanus). Winter flounder (24-67 mm TL) were the dominant piscine prey of summer flounder (n=95, 252-648 mm TL) collected in trammel nets. We observed a temporal shift in summer flounder diets from sand shrimp (Crangon septemspinosa) and winter flounder, dominant during June and early July, to blue crabs (Callinectes sapidus) and other fishes (primarily Atlantic silversides, Menidia menidia and Atlantic menhaden, Brevortia tyrannus) later in the summer. Variations in prey selection appeared to be related to changes in the spatial distribution of predators and spatio-temporal variation in prey availability. In laboratory experiments, summer flounder (271-345 mm total length, TL) preferred demersal winter flounder to a pelagic fish (Atlantic silversides) and a benthic invertebrate (sand shrimp) prey, and the vulnerability of winter flounder increased with increasing prey body size from 20 to 90 mm TL. Experiments testing habitat effects showed that mortality of winter flounder in three different size classes (20-29, 40-49, 60-69 mm TL) was not influenced by sediment grain sizes permitting differential burial of the prey. However, vegetation enhanced survival, with fish suffering lower mortality in eelgrass (Zostera marina, 15+/-0.04%) than in sea lettuce (Ulva lactuca, 38+/-0.04%) or bare sand (70+/-0.07%) when the macrophytes were planted to produce similar leaf surface areas (5000 cm(2) m(-2)). Prey vulnerability appeared to be related to the role of vision in the predator's attack strategy and prey activity levels.     

Shallow water predation risk for a juvenile flatfish (winter flounder; Pseudopleuronectes americanus, Walbaum) in a northwest Atlantic estuary

Many small fish, including several juvenile Atlantic flatfish, are most abundant in shallow areas presumable because these habitats enhance survivorship and/or growth. In this study, we investigated size-dependent depth distributions and the role of shallow habitats as predator refuges for age-0 winter flounder (Pseudopleuronectes americanus) in a northwest Atlantic estuarine nursery. Analysis of trawl surveys performed during the larval settlement period throughout the Navesink River and Sandy Hook Bay, New Jersey, showed that as fish increased in size, depth of occurrence gradually decreased, so that individuals >35 mm standard length (SL) were concentrated in habitats ∼1 m deep. Tethering in structurally simple and adjacent shallow and deep habitats showed that predation risk for flounder (30-50 mm SL) was low in shallow water (<1 m) and increased rapidly with depth. Summer flounder (Paralychthys dentatus), which were more abundant in trammel nets in deep habitats and included winter flounder in their diets, appeared to be important consumers of tethered fish. Our results indicate that following larval settlement, winter flounder emigrate from or suffer high mortality in deeper water to become concentrated in shallow habitats that can serve as predator refuges even when they lack complex physical structures. These results highlight the potential for functional habitat loss when natural and/or anthropogenic factors make shallow habitats unavailable to young fish.     

Sediment preferences and size-specific distribution of young-of-the-year Pacific halibut in an Alaska nursery

A combination of laboratory experiments and field surveys was used to test the hypotheses that responses to sediments change with fish size and that sediment grain-size is the predominant environmental factor affecting small-scale distribution in young-of-the-year (yoy) Pacific halibut Hippoglossus stenolepis . Laboratory tests showed that the smallest fish (31–40 mm L _T) chose fine sediments (muddy and fine sands), fish 51–70 mm had high selectivity (primarily medium sand), and the largest fish (80–150 mm) were not selective although they avoided the largest grain-sizes (pebbles and granules). Sediment preferences were correlated with size-dependent burial capabilities. Beam trawl collections were made over a 6 year period in Kachemak Bay, Alaska, to examine the distribution of yoy Pacific halibut (14–120 mm L _T) using small size classes (e.g. 10 mm intervals). Canonical correlation analysis showed that the per cent of sand in the sediment was a highly significant variable for all but one size and date combination. Catch per unit of effort (CPUE) for newly settled fish (<30 mm L _T) was highest on very fine sand, fish 41–80 mm were most abundant on fine sand, and the largest yoy fish (81–120 mm) were abundant over a range of sediments from fine sand to mud. Except for the smallest fish, Pacific halibut in the field were associated with sediments somewhat finer than predicted from the laboratory experiments; however, virtually all were captured where they could bury easily. The ability of flatfish to bury and shelter in sediment is related to fish size; consequently, habitat associations shift rapidly during the first year of life. Habitat models for yoy flatfishes should consider size-dependent shifts in capabilities and preferences.     

Ontogenetic dietary shifts and morphological correlates in striped mullet,Mugil cephalus

Synopsis Ontogenetic dietary shifts are well studied in fishes. However, these shifts are rarely correlated with habitat and morphology. This study investigates: (1) ontogenetic dietary changes in mullet; and if dietary shifts do occur (2) whether they are correlated with feeding location, feeding behavior and/or feeding morphology. Stomach contents from eight size classes, each with 25 individuals, were analyzed using a gravimetric sieve fractionation procedure, Czechanowski's Quantitative Index and cluster analysis. Core samples (5 mm deep) were also analyzed in the same manner. Morphological characters were measured to detect differences among the size classes. Cluster analysis showed three distinct trophic units for Mugil cephalus in Cross Bayou: (1) 20.0 mm standard length — 30.0 mm SL; (2) 30.0 mm SL — 40.0 mm SL; and (3) 40.0 mm SL — > 100 mm SL. Comparison of the gut contents to the sediment contents for each size class showed an increase in similarity as the fish grew. Mullet gut contents from fish 20–25 mm SL were only 45% similar based on the weights of the prey items to the sediment subsample prey items, while stomach contents from fish > 100 mm SL were 84% similar to the sediment subsample contents. Gill raker length and interraker spacing of four gill arches were allometric with respect to standard length. Prey particle size within size classes appeared to be more correlated with feeding behavior than gill interraker spacing because smaller particles than predicted by the interraker spacing were ingested. It is suggested that smaller fish may be browsing more selectively than larger fish which graze.     

Feeding behavior of newly settled winter flounder (Pseudopleuronectes americanus) on calanoid copepods

Field and laboratory investigations were conducted to examine feeding by newly settled winter flounder (Pseudopleuronectes americanus) on two co-occurring calanoid copepods, Eurytemora affinis and Acartia hudsonica. During the spring, these prey are present when winter flounder initiate their demersal lifestyle in estuaries of the northeastern United States. Epibenthic zooplankton were collected concurrently with winter flounder in the Navesink River estuary, NJ, in May 1998 and 1999. Although both calanoid species were in the estuary during the 2-year survey, E. affinis was consumed nearly to the exclusion of A. hudsonica by newly settled winter flounder. Annually, E. affinis and A. hudsonica had similar size distributions in field collections, indicating that species choice was not size selective. However, when preying on E. affinis, winter flounder preferred the larger sized organisms. In single species laboratory experiments, E. affinis and A. hudsonica were consumed equally by newly settled winter flounder (19-23 mm TL), but there were more strikes made toward E. affinis. Despite the lower catch efficiency, E. affinis was selected over A. hudsonica when the prey species were offered together in equal numbers. The selection for E. affinis over A. hudsonica by newly settled winter flounder may be the result of behavioral and/or morphological differences in the prey species.     

Predation by xanthid crabs on early post-settlement gastropods: the role of prey size, prey density, and habitat complexity

Small predators in marine benthic communities create a hazardous environment for newly settled invertebrates, especially for the smallest individuals. To explore the effects of predation on a newly settled gastropod, queen conch (Strombus gigas Linnaeus), by a xanthid crab (Micropanope sp.), prey size, prey density, and habitat complexity were manipulated in five laboratory experiments. All crabs >3.1 mm CW killed all conch <2 mm SL when individual crabs (<14 mm carapace width (CW)) were offered individual conch that were 2–35 days old after metamorphosis (1.2–8.8 mm shell length (SL)). Only 10% of the crabs >5.0 mm CW, however, killed conch that were >5.0 mm SL, suggesting that conch may reach a size refuge from xanthid crabs at 5 mm SL. Furthermore, when given a choice, crabs (4.8 mm CW) preferred smaller conch (2.0 mm SL) to larger (3.7 mm SL), suggesting that 1 week of additional growth in shell length is advantageous to survivorship. Proportional mortality decreased as conch density increased when crabs were offered conch at seven different densities (two to 96 individuals). Crabs proved to be effective predators regardless of the amount of seagrass structure provided in a microcosm experiment, and could consume two conch in 10 s. The high densities of xanthid crabs that occur in the wild, their effectiveness as predators, and their large appetites point to the important role that small predators may potentially play in structuring the population dynamics of their small prey immediately after settlement.     

Relationship between body size and habitat complexity preference in age-0 and -1 year winter flounder Pseudopleuronectes americanus

The interaction between body size, habitat complexity and interstice width on habitat preference of age-0 and -1 year Pseudopleuronectes americanus was examined using continuous remote video observation. The habitat choices of juvenile P. americanus were recorded over a 6 h period in tanks with four treatments: bare sand, sand with low complexity cobble, sand with intermediate complexity cobble and sand with high complexity cobble. Both age-0 and -1 year fish preferred cobble to bare sand. Within cobble treatments, age-0 year fish preferred intermediate complexity cobble, with a 1.59 ratio of interstitial space to body width. The largest age-1 year fish (123-130 mm standard length, L(S) ) preferred low complexity cobble. While a significant preference was not detected, medium age-1 year fish (83-88 mm L(S) ) tended to select low complexity cobble, whereas small age-1 year fish (73-82 mm L(S) ) tended to select low and intermediate cobble, with an interstitial space to body width ratio of 1.05. For medium and large age-1 year fish, there was an increased selection of low complexity cobble, corresponding to larger interstitial space to body size ratios. This study indicates that juvenile P. americanus prefer complex habitat to unstructured habitat and that this preference is mediated by a relationship between fish body size and the size of structure interstices. These results contribute to the growing body of knowledge of complex habitat selection and drivers of habitat choice in flatfishes.     

The effects of prey size, predator size, and sediment composition on the rate of predation of the blue crab, Callinectes Sapidus Rathbun, on the hard clam, MercenariaMercenaria (Linné)

Sediment preferences of blue crabs, Callinectes sapidus Rathbun, and predation rates on various size classes of the hard clam, Mercenaria mercenaria (Linné), in a variety of sediment types were studied in the laboratory. Blue crabs of all size classes exhibited a preference for sand, mud, and sand/mud rather than crushed oyster shell or granite gravel. Clams were more vulnerable to predation by crabs in sand and sand/mud than in crushed oyster shell or granite gravel. When crabs were given a choice of clam sizes based on carapace width (CW), small crabs (<75 mm CW) consumed 5- and 10-mm shell length (SL) clams. Medium crabs (75–125 mm CW) preferentially consumed 10-mm SL clams. Large crabs (> 125 mm CW) consumed 10- and 25-mm SL clams equally. Blue crabs did not eat clams that were >40-mm SL.     

Substratum selection by juvenile flounder Platichthys flesus (L.): effect of ephemeral filamentous macroalgae

Selection of substrata representing habitats with or without filamentous vegetation by juvenile flounder Platichthys flesus was investigated in laboratory experiments and a field study to assess the effect of the increasing occurrence of ephemeral filamentous macroalgae in shallow nursery areas and its potential effect on flounder recruitment. In the laboratory experiments, 79% of juvenile flounder (24–99 mm total length, L _T) preferred bare sand substrata in comparison to substrata with simulated filamentous vegetation. Substratum preference did not change with fish size. Field sampling using a drop trap (1 m²) in a vegetated (>50% coverage of ephemeral macroalgae) habitat in June, July and August showed small flounder (mean L _T= 37·3 mm) were caught more often in samples with less vegetation. In contrast, larger flounder (mean L _T= 57·2 and 79·7 mm in July and August, respectively) were more evenly distributed in relation to the distribution of vegetation. The results of this study indicate that the increase in ephemeral macroalgae observed in recent years potentially reduces the quantity and quality of suitable nursery habitats for juvenile flounder, which could have detrimental consequences to the overall recruitment of flounder in coastal areas.     

Factors affecting the post‐release survival of cultured juvenile Pseudopleuronectes americanus

Laboratory experiments were performed with cultured and wild juvenile winter flounder Pseudopleuronectes americanus to evaluate differences in behaviour and adaptation affecting post‐release vulnerability to predation. Studies revealed that the cryptic abilities of cultured winter flounder increased over time. Sediment‐naïve, cultured fish required a minimum of 2 days to improve their burying skills and at least 90 days for colour adaptation to match the sediment. Cultured winter flounder selected sediments consisting of small grains and colours matching their own pigment. Cultured winter flounder, regardless of their colour, were significantly more vulnerable to predation by birds. Additionally, cultured fish reacted differently than wild winter flounder when exposed to cues from a potential predator.     

Somatic growth and otolith growth in juveniles of a small subtropical flatfish, the fringed flounder, Etropus crossotus

A growth experiment was conducted with juvenile fringed flounder (Etropus crossotus) from North Inlet (South Carolina, USA) to provide information on the growth of a small, shortlived flatfish with a subtropical and tropical distribution. The fringed flounder has a maximum life span of 1.5 y and its long spawning period from March through October complicates the determination of growth rates based on length frequency data. Otoliths of juveniles with a standard length (SL) 23.1-53.0 mm were marked with Alizarin complexone and the fish were held in the laboratory for 66 days at 14, 20, 24 and 29 degrees C while being fed ad libitum. The mean somatic growth increased with temperature from 0.1 mm SL day(-1) at 14 degrees C to 0.4 mm SL day(-1) at both 24 and 29 degrees C. The maximum observed somatic growth rate was 0.7 mm SL day(1) at 29 degrees C. The number of micro-increments formed in otoliths was not significantly different from the expected value, validating formation of one increment per day. The significant relationship between increment width and somatic growth rate can be used to estimate somatic growth rates of individual wild fish based on daily increment information in their otoliths.     

Estuarine and habitat-related differences in growth rates of young-of-the-year winter flounder (Pseudopleuronectes americanus) and tautog (Tautoga onitis) in three northeastern US estuaries

Instantaneous growth rates of young-of-the-year winter flounder Pseudopleuronectes americanus (Walbaum) (12.0-60.4 mm standard length, SL) and tautog Tautoga onitis (Linnaeus) (21.4-73.8 mm total length, TL) from three estuarine systems in New Jersey (Great Bay-Little Egg Harbor and Navesink River) and Connecticut (Hammonasset River) were used in an attempt to assess the relative quality of selected nominal habitats. A series of short-term field caging experiments were conducted during 1994 and 1995 in: macroalgae (primarily, Ulva lactuca), eelgrass (Zostera marina), unvegetated areas adjacent to macroalgae and eelgrass and tidal creeks in Spartina dominated marsh. Growth rates varied with habitat, estuary and year. Comparisons across nominal habitats within and among estuaries did not show any one habitat with consistently higher growth, and growth was relatively independent of whether a habitat was vegetated or adjacent to vegetation. The growth rates of winter flounder and tautog from the Hammonasset River were not different among habitats in either year of the study. In the Great Bay-Little Egg Harbor, both winter flounder and tautog had higher growth rates in macroalgae with growth in eelgrass varying significantly between years. Conversely, in the Navesink River both species had higher growth rates in eelgrass. Environmental changes associated with temperature and dissolved oxygen appeared to influence growth rates. Winter flounder growth rate and survival was depressed in tidal marsh creeks in the three estuaries and in vegetated macroalgae habitats in the Navesink River where dissolved oxygen levels were often very low (<2 mgl(-1)) for extended periods. In summary, the growth rates of the young-of-the-year of these two species varied temporally and were dependent on the interaction of both the specific estuary and habitat in which the experiments took place. Further, habitat quality, as defined by relative growth rate, was difficult to evaluate because it can be variable and nominal habitat designations are often not sufficient to define the boundaries of a species habitat requirements.     

‘Prudent habitat choice’: a novel mechanism of size‐assortative mating

Assortative mating, an ubiquitous form of nonrandom mating, strongly impacts Darwinian fitness and can drive biological diversification. Despite its ecological and evolutionary importance, the behavioural processes underlying assortative mating are often unknown, and in particular, mechanisms not involving mate choice have been largely ignored so far. Here, we propose that assortative mating can arise from ‘prudent habitat choice’, a general mechanism that acts under natural selection, and that it can occur despite a complete mixing of phenotypes. We show that in the cichlid Eretmodus cyanostictus size‐assortative mating ensues, because individuals of weaker competitive ability ignore high‐quality but strongly competed habitat patches. Previous studies showed that in E. cyanostictus, size‐based mate preferences are absent. By field and laboratory experiments, here we showed that (i) habitat quality and body size are correlated in this species; (ii) territories with more stone cover are preferred by both sexes in the absence of competition; and (iii) smaller fish prudently occupy vacant territories of worse quality than do larger fish. Prudent habitat choice is likely to be a widespread mechanism of assortative mating, as both preferences for and dominance‐based access to high‐quality habitats are generic phenomena in animals.     

The role of fine sediment characteristics and body size on the vertical movement of a freshwater amphipod

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The relationship between body size, sediment grain size and the burying ability of juvenile plaice, Pleuronectes platessa L.

The ability of juvenile plaice (18–190 mm) to bury in sands varying in grain size from 0.062–2 mm was examined. For fish greater than approximately 30 mm in length the relationship between the proportion of the body covered with sand ( C ), total length ( L , mm) and grain size ( S , mm) can be estimated from the equation: logit C = 3.250 + 0.069 L – 6.771 S . Based on their performance in coarser sediments, fish smaller than approximately 30 mm did not bury as well as expected in the finest sediments (0.062 and 0.125 mm).     

Sediment selection in juvenile plaice and its behavioural basis

In the laboratory juvenile plaice Pleuronectes platessa (13–114 mm) were consistently found on the finest of four sediments after 24 h both in the light and the dark. Preference when assessed both by the fish's final position and by the amount of time it spent on each sediment agreed on 72% of occasions. Size did not affect preference, which was determined principally by a fish's ability to bury in a sediment. Burial reduces activity and thereby increases the time that fish spend on sediments in which they can bury. Activity level, which is dependent on light intensity and endogenous factors, therefore determines the degree of selectivity shown. Swimming duration and resting by the largest fish (71–114 mm) did not differ among sediments. Field experiments produced results similar to those obtained in the laboratory.     

Size-related mortality due to gnathiid isopod micropredation correlates with settlement size in coral reef fishes

The transition between the planktonic and the benthic habitat is a critical period for the larvae of many demersal marine organisms. Understanding the potential constraints on the timing of this habitat transition, called settlement, is important to understanding their biology. Size-specific mortality can set the limits on lifestyle and help explain ontogenetic habitat shifts. We examined whether size-based mortality risks after settlement may include micropredation by ectoparasites by testing whether survival of settlement-stage fish varies with fish size when exposed to a reef-associated micropredator. Fish (14 species) were exposed to one blood-sucking gnathiid isopod overnight, with appropriate controls; gnathiid feeding success and survival, and fish mortality were recorded relative to fish size. After adjusting for fish relatedness, we found the relationship between fish mortality and size differed with gnathiid exposure: for gnathiid-exposed fish, the mean mortality of the smallest fish was much higher (57%) than unexposed controls (10%), and decreased to ~0% for fish >12 mm standard length (SL); mortality was almost nil in controls. Thus, a predicted optimal size to switch habitat and reduce mortality risk from micropredation should be >12 mm SL. We then asked what species might be at greater risk and if the steep increase in survival at ~12 mm SL might coincide with settlement at larger sizes among fishes. Across 102 other species (32 families), 61% settled at ≥12 mm SL. After adjusting for relatedness, mean fish settlement size was 15.0 mm and this was not significantly different from 12 mm. Thus, settlement size clusters around the minimum fish size threshold our gnathiid experiment predicted would be large enough to survive a gnathiid encounter. These results suggest micropredators may contribute to size-selective mortality during settlement processes and are consistent with the hypothesis that the pelagic phase provides fish an escape from certain micropredators.     

Size selective feeding and its limitations for the black rockfish, Sebastes inermis, in a demersal fish assemblage of Onagawa Bay, northeastern Japan

The size selective feeding of the black rockfish, Sebastes inermis, in a demersal fish assemblage of Onagawa Bay, located in northeastern Japan, was studied using laboratory experiments and stomach content analysis of the dominant demersal fish coexisting with the black rockfish in Onagawa Bay. The preferred food size of the black rockfish was determined in the laboratory experiments. The average food size taken by the black rockfish (150–227 mm long) under natural conditions was much smaller than that taken by fish in laboratory experiments (148–158 mm sized). The most preferred size of food taken by fish in laboratory experiments was rarely found under natural conditions. Analysis of the competition coefficients of fish species in the fish assemblage based on stomach content analysis suggested that the degrees to which the preferred food size was found depended upon the foods availability, by competition with other fish species and by the feeding ability of the fish. Intra- and inter-specific competition in a fish assemblage might be limiting factors on the food size selection of fish under natural conditions.     

Intraspecific interactions influence size specific depth distribution in Cottus bairdi

Synopsis We tested depth selection by different sizes of mottled sculpin, Cottus bairdi, in a southern Appalachian stream. Field observations indicated that, during one hour periods, both small (<50 mm SL) and large (≥55 mm SL) individuals move within an area less than 0.50 m². Individuals of both sizes, placed in field enclosures, preferred deep microhabitat. When large fish were placed in cages with small fish, small fish initially spent more time in slope and shallow microhabitats. Average interfish distances were not correlated with their absolute size differences, suggesting C. bairdi interactions may involve both predation and competition. In streams, size-related differences in microhabitat depth may result more from intraspecific interactions than from size-specific depth preferences.