Alarm signals in Daphnia?

Daphnia magna can respond to chemical cues from freshly crushed conspecifics with various behavioural reactions. A shift in vertical distribution towards the bottom, the formation of aggregations and direct escape responses can all be induced by water-borne signals released from crushed Daphnia. The pattern and strength of the first two behavioural responses (i.e. the persistent tendency to occupy deeper strata in the experimental columns and to stay within patches) indicate that Daphnia perceive the signal from crushed conspecifics as nonspecific information, not necessarily associated with any particular kind of danger from either vertebrate or invertebrate predators. The adaptive value and possible costs associated with performing these two behavioural reactions are discussed. The adaptive value of the induced escape response was directly tested: Daphnia which had experienced the presence of a cue from crushed conspecifics avoided attacks by common bream more efficiently than naive Daphnia. The recognition of the signal originating from crushed conspecifics can be especially adaptive in encounters with unfamiliar predators and with predators that undergo ontogenetic shifts in their diet. Under natural conditions, the combination of such a signal with a predator cue can, very reliably, advertise the local scale of the predatory impact. Received: 10 October 1996 / Accepted: 20 May 1997     

Too close for comfort: spatial patterns in acorn barnacle populations

Spatial patterns in aggregations form as a result of the interplay between costs and benefits experienced by individuals. Such self-organisation of aggregations can be explained using a zonal model in which a short-range zone of repulsion and longer-range zone of attraction surrounding individuals leads to emergent pattern properties. The signal of these processes can be detected using spatial pattern analyses. Furthermore, in sessile organisms, post-settlement mortality reveals the relative costs and benefits of positions within the aggregation. Acorn barnacles are known to require contact with conspecifics for reproduction and are therefore believed to aggregate for this purpose; isolated individuals may also be more susceptible to abiotic stress and predation. At short distances, however, competition for space and resources is likely to occur. In this study spatial patterns of barnacles (Semibalanus balanoides L.) were analysed using pair-correlation functions. Individuals were dispersed at distances below 0.30 cm, but peak relative density occurred at a distance of 0.36 cm from conspecifics. This is much closer than required for reproductive access, implying a strong aggregative drive, up to the point of physical contact with neighbours. Nevertheless, analysis of dead barnacles illustrated that such proximity carries a cost as barnacles with many neighbours were more likely to have died. The inferences obtained from these patterns are that barnacles aggregate as closely as they can, and that local neighbourhood competition is a powerful determinant of mortality. These processes give rise to the observed pattern properties.     

A first insight into the scanning behaviour of the presocial blow fly larvae

Aggregation of necrophagous larvae has several benefits: the sharing of salivary enzymes (exodigestion), temperature regulation, protection from predators and parasites, etc., and is well developed in blow flies (Diptera: Calliphoridae). The present study focuses on the aggregation mechanism used by the necrophagous larvae of Lucilia sericata Meigen, the common green bottle fly. The ability of single larva to detect and follow a signal (trail) created by conspecifics is investigated initially. A circular ring is drawn in a Petri dish where 20 starved larvae have crawled for a period of 30 min. A naïve (test) larva is then placed in the dish and video‐tracked. Naïve larvae are able to detect the boundaries of the larvae‐crawled area and stay preferentially within this conspecific‐marked zone. In a second step, the orientation of larvae by scanning, a dedicated, ground‐signal detection behaviour of dipteran larvae, is analyzed. Four experimental conditions are tested: control, presence of food, conspecifics, and food + conspecifics. When conspecifics have been previously present in a given area, the scanning behaviour by naïve larvae in this area decreases (both in number and frequency of scans). Accordingly, scanning by necrophagous larvae of L. sericata should be considered not only as locomotion behaviour, but also as a potential way to detect signals from conspecifics for the purpose of aggregation. The chemical composition of the attractant(s), the behavioural effects (attraction or retention) and the implication of larval signalling in the aggregation process are new fields to explore.     

Space Partitioning within Populations of Sea Anemones (Genus: Anthopleura) in the California Rocky Intertidal Zone

The spatial patterns of sea anemones (genus: Anthopleura) were analyzed off the rocky intertidal of California. A. elegantissima partitions space in a regular or even pattern and interferes with conspecifics. Its associate A. xanthogrammica shows no signs of interference with conspecifics or congenerics and displays a clumped spatial distribution.     

Early Learning and the Recognition of Conspecific Cocoons by Carpenter Ants (Camponotus spp.)

Groups of newly-eclosed workers of two carpenter ant species (Camponotus floridanus and C. tortuganus) were reared in the presence of conspecific cocoons, cocoons of the other species, or were kept without cocoons. Groups of older workers (> 20 days), previously exposed to conspecific brood in their natal nest, were familiarized with either conspecific or heterospecific cocoons. After 14 days of exposure, groups were subjected to short-term (5 min) and long-term (10 day) preference tests. Young and older workers retrieved and retained many cocoons of both species, familiar and unfamiliar. However, a pattern of non-exclusive discrimination emerged: (1) Young workers exposed to conspecific cocoons picked up and retrieved conspecifics before unfamiliar heterospecifics, and retained conspecifics longer. (2) Young workers generally required experience with conspecifics to develop this preference, as those exposed to only heterospecifics and those deprived of cocoons were impartial in short-term tests. However, in long-term tests (5–10 days), naive young workers significantly preferred unfamiliar conspecifics. (3) Older workers preferred conspecific cocoons, whether familiar or unfamiliar, in short-term tests, but their tolerance for heterospecifics in the longer term (5 days) could be increased by recent familiarization. In no case did young or older ants significantly prefer familiar heterospecifics to conspecifics. These results confirm a role for early learning in brood recognition by carpenter ants, but suggest that it is less important than in Formica species studied by previous authors.     

Invasive species of crayfish use a broader range of predation-risk cues than native species

The responses of invasive and native species of crayfish to conspecific and heterospecific alarm odors were recorded in the laboratory. Individuals of the North American invasive Procambarus clarkii responded just as strongly to odors from crushed Austropotomobius pallipes as they did to crushed conspecifics. The North American invasive Orconectes limosus also responded as strongly to P. clarkii odor as to conspecific odor. The native Italian species A. pallipes responded more strongly to conspecific alarm than to heterospecific alarm from P. clarkii. The pattern of invasive species of crayfish using a broader range of danger signals than displaced native species appears to be robust.     

Male tree weta are attracted to cuticular scent cues but do not discriminate according to sex or among two closely related species

Recognition of conspecifics is an essential precursor of successful mating. Where related species coexist, species discrimination might be important, but because related species are similar, species signal recognition may actually be low. Chemical cues such as cuticular hydrocarbons (CHCs) are frequently used by insects to identify suitable sexual partners. We predicted that New Zealand tree weta (Hemideina spp.), a genus of nocturnal ensiferan Orthoptera that live both allopatrically and sympatrically, use chemical signals from either frass or CHCs to find mates. In a series of six laboratory trials using both H. thoracica and H. crassidens, we found that male tree weta, but not female tree weta, occupied cavities primed with female cuticular cues more often than cavities without. However, males did not discriminate between chemical cues of male and female conspecifics, or between conspecifics and heterospecifics. In field trials, tree weta did not occupy artificial cavities primed with either female frass or female cuticular cues more often than unscented cavities. However, in both trials weta preferentially returned to cavities that had already been occupied earlier in the trials. A final field trial confirmed the presence of mixed species harems during the mating season in one region of sympatry. Our results suggest that selection on sex and species specific chemical cues that could be used to find conspecific mates is weak. Mixed species aggregations suggest that identification of conspecific mating cues has not evolved to be species specific. We infer that for male tree weta, the cost of mating with heterospecifics is likely less than not mating at all.     

Effect of predation risk on grouping pattern and whistling behaviour in a wild mouflonOvis aries population

We studied the effect of predation risk on grouping pattern and whistling behaviour in a free-ranging mouflonOvis aries Linnaeus, 1758 population in Sardinia. Direct observations were carried out from July 2005 to June 2007 (n _groups=881,n _mouflon=3477). In our study site, the rut occurred in October and November, when social sexual segregation disappeared, while lambing peaked in April. Groups with lambs (mean ± SE: female groups with lambs 4.98 ± 0.23, mixed groups 6.49 ± 0.29) were larger than male groups (2.01 ± 0.10) and female groups without lambs (2.77 ± 0.11), especially during the lambing season. This reflected the anti-predator tactics adopted by mothers so as to benefit from the dilution effect. Also male aggregations increased in size during the lambing season, as a consequence of the gradual decrease of rutting activities and consequently of male-male aggressiveness. Among the anti-predator tactics adopted by mouflon, whistling behaviour seemed to be a warning signal directed to predators and not to conspecifics. This was because whistling was shown by smaller aggregations only, regardless of group type, presence of lamb and habitat occupied. Smaller groups cannot profit from the dilution effect and therefore may be more motivated to signal the predator that it has been spotted.     

Firebrats,Thermobia domestica,aggregate in response to the microbes Enterobacter cloacae and Mycotypha microspora

The firebrat, Thermobia domestica (Packard) (Thysanura: Lepismatidae), aggregates in response to the faeces of conspecifics as well as shelters previously inhabited by conspecifics. Our objective was to determine the source of the aggregation signal. Filter paper previously exposed to firebrats induced strong arrestment of firebrats. Polar solvents (water, methanol, acetonitrile) and less polar solvents (hexane, dichloromethane, ethyl ether), alone or in combination, failed to extract the aggregation signal from firebrat‐exposed paper. Moreover, solvent‐extracted paper continued to induce firebrat arrestment. In contrast, the aggregation signal could be obtained by physical extraction (freeze/thawing or ultrasonication) of firebrat‐exposed paper submerged in water. Five fungal species and four bacterial species were isolated from ultrasonicant solutions on potato dextrose‐, nutrient‐, and GlcNAc‐agar. Of the nine isolated microbes tested, only the fungus Mycotypha microspora Fenner (Mucorales) and the bacterium Enterobacter cloacae (Jordan) Hormaeche & Edwards (Enterobacteriaceae) induced arrestment of firebrats in bioassays. Our data support the conclusion that firebrats do not form aggregations in response to pheromones; instead, they aggregate in the presence of specific microbes or their metabolites.     

Cannibalism of parasitoid‐attacked conspecifics in a non‐carnivorous caterpillar

Cannibalism, the killing and consumption of conspecifics, can even occur in insect species typically considered to be non‐carnivorous. Of particular interest is the cannibalism of parasitoid‐attacked conspecifics, which could reduce parasitism levels in subsequent generations for that conspecific population. This study reports on the occurrence and some of the consequences of cannibalism in parasitoid‐attacked obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae). We show that larvae of C. rosaceana, which is considered to be an herbivorous caterpillar species, did not prey upon live conspecifics, but readily consumed conspecifics attacked by Habrobracon gelechiae Ashmead (Hymenoptera: Braconidae). Further examination found that C. rosaceana larvae feeding on parasitoid‐attacked conspecifics, since their fourth instar, suffered a higher mortality and reduction in body size than those fed on plant material only. The cannibalism of attacked conspecifics did not appear to offer any nutrient benefits for the cannibal. To our best knowledge, this is the first empirical example of the occurrence and some of the consequences of cannibalism by a non‐carnivorous insect on its parasitoid‐attacked conspecifics. We discuss the adaptive significance of such cannibalism on parasitoid‐attacked conspecifics with respect to a trans‐generational fitness gain for the population through the killing of the parasitoids, thereby reducing parasitism in subsequent generations.     

Do Golden Egg Bugs (Phyllomorpha laciniata: Heteroptera, Coreidae) Require Conspecifics for Oviposition?

Golden egg bug (Phyllomorpha laciniata) females lay eggs on the bodies of conspecifics of both sexes. We investigated to what extent reproduction depended on the availability of conspecifics as oviposition substrate and the acceptability of the host plant as an alternative oviposition substrate in the absence of conspecifics. Mated females were placed in experimental enclosures each containing a sprig of fresh host plant. Each experimental female was subjected to one of three treatments: isolated from conspecifics (solitary), paired with another female, or paired with a male. Solitary females laid a few eggs on the host plant but then stopped laying eggs, and solitary females laid significantly fewer eggs than those enclosed with another female or a male. Females enclosed with a male laid no more eggs than those enclosed with a female. When two previously isolated females were later enclosed together, they soon renewed oviposition. Females in nature contained significantly more oviducal eggs than did females that were enclosed with other females for a short period. Thus the availability of suitable conspecifics as oviposition substrate stimulates the deposition of mature eggs, and reproduction depends on the presence of conspecifics of either sex as oviposition substrate.     

Detection of the Sexual Identity of Conspecifics through Volatile Chemical Signals in a Territorial Salamander

Territorial red‐backed salamanders (Plethodon cinereus) have been shown to use nonvolatile chemical signals in both territorial defense and to convey a variety of information to conspecifics. We investigated whether or not red‐backed salamanders could determine the sexual identity of conspecifics through volatile chemical signals, and we explored their use in the context of territorial defense. We exposed male and female red‐backed salamanders to four experimental treatments (i.e. filter papers that had been scent marked by male or female conspecifics for 1 and 5 d) and two control treatments (i.e. unscented filter papers for 1 and 5 d tests). The focal salamanders were prevented from physically accessing the scent marked filter papers and, presumably, some of the substrate scent marks had volatile components that were detected and interpreted by the focal salamanders. Both male and female red‐backed salamanders spent significantly more time in threat displays when they were exposed to volatile chemical signals from same‐sex conspecifics than they did toward similar signals from opposite‐sex conspecifics. A similar statistical pattern was observed for the amount of chemosensory sampling exhibited by focal red‐backed salamanders. From these results, we infer that red‐backed salamanders can determine the sexual identity of conspecifics through volatile chemical signals, some of which may be used in territorial defense. Further, such airborne pheromones may influence the spatial organization of salamander territories on the forest floor.     

Do damselfly larvae recognize and differentially respond to distinct categories of macroinvertebrates?

Zygopteran larvae normally encounter other aquatic macroinvertebrates that are predators, competitors, and prey and should therefore demonstrate varied responses when faced with different categories of opponent. In a laboratory experiment individual final-instarIschnura posita (Hagen) larvae were observed in interactions with six categories of invertebrate opponents. The opponent categories were a nonconspecific damselfly and a small crayfish, which represented threatening opponents because they were larger than theI. posita subject larva, and a mayfly and a stonefly, which constituted nonthreatening opponents because they were smaller than the subject larva. The levels of threat posed by conspecific larvae of final and penultimate instar were inferred by comparison to the other opponent categories. Multivariate analysis showedI. posita's response differed between the two larger opponents, but responses were statistically indistinguishable between the two smaller opponents. Larvae retreated, moved around the stalk, and struck their opponents with their lamellae more often in the presence of a crayfish than the nonconspecific zygopteran. In contrast, they assumed an S-bend posture frequently with the zygopteran. Responses toward final-instar conspecifics differed from responses toward the larger opponents. Effectively, larvae wagged their abdomens only in the presence of final-instar conspecifics and retreated and moved around the stalk less frequently in these trials. Responses toward the smaller conspecifics differed from the responses to the small opponents. Larvae struck penultimate-instar conspecifics with their lamellae more frequently than the other small opponents. Our results suggest that larval zygopteran behaviors (such as S-bend and SCS) that have previously been described as intraspecific displays are of a more general nature and used toward a variety of opponents, whereas wag is unique to intraspecific interactions inI. posita.     

Local enhancement of arrivals of Ceratitis capitata females on fruit mimics

Mature female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), isolated from conspecifics for 7 days prior to testing in laboratory cages, were attracted to yellow plastic fruit-mimicking hemispheres or spheres in significantly greater numbers when groups of conspecific females were present than absent on fruit mimics. Attraction of females to conspecifics occurred for females deprived of protein or continuously provided with protein since eclosion, for fruit mimics that were unbaited or baited with odor of food or fruit, and under single-choice as well as multiple-choice test conditions. Attraction proved just as great to fruit mimics having dead conspecifics as to mimics having live conspecifics, was evident also when fruit mimics were orange or white, and occurred also in response to presence of live Bactrocera dorsalis (Hendel)females on fruit mimics. We discuss our findings in relation to local enhancement, a form of social facilitation in insects and other animals.     

Pursuit of Heterospecific Targets by Territorial Amberwing Dragonflies (Perithemis tenera Say): A Case of Mistaken Identity

Although they are defending mating territories, territory residents of a wide variety of insect species have been observed to pursue heterospecifics in addition to the conspecifics that intrude on their territories. One species that has such heterospecific pursuits is the Eastern amberwing dragonfly (Perithemis tenera Say) (Anisoptera: Libellulidae). In this study, we tested five alternative hypotheses for the function of heterospecific pursuits in amberwings: competition for resources, prevention of interference while mating, predator deterrence, foraging, and mistaken identity. Resident males pursued both male and female conspecifics, as well as a species of horse fly (Tabanus spp.) and butterfly (Ancyloxypha numitor). Other intruding odonates, including Epitheca princeps, Erythemis simplicicollis, Libellula luctuosa, Pachydiplax longipennis, and Plathemis lydia, were relatively ignored. Because the horse fly and butterfly were similar to amberwings in body size, color, and flight height, and because they are not predators or prey of amberwings, we concluded that the pursuit of these heterospecifics was due to mistaken identity. The characteristics of the horsefly and butterfly likely correspond to the cues that the male amberwings use to identify conspecifics, and the relative rarity of intrusions by these two species (as well as by female amberwings) probably made it more costly to discriminate and pursue only conspecifics than to make some mistaken pursuits.     

Pathogens as a factor limiting the spread of cannibalism in tiger salamanders

Summary Intraspecific predation is taxonomically widespread, but few species routinely prey on conspecifics. This is surprising as conspecifics could be a valuable resource for animals limited by food. A potential cost of cannibalism that has been largely unexplored is that it may enhance the risk of acquiring debilitating pathogens or toxins from conspecifics. We examined how pathogens affect variation in the incidence of cannibalism in tiger salamander larvae (Ambystoma tigrinum nebulosum), which occur as two environmentally-induced morphs, typicals and cannibals. Salamanders from one population were more likely than those in another to develop into cannibals, even when reared under identical conditions. Variation in the propensity to become a cannibal may be caused by variation in pathogen density. In the population with cannibals at low frequency, bacterial blooms in late summer correlated with massive die-offs of salamanders. The frequency of cannibals correlated significantly negatively with bacterial density in ten different natural lakes. In the laboratory, cannibals exposed to a diseased conspecific always preyed on the sick animal. As a result, cannibals wre more likely to acquire and die from disease than were typicals that were similarly exposed, or cannibals that were exposed to healthy conspecifics. Since conspecifics often share lethal pathogens, enhanced risk of disease may explain why cannibalism is generally infrequent. Pathogens may constrain not only the tendency to be behaviorally cannibalistic, but also the propensity to develop specialized cannibal morphologies.     

Distress call-induced gene expression in the brain of the Indian short-nosed fruit bat, Cynopterus sphinx

Individuals in distress emit audible vocalizations to either warn or inform conspecifics. The Indian short-nosed fruit bat, Cynopterus sphinx, emits distress calls soon after becoming entangled in mist nets, which appear to attract conspecifics. Phase I of these distress calls is longer and louder, and includes a secondary peak, compared to phase II. Activity-dependent expression of egr-1 was examined in free-ranging C. sphinx following the emissions and responses to a distress call. We found that the level of expression of egr-1 was higher in bats that emitted a distress call, in adults that responded, and in pups than in silent bats. Up-regulated cDNA was amplified to identify the target gene (TOE1) of the protein Egr-1. The observed expression pattern Toe1 was similar to that of egr-1. These findings suggest that the neuronal activity related to recognition of a distress call and an auditory feedback mechanism induces the expression of Egr-1. Co-expression of egr-1 with Toe1 may play a role in initial triggering of the genetic mechanism that could be involved in the consolidation or stabilization of distress call memories.     

On the genetic diversity of spiny mice (genus Acomys) and gerbils (genus Gerbillus) in the Arabian Peninsula

Using non-destructive sampling we provide further genetic characterisations for spiny mice (Acomys dimidiatus/cahirinus) and gerbils (Gerbillus sp.) in three regions in Saudi Arabia. All individuals were sequenced for a fragment of the cytochrome b gene, and compared against available conspecifics and closely related taxa. We confirm the existence of a second Acomys dimidiatus/cahirinus lineage specific to the Arabian Peninsula as seen previously. The Arabian Gerbillus nanus is shown to group with Middle Eastern rather than African conspecifics. A second cryptic Gerbillus lineage was also sampled across multiple locations, which may be an uncharacterised G. dasyurus.     

Phylogenetic conservatism in plant‐soil feedback and its implications for plant abundance

We examined whether plant‐soil feedback and plant‐field abundance were phylogenetically conserved. For 57 co‐occurring native and exotic plant species from an old field in Canada, we collected a data set on the effects of three soil biota treatments on plant growth: net whole‐soil feedback (combined effects of mutualists and antagonists), feedback with arbuscular mycorrhizal fungi (AMF) collected from soils of conspecific plants, and feedback with Glomus etunicatum, a dominant mycorrhizal fungus. We found phylogenetic signal in both net whole‐soil feedback and feedback with AMF of conspecifics; conservatism was especially strong among native plants but absent among exotics. The abundance of plants in the field was also conserved, a pattern underlain by shared plant responses to soil biota. We conclude that soil biota influence the abundance of close plant relatives in nature.     

Use of olfaction for sexual recognition in the subterranean rodent<Emphasis Type="Italic">Ctenomys talarum</Emphasis>

The ability of the tuco-tucoCtenomys talarum Thomas, 1898 to recognize sex by olfactory cues contained in urine, faeces and soiled shavings was tested by using preference tests. Nonbreeding tuco-tucos selected odours from opposite-sex rather than same-sex conspecifics. This pattern differed between sexes: females spent more time sniffing male than female odours for all scent sources whereas males did not show any difference in the time they spent investigating odours of each sex for each tested odour sources. Dissimilarities in odour selections between sexes may be attributed to a different combination of factors involved in olfactory interest for each sex. The function of gender cues recognition is discussed.