Cryptic prey colouration increases search time in brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>): effects of learning and body size

Little is known about how cryptic colouration influences prey search in near-surface aquatic habitats, although such knowledge is critical for understanding the adaptive value of colour crypsis as well as the perceptive constraints influencing foraging behaviour in these environments. This study had two main aims: (1) to investigate how background colour matching by prey affects foraging efficiency by brown trout parr and (2) to investigate how foraging ability on cryptic and conspicuous prey is affected by fish size at age (reflecting dominance). We addressed these questions by training wild brown trout parr to forage individually on live brown-coloured maggots on a cryptic (brown) or conspicuous (green) background. A separate experiment confirmed the absence of trout preference for brown or green substrate. The results show that prey background colour matching increases search time in brown trout. Search time generally decreased by learning, but conspicuous prey remained an easier prey to find throughout the six training trials. Thus, perceptive constraints appear to limit search efficiency for cryptic prey, suggesting that cryptic colouration can confer survival benefits to prey in natural environments. Smaller fish generally found conspicuous prey faster than larger individuals, whereas search time for cryptic prey was not influenced by body size. This suggests that smaller individuals compensate for inferior competitive ability by increasing foraging activity rather than improving cognitive ability. The technique of varying cognitive demands in behavioural tasks could be used more in future studies aimed at distinguishing motivational effects from cognitive explanations for variation in behavioural performance.     

The feeding behaviour of Schistocerca gregaria,the desert locust,on two starch mutants of Arabidopsis thaliana

Summary. Schistocerca gregaria, the desert locust, has been shown to regulate its dietary intake with respect to specific macronutrients in synthetic foods. This study examined the nutrients in the leaves of two starch mutants of Arabidopsis thaliana, and then compared the feeding behaviour of locusts on the two starch mutants. The high-starch mutant had c. 25 times more starch than the no-starch mutant. Newly molted 5th stadium locusts were preconditioned for 3 days on one of the mutants, and then observed for 90 min while exposed to the same or the alternative mutant. Locusts pretreated with the no-starch mutant fed longer during the first meal on high-starch mutants, spent more time feeding, and had the smaller latency to begin a meal when compared to the locusts pretreated on the high-starch mutant. The results of the study are interpreted in light of an integrative model of nutrient balancing. Received 16 November 1999; accepted 23 December 1999     

Toxin accumulation and feeding behaviour of the planktonic copepod Calanus finmarchicus exposed to the red-tide dinoflagellate Alexandrium excavatum

The planktonic copepod Calanus finmarchicus is a dominant member of the zooplankton community in the lower St. Lawrence Estuary in eastern Canada. Blooms of the toxic marine dinoflagellate Alexandrium excavatum which produces high cellular levels of paralytic shellfish poisoning (PSP) toxins, occur during the period of high C. finmarchicus production in summer in this region. To study the feeding behaviour of C. finmarchicus in the presence of Alexandrium spp., experiments were conducted in which female adult copepods collected from the St. Lawrence Estuary between May and September 1991 were exposed under controlled conditions to two toxic isolates of A. excavatum (Pr18b and Pr11f) from the estuary and to a non-toxic control (PLY 173) of a closely related species, A. tamarense isolated from the Tamar Estuary, Plymouth, U.K. Clearance rates on non-toxic A. tamarense cells averaged 5.5 ml ind^-1 h^-1 but were nearzero with either toxic isolate. When presented with a mixture of A. excavatum and the non-toxic diatom Thalassiosira weissflogii in varying proportions, C. finmarchicus fed upon the diatom but avoided the toxic dinoflagellate. Although feeding rates on A. excavatum were very low, toxin analysis by high-performance liquid chromatography with fluorescence detection (HPLC-FD) revealed that the PSP toxins were accumulated in copepods exposed to toxigenic dinoflagellates.The toxin composition in copepods was similar to that of the toxic dinoflagellate, but not necessarily identical, particularly after short-term (2-h) exposure, when relatively elevated levels of N-sulfocarbamoyl toxins were detected. The evidence suggests that C. finmarchicus ingests toxic dinoflagellate cells, either mistakenly or during exploratory bouts of feeding, and accumulates PSP toxins in its gut system and perhaps in other tissues.     

Locomotor impairment and defense in gravid lizards (Eumeces laticeps): behavioral shift in activity may offset costs of reproduction in an active forager

Summary Female Eumeces laticeps experience a substantial decrease in running speed (ca. 25%) and an even greater loss of endurance (slightly over 50%) while gravid. Because some widely foraging lizards, including E. laticeps, rely primarily on running to escape predators, the decreases in speed and stamina may contribute to an increased risk of predation. However, observations suggest that gravid females become less active or conspicuous on the surface. Ambush foraging lizards rely relatively more on crypsis associated with immobility to avoid predation and thus can have greater average relative clutch mass (RCM) than active foragers. Behavioral compensation for locomotor impairment by becoming less active or conspicuous may allow some species the advantages inherent in both high relative clutch mass when gravid and the increased energetic profitability of active foraging when not gravid. As females gain weight during the breeding season, they may forage actively until the risk due to increasing locomotor impairment becomes too great and then change defensive strategy to greater reliance on crypsis. Without such a shift, widely foraging squamate reptiles may be less able than ambush foragers to exploit life-historical strategies demanding high current investment in reproduction.     

Phenotype-environment matching in the shore crab (Carcinus maenas)

The shore crab (Carcinus maenas) exhibits a range of carapace pattern polymorphisms, but little is known regarding their function or maintenance. If patterns represent some form of crypsis, then associations between carapace colouration and substrate are expected; to determine whether such relationships exist, frequency of crab morphs and quantity of substrate type were measured from fifteen 10×40 m² quadrats at each of three sites along the southern shore of the Firth of Forth, Scotland. Five thousand one hundred and thirty-seven crabs and 3.6 km of line intercept transect data were collected during a 9-week period. Crab abundance, relative frequency of morphs and substrate type varied significantly among the three sites. Plain crabs were strongly associated with macro-algal substrates whereas patterned crabs were associated with mussel beds. This pronounced phenotype-environment matching, as well as various characteristics of the carapace patterns themselves, suggests that patterned crabs are cryptic on polychromatic backgrounds. The frequency of patterned crabs and the percentage of white pigment on the carapace both declined significantly with carapace width. The loss of pattern coincides with an ontogenetic shift in habitat use and we present evidence to suggest that individual crabs lose their pigment, rather than larger patterned crabs being preferentially removed from the population by predators. Throughout their life history, shore crabs encounter high variation in predation, food supply, and physical habitat; to survive they have evolved a strategy that includes elements of pattern polymorphism, crypsis, ontogenetic shifts, and plastic responses.     

Seasonal ontogenetic colour plasticity in the adult striated shieldbug <Emphasis Type="Italic">Graphosoma lineatum</Emphasis> (Heteroptera) and its effect on detectability

Camouflage and warning colouration are two important forms of protective colouration. We have studied the detectability of two seasonal colourations in the aposematic striated shieldbug, Graphosoma lineatum. The typical colouration of this insect is red and black, which is also the colouration of the reproductive post-hibernation bugs in our study area in south central Sweden. However, the majority of newly eclosed adults in late summer exhibit a ‘pale’ (light brownish, non-red) and black striation, and these bugs appear quite cryptic to the human eye when sitting on the dried stems and umbels of their host plants. In experiments using photographs of prey in the late-summer habitat shown on a computer screen, we compared the time to detection by human subjects of bugs, which had been manipulated to show either of the two typical seasonal colourations. Time to detection was significantly longer for the pale and black than for the red-and-black striation in images with the bug photographed at two different distances. This indicates that the pale pre-hibernation striation may have a cryptic function. In a separate experiment, we tested detectability of striated and non-striated manipulations of bug pre-hibernation colouration against the late-summer background, and found that time to detection was significantly longer for the striated bugs. We discuss potential functional explanations for the seasonal ontogenetic colour plasticity and suggest that the epidermal pale colour in the late summer provides a benefit of increased camouflage.     

Evaluating the coastal environment for marine birds

The marine environment is being increasingly exploited by fisheries and the oil and gas industry. Conservationists urgently need the ability to identify the processes that determine patterns of abundance of marine species. We describe a preliminary Geographic Information System (GIS) in which spatial data on environmental variables (seabird colony locations, sea depth and seabed sediments) are integrated with realistic energy constraints faced by marine birds during the breeding season. A simple foraging model predicts the spatial variation in the quality of given locations as potential feeding sites under different feeding conditions and stages of the breeding cycle. We show how the approach can be used to help managers identify key marine areas and assess the impacts of environmental change or damage.     

Effects of furoquinoline alkaloids on the growth and feeding of two polyphagous lepidopterans

Summary. The furoquinoline alkaloids skimmianine and dictamnine were tested for effects on the feeding and growth of larvae of the generalist lepidopteran, Spodoptera litura. Skimmianine was further investigated for toxic and phototoxic effects on the larvae of Trichoplusia ni, also a generalist species. In feeding experiments with S. litura, growth and consumption of larvae decreased with increasing concentration of furoquinolines; skimmianine caused a greater reduction in growth than dictamnine. In T. ni, dietary skimmianine reduced growth and consumption; when administered topically, it significantly reduced consumption but without a concomitant reduction in growth. Phototoxicity of skimmianine was not apparent in T. ni because UV light failed to increase the negative effects of the alkaloid on larvae.     

Female-specific colouration, carotenoids and reproductive investment in a dichromatic species, the upland goose Chloephaga picta leucoptera

Although studies on the evolution and function of female ornaments have become more numerous in the last years, the majority of these studies were carried out in cases where female ornaments were a smaller and duller version of the ornaments found in males. There are substantially fewer studies on species with female-specific ornaments. However, no study so far investigated the potential of female-specific colouration as a quality signal in birds with conventional sex roles. We studied female-specific ornamentation in a strongly sexually dichromatic species, the upland goose Chloephaga picta leucoptera, in two consecutive years. Male upland geese have white head and breast feathers and black legs, whereas females have reddish-brown head and breast feathers and conspicuous yellow-orange legs. We found that female-specific colouration in upland geese can reliably indicate different aspects of female phenotypic quality. Females with more orange coloured legs and more red-like head colours had higher clutch and egg volumes than females with a paler leg and head colouration, and a more reddish plumage colouration was related to a higher body condition. These relationships provide the theoretic possibility for males to assess female phenotypic quality on the basis of colouration. Furthermore, the females with a more orange-like tarsus colouration had higher plasma carotenoid levels. Both tarsus colouration and carotenoid concentrations of individual females were highly correlated across years, indicating that tarsus colour is a stable signal. Despite this correlation, small individual differences in plasma carotenoid concentrations between the two study years were related to differences in tarsus colouration. We thus show for the first time in a wild bird and under natural conditions that carotenoid-based integument colouration remains consistent between individuals in consecutive years and is also a dynamic trait reflecting individual changes in carotenoid levels. In this species, where pairs form life-long bonds, the honesty of the carotenoid-based integument colouration suggests that it may be a sexually selected female ornament that has evolved through male mate choice.     

How do slugs cope with toxic alkaloids?

Summary. Arion lusitanicus and other slugs are able to feed on a variety of plants, even those containing toxic secondary metabolites, such as alkaloids. Alkaloids, like sparteine, lupanine, quinidine and atropine are avoided in no-choice feeding experiments only when they are present in concentrations above 1 mg/g food pellet. Alkaloids (sparteine, lupanine, cytisine, quinidine, atropine, senecionine, eserine, and harmaline) are toxic to Arion lusitanicus when injected. LD₅₀ values are 10 to 20 fold higher than in vertebrates indicating that slugs have a high tolerance towards food toxins. The tolerance is higher in young animals than in adult slugs. Injected alkaloids are rapidly detoxified within 72 h. Tolerance and detoxification can be induced by feeding slugs on non-lethal doses of lupin alkaloids. Using isolated microsomal preparations from the digestive gland, active detoxification was observed in vitro. Evidence is presented that cytochrome p450 plays an important role in detoxification of the applied alkaloids. A powerful and inducible detoxification systems appears to be the main mechanism that allows slugs to feed on plants rich in secondary metabolites (when no other food is available), that are usually avoided by other herbivores.     

Pyrrolizidine alkaloids on three trophic levels – evidence for toxic and deterrent effects on phytophages and predators

Summary. Pyrrolizidine alkaloids (PAs) present a model system in the investigation of tritrophic interactions mediated by plant secondary compounds. However, their toxicity for insect herbivores has never been experimentally proven. Here, we demonstrate the toxic effects of a PA on growth and survival of the eri silk moth Philosamia ricini. In a feeding experiment, larvae of this generalist herbivore fed with an artificial PA diet gained weight significantly slower than control animals, and died as pupae. We suggest that derivatives of the ingested PA N-oxide damage developmental functions during metamorphosis. A tracer test with [¹⁴C]senecionine N-oxide revealed that the caterpillars lack adaptations that would prevent conversion of the chemical into the pro-toxic free base. In contrast, the PA adapted leaf beetle Longitarsus anchusae accumulates PAs as N-oxides. We tested the purpose of sequestration in this species as defence against predators. Through a series of prey choice experiments with three carabid predator species, chemically non-protected bark beetle pupae were chosen almost uniformly over L. anchusae pupae. In a following choice test with one of these predators, artificially PA-treated mealworm segments deterred the predator from feeding. Overall the study corroborates the immediate toxic effect of PAs on non-adapted herbivores and the protective effect that adapted insects may gain by sequestering them. It thereby underlines the potential for PAs to play a central role in multitrophic interactions between plants, phytophages and their predators.     

Chemical determinants of resistance in winter-dormant seedlings of European white birch (Betula pendula) to browsing by the mountain hare

Summary A sample of one-year-old seedlings of European white birch (Betula pendula) was analyzed to determine the content of sugars, phenolics and terpenoid compounds. Two vertical segments of each seedling were analyzed separately. The number of resin droplets, which correlates strongly and negatively with feeding by the mountain hare, was also counted on the bark of experimental seedlings. The variation in the palatability of birch seedlings to mountain hare was determined primarily by the most abundant terpenoid, papyriferic acid. On the other hand, the resistance to hare feeding at the seedling bases, which are frequently attacked by voles, appeared to be dependent on other components, apparently phenolic substances. Sugars did not affect the resistance of the tested seedlings. The variation among experimental seedlings was much greater for secondary substances, especially terpenoid compounds, than for sugars. It is suggested that this high variation in protective compounds may be an adaptive trait selected for by the feeding of generalist herbivores.     

Behavioral attributes and parental care of Varroa mites parasitizing honeybee brood

Varroa jacobsoni, an ectoparasite of the Asian honeybeeApis cerana, has been introduced world-wide, and is currently decimating colonies of the European honeybeeApis mellifera.Varroa's reproductive cycle is tuned to that of drone cells, those mainly parasitized in the original host. We describe here how a single fertilized female, infesting a brood cell, can produce two to four adult fertilized females within the limited time span of bee development (270 h in worker and 320 h in drone cells), despite the disturbance caused by cocoon spinning and subsequent morphological changes of the bee. From observations on transparent artificial cells we were able to show how the mite combats these problems with specialized behaviors that avoid destruction by the developing bee, prepares a feeding site for the nymphs on the bee pupa, and constructs a fecal accumulation on the cell wall which serves as a rendezvous site for matings between its offspring. The proximity of the fecal accumulation to the feeding site facilitates feeding by the maturing progeny. However, communal use of the feeding site leads to competition between individuals, and protonymphs are most disadvantaged. This competition is somewhat compensated by the timing of oviposition by the mites. Use of a common rendezvous and feeding site by two or moreVarroa mothers in multiinfested cells may have developed from the parental care afforded to them as nymphs.     

Milkweeds,monarch butterflies and the ecological significance of cardenolides

Summary The contribution of Miriam Rothschild to the monarch cardenolide story is reviewed in the light of the 1914 challenge by the evolutionary biologist, E.B. Poulton for North American chemists to explain the chemical basis of unpalatability in monarch butterflies and their milkweed host plants. This challenge had lain unaccepted for nearly 50 years until Miriam Rothschild took up the gauntlet and showed with the help of many able colleagues that monarchs are aposematically coloured because they sequester toxic cardenolides from milkweed host plants for use as a defence against predators. By virtue of Dr Rothschild's inspiration and industry, and subsequently that of Lincoln Brower and his colleagues, this tritrophic interaction has become a familiar paradigm for the evolution of chemical defences and warning colouration. We now know that the cardenolide contents of different milkweeds vary quantitatively, qualitatively and spatially, both within and among species and we are starting to appreciate the implications of such variation. However, as Dr Rothschild has pointed out in her publications, cardenolides have sometimes blinded us to reality and it is curious how little evidence there is for a defensive function to cardenolides in plants — especially against adapted specialists such as the monarch. Thus the review will conclude with a discussion of the significance of temporal variation and induction of cardenolide production in plants, the lethal plant defence paradox and an emphasis on the dynamics of the cardenolide-mediated interaction between milkweeds and monarch larvae.     

Copper tolerance in a population of Silene vulgaris ssp. maritima (A. & D. Love) at Dolfrwynog Bog near Dolgellau,North Wales

Clones of Silene vulgaris ssp. maritima individuals from Dolfrwynog Bog, North Wales and from a coastal control site were propagated under standardised environmental conditions. After five months, cuttings were taken and tested for copper tolerance by means of root extension analysis in solutions of various different copper concentrations. Indices of tolerance suggested that individuals from the Dolfrwynog population were more resistant to elevated copper concentrations than were the control population individuals. Due to the initial five months under standardised conditions, it was assumed that the cloned plant material would have lost all plastic adaptive responses to its native environment. If this assumption was valid then the differences in copper tolerance observed between the two populations had a genetic basis and the Dolfrwynog Bog population is a copper tolerant ecotype.     

Defence effectiveness of easy bleeding sawfly larvae towards invertebrate and avian predators

Summary. Easy bleeding is a phenomenon discovered in some tenthredinid insects which possess a particularly low mechanical resistance of the integument, leading under mechanical stress to haemolymph exudation. It has a defensive effect against ants and wasps through harmful plant compounds which are sequestered in the haemolymph. Here we describe etho-ecological and some chemical aspects of the defence of easy bleeders and specify the range of predators to which easy bleeding might be effective. Beside a high haemolymph deterrence associated with low integument resistance across sawfly species, we also detected toxicity of the haemolymph of some species to workers of the ant Myrmica rubra. The behaviour of easy bleeders is to move slowly and, once disturbed, to become motionless, thereby probably impeding the tendency of a predator to attack. This behaviour had no beneficial effect for easy bleeders when attacked by the predatory bug Podisus maculiventris. Bugs could successfully and without harm prey on sawfly larvae without evoking easy bleeding. For the easy bleeder Athalia rosae, host plants with different secondary metabolite profiles, and, consequently, changes in haemolymph chemistry only slightly affected the feeding behaviour of the bugs. To test the effectiveness of easy bleeding towards a vertebrate predator, easy bleeders were offered to birds, Sturnus vulgaris. The body colouration of the sawfly larvae was of prime importance in determining the predators response when testing birds in a group. It is likely that easy bleeding is a defence strategy directed primarily towards foraging insects with biting-chewing mandibles and that it is much less active towards predatory insects with piercing-sucking mandibles as well as birds. The involvement of chemical and/or physical cues in the strategy is discussed with respect to these types of predators.     

Burrowing behaviour inferred from feeding traces of the opheliid polychaete <Emphasis Type="Italic">Euzonus</Emphasis> sp. as response to beach morphodynamics

The distribution of the opheliid polychate Euzonus sp. and the orientation of its feeding traces were examined in a wave-dominated Pacific sandy beach of central Japan by periodical sampling from June to December 2006. This species occurred constantly within sediments at mid-intertidal level, although its horizontal distributions were variable depending on changes of the beach profiles. These facts indicate that Euzonus sp. moved horizontally seaward or landward within the substratum in response to the shifting beach face due to changes of wave conditions from the ocean. By analysing the orientation of feeding traces, the adaptive burrowing behaviour of this species in response to beach topographical changes was revealed: Euzonus worms burrowed horizontally to various directions in a relatively stable condition of beach topography, whereas they moved preferentially landward under a heavy erosion of the beach face by a storm. These findings indicate that the burrowing behaviour prevents Euzonus sp. from excessive burial and washing out due to the beach morphodynamics.     

Schooling behavior of Menidia medidia in the presence of the insecticide Sevin (Carbaryl)

A single application of 100 parts per billion Sevin in seawater caused disruption of schooling behavior of Menidia menidia in an artificial setting. Although the change, primarily a spreading out of the school over a larger area, was observed after 24 h exposure, placing the fish in clean water did not bring about a return of normal schooling patterns until 72 h. This is attributed to the accumulation of 1-naphthol, a toxic product of hydrolysis of Sevin. The adaptive significance of changes in schooling behavior is discussed.     

Sequestration of phorbolesters by the aposematically coloured bug Pachycoris klugii (Heteroptera: Scutelleridae) feeding on Jatropha curcas (Euphorbiaceae)

Summary. The phorbol ester DHPB has been detected in 5th instars and adults of Pachycoris klugii which feed on Jatropha curcas, a producer of phorbol esters with mollusc-, insect-, and vertebrate toxicity. DHPB from Pachycoris activates protein kinase C (PKC) which appears to be the main molecular target for phorbol esters. Phorbol esters of J. curcas exhibit a wide range of acute toxic effects in vertebrates and insects. It is therefore likely that the sequestration of DHPB, which would explain the aposematic colouration of the bugs, confers chemical protection to P. klugii against vertebrate predators. Received 26 April 2000; accepted 31 May 2000     

Costs of reproduction in lizards: escape tactics and susceptibility to predation

Summary Increased basking and reduced agility in gravid female southern water skinks (Eulamprus tympanum) suggest that they will be more vulnerable to predators. However, gravid females shift their anti-predator tactics towards crypsis, by allowing potential predators (such as a human observer) to approach more closely than do males and non-gravid females. Gravid females were taken no more frequently than were non-gravid females or males when exposed to two types of natural predators, birds (kookaburra, Dacelo gigas) or snakes (common blacksnakes, Pseudechis porphyriacus) in field enclosures. Both these results suggest that the vulnerability of potential prey in this system is determined by the predator's probability of detecting a potential prey item, not its probability of capturing the prey item after detection. Hence, laboratory-based measures of performance demonstrating reduced escape speed may sometimes have little relevance to actual fitness under field conditions, if the probability that an animal will be taken by a predator depends primarily on whether or not it is seen, rather than on how quickly it can escape. Correspondence to: L. Schwarzkopf