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1.
Individual fish commonly leave the relative safety of the shoal to approach potential predators at a distance. Not all members of a shoal are equally likely to initiate such predator inspection visits. Here, we show for the first time that the current hunger state of individual fish strongly influences their predator inspection behaviour, as well as their foraging rate, in the face of predation hazard. When all members of threespine stickleback (Gasterosteus aculeatus) test shoals were in a similar hunger state, they were equally likely to inspect a trout predator model alone and did not differ in the frequency of their inspection visits or foraging rate. However, when individual sticklebacks in a shoal differed in their hunger state, the food-deprived (i.e. hungrier) member of the shoal fed at a higher rate, was significantly more likely to initiate solitary predator inspection visits, and inspected the predator model significantly more often than its less hungry (i.e. well-fed) shoal mates. Individual fish which inspected the predator model more frequently also tended to have higher feeding rates. The results indicate that the hungrier fish in a shoal are more willing to take greater risks to inspect a potential threat at a distance, compared with their well-fed shoal mates, and suggest that they may gain a foraging benefit in doing so. If marked asymmetries in hunger state exist among members of fish shoals, then mutual cooperation during predator inspection visits may be difficult to achieve because well-fed individuals are not as likely to initiate or participate in inspection visits as are hungry individuals.Correspondence to: J.-G.J. Godin  相似文献   

2.
The metabolic enzyme activities were determined in larvae of red drum, Sciaenops ocellatus, and lane snapper, Lutjanus synagris, to determine the effect of temperature and nutrition on metabolic enzyme activities and to evaluate if metabolic enzyme activities are useful in assessing the feeding condition of larval fish. During experiments conducted during the spring of 1990, lactate dehydrogenase (LDH) activities in both red drum and lane snapper were approximately an order of magnitude lower than values typical for adult fish; LDH and citrate synthase (CS) activities increased during early developmental stages, but nutritional effects were apparent. Clear differences (up to 4-fold) between well-fed and starving fish were evident in both LDH and CS activity in red drum. Differences between well-fed and poorly fed larvae were evident until 9 d after hatching. Lane snapper larvae reared at a 25°C had significantly lower LDH activities than larvae reared at 28°C.  相似文献   

3.
Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.  相似文献   

4.
Summary Thermal preferences of well-fed and food-limited fire ant colonies (Solenopsis invicta) were studied in relation to colony growth and metabolic costs. The growth curve for well-fed colonies was strongly skewed toward warmer temperatures with maximal growth occurring near 32° C (Fig. 2A). The growth curve for food-limited colonies was skewed toward cooler temperatures with maximal colony size occurring around 25° C (Fig. 2B). Food-limited colonies apparently grew larger at cooler temperatures because metabolic costs of workers were reduced. A series of binary choice tests confirmed three predictions concerning fire ant thermal preferences (Figs. 3–4). First, well-fed colonies preferred brood temperatures very near the optimum for colony growth (31° C versus 32° C). Colonies were also able to select appropriate suboptimal growth temperatures when the optimal range was unavailable. Secondly, as predicted, a large percentage of colony workers ( 30% in well-fed colonies) consistently chose cooler temperatures than those selected for the brood. This strategy probably increases longevity of workers not directly associated with brood care. Thirdly, food-limited colonies preferred cooler temperatures than well-fed colonies. Metabolic costs of food-limited colonies were reduced by approximately 7% because of (1) slightly cooler brood temperatures (30° C versus 31° C) and because (2) an additional 20–30% of the workers selected cooler temperatures. The addition of excess food reversed food-limited thermal preferences within 12 h for the brood (Fig. 5) and several days for the workers. Contrary to expectations, thermal preferences for brood in food-limited colonies did not match the food-limited growth curve, perhaps because fire ant colonies can choose to rear brood at warm temperatures while maintaining accumulated colony biomass at cooler temperatures. Correspondence to: S.D. Porter  相似文献   

5.
Fish school density and volume   总被引:3,自引:0,他引:3  
All the fish in a school occupy a volume estimated as N·BL3, where N is the number of fish and BL is their mean body length. We present extensive data from our experiments on cruising schools of saithe (Pollachius virens), herring (Clupea harengus) and cod (Gadus morhua) to validate this formula. Two methods of calculating the volumes of schools are described. One method is aggregative and depends on measuring the envelope of free space around a schooling fish, whereas the other is based on the dimensions of the school as a whole. The whole-school method is more reliable since it includes lacunae between the sub-units which exist in schools. For this method, we derive a computation which eliminates bias from outliers. The most realistic theoretical aggregative packing model predicts a volume per fish of 0.6 BL3. In saithe, the envelope of free space is approximately an ellipsoid, which, although it becomes more compressed at higher swimming speeds, yields a volume close to 0.7 BL3. From the whole-school method we calculate average volumes of 1.4 BL3 for saithe and 0.7 BL3 for herring. Increase in swimming speed produces more compact schools in saithe, but changes in arousal level can generate equally large differences. Changes in volume were not adequately explained by changes in nearest neighbour distance, giving support to the whole-school method.  相似文献   

6.
Many species base their choice of mates on multiple signals which provide them with different kinds of information. Choosers may assess the signals together to evaluate the overall quality of potential mates, but individuals often pay attention to different signals in different contexts. In Rhinogobius brunneus, a fish displaying exclusive male parental care, females generally prefer males showing larger first dorsal fins (FDF) and more active courtship displays as mates. Females choosing a mate usually initially assess the FDF and later utilize courtship for the final decision. In our experiments, females with different hunger states used different signals when selecting mates. Females in both hunger states preferred males with larger FDF in the first stage. In the second stage, well-fed females showed highly repeatable choice, whereas poorly fed females responded only to variation in the courtship activity of males. The males preferred by poorly fed females exhibited significantly higher offspring survival than nonpreferred males. Under conditions of food shortage, males allocate more energy to future reproduction at the expense of the present brood, and females may prioritize signals predictive of offspring survivorship over signals reflecting other aspects in male quality to minimize the losses in direct benefits. We conclude that R. brunneus females may employ information from both signals but dynamically adjust their prioritization of each signal to current conditions to ensure the choice that is currently most adaptive.  相似文献   

7.
We studied mate attraction by females of the praying mantid, Tenodera aridifolia sinensis, testing honest signaling of mate availability versus deceptive signaling to attract males for sexual cannibalism. We experimentally varied female diet and mating history and measured the rate of attraction of a wild population of males to caged females. Honest signaling theory predicts that virgin females will attract males at the greatest rate whereas deceptive signaling predicts that hungry females (which are more likely to cannibalize males) will attract more males, particularly among non-virgin females. Our results show that hungry females did not attract more males than well-fed females. Indeed, the opposite was true: hungry females attracted significantly fewer males. Moreover, hungry females were no more likely than well-fed females to attract males subsequent to mating, and mated females attracted males at a lower rate than did virgin females. We also observed female T. aridifolia sinensis and male Mantis religiosa arriving at the caged females and we discuss the significance of these observations. The results refute the hypothesis of deceptive signaling and show that mate attraction signals of female T. aridifolia sinensis are honest indicators of female mate availability and a lower risk of sexual cannibalism.  相似文献   

8.
Gut fullness of larval fishes was used to determine the nutritional significance of an estuarine plume front off Botany Bay, SE Australia, on three days in March/April 1990. Fishes were captured in three different water masses (estuarine plume, front and shelf water), each separated by 200 m, using a 260 m mesh purse-seine net. Overall, the gut-fullness index (GFI) of 260 fish larvae combined from eight families (Gerreidae, Mugilidae, Mullidae, Sparidae, Blenniidae, Kyphosidae, Monodactylidae, Pomacentridae), was significantly greater in the plume and front water on two of the sampling occasions, but no difference was detected on the third occasion. Trends in GFI among families were inconsistent with respect to the front. The mugilid (Liza argentea) fed equally and abundantly in all water masses (gut fullness>90%), while the kyphosid (Kyphosus spp.) had a significantly greater GFI in the plume compared to the shelf water. In general, the response of GFI to the front varied between dates and amongst taxa. Diet analysis showed that mugilids selected for copepod nauplii within the plume and shelf water [alpha selectivity index (ASI) of (0.22 to 0.98)] and for harpacticoid copepods within the front (ASI=0.92), representing characteristic diets identified by canonical discriminant analysis. Kyphosids selected positively for copepod nauplii in all three water masses on both occasions (ASI>0.96). In general, diets of larval fish were taxon-specific, and responded variably to the three habitats. Comparison of diets across water masses was complicated by the large number of families, which rarely occurred in all water masses. The nutritional significance of an estuarine plume front varies between species of larval fish, and there are no obvious trends that can be applied to the larval fish community in general.  相似文献   

9.
A feeding experiment was conducted on the pelagic stages of the tropical goatfish Upeneus tragula (family Mullidae) to examine how food quantity affects growth characteristics and potential success at settlement. Pelagic goatfish were collected from aggregation rafts 3 nautical miles west of Lizard Island on the northern Great Barrier Reef during December 1990. Three tanks in each of four feeding regimes were stocked with 25 pelagic goatfish between 20 and 23 mm standard length (SL). The four feeding regimes were: fed ad lib. (fed): fed once per day (f1pd); starved every second day (stld); starved for 3 d and re-fed (st3d). Fishes were fed 36 to 48 h old Artemia sp. nauplii (Ocean Star strain). Fish were removed from tanks when they underwent metamorphosis, changed pigmentation and settled to the bottom of the tanks. Morphology, muscle development, time taken to settle and biochemical condition were examined. Growth attributes of the treated fish were compared to fish which settled within 24 h of capture (field). All attributes examined were significantly influenced by the feeding treatments. Fish within the fed and st3d treatments were significantly larger and heavier than fish in the less well-fed treatments (f1pd, st1d). Similarly, concentrations of total lipid, carbohydrate and protein in the settled fish were significantly higher in the fed and st3d treatments compared with the st1d and f1pd treatments. For all these morphological and biochemical attributes the st1d and f1pd fish did not significantly differ from the field fishes, but did differ from the fed and st3d fishes. Water content was significantly higher in the f1pd, st1d and field fishes compared with the fed and st3d fish. Furthermore, the average time taken to settle followed the pattern: fed (14d)U. tragula are physiologically well suited to exploiting a patchy food source, and that food availability within the pelagic stages can have a major influence on the growth characteristics of this reef fish at settlement. The ramifications of this finding are discussed in relation to survival and success once fish have recruited to the reef population.  相似文献   

10.
Summary Solitary and schooling banded killifish (Fundulus diaphanus Lesueur) were attacked with a fish predator model in the littoral zone of a lake and their flight reaction distance to the model recorded. Reaction distance of solitary fish did not differ significantly from that of schooling fish, which was statistically constant over a wide range of observed school sizes. The results fo not support the proposed early predator warning function of social groups, which predicts an asymptotically increasing flight reaction distance with increasing group size. However, fish in larger and safer groups may detect an approaching predator sooner but delay their flight response, thereby permitting their individual members to spend more time assessing the nature of the threat before escaping. Such a strategy would reduce costly false alarms. Relative variation in recorded reaction distances was correlated negatively with school size, suggesting that schooling reduces variability in latency time to escape predators. This may be evidence for the above behavioural strategy. Further, the speed of transmission of the flight response within the school greatly exceeded the speed of approach of the predator model. This phenomenon enables schooling individuals to initiate avoidance behaviour before the approaching predator can be detected and before it reaches them.  相似文献   

11.
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)  相似文献   

12.
Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1-1 (about 250 g C 1-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h-1.  相似文献   

13.
The behaviour of juvenile herring (Clupea harengus L.) feeding on Artemia sp. nauplii in both light and dark was recorded using an infra-red-sensitive television-recording technique. In the light, two modes of feeding were observed, particle biting and filtering, but in the dark only filtering was observed. Marked differences in swimming behaviour were seen between light and dark. In the light, the fish continued to school while feeding in both modes; in the dark, the school dispersed, the fish swam slower in tight circular paths and fed only by filtering. In the dark, filtering fish swam faster (0.11 m s-1) than non-filtering fish (0.07 m s-1). In the light, no difference in speed was measured between filtering and non-filtering fish (0.34 m s-1). Owing to the lower filtering speeds in the dark, the removal rate of nauplii from the water was much lower than in light, except in the highest prey concentrations. This suggests that if night-time filter-feeding takes place in the sea, it will be of importance only when exploiting dense patches of food.  相似文献   

14.
Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.  相似文献   

15.
An infra-red sensitive video-recording technique was used to study the effects of darkness and light intensities from 0.0001 to 270 photopic lx on the feeding behaviour of herring (Clupea harengus L.). When offered natural zooplankton, consisting of a mixture ofCalanus finmarchicus, Euchaeta norvegica, Oithona similis, Balanus sp. nauplii, and crustacean nauplii as prey, the fish fed by biting (snapping) at light intensities above a threshold of 0.001 lx and were size-selective, taking the larger organisms first. When fed on pure cultures of CaliforniaArtemia sp. nauplii (San Francisco Bay brand), the threshold light intensity was 0.01 lx. Swimming speed increased with increasing light intensity when the fish were actively feeding by biting. When the fish were filter-feeding on high densities ofArtemia sp. nauplii in the light, they continued to school and swimming speed was not related to light intensity.  相似文献   

16.
We examined the behavioral and developmental responses of Xenopus laevis larvae to their mirror images in three experiments. The mirrors allowed us to visually simulate increased density, without the tadpoles’ behavior being confounded by chemical cues from additional tadpoles. In the first experiment, we demonstrated that Xenopus tadpoles have a right eye preference for mirrors, contrary to the left eye preference of all other anuran species studied to date. This lateralized eye use disappeared, however, as tadpoles approached metamorphosis. Next, we examined how mirrored aquaria walls affected tadpole growth and development. We found that tadpoles raised in aquaria with partially mirrored walls showed depressed growth compared to tadpoles raised without mirrors, despite the fact that Xenopus larvae normally thrive when raised in visual contact with conspecifics. The tadpoles raised with mirrors had, though not significantly, proportionally larger bodies relative to their tail length (d = 0.51). This suggests that a phenotypically plastic response in body proportions was induced in these tadpoles solely by the sight of other tadpoles. The third experiment established that X. laevis tadpoles are more active in front of a mirror; i.e., they turn more often and spend more time in front of mirrored surfaces. We consider this increased activity to be an aberrant behavior of the tadpoles, which were attempting to school with their own images. We suggest that this extra activity reduced the amount of energy available for growth, accounting for the depressed growth seen in our second experiment.  相似文献   

17.
Pyjama sharks (Poroderma africanum) were exposed to a wide range of salinities, over which blood serum was analysed for osmolarity, chloride and urea concentrations. Fish were divided into two groups, those fed twice weekly (high intake), and those fed once a month (low intake). Both groups were exposed to the same salinity range. High intake fish showed the characteristic elasmobranch osmolarity picture, with serum values slightly hyper-osmotic at all times. Low intake fish, however, showed a degree of hypo-osmotic regulation. Serum values for both groups overlapped at very low salinities. Serum urea was also affected by diet, so that again two distinct sets of values were produced, again with overlap at the lower salinities. When previously well-fed fish were starved over a period of one month, serum urea and osmolarity decreased simultaneously. Consequently, it is felt that serum osmolarity is directly related to serum urea levels. Serum chloride was not found to be affected by diet, both groups showing the same change in blood values when exposed to the same change in salinity. It is shown, however, that a reduction in food intake, over a period of more than a fortnight, can reduce metabolic urea to the extent of depressing serum osmolarity and, hence, shift the ionic and osmotic equilibrium between the fish and the sea water. This may result in varying degrees of hypoosmotic regulation.  相似文献   

18.
Closure of areas to fishing is expected to result in an increase in the abundance of targeted species; however, changes to populations of species not targeted by fishermen will depend upon their role in the ecosystem and their relationship with targeted species. The effects of protection on targeted and non-targeted reef fish species at the Houtman Abrolhos Islands, Western Australia were studied using baited remote underwater stereo–video cameras. Video images were collected from shallow (8–12 m) and deep (22–26 m) reef sites inside a Marine Protected Area (MPA) at each of three island groups and from three replicate fished locations at each of these groups that span a temperate-tropical transition area. The MPAs were established in 1994 and vary in size from 13.72 km2 at the Pelsaert group in the south to 22.29 km2 at the Easter group to 27.44 km2 at the Wallabi group in the north. The relative abundances of 137 fish species from 42 families were recorded. Large differences in fish assemblage structure existed between MPA and fished locations, and also between shallow and deep regions. Targeted fish species Plectropomus leopardus, Lethrinus miniatus, Lethrinus nebulosus, Pagrus auratus and Glaucosoma hebraicum were more abundant inside MPAs than in areas open to fishing. Their abundance inside MPAs was between 1.13 and 8 times greater than their abundance at fished locations. For non-targeted fish species many were more abundant in areas open to fishing, e.g. Coris auricularis, Thalassoma lutescens, Thalassoma lunare, Dascyllus trimaculatus, however others were conversely more abundant inside MPAs, e.g. Gymnothorax spp, Kyphosus sydneyanus, Scarus microhinos, Chromis westaustralis, Chaetodon spp. This study demonstrates that the removal of abundant targeted species from an ecosystem by fishing can indirectly impact non-fished species and alter the trophic structure of fish assemblages. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.  相似文献   

19.
The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe  相似文献   

20.
The swimbladders of three species of myctophid fishes (Symbolophorus californiensis, Tarletonbeania crenularis and Diaphus theta, collected in the Southern California Bight between 1963 and 1978), previously observed to have both inflated and noninflated swimbladders as adults, were examined to determine if the swimbladders either vary in their inflation on a diel basis or cease to be inflated and possibly become nonfunctional in some individuals. The swimbladders of two lanternfishes reported to lack inflated swimbladders as adults (Lampanyctus regalis and L. ritteri) were included in the study for comparison. Only the swimbladder of L. regalis did not increase in size with increasing fish size. In the remaining four species, both the lengths and volumes of noninflated swimbladders were positively correlated with fish standard length (SL). Although the swimbladder continued to increase in size, inflated swimbladders were not found in L. ritteri>2 mm SL. Inflated and noninflated swimbladders occurred in overlapping size ranges of S. californiensis, T. crenularis, and D. theta. Only 11% of the D. theta swimbladders were inflated, and the occurrence of inflated swimbladders dit not differ significantly with day vs night capture or fish size. Although the frequency of occurrence of inflated swimbladders decreased significantly in larger S. californiensis, both S. californiensis and T. crenularis caught at the surface at night with a neuston net had significantly higher proportions of inflated swimbladders than did those collected below the surface with either daytime or night-time trawls. The swimbladder does not appear to become nonfunctional in either species. Rather than maintaining either constant volumes or constant masses of gas in their swimbladders during their vertical migrations, both species may inflate their swimbladders only while in the surface waters at night, and have noninflated swimbladders while at other depths of their vertical ranges.  相似文献   

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