Hemolymph ion composition and volume changes in the supralittoral isopod Ligia pallasii Brandt, during molt

We analyzed ion composition and volume of the hemolymph of Ligia pallasii in four different stages of the molt cycle using capillary electrophoresis and ³H-inulin. The main ions in the hemolymph were Na⁺, K⁺, Mg²⁺, Ca²⁺, and Cl⁻. The Ca²⁺concentration increased significantly during the molt by 47% from intermolt to intramolt and by 37% from intermolt to postmolt, probably due to resorption of Ca²⁺ from the cuticle and sternal CaCO₃ deposits. The K⁺ concentration increased significantly by 20% during molt. The hemolymph volume normalized to the dry mass of the animals decreased by 36% from intermolt to late premolt. This was due to a reduction in the hemolymph volume and to an increase in dry mass of the animals during premolt. A sudden increase in the hemolymph volume occurring between late premolt and intramolt served to expand the cuticle. Since the Na⁺, K⁺, Mg²⁺, and Cl⁻ concentrations did not change significantly from late premolt to intramolt, the increase in hemolymph volume suggests an uptake of seawater rather than freshwater. Accepted: 7 March 2000     

Cold adaptation of the terrestrial isopod,Porcellio scaber,to subnivean environments

The terrestrial isopod, Porcellio scaber, was susceptible to subzero temperature: both freezing and chilling were injurious. The level of cold hardiness against chilling and freezing showed different patterns in their seasonal variation. The lower lethal temperature causing 50% mortality, an indicator of the tolerance to chilling, ranged from-1.37°C in August to-4.58°C in December. The whole body supercooling point, the absolute limit of freeze avoidance, was kept at about-7°C throughout the year. The winter decrease in lower lethal temperature was concomitant with an accumulation of low molecular weight carbohydrates which are possible protective reagents against chilling injury, whereas the less seasonally variable supercooling point seemed to be associated with the year-round presence of gut content. Food derivatives may act as efficient ice nucleators. The different trend in seasonal changes between lower lethal temperature and supercooling point may be related to the microclimate of the hibernacula in subnivean environments, where the winter temperature became lower than the lower lethal temperature in the summer active phase, but remained higher than the summer supercooling point.Abbreviations LLT₅₀ lower lethal temperature inducing 50% mortality - SCP supercooling point - T _a ambient air temperature - T _s soil surface temperature     

Why selection favors protandrous sex change for the parasitic isopod, Ichthyoxenus fushanensis (Isopoda: Cymothoidae)

A flesh burrowing parasitic isopod, Ichthyoxenus fushanensis, was found infecting the body cavity of a freshwater fish, Varicorhinus bacbatulus, in pairs. The marked sexual size dimorphism, with much larger females than males, and the presence of penes vestige on mature females suggest a protandrous sex change in I. fushanensis. Here we investigate the question of why selection favors protandrous sex change for I. fushanensis, by analyzing the interactions among clutch size, female size, male size, and their host size. The number of manca, the first free-living juvenile stage released, per brood was closely related to the size of the female. Excluding the effects of interaction among causal variables, the negative correlation of male size alone on clutch size suggests that a small male did not limit an individual's mating and fertilization success. When the effect of host size is removed statistically, there exists a significant negative relationship between the sizes of paired males and females. This indicates that the resources available from host fish are limited, and that competition exists between paired male and female resulting in a trade-off of body size. Due to the very low success rate of hunting for a host of mancas, a female with larger body size and higher fecundity has a fitness advantage. To augment the clutch size, a productive combination is a smaller male and a larger female in a host. The constraints of the limited resources and the trade-off between the sizes of paired male and female may favor I. fushanensis to adopt the reproductive strategy of protandrous sex change resulting in a larger female and hence more mancas. The pattern of the interactions among male, female, and the number of mancas, may be considered as a selective force for I. fushanensis protandrous sex change, where the available resources are constrained by the size of the host. This revised version was published online in July 2006 with corrections to the Cover Date.     

Colour choices of naive bumble bees and their implications for colour perception

The innate preferences of inexperienced bumble bees, Bombus terrestris, for floral colour stimuli were studied using artificial flowers. The artificial flowers provided a colour pattern and consisted of a star-shaped corolla and of central colour patches similar to the nectar guide of natural flowers. The innate choice behaviour was assessed in terms of the number of approach flights from some distance towards the artificial flowers and the percentage of approach flights terminating in antennal contact with the floral guide. The colours of the floral guide, the corolla and the background were varied. It was shown that the innate flower colour preference in bumble bees has two components. 1. The frequency of approaches from a distance is correlated with the colour difference between the corolla and the background against which it is presented. If the corolla colour was constant but its background colour varied, the relative attractiveness of the corolla increased with its colour difference to the background. The colour difference assessment underlying this behaviour on a perceptual basis can be attained by means of colour opponent coding, a system well-established in Hymenoptera. 2. The frequency of antennal contacts with the floral guides relative to that of approach flights cannot be accounted for by colour opponent coding alone. Whether the approach flights are interrupted, or whether they end in an antennal contact with the nectar guide is strongly dependent on the direction (sign) of the colour difference, not only its magnitude. The choice behaviour requires a unique perceptual dimension, possibly that of colour saturation or that of hue perception comparable to components of colour perception in humans.     

Predator avoidance behaviour in the buffy-headed marmoset,Callithrix flaviceps

The predator avoidance behaviour of a free-ranging group of buffy-headed marmosets,Callithrix flaviceps, was recorded in detail during the course of a long-term study of behavioural ecology at the Fazenda Montes Claros, southeastern Brazil. Four distinct patterns of predator avoidance behaviour, each with specific vocalisations, were recognised and are described here. The selection and use of sleeping sites by the study group are also described. An analysis of the records indicates that these small monkeys are generally most vulnerable to predation by aerial raptors. Variations in the frequency of alarm calls also indicate that the marmosets tend to be more vigilant at higher levels in the forest and when the leaf cover is less extensive. The implications of group size and social structure for both the evolution and the efficacy of the anti-predator behaviour of marmosets are also discussed.     

Acceleratory nervous regulation of juvenile myogenic hearts in the isopod crustacean Ligia exotica

We examined regulation of the myogenic heart by two identified cardioacceleratory neurons (CA1, CA2) in early juveniles of the isopod Ligia exotica. Repetitive stimulation of either the CA1 or CA2 axon increased the frequency and plateau amplitude of the action potential and decreased the maximum hyperpolarization of the cardiac muscle. These effects were larger with increasing stimulus frequency. The rate of increase in the frequency caused by CA1 stimulation was significantly larger than that by CA2. No impulse activity of the cardiac ganglion was induced by acceleratory nerve stimulation. The frequency of the muscle activity was decreased by injection of a hyperpolarizing current into the muscle during stimulation of the acceleratory nerve. In a quiescent heart, acceleratory nerve stimulation caused an overall depolarization in the muscle membrane and the amplitude of the depolarization induced by CA1 stimulation was significantly larger than that by CA2. These results suggest that CA1 and CA2 neurons regulate the myogenic heart affecting directly the cardiac muscle; the CA1 neuron produces more potent effects than does the CA2 neuron.     

Floral colour change in Pedicularis monbeigiana (Orobanchaceae)

We examined the effects of the retention of colour-changed flowers on long- and short-distance attractiveness of bumblebees and the likelihood of successive flower visits by bumblebees in Pedicularis monbeigiana. The lower lip changed colour with age from white to purple. Hand geitonogamous pollination significantly reduced seed production. No pollen limitation occurred in this species. Purple-phase flowers contributed minimally to pollinator attractiveness at long distance. The combination of less reproductive flowers with a lower amount of reward and floral colour change enabled plants to direct pollinators to reproductive, highly rewarding white flowers at close range. A high percentage of purple-phase flowers in an inflorescence was associated with a marked reduction in the frequency of successive flower visits to individual plants. We suggest floral colour change in P. monbeigiana may serve as a mechanism for enhancing inter-individual pollen transfer and reducing intra-individual pollen transfer.     

Turn the temperature to turquoise: cues for colour change in the male chameleon grasshopper (Kosciuscola tristis) (Orthoptera: Acrididae)

Rapid, reversible colour change is unusual in animals, but is a feature of male chameleon grasshoppers (Kosciuscola tristis). Understanding what triggers this colour change is paramount to developing hypotheses explaining its evolutionary significance. In a series of manipulative experiments the author quantified the effects of temperature, and time of day, as well as internal body temperature, on the colour of male K. tristis. The results suggest that male chameleon grasshoppers change colour primarily in response to temperature and that the rate of colour change varies considerably, with the change from black to turquoise occurring up to 10 times faster than the reverse. Body temperature changed quickly (within 10 min) in response to changes in ambient temperature, but colour change did not match this speed and thus colour is decoupled from internal temperature. This indicates that male colour change is driven primarily by ambient temperature but that their colour does not necessarily reflect current internal temperature. I propose several functional hypotheses for male colour change in K. tristis.     

The adaptive significance of ontogenetic colour change in a tropical python

Ontogenetic colour change is typically associated with changes in size, vulnerability or habitat, but assessment of its functional significance requires quantification of the colour signals from the receivers' perspective. The tropical python, Morelia viridis, is an ideal species to establish the functional significance of ontogenetic colour change. Neonates hatch either yellow or red and both the morphs change to green with age. Here, we show that colour change from red or yellow to green provides camouflage from visually oriented avian predators in the different habitats used by juveniles and adults. This reflects changes in foraging behaviour and vulnerability as individuals mature and provides a rare demonstration of the adaptive value of ontogenetic colour change.     

The role of omnivorous crayfish in littoral communities

Large omnivorous predators may play particularly important roles determining the structure of communities because of their broad diets and simultaneous effects on multiple trophic levels. From June 2001 to June 2002 we quantified community structure and ecosystem attributes of six newly establishing freshwater ponds (660 m² each) after populations of omnivorous crayfish (Orconectes virilis) were introduced to three of the ponds. Crayfish preyed heavily on fish eggs in this experiment, which reduced recruitment of young-of-year fish. This effect indirectly enhanced zooplankton biomass in crayfish ponds. Phytoplankton abundance exhibited a more complex pattern and was probably influenced by non-trophic (e.g., bioturbation) effects of crayfish. Peak dissolved oxygen levels were lower in the crayfish ponds indicating that they had lower primary production: respiration ratios. Metaphytic algae were strongly affected by crayfish presence; filamentous greens quickly disappeared and the blue-green Gleotrichia (a less preferred food item) eventually dominated the composition in crayfish ponds. Chara vulgaris and vascular macrophytes established 34% cover in control ponds by June 2002, but were not able to establish in crayfish ponds. Two important periphyton herbivores (tadpoles and gastropods) were absent or significantly reduced in the crayfish ponds, but periphyton differences were temporally variable and not easily explained by a simple trophic cascade (i.e., crayfish—snails and tadpoles—periphyton). Our results indicate that crayfish can have dramatic direct and indirect impacts on littoral pond communities via feeding links with multiple trophic levels (i.e., fish, invertebrates, and plants) and non-trophic activities (bioturbation). Although the effects of omnivorous crayfish on littoral communities can be large, their complex effects do not fit neatly into current theories about trophic interactions or freshwater community structure.     

An intra-species demonstration of the independence of distance and time in turn alternation of the terrestrial isopod, Porcellio scaber

In a study of turn alternation in the terrestrial isopod Porcellio scaber (woodlouse), smaller individuals (<11 mm long) took significantly longer to traverse a 60-mm open-ended post-forced-turn runway following a 90 degrees forced turn, than larger subjects (>11 mm long). However, there were no significant differences between the two groups in their probability of alternating, or in the magnitude of alternating turns as reflected in the size of the free-turn angles they turned on emerging from the runway. Nor was there a significant relationship for either group between the time taken to traverse the runway and either the probability of alternating or the size of a free-turn angle. In a second experiment, significant alternation occurred after woodlice emerged from runways that were 30 and 60 mm long, but not when they were 100 or 145 mm long. Their free-turn angles also became smaller as the runway length increased. Overall, the results of the three experiments supported the long-held view that, for reasons not yet understood, distance and time cannot necessarily be equated in the determination of invertebrate turn alternation.     

Infection characteristics of a trematode in an estuarine isopod: influence of substratum

The estuarine isopod Cyathura carinata is a second intermediate host to microphallid trematodes, which use mud snails Hydrobia spp. and shorebirds as respectively first intermediate and final hosts. To identify processes responsible for infection patterns observed in C. carinata, a short-term microcosm experiment was conducted with both macroinvertebrates and one of their common parasites – Maritrema subdolum. Fine sand collected from two different shallow water sites was used to test if sediment type could affect infection rates. After 7 days at 25 °C, C. carinata from the substratum with the highest proportion of particles <125 m were more surface active and obtained significantly more M. subdolum individuals than isopods from the other sediment type. No parasite-induced effects on the hosts were found during this short-term experiment. The distribution pattern of microphallid cysts and mesocercariae inside the isopods revealed that M. subdolum cercariae primarily penetrated through the pleopods and afterwards located themselves in the middle-posterior region of the hosts body. Even if it was not possible to identify the factor responsible for the observed infection patterns (cercariae production and/or host behaviour), the results of this experiment indicate that small-scale factors, such as differences in substratum and associated features, may have considerable impact on infections of host populations.     

Genetic and environmental variation in performance of a marine isopod: effects of eutrophication

Environmental variation in food resources modifies performance of herbivores, in addition to genetic variation and maternal effects. In marine benthic habitats, eutrophication may modify herbivores diets by changing host species composition or nutritional quality of algae for herbivores. We studied experimentally the effects of diet breadth and nutrient availability for the host algae on fitness components of the herbivorous isopod Idotea baltica. We fed the adult isopods with the brown algae Fucus vesiculosus and Pilayella littoralis and juveniles with the green alga Cladophora glomerata. By using half-sib families, we were able to separate the genetic, environmental and maternal effects on intermolt duration and size of the juveniles. The mothers confined to the diet consisting of both Fucus and Pilayella grew better and produced larger egg mass than those having consumed Fucus alone. Nutrient enhancement of algae did not influence the performance of the adult herbivores. However, the juveniles achieved twice the weight as well as shorter intermolt duration when consuming nutrient-treated C. glomerata. Mothers nutrition, either nutrient enrichment of her food algae or diet breadth, did not affect juvenile survival or growth as such, but we found evidence that the broader diet consumed by the mother mediated offspring performance by further accelerating growth rate of the offspring that fed on nutrient-treated alga. Intermolt duration was a highly heritable trait, but size showed very low heritability. Instead, maternal effects on size were substantial, suggesting that differences among mothers in their egg-provisioning ability strongly affect weight gain of their offspring. A high amount of additive genetic variance in intermolt duration implies potential for quick evolutionary responses in the growth schedule in the face of changes in the selective environment. We conclude that eutrophication, in addition to improving growth and reproduction of I. baltica by enhancing food quality and by providing opportunity for broader, more profitable diets, may act as a selective agent on life-history traits. Eutrophication of coastal waters is thus likely to reflect in herbivore density, population dynamics and eventually, grazing pressure of littoral macroalgae.     

Ecological mechanisms for coexistence of colour polymorphism in a coral-reef fish: an experimental evaluation

The evolution of different colour morphs and how they are maintained in animal populations is poorly understood. We investigated the mechanisms maintaining yellow and brown morphs of a coral-reef fish, Pseudochromis fuscus, at Lizard Island, on the Great Barrier Reef. Histological examination of the gonads revealed that colour morphs were not sex-limited, therefore sexual selection does not appear to promote dichromatism in this species. The field distributions of the two colour morphs were spatially segregated, limiting the opportunity for negative frequency-dependent selection to operate. Our results support another ecological mechanism of coexistence. The yellow morph occurred in deeper areas, usually close to the reef edge, where there was a proportionally high cover of live branching corals. In contrast, the brown morph occurred in shallower areas, more distant from the reef edge, that were proportionally low in live branching corals. Within these habitats, each colour morph of P. fuscus displayed a close association with similar coloured damselfishes from the genus Pomacentrus. The yellow morph was associated with predominantly yellow damselfishes (P. moluccensis and P. amboinensis) and the brown morph with darker coloured species (P. adelus and P. chrysurus). Multiple-choice experiments in the laboratory revealed that: (1) each colour morph of P. fuscus preferentially selected habitat patches occupied by damselfishes with the same colouration; and (2) differences in microhabitat use between the two colour morphs of P. fuscus were due to the presence of different coloured damselfishes in these microhabitats. P. fuscus is a predator of newly recruited damselfishes and the striking resemblance between each morph of P. fuscus and the damselfish with which it was associated, suggests that aggressive mimicry may promote coexistence of P. fuscus colour morphs.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Using colour spectral data in studies of geographic variation and taxonomy of birds: examples with two hummingbird genera, Anthracothorax and Eulampis

I propose a methodology to obtain and compare integral information on bird plumage coloration, using colour spectral data to conduct studies on geographic variation and taxonomy of different bird groups. I used principal component analysis and discriminant function analysis to compare groups of individuals by plumage coloration. As examples of the application of the methodology, I compared populations within the genus Eulampis and Anthracothorax. The results indicate possible taxonomic inadequacies and reveal situations that deserve further analysis, demonstrating the potential of the methodology in this area.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

The predatory behaviour of common kestrels facing various types of prey

Nine rehabilitated adult common kestrels (Falco tinnunculus) were tested for predatory behaviour. Tests were carried out individually on four different prey: two invertebrates (grasshopper and earthworm) and two vertebrates (lizard and laboratory mouse). The birds were offered prey randomly once daily, not necessarily on subsequent days. The latency to attack was similar. In contrast to predation tests with vertebrates, kestrels preying on invertebrates landed mostly (P<0.01 for earthworm and P<0.05 for grasshopper) a few centimetres from the prey itself, grasping it after a few steps. Earthworms were gripped almost invariably with the beak, but other prey with the toes. The birds struck the prey with their beak at highly variable rates (P<0.001): vertebrates mostly with the beak, while earthworms were never struck. The latency to ingest prey varied greatly (P<0.001) between prey. Captivity had only limited importance; it did not affect the behaviour sequence on any prey, but just induced kestrels in captivity for long time to attack mice with shorter latency and to strike them more with the beak. It is then demonstrated that the common kestrel has a strong behavioural plasticity even within the predatory sequence, which is then not stereotyped, and is able to modify its behaviour patterns in relation to different prey.An erratum to this article can be found at     

Biology and behaviour of reintroduced gibbons

We observed and recorded the behaviours of gibbons undergoing rehabilitation, before and after release, to document the behavioural and social changes of gibbons in the rehabilitation program and develop criteria for determining the suitability of a pair of gibbons for release. Hylobates albibarbis were observed at the Kalaweit Gibbon Rehabilitation Project in Kalimantan, Indonesia. Data were collected on animals both pre- and post-release and on wild gibbons for comparison. Data presented here show that reintroduced gibbons are capable of surviving without human intervention. In addition, their behaviour is similar to that of wild gibbons in terms of activity budgets, position in the canopy, body posture, pair association (PA) and diet. Prior to this study, no attempt has been made to quantify the rehabilitation process for gibbons, and rehabilitation project personnel require data reporting all aspects of a release so that improvements can be made. It is important to report these data for the benefit of future releases. Criteria, based on the behaviour of wild gibbons, are proposed to assist rehabilitation centers in assessing the suitability of gibbon pairs for release.     

Mate guarding behaviour in the supralittoral isopod,Ligia dentipes (Oniscidea) from the Andaman and Nicobar Islands

Precopulatory mate guarding is a characteristic feature in the mating behaviour of many Malacostraca, and a necessary prerequisite for those species in which female receptivity for males is restricted to a short period of time after the pubertal/reproductive moult. This study deals with the pre-mate guarding behaviour of the semi-terrestrial isopod Ligia dentipes living in the crevices of coral boulders and rocks in the supralittoral region of the Andaman Islands. As in other isopods, moulting in L. dentipes is biphasic, in which the posterior body part invariably moults first. The guarding male aids the female partner in the removal of the moulted exoskeleton. Mating occurs immediately after the posterior body exuviates. The male leaves the female after copulation and goes in search of another receptive female, demonstrating a polygamous mating system in these isopods. The mated females also re-mate with several other males without mate guarding. Females that had mated several times produced more young, compared to females mated only once in the laboratory. Female receptivity ceases following moulting of the anterior half. Intrasexual encounters among males lead to the large males acquiring receptive females. This study reveals interesting deviations from the general pattern of mate guarding already reported in other isopods and decapods. The evolutionary and ecological significances of mate guarding, intrasexual and intersexual conflicts, found in these semi-terrestrial isopods, are discussed.     

Observations with a different approach to morphological colour variation in an isopod population (Crustacea: Isopoda)

Morphological colour variation in Idotea baltica basteri Audouin, 1827 (Isopoda: Crustacea) is reported for the first time on populations from a brackish water lake (Kucukcekmece Lagoon) in Turkey. According to pigmentation characteristics, the isopods were described and sorted to ten different colour morphs which are named as maculata, maculata-lineata, uniformis, uniformis-lineata, immaculatum, immaculatum-lineata, nigrum, nigrum-lineata, albafusca, and albafusca-fasciatum. The similarity of the morphs was compared by defining digit codes to some of the visual pigmentation characteristics and calculating the Bray-Curtis similarity index of the colour morphs. The albafusca and albafusca-fasciatum forms were found to be the most similar to each other (similarity 96%), whereas immaculatum forms were considered the most different from maculata-lineata (similarity 50%). The morphs named maculata and maculata-lineata, having the highest abundance with 70%, belong to the most successful forms which may achieve crypsis by background matching in their living habitat in Kucukcekmece Lagoon which has sandy and rocky bottom surface area.     

The effect of change of background colour on the ultrastructure of the pars intermedia of the pituitary of the eel (Anguilla anguilla)

Summary Colour change in the eel resulted in marked ultrastructural changes in the pre-dominating (Type II) secretory cells of the pars intermedia of the pituitary. The effects on these cells of transferring eels from white to black backgrounds for periods of up to 56 days were: a) hypertrophy of the rough endoplasmic reticulum, which increased from 12 to 35% of the cytoplasmic volume; b) loss of secretory granules which decreased from 38 to 5% of the cytoplasmic volume; c) development of a system of fine (25–35 nm) tubules located especially at the secretory poles of the cells but also found in the region of the Golgi apparatus. The tubules were seen to connect with the plasma membrane, with the limiting membrane of the secretory granules, and in one instance to connect a granule with the plasma membrane. After glutaraldehyde fixation at pH 5, electron dense material similar to that found in the secretory granules was observed in the lumen of many of the tubules.The changes that occurred in black background eels are taken to indicate that the Type II cells of the pars intermedia are responsible for MSH secretion, particularly since these changes were reversed by returning eels to white backgrounds. The cytoplasmic tubules found in Type II cells may indicate a process for MSH release which does not involve granule extrusion, but rather direct transport of material from the Golgi apparatus to the cell membrane.The electron microscope facilities used in this investigation were funded by the Medical Research Council.