Effects of movement and eating on chemosensory tongue-flicking and on labial-licking in the leopard gecko (Eublepharis macularius)

Summary Two forms of lingual protrusion, tongueflicking and labial-licking, were differentially affected by combinations of movement and eating conditions in a eublepharid gecko (Eublepharis macularius). Tongue-flicking, in which the tongue contacts substrates beyond the lizard's body, occurred at increased rates during locomotion and during locomotion was significantly more frequent after eating than in a baseline condition. Labial-licking, in which a protruded portion of the tongue touches the labial, mental or rostral scales that surround the mouth, increased after eating. Unlike tongue-flick rates, by far the highest labial-lick rates were observed in stationary lizards after eating. The elevated tongue-flicking rates during movement after eating may be a manifestation of a postingestive chemosensory search for prey. In addition to grooming, several possible chemosensory functions of labial-licking are discussed, including gustatory sampling, sampling prey chemicals on the labials for transfer to the vomeronasal system, and redistribution of chemicals on the tongue to enhance transfer. It is suggested that labial-licking might help motionless lizards maintain vigilance for visual prey stimuli associated with the specific chemical prey cues. Another possible explanation for the increased labial-lick rate while motionless after eating is that prey chemicals induce tongue-flicking, but that the distance protruded is lessened and the tongue does not contact environmental substrates. Tongue-flicking while stationary is unlikely to lead to detection of additional prey and might incur detection by the lizard's predators or prey.     

Ontogenetic change in the radula of Conus magus (Gastropoda)

Specimens of the Indo-Pacific piscivorous gastropod Conus magus Linnaeus, 1758 were either collected by SCUBA in the field in Palau (1983–1984) or raised in the laboratory from egg capsules. C. magus captures adult fish with a specially modified radula tooth. Radula morphology was analyzed in a size range of individuals from 4.1 to 43.7 mm shell length, which encompassed early postmetamorphic juveniles to adults. Post-metamorphic C. magus juveniles are too small to consume fish, and all individuals below 9 mm possess a juvenile radula tooth totally different from that of the adult and resembling that of some vermivorous Conus species. The only food remains found in the digestive tracts of juveniles were the setae of syllid polychaetes. All individuals above 10.5 mm possessed adult teeth and had only fish remains in the gut. Two specimens, 10.1 and 9.2 mm in shell length, had intermediate-type radula teeth.     

Ontogenetic change in the diet ofAplodactylus punctatus (Pisces: Aplodactylidae): an ecophysiological explanation

Aplodactylus punctatus is a temperate berbivorous fish that changes from an omnivorous to a herbivorous diet and increases its ability ot assimilate algae as it grows. To investigate whether this dietary shift is related to size-specific differences in energetic demands imposed by metabolism and the amount of assimilated energy, oxygen consumption ( ) was determined experimentally in 12 specimens ranging in size from 62 to 545 g. increased allometrically with body size from 8.41 to 55.95 mg O₂ individual^-1 h^-1. Individual energetic requirements were 2.8 to 33.7 kJ d^-1. The assimilated energy was estimated, taking into consideration: (1) the energetic value of the most important alga in the diet (Lessonia trabeculata); (2) size-specific differences in assimilation rates for fish fed on this alga; (3) size-specific differences in throughput time and in the amount of food in a full gut. Comparison of the energy required and the assimilated energy revealed that fishes of < 22 to 29 cm total length had a negative energetic balance when consuming algae exclusively. This may explain the reliance of smallA. punctatus on more easily-digested invertebrates. The largest individuals can meet their energetic demands by consuming algae alone, apparently because of their higher assimilation capability. InA. punctatus, changing energetic requirements and capacities for algal assimilation may be responsible for the observed ontogenetic change in diet.     

On the evolution of nest raiding and male defensive behaviour in sticklebacks (Pisces: Gasterosteidae)

Summary We develop a model to explain the evolution of nest raiding by female sticklebacks based on enhanced female reproductive success derived by creating mating opportunities with the male whose nest has been destroyed. Our model depends critically on the shape of the function relating percentage hatch success to the number of eggs in a male's nest, and the probability that the female will later mate with the male whose nest was raided. Raiding by females will be favoured when the number of eggs in the nest is high providing high egg numbers reduces the percentage hatch rate. High probabilities of mating with the male in question also favour raiding. A second model suggests that males cannot reduce the probability of being raided by limiting the number of eggs in the nest. Instead selection should favour their increasing the number of eggs in the nest. Tests of these models are suggested and their relation to other factors which may influence the evolution of cannibalism are discussed.     

Offensive and defensive sperm competition roles in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

Sperm competition is predicted to generate opposing selection pressures on males. On one hand, selection should favour ‘defensive’ adaptations that protect a male’s ejaculate from displacement, while, on the other hand, selection should favour ‘offensive’ adaptations that overcome paternity assurance mechanisms of rivals. Here, we use the sterile male technique to assess sperm precedence when a male dung beetle Onthophagus taurus mates in both a defensive (first male) and an offensive (second male) role. Significant variation in a male’s sperm precedence (both P ₁ and P ₂) was detected, and an individual’s defensive (P ₁) and offensive (P ₂) abilities were positively correlated. Thus, it appears that sexual selection simultaneously selects for ‘defensive’ and ‘offensive’ adaptations in O. taurus. We discuss a variety of male traits in O. taurus that potentially contribute to a male’s ability to be successful when mating in an ‘offensive’ and a ‘defensive’ role.     

Ontogenetic habitat shift and risk of cannibalism in the common chameleon (Chamaeleo chamaeleon)

Ontogenetic habitat shifts have been documented in numerous fish and amphibians and in some reptiles. Intraspecific competition together with differential predation, prey size, social interactions, size-related thermal requirements, and morphological constraints on movement are often implicated in this ontogenetic habitat separation. In the current study, we combined field observation with experiments in seminatural arenas to test various hypotheses regarding the ontogenetic habitat shift that we have documented in the common chameleon. Juveniles (mean, 1 g) occupied low grasses and the adults (mean, 35 g) were found on bushes and trees. Overlap in habitat use between these two age classes was minimal. Our field experiments showed that juveniles actively avoid the presence of adults by concealment or flight. Adults readily attacked and consumed juveniles, regardless of their own mass. These results suggest that the risk of cannibalism towards juveniles is an important selective force behind the ontogenetic habitat shift observed in the common chameleon and may be important in other species too.     

On the biology of Berthellina citrina (Gastropoda: Opisthobranchia) and its defensive acid secretion

The opisthobranch gastropod Berthellina citrina (Ruppel and Leuckart, 1828¹) was collected from shallow water in the Gulf of Elat (Red Sea), where it is encountered only on sandy substrata overlaid with stones. The mollusc is nocturnal, feeds on various types of sponges, and hides during the day under stones. It releases an acid secretion (pH 1) containing sulphate and chlorine ions. This secretion is released, as a response to stimulation, from the entire body surface. In histological sections stained by standard dyes, the secretory cells remain colorless. In controlled laboratory experiments, it was found that this acid secretion protects B. citrina against all tested sea anemones, fishes, and crustaceans.     

Development of the escape response in larval walleye pollock (Theragra chalcogramma)

The development of the escape response of walleye pollock (Theragra chalcogramma) larvae from attacks by macrozooplanktonic and small-fish predators was quantified in laboratory experiments. Behavior was recorded using video cameras with silhouette illumination from infrared-emitting diodes and by visual observation. Laboratory-reared larvae of 1, 3, 8, 10, 12, 18, 22, 27, 42 days post-hatching, ranging in size from 4 mm to 10 mm total length, were used in the experiments. Even the youngest larvae were observed to exhibit a fast startle response. The percentage of successful larval escapes from the different predators increased as the larvae developed. Euphausiids (Thysanoessa raschii) and amphipods (Calliopiella pratti) often touched larvae but the larvae were usually able to escape and no successful captures of larvae over 22 days old were observed. Although successful escape from initial attacks by three-spine sticklebacks (Gasterosteus aculeatus) increased ontogenetically, sticklebacks were able to consume most larvae, even of the oldest age group, by repeated attacks. Day-old larvae had the lowest percent of escapes after encounters with jellyfish (Sarsia sp.), but the percentage of escapes increased dramatically for 3-day-old larvae. Escape speeds after an attack also increased with age, and tended to be higher after stickleback attacks and lower after jellyfish attacks. This study revealed that the escape response of larval pollock to attack by predators improves rapidly with development during the early larval stage.     

Chemical ecology and origin of defensive compounds in the Antarctic nudibranch Austrodoris kerguelenensis (Opisthobranchia: Gastropoda)

The common Antarctic nudibranch Austrodoris kerguelenensis (Bergh) contains diterpene diacylglycerides only present in its external body parts. These compounds provide a chemical defense against sympatric predators, such as the seastar Odontaster validus Koehler. Bioassays conducted with O. validus revealed that live nudibranchs, mantle tissue and Et₂O extract of the A. kerguelenensis mantle deterred feeding by the seastar. Further bioassays testing organic fractions of the Et₂O mantle extract showed that the diterpene diacylglycerides, as well as corresponding monoacylglycerides and monoacylglycerides of regular fatty acids, were responsible for the feeding deterrence in O. validus. We suggest that A. kerguelenensis derives the bioactive diacylglycerides by de novo biosynthesis rather than by sequestration from its sponge diet, since the mollusk does not contain active metabolites in the viscera, and neither the active compounds nor precursors were detected in the sponge diet. Furthermore, A. kerguelenensis did not show a strong chemodetection or feeding preference for its main diet, hexactinellid sponges, in Y-maze and food choice experiments, respectively.     

Locomotion versus spawning: escape responses during and after spawning in the scallop Argopecten purpuratus

The energetic cost of spawning and the endogenous factors that modulate spawning could modify escape response performance, leading to a conflict between the requirements of two fundamental components of fitness: reproduction and survival. We examined whether spawning changed force production during escape responses by the functionally hermaphroditic scallop, Argopecten purpuratus, and whether the response of smooth (tonic) and striated (phasic) muscles differed. Force production during escape responses by mature scallops was compared before induction of spawning, during spawning and after completion of spawning. Maximum tonic force and the area under the force curve (total force recorded) were diminished during gamete release, whereas phasic force production (maximum and mean force) increased after spawning was completed. The number and frequency of phasic contractions did not change during the spawning process, suggesting that spawning did not limit fuel availability for phasic contractions. The decrease in tonic force during spawning and the increased phasic force production after spawning may reflect changes in monoamine levels during gamete release. Whereas the spawning process modified force production during escape responses, the changes would, if anything, enhance escape performance during an initial encounter between a scallop and a predatory sea star.     

Mammalian breath: trigger of defensive chemical response in a tenebrionid beetle (Bolitotherus cornutus)

Summary The tenebrionid beetle Bolitotherus cornutus everts a pair of quinone-producing defensive glands in response to mammalian breath. Experiments with a controlled airstream indicate that the beetle recognizes breath on the basis of temperature, humidity, and airflow dynamics. Under attack by mice the beetle everts the glands immediately upon being mouthed and may secure its release as a result. Against ants the beetle is protected by its tough exoskeleton and usually refrains from everting the glands. Other arthropods also show defensive responses when breathed upon.Paper No. 77 of the series Defense Mechanisms of Arthropods     

Studies on the defensive behaviour ofHypselodoris species (Gastropoda: Nudibranchia): Ultrastructure and chemical analysis of mantle dermal formations (MDFs)

The mantle dermal formations (MDFs) ofHypselodoris webbi (D'orbigny, 1839), andH. cantabrica (Bouchet and Ortea, 1980) are globular sub-epidermal structures located in the cephalic and caudal regions. Histologically they consist of an accumulation of vacuolate cells surrounded by a basal lamina and an outer muscular capsule. Chemical analysis ofH. webbi MDFs reveals the presence of high concentrations of longifolin, a well known deterrent furanosesquiterpenoid that had been previously isolated from this species. In the present paper it is demonstrated that the great majority of longifolin accumulated in the mantle ofH. webbi is stored in the MDF vacuolar cells. This finding strongly suggests that such structures act as chemical weapons against predation, mainly protecting vital organs such as the head, rhinophores and gills.     

Antipredator activity and endogenous biosynthesis of defensive secretion in larval and pupal Delphastus catalinae (Horn) (Coleoptera: Coccinellidae)

Delphastus catalinae (Horn) is a predatory ladybird beetle (Coccinellidae) commonly used as a biocontrol agent against greenhouse infestation by whiteflies. It belongs to the basal subfamily Microweisinae, a group for which chemical defenses have not been previously investigated. The larval and pupal stages of D. catalinae possess minute secretory hairs that produce droplets containing compounds of both isoprenoid and polyketide origin. Bioassays with the predatory ant Crematogaster lineolata showed both the larval and pupal secretions to be deterrent. Moreover, isolated secretion components, from both classes of compounds, displayed antipredator activity against the ant. Experiments with D. catalinae larvae fed isotopically labeled glucose showed ¹³C-incorporation into both categories of compounds within the pupal secretion, demonstrating that these antipredator compounds, which differ from the typical nitrogenous defensive molecules of coccinellids, are biosynthesized endogenously. This suggests that the wide use of alkaloids by more derived coccinellids may have arisen after their divergence from the more basal Microweisinae.     

Old endemics and new invaders: alternative strategies of passerines for living in the Australian environment

Life history parameters of ”old endemics” and ”new invaders” among the Australian passerines were compared, controlling for phylogeny by using current, phylogenetic research methods. Old endemics were found to have significantly smaller clutches and demonstrated significantly more cooperative breeding than the new invaders, whose fledging periods are significantly longer than those of the old endemics. We argue that breeding under conditions of a small annual food increment during the breeding season, as is probably the situtation in Australia, is possible only when clutch size is small, or while breeding cooperatively, or by extending the fledging period. Our data show that the old endemics use the first two options, while the new invaders employ the third. Received: 10 February 1999 / Received in revised form: 10 November 1999 / Accepted: 20 November 1999     

The enantiomeric composition of monoterpene hydrocarbons in the defensive secretions ofNasutitermes termites (Isoptera): Inter- and intraspecific variations

Summary Frontal gland secretions of termite soldiers of the speciesNasutitermes costalis, N. ephratae, N. nigriceps, N. rippertii, andN. hubbardii were analyzed. The chiral compositions of the monoterpene hydrocarbons were investigated by means of two-dimensional gas chromatography. Two of them, 3-carene and-pinene were present in the form of (–)-enantiomers of high purity in the majority of the samples. The other monoterpenes,-pinene, camphene, sabinene, limonene, and-phellandrene, exhibited a great variation in the enantiopurity between different species.-Pinene and limonene varied significantly even between different colonies within the species.N. costalis was the only species with an invariable composition of the secretion, independent of the locality or origin.     

Ontogenetic variation in digestive proteinase activity of larvae and postlarvae of the pink shrimp Farfantepenaeus paulensis (Crustacea: Decapoda: Penaeidae)

Proteinase (endopeptidase), trypsin-like and chymotrypsin-like activities were examined throughout the ontogenetic development of cultured Farfantepenaeus paulensis. Whole individuals from different larval and postlarval stages, and the hepatopancreas of adults were homogenized and assayed to quantify the enzyme activities of specific substrates. Proteinase activity was identified by substrate-SDS-polyacrylamide gel electrophoresis. Specific inhibitors for trypsin (TLCK), chymotrypsin (TPCK) and serine proteinases (PMSF) were used to identify activity zones of these enzymes in gels. Protein-specific activity of total proteinases, trypsin and chymotrypsin was negligible at the egg stage and at Nauplius III, increasing in the first protozoeal substage (PZ I), and reaching a peak at PZ III; it decreased again in the subsequent postlarval substages. Different patterns of proteinase activity were observed in SDS-PAGE zymograms during ontogenetic development. Active bands of 14.6, 16.4, 17.5, 19.5, 22.5, 23.9, 25.8, 28.9, 32.0, 34.4, 37.7, and 42.2 kdaltons were detected in the adult hepatopancreas. Proteolytic activity was detected on gels in PZ I, and intense activity zones of 16.4, 17.5 and 19.5 kdaltons were found up to Mysis I (M I). Intense bands of 39.1 and 53.5 kdaltons were observed only at PZ III and M I. Band-activity intensity decreased after metamorphosis to the postlarval stage (PL). The chymotrypsin inhibitor TPCK had no effect on the proteinase bands. Active zones in gel inhibited with both TLCK and PMSF were considered to represent trypsin. The inhibitory effect of PMSF alone on proteinase extracts indicated chymotrypsin activity. TLCK and PMSF inhibition also varied during ontogenetic development. The inhibition of bands recorded between 14.6 and 21.7 kdaltons suggested the presence of low molecular weight trypsin in F. paulensis. The 39.1 kdaltons band observed at PZ III and M I were trypsin-like. On the other hand, bands of 28.9, 32 and 37.7 kdaltons from the adult hepatopancreas seem to represent a chymotrypsin. We conclude that the recorded variation in enzyme activity may be associated with morphological and behavioral changes during penaeid ontogenetic development. The higher enzyme activity at PZ II, PZ III and M I may reflect the increased energy turnover associated with intense swimming behavior and food ingestion. Received: 24 September 1998 / Accepted: 20 August 1999     

Quantitative composition of the defensive secretion ofBledius species (Coleoptera: Staphylinidae: Oxytelinae) is adapted to naturally occurring predators

Summary The adaptation of defensive secretions to their target organisms was examined for the abdominal gland secretions ofBledius furcatus, B. spectabilis andB. arenarius. Therefore the target organisms of the secretion of theseBledius species (i.e. their predators) had to be identified. At the collection sites examined these were the earwigLabidura riparia, the antCataglyphis bicolor, the flyLispe candicans, different carabids of the generaPogonus, Dichirotrichus, Dyschirius, Bembidion andCalathus and the wading birdsHaematopus ostralegus andCalidris alba. The secretion of the abdominal glands contains the toxin ptoluquinone dissolved in either-dodecalactone and 1-undecene (B. furcatus andB. spectabilis) or in octanoic acid and octyloctanoate (B. arenarius). The ratio of these solvents is species-specific. Application experiments using some of the natural insect predators (L. riparia, C. bicolor, Pogonus, Di. gustavii, Dyschirius) revealed that these solvent ratios provided a more effective deterrent than other possible ratios. Thus by combining the solvents in certain ratios, the capability of cuticular penetration and therefore the effectiveness of the defensive secretions are adapted to their natural targets.     

Ontogenetic variation in metabolism, biochemical composition and energy content during the early life stages of Farfantepenaeus paulensis (Crustacea: Decapoda: Penaeidae)

Dry weight (DW), oxygen consumption, ammonia-N excretion, proximate biochemical composition (total protein, carbohydrate, lipid, water and ash), and energy content (estimated from biochemical composition and by wet combustion) were determined in early developmental stages of cultured Farfantepenaeus paulensis. Pooled samples from embryonic, larval and postlarval stages (at 26 ± 1 °C and 34 ± 1‰) were used for measurements. The study focused on physiological and biochemical processes during transitional periods of ontogeny, such as hatching, lecithotrophic and planktotrophic stages, metamorphosis, and the attainment of a benthic existence in postlarva. DW showed higher increment between protozoea I (PZ I) and mysis I (M I) than in the next mysid and postlarval stages. Individual rates of oxygen consumption and ammonia-N excretion increased, while weight-specific rates presented significant reduction throughout development. Higher weight-specific oxygen consumption was registered in nauplius III (N III) and PZ I, following a decrease in subsequent stages. Postlarval stages PL V–VI and PL X–XII exhibited the lowest values among the stages studied. Weight-specific excretion was high in N III and protozoeal stages, with maximum values in PZ II, while the following stages were marked by lower rates. O:N ratios indicated higher protein catabolism in the stages between egg and M I and a shift to more lipid utilization close to metamorphosis. Water content was higher in the protozoeal stages and decreased afterwards. Higher percentages of protein, lipid and carbohydrate (%DW) were observed in egg and nauplius stages. Protein and lipid decreased from the egg through the naupliar and protozoeal stages, rising again in mysis stages. Lipid content (%DW) decreased in PL V–VI and PL X–XII. Lipid:protein ratios showed an increase of the importance of lipid between PZ III and M II. Carbohydrates represented a minor fraction of body composition, and ash percentages increased from egg to a maximum in PZ II, decreasing in subsequent stages. Energy content determined by wet combustion or calculated by energy equivalents presented the same trend throughout development, varying similarly to protein. Protein was the main energy contributor to body energy in all stages, while the importance of lipid was higher in egg and early naupliar stages. Trends observed in metabolic rates and body composition may be associated to morphological and behavioral changes during the early stages of penaeid development, such as the transition from herbivory to omnivory, and the adoption of a benthic existence. Different ontogenetic energy strategies contribute to succeed through such diverse type of development. Received: 4 July 2000 / Accepted: 6 December 2000