The functionality of female reciprocal calls in the Iberian midwife toad (<Emphasis Type="Italic">Alytes cisternasii</Emphasis>): female-female acoustic competition?

Female midwife toads (genus Alytes) emit highly variable reciprocal calls of unclear function prior to and during courtship. In some species, female-female competition, expressed as physical fighting, has been reported. Males of Majorcan midwife toads (Alytes muletensis) show phonotactic response to female calls, and females of Iberian midwife toads (Alytes cisternasii) respond differently according to the male call characteristics. In this study, I test the hypothesis of female-female acoustic competition as an additional function of female reciprocal calls. Playback tests indicate that female calls are not clearly involved in female acoustic competition in the Iberian midwife toad, therefore female calls could be directed at males rather than towards competitive females.     

Vocal acrobatics in a Chinese frog, <Emphasis Type="Italic">Amolops tormotus</Emphasis>

Although amphibians are highly vocal, they generally emit only a limited number of acoustic communication signals. We report here the extraordinarily rich vocal repertoire of Amolops tormotus, a ranid species in China. These frogs produce countless vocalizations, some of which share features of birdsong or primate calls, e.g., ultrasonic frequency components, multiple upward and downward FM sweeps, and sudden onset and offset of selective harmonic components within a call note. Frame-by-frame video analysis of the frog's calling behavior suggests the presence of two pairs of vocal sacs that may contribute to the remarkable call-note complexity in this species. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00114-002-0335-x.     

Where to find a mate? Resource-based sexual communication of webbing clothes moth

Mate location in moths typically entails long-range attraction of males to female-produced pheromone. Here, we show that male and female webbing clothes moths, Tineola bisselliella, seek larval habitats (dry carrion, animal lairs, etc) to encounter mates. With males seeking, and arriving at, larval habitat earlier at night than females, male-produced pheromonal and sonic signals enhance the habitat's attractiveness to females. This resource-based mating strategy of T. bisselliella differs from that known for most other moths. It may have evolved in response to larval habitats that are patchy and temporary, but that disseminate attractive semiochemicals so abundantly that T. bisselliella encounter them more readily than their own pheromones.     

Phylogeny of cooperatively breeding cuckoos (Cuculidae,Crotophaginae) based on mitochondrial gene sequences

The Crotophaginae is a subfamily of New World cuckoos comprising the monotypic genus Guira and three ani species (Crotophaga). All exhibit a rare form of cooperative breeding known as plural female joint-nesting, whereby two or more females lay eggs in a single nest. I reconstructed the phylogeny of Crotophaginae using the mitochondrial genes cytochrome oxidase I, II, and III, ATPase 6 and 8, and cytochrome b. The subfamily was monophyletic, implying a single origin of cooperative breeding in New World cuckoos. Crotophaga was also monophyletic with Guira as its sister taxon. Within Crotophaga, the smooth-billed (C. ani) and groove-billed (C. sulcirostris) anis formed the internal clade with the greater ani (C. major) basal to this pair. This phylogeny is consistent with differences in reproductive patterns and social organization exhibited by crotophagine cuckoos, and will serve as a framework for future study of the evolution of cooperative breeding in this subfamily.     

Phonotaxis during walking and flight: are differences in selectivity due to predation pressure?

Female selectivity was tested in Tettigonia viridissima during two different phonotaxis situations; compensated walking and tethered flight. For two of the three temporal parameters that are important for call recognition in T. viridissima, selectivity was similar in the two situations. Selectivity for the third parameter (minimum interval duration between the double pulses) was much higher during walking than during flight: walking females responded only to stimuli with intervals of 28 ms or longer, while call models with intervals of 18 ms were attractive during flight. One interneuron (TN-1) is probably involved in filtering the minimum interval duration. As this neuron is also the most likely candidate for transmitting bat calls during flight, it is suggested that the selectivity differences between walking and flying might be due to the need for detecting predator signals during flight, when TN-1 would be occupied listening for bats. With TN-1 unavailable for song processing during flight, temporal selectivity for the minimum interval duration should be reduced, as was found here.     

The use of olfaction in the foraging behaviour of the golden-mantled flying fox,<Emphasis Type="Italic"> Pteropus pumilus</Emphasis>, and the greater musky fruit bat,<Emphasis Type="Italic"> Ptenochirus jagori</Emphasis> (Megachiroptera: Pteropodidae)

Double-choice experiments with three adult males of the little golden-mantled flying fox, Pteropus pumilus, and ten adult greater musky fruit bats, Ptenochirus jagori (both Megachiroptera: Pteropodidae), demonstrate that they are able to discriminate accurately between an empty dish and a dish containing fruits of one of several species by odour alone. Tests were run using fruits of six fruit species for Pteropus pumilus and five fruit species for Ptenochirus jagori. The fruit species used are known to be consumed in the wild by Ptenochirus jagori and are, with two exceptions, species of the natural rain-forest habitat. This is the first study to show that fruit bats are also able to assess the ripeness of a fruit exclusively by its odour. The bats preferred ripe over unripe fruits of the same species. Thus, both Pteropus pumilus and Ptenochirus jagori can not only locate fruits by their odour but can also discriminate between ripe and unripe fruits of the same species by olfaction. The results confirm and expand earlier findings on the role of olfactory cues in the orientation of foraging pteropodids.     

Delayed onset of vocal recognition in Australian sea lion pups (Neophoca cinerea)

In pinnipeds, maternal care strategies and colony density may influence a species’ individual recognition system. We examined the onset of vocal recognition of mothers by Australian sea lion pups (Neophoca cinerea). At 2 months of age, pups responded significantly more to the calls of their own mothers than alien female calls demonstrating a finely tuned recognition system. However, newborn pups did not respond differentially to the calls of their mother from alien female calls suggesting that vocal recognition had not yet developed or is not yet expressed. These findings are in stark contrast to other otariid species where pups learn their mother’s voice before their first separation. Variance in colony density, pup movements, and natal site fidelity may have reduced selective pressures on call recognition in young sea lions, or alternatively, another sensory system may be used for recognition in the early stage of life.     

A pear-derived kairomone with pheromonal potency that attracts male and female codling moth, Cydia pomonella (L.)

Ethyl (2E, 4Z)-2,4-decadienoate, a pear-derived volatile, is a species-specific, durable, and highly potent attractant to the codling moth (CM), Cydia pomonella (L.), a serious pest of walnuts, apples, and pears worldwide. This kairomone attracts both CM males and virgin and mated females. It is highly attractive to CM in both walnut and apple orchard contexts, but has shown limited effectiveness in a pear orchard context. Rubber septa lures loaded with ethyl (2E, 4Z)-2,4-decadienoate remained attractive for several months under field conditions. At the same low microgram load rates on septa, the combined gender capture of CM in kairomone-baited traps was similar to the capture rate of males in traps baited with codlemone, the major sex pheromone component. The particular attribute of attracting CM females renders this kairomone a novel tool for monitoring population flight and mating-ovipositional status, and potentially a major new weapon for directly controlling CM populations.     

Unusually high-pitched neonate distress calls of the open-habitat Mongolian gazelle (<Emphasis Type="Italic">Procapra gutturosa</Emphasis>) and their anatomical and hormonal predictors

In neonate ruminants, the acoustic structure of vocalizations may depend on sex, vocal anatomy, hormonal profiles and body mass and on environmental factors. In neonate wild-living Mongolian gazelles Procapra gutturosa, hand-captured during biomedical monitoring in the Daurian steppes at the Russian-Mongolian border, we spectrographically analysed distress calls and measured body mass of 22 individuals (6 males, 16 females). For 20 (5 male, 15 female) of these individuals, serum testosterone levels were also analysed. In addition, we measured relevant dimensions of the vocal apparatus (larynx, vocal folds, vocal tract) in one stillborn male Mongolian gazelle specimen. Neonate distress calls of either sex were high in maximum fundamental frequency (800–900 Hz), but the beginning and minimum fundamental frequencies were significantly lower in males than in females. Body mass was larger in males than in females. The levels of serum testosterone were marginally higher in males. No correlations were found between either body mass or serum testosterone values and any acoustic variable for males and females analysed together or separately. We discuss that the high-frequency calls of neonate Mongolian gazelles are more typical for closed-habitat neonate ruminants, whereas other open-habitat neonate ruminants (goitred gazelle Gazella subgutturosa, saiga antelope Saiga tatarica and reindeer Rangifer tarandus) produce low-frequency (<200 Hz) distress calls. Proximate cause for the high fundamental frequency of distress calls of neonate Mongolian gazelles is their very short, atypical vocal folds (4 mm) compared to the 7-mm vocal folds of neonate goitred gazelles, producing distress calls as low as 120 Hz.     

A "spare" compensates for the risk of destruction of the elongated penis of earwigs (Insecta: Dermaptera)

Male animals in several groups have multiple intromittent organs that outnumber the corresponding female gonopore. In Dermaptera (earwigs), males of the family Anisolabididae have paired, elongated male intromittent organs (virgae), while females have a single sperm-storage organ (spermatheca). Several authors have assumed that one of the paired virgae is non-functional, because it points in the "wrong" direction. We investigated the mating success of handicapped males of Euborellia plebeja in which one of their paired virgae was removed experimentally. These handicapped males succeeded in inseminating a mate. Males with genital damage are found in the field, suggesting that the "spare" functions under natural conditions. Based on phylogenetic information on earwigs, we discuss possible evolutionary scenarios for this genital peculiarity.     

A novel sex-specific and inducible monoterpene synthase activity associated with a pine bark beetle,the pine engraver,<Emphasis Type="Italic"> Ips pini</Emphasis>

Ecological interactions of conifers and coniferophagous bark beetles are determined in part by terpenoids (isoprenoids), which are major defense metabolites of conifer oleoresin. Curiously, similar compounds are important aggregation pheromones of conifer-attacking bark beetles. Terpene synthases are responsible for generating the enormous variety of terpenoid carbon skeletons found in nature. These catalysts convert short-chain prenyl diphosphates to a diverse assortment of hemiterpene, monoterpene, sesquiterpene, and diterpene natural products. While terpene synthases have frequently been characterized from plant and microbial sources, they have not yet been described in animals. Here we report the discovery of a monoterpene synthase activity in an insect, the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae). Cell-free assays of I. pini revealed that geranyl diphosphate (GDP) is converted to the acyclic monoterpene myrcene in whole-body extracts from males, but not females. Furthermore, the monoterpene synthase activity in males can be induced by prior treatment with juvenile hormone III (JH III) or by feeding on phloem from the host trees, Jeffrey pine (Pinus jeffreyi Grev. &; Balf.) or red pine (Pinus resinosa Ait.). The sex-specificity and endocrine induction of this activity argue for its involvement in the biosynthesis of monoterpenoid pheromones mediated by enzymes from insect tissue. This discovery is the first example of a monoterpene synthase in the Metazoa and evokes exciting new questions about the origin, evolution, and occurrence of terpene synthases.     

Alcoholism in cockchafers: orientation of male Melolontha melolontha towards green leaf alcohols

Chemical orientation of the European cockchafer, Melolontha melolontha L., a serious pest in agriculture and horticulture, was investigated by field tests and electrophysiological experiments using plant volatiles. In total, 16 typical plant volatiles were shown to elicit electrophysiological responses in male cockchafers. Funnel trap field bioassays revealed that green leaf alcohols (i.e. (Z)-3-hexen-1-ol, (E)-2-hexen-1-ol and 1-hexanol) attracted males, whereas the corresponding aldehydes and acetates were behaviourally inactive. Furthermore, male cockchafers were attracted by volatiles from mechanically damaged leaves of Fagus sylvatica L., Quercus robur L. and Carpinus betulus L. However, volatiles emitted by damaged leaves of F. sylvatica attracted significantly more males than those from the other host plants. Odour from intact F. sylvatica leaves was not attractive to M. melolontha males. Females were not attracted by any of the tested volatile sources. The results suggest that plant volatiles play a similar role as a sexual kairomone in mate finding of M. melolontha, as has been shown for the forest cockchafer, Melolontha hippocastani F. Nevertheless, both species show remarkable differences in their reaction to green leaf alcohols.     

Split-second escape decisions in blue tits (Parus caeruleus)

Bird mortality is heavily affected by birds of prey. Under attack, take-off is crucial for survival and even minor mistakes in initial escape response can have devastating consequences. Birds may respond differently depending on the character of the predator's attack and these split-second decisions were studied using a model merlin (Falco columbarius) that attacked feeding blue tits (Parus caeruleus) from two different attack angles in two different speeds. When attacked from a low attack angle they took off more steeply than when attacked from a high angle. This is the first study to show that escape behaviour also depends on predator attack speed. The blue tits responded to a high-speed attack by dodging sideways more often than when attacked at a low speed. Escape speed was not significantly affected by the different treatments. Although they have only a split-second before escaping an attack, blue tits do adjust their escape strategy to the prevailing attack conditions.     

A secondary copulatory structure in a female insect: a clasp for a nuptial meal?

Secondary copulatory structures are well-known in male dragonflies and spiders. Here I report a secondary copulatory organ in female ground weta, Hemiandrus pallitarsis (Ensifera, Orthoptera - crickets and allies). The organ, located on the underside of the abdomen, appears to secure the male's genitalia during the transfer of a spermatophylax nuptial meal to this location, an area quite separate from the female's primary copulatory structures, where the sperm ampulla is attached.     

Monogamy in a new species of wild guinea pigs (Galea sp.)

A new species of wild guinea pigs from Bolivia was recently discovered: Galea sp. So far no data are available concerning its social and mating systems. We therefore investigated behavioural compatibility, sexual dimorphism, paternity in a male-competing situation, testicular and epididymal size, and sperm morphology and motility. The results showed a complete intrasexual and moderate intersexual behavioural incompatibility between unfamiliar animals, a reversed sexual dimorphism with females larger than males, and litters always fathered by single males. Furthermore, both the reproductive organ weights and the percentage of motile sperm and sperm with intact acrosomes were significantly lower compared with the promiscuous congeneric Galea musteloides. In summary, these data strongly suggest a monogamous social and mating system for the recently discovered Galea sp.     

Feminization of genetic males by a symbiotic bacterium in a butterfly,Eurema hecabe (Lepidoptera: Pieridae)

Wolbachia are symbiotic bacteria found in many arthropods and filarian nematodes. They often manipulate the reproduction of host arthropods. In the present study, female-biased sex-ratio distortion in the butterfly Eurema hecabe was investigated. Breeding experiments showed that this distorted sex ratio is maternally inherited. When treated with tetracycline, adult females of the thelygenic line produced male progeny only. After PCR using Wolbachia-specific primers for the ftsZ gene a positive result was seen in the thelygenic females, but not in male progeny from tetracycline-treated females, or individuals from a Tokyo population with normal sex ratio and reproduction. Cytological observations showed that thelygenic females lack the sex chromatin body (W chromosome). The results strongly suggest that the sex-ratio distortion in E. hecabe is due to feminization of genetic males by Wolbachia.     

Cross-cultural and cross-ecotype production of a killer whale ‘excitement’ call suggests universality

Facial and vocal expressions of emotion have been found in a number of social mammal species and are thought to have evolved to aid social communication. There has been much debate about whether such signals are culturally inherited or are truly biologically innate. Evidence for the innateness of such signals can come from cross-cultural studies. Previous studies have identified a vocalisation (the V4 or ‘excitement’ call) associated with high arousal behaviours in a population of killer whales in British Columbia, Canada. In this study, we compared recordings from three different socially and reproductively isolated ecotypes of killer whales, including five vocal clans of one ecotype, each clan having discrete culturally transmitted vocal traditions. The V4 call was found in recordings of each ecotype and each vocal clan. Nine independent observers reproduced our classification of the V4 call from each population with high inter-observer agreement. Our results suggest the V4 call may be universal in Pacific killer whale populations and that transmission of this call is independent of cultural tradition or ecotype. We argue that such universality is more consistent with an innate vocalisation than one acquired through social learning and may be linked to its apparent function of motivational expression.     

The structure of stereotyped calls reflects kinship and social affiliation in resident killer whales (Orcinus orca)

A few species of mammals produce group-specific vocalisations that are passed on by learning, but the function of learned vocal variation remains poorly understood. Resident killer whales live in stable matrilineal groups with repertoires of seven to 17 stereotyped call types. Some types are shared among matrilines, but their structure typically shows matriline-specific differences. Our objective was to analyse calls of nine killer whale matrilines in British Columbia to test whether call similarity primarily reflects social or genetic relationships. Recordings were made in 1985–1995 in the presence of focal matrilines that were either alone or with groups with non-overlapping repertoires. We used neural network discrimination performance to measure the similarity of call types produced by different matrilines and determined matriline association rates from 757 encounters with one or more focal matrilines. Relatedness was measured by comparing variation at 11 microsatellite loci for the oldest female in each group. Call similarity was positively correlated with association rates for two of the three call types analysed. Similarity of the N4 call type was also correlated with matriarch relatedness. No relationship between relatedness and association frequency was detected. These results show that call structure reflects relatedness and social affiliation, but not because related groups spend more time together. Instead, call structure appears to play a role in kin recognition and shapes the association behaviour of killer whale groups. Our results therefore support the hypothesis that increasing social complexity plays a role in the evolution of learned vocalisations in some mammalian species.     

Response of red deer stags (Cervus elaphus) to playback of harsh versus common roars

Red deer stags (Cervus elaphus) give two distinct types of roars during the breeding season, the “common roar” and the “harsh roar.” Harsh roars are more frequent during contexts of intense competition, and characterized by a set of features that increase their perceptual salience, suggesting that they signal heightened arousal. While common roars have been shown to encode size information and mediate both male competition and female choice, to our knowledge, the specific function of harsh roars during male competition has not yet been studied. Here, we investigate the hypothesis that the specific structure of male harsh roars signals high arousal to competitors. We contrast the behavioral responses of free ranging, harem-holding stags to the playback of harsh roars from an unfamiliar competitor with their response to the playback of common roars from the same animal. We show that males react less strongly to sequences of harsh roars than to sequences of common roars, possibly because they are reluctant to escalate conflicts with highly motivated and threatening unfamiliar males in the absence of visual information. While future work should investigate the response of stags to harsh roars from familiar opponents, our observations remain consistent with the hypothesis that harsh roars may signal motivation during male competition, and illustrate how intrasexual selection can contribute to the diversification of male vocal signals.     

A colony-level response to disease control in a leaf-cutting ant

Parasites and pathogens often impose significant costs on their hosts. This is particularly true for social organisms, where the genetic structure of groups and the accumulation of contaminated waste facilitate disease transmission. In response, hosts have evolved many mechanisms of defence against parasites. Here we present evidence that Atta colombica, a leaf-cutting ant, may combat Escovopsis, a dangerous parasite of Atta's garden fungus, through a colony-level behavioural response. In A. colombica, garden waste is removed from within the colony and transported to the midden - an external waste dump - where it is processed by a group of midden workers. We found that colonies infected with Escovopsis have higher numbers of workers on the midden, where Escovopsis is deposited. Further, midden workers are highly effective in dispersing newly deposited waste away from the dumping site. Thus, the colony-level task allocation strategies of the Atta superorganism may change in response to the threat of disease to a third, essential party.