The trade-off between feeding, mate seeking and predator avoidance in copepods: behavioural responses to chemical cues

The behavioural responses of adult Temora longicornis to chemicalsignals from predators and potential sexual partners was studiedusing an automated three-dimensional (3-D) tracking system anda 2-D high-resolution filming set-up. The animals adapted theirswimming speed and turning behaviour in reaction to chemicalsignals to increase or decrease encounter probability with otherorganisms. Female copepods were shown to react to the smellof males with little ‘hops’, quite distinct fromboth their normal smooth swimming motion and escape responses.These hops are likely to be signals intended to increase theencounter probability with males. Swimming behaviour was shownto be a trade-off between conflicting motivations such as predatoravoidance on the one hand, and the search for food and sexualpartners on the other.     

Detection and capture of inert particles by calanoid copepods: the role of the feeding current

Although there is a scarcity of supporting empirical evidence,it has long been suspected that calanoid copepods use mechanoreceptionto detect the presence and location of potential prey itemsentrained in the feeding current. In this study, we documentthe first observations showing a freely swimming calanoid copepod,Skistodiaptomus oregonensis, attacking prey-sized, non-motile,inert particles entrained in the feeding current before theparticles contact the copepod's sensory appendages. Feedingcurrent geometry, fluid velocities and associated behavioursthat characterize these interactions are described. The resultsof this study show how copepod swimming behaviour, coupled witha low-velocity feeding current, not only increases copepod encounterrates with inert prey by increasing direct contact rates, butalso increases the probability of detecting and capturing remotelylocated prey that have well-developed escape responses. In turbulentregimes, a far-reaching, low-velocity feeding current shouldincrease encounter rates, but only if coupled with behavioursthat quickly minimize separation distances once prey is detected.     

Encounter probabilities between male and female rotifers

The relationship between population density and sexual reproduction is explored for planktonic rotifer populations. It has been argued that selection favors sex at population maxima because of favorable probabilities of encounter. Factors determining the probability of encounter are examined. Experimentally determined parameters for males and females (swimming speed, duration of fertility, net fecundity, and encounter distance) are used to investigate density effects on the probability of encounter. As population density increases, the probability of encounter also increases sigmoidally. The probability of a female encountering a male during her fertile period begins to increase rapidly at ≈ 50 males/1. The increase continues until it becomes asymptotic at ≈ 1000 males/1, where the probability of a female encountering, at least one male during her fertile period approaches one. The duration of female fertility is less than that of males, causing the female curve to be displaced toward higher population densities. Critical density, the population density where R_o′ is exactly equal to one, for males and females is 11.7 and 141 individuals/1, respectively. The precision of the model was evaluated by observing the frequency of male-female encounters during 212 5-min observation periods. The observed encounter rate was 1.49/min, compared to a predicted rate of 1.53/ min. These values are not significantly different as determined by t-test. The constraints placed on the timing of sex by population density are discussed.     

Experimental evidence of habitat selection and territoriality in the Amazonian whip spider Heterophrynus longicornis (Arachnida, Amblypygi)

Animals that select and defend suitable habitats against conspecifics may be favored by maximizing prey encounter rate, gaining protection, or securing matings. However, the identification of habitat selection and territoriality may be hindered in observational studies in sedentary species with low-density populations, such as the whip spider Heterophrynus longicornis. To circumvent such difficulties, we adopted an experimental field approach to evaluate if H. longicornis selects and defends habitat in Central Amazon. To evaluate whether individuals perform habitat selection, we monitored the permanence of 29 experimentally released individuals in buttressed trees with a wide variation in diameter at breast height (DBH), with and without burrows at their bases. To evaluate whether individuals are territorial, we experimentally removed 21 individuals from their previously occupied trees and monitored the recolonization of these trees. If H. longicornis were territorial, we predicted that the recolonizers would be smaller than the removed individuals. We found 12 individuals in the trees where they were released on subsequent days, none of them in trees without burrows. The individual permanence was related to the presence of burrows, and not to DBH. There was recolonization by smaller males and females on ten trees from which the amblypygids were removed. In combination, the permanence of individuals in trees with burrows, the rapid recolonization of experimentally vacated trees by smaller individuals, and the preponderance of one individual per tree, suggest that both male and female H. longicornis perform habitat selection and also that after selecting a site, they defend it against conspecifics.     

The effect of size and temperature on the frequency of limb beat of Temora longicornis Müller (Crustacea : Copepoda)

Copepods normally swim by rhythmically beating the cephalic limbs, so records of antennal movements represent their activity. The limb beat rate of Temora longicornis Müller was determined in relation to several factors. There was an inverse relationship between swimming rate and body size, and activity increased with environmental temperature up to 20–25°C. Copepods readily acclimated, as those kept at 15°C were less active than those kept at 5°C. The summer population was also less active in the low temperature range, but swimming reached a higher rate at higher temperatures than were tolerated by the winter population. No difference in rate of limb beat was found between similar sized males and females over a wide range of temperatures.     

Life table and predation of Neoseiulus longispinosus (Acari: Phytoseiidae) on Oligonychus coffeae (Acari: Tetranychidae) infesting tea

Life table and predation of the predatory mite Neoseiulus longispinosus (Evans) on the red spider mite (RSM), Oligonychus coffeae (Nietner), a major pest of tea in India, were studied in the laboratory. Developmental time from egg to adult varied from 4 to 14 days at 30 to 15 °C, respectively; at 35 °C no larva survived. Survival of immature stages was more than 94 % at all temperatures. Threshold temperature for development of immature stages of females and males was 10 and 9.9 °C, respectively, and thermal constant was 84.03 degree-days for females and 80 for males. Sex ratio was female biased and temperature (20–30 °C) had no clear effect on sex determination. Egg hatchability was 73 % at 35 °C and >97 % at lower temperatures. Average number of eggs laid per female/day was higher at 30 °C than at 20 or 25 °C. The highest net reproductive rate (R ₀) was 40.7, at 20 °C. Mean generation time (T) decreased from 28 to 13 days with temperature increasing from 20 to 30 °C. Weekly multiplication (6.5) and intrinsic rate of natural increase (r _m ) (0.268) were highest at 30 °C. Males lived longer than females at every temperature tested. Longevity was highest at 20 °C (50 days for females and 55 for males). Survival and longevity were adversely affected by temperature above 30 °C. Daily consumption of prey increased with the advancement of predator’s life stages; adult females consumed the highest numbers of prey items, preferably larvae and nymphs.     

Dietary overlap among early juvenile stages in an Antarctic notothenioid fish assemblage at Potter Cove,South Shetland Islands

To date, studies of food overlap in Antarctic fish have been performed on a mixture of late juvenile and adult stages, leaving the young immature specimens (TL ≤ 10 cm) practically unexplored. We studied diet overlap and potential competition among early juvenile individuals in a coastal notothenioid community at Potter Cove, by analysing the stomach contents of 225 fish of 5 species collected in the summer of 2009–2010. We used frequency of occurrence (F %) and the coefficient “Q” for diet evaluation and the method of Tyler and the similarity index “S” for food overlap. Amphipods of the suborder Gammaridea were the main (Q > 2.900) and most frequent (% F) prey for all species, although Notothenia coriiceps also consumed gastropods of the family Littorinidae, mostly Laevilitorina antarctica. Secondary prey were algae for Notothenia rossii and N. coriiceps, calanoid (pelagic) and harpacticoid (benthic) copepods for Trematomus newnesi and the latter copepods and isopods of the family Munnidae for Lepidonotothen nudifrons. The reoccurrence of prey among fish species was 39.6 % and food overlap between 90 % of species pairs was under 58 %. Because similarly low values of diet overlap were reported for intermediate/advanced juveniles and adults of the same species at the same site, we conclude that there is no difference in the degree of interspecific food overlap and therefore potential competition between the immature and mature fraction of the fish community. Food competition is avoided by resource partitioning along a depth gradient or by different prey species.     

Females as intraguild predators of males in cross-pairing experiments with phytoseiid mites

Studies on intraguild interactions between phytoseiid species have shown that intraguild predation occurs and is most commonly manifested as adult females of one species feeding on juveniles of another. Whether such intraguild interactions can also occur between adult females of one species and adult males of another, is not known. Herein, we report on intraguild interactions between adults of the two sexes in cross-pairing experiments involving three related phytoseiid species (Neoseiulus paspalivorus, N. baraki and N. neobaraki) that are potential candidates for controlling the coconut mite Aceria guerreronis, a serious pest of coconut palms in tropical countries. For comparative reasons, the experiments were repeated with larvae instead of males, and with only males or only females of two different species together. In the presence of an ample supply of prey, females of N. neobaraki never fed on individuals of their own species, yet appeared to be very aggressive against males, as well as larvae of the other two phytoseiid species. They also fed on females of N. paspalivorus, but rarely on females of N. baraki. Males of N. neobaraki did not suffer mortality when together with females of either of the two other phytoseiid species. Males of N. baraki did not suffer predation from females of N. paspalivorus, but males of N. paspalivorus suffered some mortality (15 %) from N. baraki females. Larvae of each of the three species were vulnerable to intraguild predation by heterospecific adult females, except for N. neobaraki larvae when together with N. baraki females. The absence or presence of intraguild predation is largely explained by the size ratios of the individuals that were put together: large individuals feed on smaller ones, but never the reverse. For each sex, size declines in the following order: N. neobaraki > N. baraki > N. paspalivorus. Moreover, for each species, females are larger than males and males are larger than larvae. Strikingly, however, females did not kill males and larvae of their own species. We propose that niche competition between related phytoseiid species is not only determined by intraguild predation on heterospecific larvae, but also by imposing great mortality on males from the intraguild prey because phytoseiid females being pseudo-arrhenotokous require insemination to produce offspring of both sexes.     

Spring diet of common eiders (Somateria mollissima) in Breiðafjörður, West Iceland, indicates non-bivalve preferences

Breiðafjörður is an important molting, breeding, and wintering area for about 25 % of the Icelandic common eider (Somateria mollissima) population. However, feeding habits of eiders in this area have not been investigated until now. Prey selection was analyzed from 192 stomach samples (esophagus and proventriculus) collected in spring 2007–2010. Thirty-five prey species were identified; the highest percentage occurrences were of gastropods (79 %), chitons (polyplacophorans) (58 %), crustaceans (43 %), bivalves (26 %), and echinoderms (8 %). The most common food species was the mottled red chiton Tonicella marmorea (58 %), followed by the common whelk Buccinum undatum (40 %), the spider crab Hyas arenarius (39 %), and the chink shell Lacuna vincta (35 %). The majority of the food items was of small size and consumed in high quantity. The chitons and mussels were of similar average sizes (11.7 and 13.4 mm, respectively), which might suggest that prey size could be as important as species in food selection. There were some inter-annual differences in dominant prey classes between years. For example T. marmorea was found in 60–70 % of birds in the years 2007 and 2009 but only in 30 % of the birds in the other years investigated. Diets of males and females were equally diverse and similar when all months and years were pooled. Prey selection was highly variable but most individuals focused on few or a single species in the hours prior to collection. Results indicate that the most common prey species for common eiders is a chiton and not blue mussels as reported elsewhere.     

Why eat extrafloral nectar? Understanding food selection by Coleomegilla maculata (Coleoptera: Coccinellidae)

Methods of increasing predator abundance within a habitat include the incorporation of non-prey food items, yet the influence of this on predation intensity toward herbivores remains unknown. In order to gain an understanding of nectar feeding in the predaceous beetle, Coleomegilla maculata (DeGeer), laboratory studies were conducted evaluating prey consumption in the presence of extrafloral nectaries. The physiology of beetles with access to prey only and a mixed diet were compared. To elucidate results of beetle physiology, Y-tube olfactometer studies were conducted and preferences between food types evaluated. Coleomegilla maculata females consumed 9 % fewer aphids when nectar was available. Lipid and glycogen content, as well as oocyte volume were not increased upon consumption of a mixed diet. Evaluation of predator behavior when offered both food resources together and separately demonstrated that extrafloral nectaries are attractive.     

Bioenergetic model of planktivorous fish feeding, growth and metabolism: theoretical optimum swimming speed of fish larvae

The feeding activity of an individual fish larva is described by an equation which includes parameters for the area successfully searched, probability of food capture multiplied by the cross-sectional perceptive visual field, larval swimming speed and the time required to consume a unit of food energy. The proportion of ingested food energy used for metabolism increases exponentially with increasing swimming speed. The model predicts that food consumption rate increases asymptotically whereas metabolic rate increases exponentially. This results in a predicted growth rate curve that reaches a maximum at a certain swimming speed and decreases at both higher and lower speeds. The model can be used to predict the influence of type of prey, prey density, water temperature etc. on larval growth. An expression describing how many hours per day fish larvae must forage in order to grow at a certain daily body weight gain allows the limits of environmental conditions for positive, zero and negative growth rate to be set. Results of simulations demonstrated that the optimum swimming speed for maximum growth of coregonid larvae increased with an increase in food density, decrease in water temperature or decrease of prey vulnerability. At optimum ‘theoretical’ swimming speed an increase in water temperature from 5 to 17° C required the food density to be increased from 20 to 80 copepods l^?1 in order to maintain a daily growth increment of 2%. The minimum Artemia density required for maintenance metabolism increased from 10 to 30 items 1¹ over the same temperature increase from 5 to 17° C, and food densities required for 8% growth rates were 26 and 56 Artemia nauplii l^?1 at 5 and 17° C, respectively. Contrary to previous findings, results of the present study suggest that metabolic rates of actively feeding fish larvae may be from 5 to 50 times the standard metabolic rate: earlier studies suggested that a factor of 2–3 may be generally applicable.     

Increased Frequency of Intergroup Encounters in Wild Bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>) Around the Yearly Peak in Fruit Abundance at Wamba

Intergroup interactions in primates vary from nonagonistic to severely aggressive. Food resources and fertile females cause intergroup aggression when groups defend resources and mates from other groups. Peaceful intermingling during intergroup encounters is rare but has been reported in several primates, including bonobos (Pan paniscus). Although intergroup encounters in bonobos occur at both nonprovisioned and provisioned sites, provisioning may be one factor responsible for frequent intergroup encounters. We studied intergroup encounters between one bonobo group (PE) and its neighboring groups, one semihabituated and two habituated groups, under nonprovisioned conditions in 2010–2015 at Wamba, Democratic Republic of Congo. We examined whether fruit abundance and females with sexual swellings influenced encounter frequency. PE group encountered the other groups on 7.1 days (mean, range: 0–19) per month. Up to four different groups met simultaneously at one location. Encounter frequency was highest around the yearly peak in fruit abundance, suggesting that reduced food competition is linked to increased encounter frequency. During periods of relatively low fruit abundance, the probability of an encounter was higher when a higher number of adult females with maximum sexual swellings were present, suggesting that the attractiveness of females with sexual swellings promoted group encounters during periods of relatively low fruit abundance. These findings suggest that both competition for food and the availability of females with sexual swellings influence encounter frequency in bonobos.     

Rapid response of brook trout to removal of its intraguild prey, yellow perch

Brook trout (BT) and yellow perch (YP) interact as intraguild predator and prey: while adult BT consume YP, juvenile BT and YP compete for food resources, i.e. zooplankton (mainly a pelagic resource) and zoobenthos (mainly littoral). YP appear to dominate the interaction, reducing BT abundance in lakes with YP present. We removed 90 % of YP from a 35 ha oligotrophic lake to test whether the removal of the dominant competitor would lead to an increase in growth and recruitment of juvenile BT. This manipulation shifted the YP:BT biomass ratio from 11:1 to 1.5:1. The food web response was evaluated using the pre-removal condition as the reference. The abundance of cladoceran zooplankton increased, especially large-bodied Dapnia catawba, and the body size of Daphnia dubia increased significantly, responses indicative of reduced zooplanktivory. Some benthic taxa (chironomid and Ephemeroptera larvae) also increased following YP removal. BT responded rapidly: mean length at age-1 increased from 11.6 cm to 12.8 cm (~35 % weight increase), the condition of 15–25 cm trout improved, and stable isotopes indicated increased consumption of littoral-linked prey by 20 to 30 cm BT. YP, despite their reduced abundance, remained the primary prey of large BT (>?25 cm). The suite of changes observed in response to YP removal support the view that although YP are the primary prey of large BT, they exert a strong competitive effect on juvenile BT. Further study would be required in order to assess whether piscivorous BT can establish persistent top–down control of YP due to enhanced juvenile BT recruitment.     

Sexual cannibalism: high incidence in a natural population with benefits to females

Background

Sexual cannibalism may be a form of extreme sexual conflict in which females benefit more from feeding on males than mating with them, and males avoid aggressive, cannibalistic females in order to increase net fitness. A thorough understanding of the adaptive significance of sexual cannibalism is hindered by our ignorance of its prevalence in nature. Furthermore, there are serious doubts about the food value of males, probably because most studies that attempt to document benefits of sexual cannibalism to the female have been conducted in the laboratory with non-natural alternative prey. Thus, to understand more fully the ecology and evolution of sexual cannibalism, field experiments are needed to document the prevalence of sexual cannibalism and its benefits to females.

Methodology/Principal Findings

We conducted field experiments with the Mediterranean tarantula (Lycosa tarantula), a burrowing wolf spider, to address these issues. At natural rates of encounter with males, approximately a third of L. tarantula females cannibalized the male. The rate of sexual cannibalism increased with male availability, and females were more likely to kill and consume an approaching male if they had previously mated with another male. We show that females benefit from feeding on a male by breeding earlier, producing 30% more offspring per egg sac, and producing progeny of higher body condition. Offspring of sexually cannibalistic females dispersed earlier and were larger later in the season than spiderlings of non-cannibalistic females.

Conclusions/Significance

In nature a substantial fraction of female L. tarantula kill and consume approaching males instead of mating with them. This behaviour is more likely to occur if the female has mated previously. Cannibalistic females have higher rates of reproduction, and produce higher-quality offspring, than non-cannibalistic females. Our findings further suggest that female L. tarantula are nutrient-limited in nature and that males are high-quality prey. The results of these field experiments support the hypothesis that sexual cannibalism is adaptive to females.     

Living in the fast lane: effects of cost of locomotion on foraging behaviour in juvenile Atlantic salmon

Stream-dwelling, juvenile Atlantic salmon, Salmo salar L., feed mainly on drifting invertebrates, usually by swimming upstream from a stationary position to intercept individual prey items. Laboratory experiments tested the prediction that individual salmon should reduce the distance over which they would travel (attack distance) to intercept drifting food items as the energy cost of swimming increases with increasing current velocity. Attack distance varied inversely with current velocity as expected. The fish's average speed of upstream movement relative to the substrate remained constant and the duration of individual attacks therefore declined as current velocity increased. Calculated reaction distances and a second ecperiment using tethered prey drifting at speeds independent of current velocity confirmed that these relationships were due to fish actually delaying attacks on perceived prey for longer periods as current velocity increased. Using estimated metabolic rates for burst swimming, it appears that energy expenditure per attack varies little with current velocity. Therefore, by reducing their reaction and attack distances in response to increasing current velocity, the fish reduced their energy cost of travel per attack.     

Properties of food loads delivered to nestlings in the pied flycatcher: effects of clutch size manipulation, year, and sex

Experimental manipulation of the number of altricial offspring is supposed to modify parental expenditure in birds. In addition to the observed increase in parental feeding rate, it is also possible that the choice of prey or the size of load may change with the changing demand for food. Sexual differences in the provisioning response are also expected, on the basis of earlier studies. We examined the effect of brood size manipulation on choice of prey brought to nestlings and load size in the pied flycatcher. The composition and size of loads differed between years, possibly depending on varying availability of different prey types. Males responded to brood size enlargement by gathering heavier loads, whereas females showed no response. The alteration of load size in males was not explained by a larger number of prey items or mean prey size, but was a combination of these components. It is likely that males also increased their work rate in response to increased food demand at the nest. The absence of response in females might be because they are unable to increase work rate any further, or because food delivery rate in females can not be optimized by changing load properties. Received: 18 December 1997 / Accepted: 1 March 1998     

A meal or a male: the ‘whispers’ of black widow males do not trigger a predatory response in females

Introduction

Female spiders are fine-tuned to detect and quickly respond to prey vibrations, presenting a challenge to courting males who must attract a female’s attention but not be mistaken for prey. This is likely particularly important at the onset of courtship when a male enters a female’s web. In web-dwelling spiders, little is known about how males solve this conundrum, or about their courtship signals. Here we used laser Doppler vibrometry to study the vibrations produced by males and prey (house flies and crickets) on tangle webs of the western black widow Latrodectus hesperus and on sheet webs of the hobo spider Tegenaria agrestis. We recorded the vibrations at the location typically occupied by a hunting female spider. We compared the vibrations produced by males and prey in terms of their waveform, dominant frequency, frequency bandwidth, amplitude and duration. We also played back recorded male and prey vibrations through the webs of female L. hesperus to determine the vibratory parameters that trigger a predatory response in females.

Results

We found overlap in waveform between male and prey vibrations in both L. hesperus and T. agrestis. In both species, male vibrations were continuous, of long duration (on average 6.35 s for T. agrestis and 9.31 s for L. hesperus), and lacked complex temporal patterning such as repeated motifs or syllables. Prey vibrations were shorter (1.38 - 2.59 s), sporadic and often percussive. Based on the parameters measured, courtship signals of male L. hesperus differed more markedly from prey cues than did those of T. agrestis. Courtship vibrations of L. hesperus males differed from prey vibrations in terms of dominant frequency, amplitude and duration. Vibrations of T. agrestis males differed from prey in terms of duration only. During a playback experiment, L. hesperus females did not respond aggressively to low-amplitude vibrations irrespective of whether the playback recording was from a prey or a male.

Conclusions

Unlike courtship signals of other spider species, the courtship signals of L. hesperus and T. agrestis males do not have complex temporal patterning. The low-amplitude ‘whispers’ of L. hesperus males at the onset of courtship are less likely to trigger a predatory response in females than the high-amplitude vibrations of struggling prey.     

Dietary habits of the polkadot skate <Emphasis Type="Italic">Dipturus chinensis</Emphasis> in the East China Sea

We investigated the dietary habits of the polkadot skate, Dipturus chinensis (Japanese name: Gangi-ei), in the East China Sea by analyzing the stomach contents of 529 individuals. Of these, 18 (3.40%) had empty stomachs. The mean and maximum fullness indices (FIs) were 0.67% and 6.12%, respectively. The mean FI of female skates > 500 mm in total length (TL) was higher than that of males and smaller females. The most dominant prey category was crustaceans, which included shrimps, crabs, and anomurans, followed by fishes. Cephalopods were additionally observed. Dipturus chinensis fed on a wide variety of crustacean taxa, and among them, Caridea and Galatheidae were particularly common. Fishes that could be identified were mostly demersal fishes, such as Callionymidae, Triglidae, and Rajidae. Some juvenile D. chinensis were also observed, indicating that cannibalism occurs in this species. Even though dental sexual dimorphism was observed in large size classes (mainly > 500 mm TL), the diet of males and females significantly overlapped. Size-related dietary shifts from small crustaceans, such as shrimps and anomurans, to relatively fast-swimming prey, such as fishes, crabs, and cephalopods, were observed in all seasons. The composition of these prey categories varied among seasons. Shannon–Wiener diversity indices calculated for each size class ranged from 2.01 to 2.51, indicating D. chinensis are generalist predators. As size increased, trophic level increased from 3.58 to 3.85, indicating that larger specimens are at a slightly higher trophic level than smaller specimens. These results suggest that D. chinensis are middle-trophic-level generalist predators that utilize diverse prey depending on their size and the season, and thus play an influential role in the demersal ecosystem of the East China Sea.     

Ontogeny of feeding in two native and one alien fish species from the Murray-Darling Basin, Australia

Investigations into the feeding of the early stages of fishes can provide insights into processes influencing recruitment. In this study, we examined ontogenetic changes in morphology and feeding behaviour of two native Australian freshwater species, Murray cod, Maccullochella peelii peelii, and golden perch, Macquaria ambigua, and the alien species, common carp, Cyprinus carpio. Murray cod free embryos are large and well developed at the onset of feeding, whereas the other two species begin exogenous feeding much younger and are smaller and less-developed. Carp commence exogenous feeding 3 days earlier than golden perch, and show more advanced development of the eyes and ingestive apparatus. We conducted feeding experiments, presenting larvae of the three species with a standardised prey mix (comprising equal numbers of small calanoid copepods, large calanoid copepods, small Daphnia, and large Daphnia). Larvae of most tested ages and species showed a preference for mid-sized prey (300–500 μm wide). This was true even when their gapes substantially exceeded the largest prey offered. Daphnia were consumed more than similar-sized copepods. The results of this study suggest that survival through their larval period will be threatened in all three species if catchable prey <500 μm in width are not available throughout such time. They also suggest that interspecific competition for prey may occur, especially when larvae are very young. The precocious development of structures involved in feeding and the extended transition from endogenous to exogenous feeding of early carp larvae are likely to have contributed to the success of this species since its introduction to Australia.