Cues for Mate Recognition and the Effect of Prior Experience on Mate Recognition in Enallagma Damselflies

In many coenagrionid damselflies, sexually mature females exhibit color polymorphisms, with some females resembling conspecific males. Although it has been suggested that the latter function as male mimics, this does not appear to be the case for those in the genus Enallagma. We found that sexually dimorphic coloration of the female abdomen and thorax are important cues for sexual recognition by males. We demonstrate for the first time in the Odonata, that males learn to recognize andromorphs as potential mates. After 2 days in an enclosure, sexually mature males exposed to only andromorphic females initiated more sexual interactions with tethered andromorphs than with heteromorphs, the majority morph in the natural population. Exposure to only heteromorphic females tended to decrease males' sexual responses to andromorphs, but not significantly so. Because the frequency of female morphs often varies within a population, learned mate recognition would be advantageous for males that search for mates. Our results lead to a novel, frequency-dependent hypothesis for the occurrence and maintenance of female-limited color polymorphisms.     

Age-Specific Behavior and Habitat Selection of Adult Male Damselflies, Calopteryx maculata (Odonata: Calopterygidae)

We compared the age, movement, and time–activity budgets of male Calopteryx maculata damselflies occupying off-stream tree-fall gaps with those at stream sites within a 10-ha woodland. All males collected at off-stream sites were younger than males collected at stream sites—as indicated by their significantly higher wing transmittance. Thirty-three percent of teneral males marked at off-stream gaps moved to stream sites within 4 days (mean distance = 140 m), while mature males marked at stream sites never left the stream. In contrast to stream site males, off-stream males spent significantly more time capturing prey and never engaged in aggressive interactions with other males. Behavioral differences were not due to variations in the operative body temperature. However, malaise trapping revealed a greater frequency of suitable prey in forest light gaps. Our findings support the idea that teneral male Calopteryx leave their emergence sites along the stream for off-stream light gaps to forage without interference and build the energy reserves necessary to attain and hold streamside territories.     

Oviposition habitat selection by the mosquito, Culiseta longiareolata: effects of conspecifics, food and green toad tadpoles

Abstract. 1. Ovipositing females should choose egg-laying sites which maximize the growth and survivorship of their offspring. Accordingly, they should avoid habitats with high densities of competitors and predators and should choose sites with adequate food. We tested experimentally the oviposition response of the mosquito, Culiseta longiareolata Macquart, to extra food and two species which are both potential competitors and predators of its offspring: conspecific larvae and green toad ( Bufo viridis ) tadpoles.
2. We conducted the experiment in outdoor artificial pools where wild C. longiareolata females were presented with choices of all combinations of the presence or absence of: (1) C.longiareolata larvae, (2) B.viridis tadpoles, and (3) extra food (a mixture of ground fish food and mouse chow).
3. Oviposition rates averaged approximately 4 times greater in response to extra food.
4. The presence of the tadpoles, particularly later in the experiment when they were larger, caused a significant reduction (approximately 50%) in oviposition rate.
5. We did not detect a statistically significant effect of conspecific larvae on oviposition habitat selection.
6. Our results indicate that Culiseta females choose habitats for oviposition based on food availability. They also tend to avoid habitats containing Bufo tadpoles, presumably to avoid predation and/or competition.     

Aggressive Interactions and Risk of Fish Predation for Larval Damselflies

Larval damselflies frequently engage in aggressive interactions that may increase their risk of fish predation. To test this we analyzed the behavior of larval Ischnura verticalis exposed to both conspecifics and fish predators. Larvae in the presence of conspecifics oriented, struck, and swam more but crawled less compared to solitary larvae; the presence of fish reduced, or tended to reduce, all behaviors. Fish struck more at interacting larvae compared to noninteracting larvae. Increased attack rate by fish likely reflects the increase in the very active swimming behavior by larvae and suggests a conflict between antipredator behaviors. Swimming is an appropriate response to avoid predation by odonate larvae which normally ambush prey but is clearly dangerous when fast-swimming fish that cue in on movement are nearby.     

Experimental evidence for predation risk sensitive oviposition by a mosquito, Culiseta longiareolata

1. Females should choose to oviposit in habitats where risk of predation and competition are low. The ovipositional responses of a mosquito, Culiseta longiareolata, to a predator and to species sharing the same trophic level as this mosquito (controphic species) were assessed experimentally in outdoor artificial pools. 2. The predator, larval Anax imperator, which strongly reduced larval C. longiareolata survival, resulted in a 52% reduction of C. longiareolata egg rafts. The controphic species (primarily Daphnia magna), which had a small but statistically significant negative effect on the survival of C. longiareolata larvae, did not have a statistically significant influence on the number of egg rafts. 3. Laboratory trials indicated that only a small fraction of the reduced number of egg rafts seen in predator pools may be due to consumption of the egg rafts by A. imperator. 4. The experimental evidence indicates that the reduced number of C. longiareolata egg rafts found in the presence of A. imperator is due largely to oviposition habitat selection, i.e. C. longiareolata females choose pools with low risk of predation for their offspring.     

Use of winter habitat by roe deer at a northern latitude where Eurasian lynx are present

Winter climate at northern latitudes is a challenge to small-bodied ungulates, and they modify behaviour to save energy and to increase the likelihood of survival. Also, the ongoing expansion of large carnivores in several European countries can lead to the recovery of (potentially energetically costly) anti-predator behaviours. In an area recently recolonized by Eurasian lynx Lynx lynx , we snow-tracked radio-collared roe deer Capreolus capreolus in order to investigate their bedding and feeding behaviour during winter, and assess how environmental factors affect their habitat use. We also tested the prediction that roe deer use more open sites than locally available in areas with a stalking predator such as the lynx. Our results showed that both bed sites and foraging sites had more cover, compared with random sites. Most of the variation in canopy cover and in the distance and foraging sites between bed sites and foraging sites was explained by prevailing weather. As the winter progressed, the presumed depletion of fat reserves promoted the use of more canopy cover at foraging sites by night, less by day and a decrease in the distance between beds, foraging sites and human activities. Males used artificial feeding sites less often and bedded further from humans than females. The data fit the hypothesis of tighter energy budgets for family groups (females with fawns) or that males are more cautious towards humans. There was no support for the hypothesis that roe deer used more open habitat than locally available in order to reduce their vulnerability to lynx predation. Owing to severe winter conditions and the danger of starvation, roe deer seem to be forced to accept a high risk when predators are present, not changing their main pattern of habitat use from comparative areas where predators are absent.     

Oviposition in three species of limnephiloid caddisflies (Trichoptera): hierarchical influences on site selection

1. Oviposition site selection was studied in three trichopteran species; an undescribed species of Hydatophylax (Limnephilidae), Onocosmoecus unicolor (Limnephilidae) and Neophylax rickeri (Uenoidae), in two Coastal Range streams in California, U.S.A. 2. Hydatophylax sp. egg masses were generally found at or above the water surface on substrata near the bank in pools, where undercut banks, overhanging vegetation and rocks shaded attachment sites and provided wind protection. Onocosmoecus unicolor females deposited their egg masses exclusively above the water on moist emergent wood. The egg masses of N. rickeri were almost exclusively found in high velocity areas of riffles, where females oviposited under water and attached their egg masses to the submersed undersides of unembedded, protruding stones with large emergent surfaces. 3. For Hydatophylax sp. and N. rickeri, a hierarchical selection scheme is proposed in which females use cues at three different spatial levels (stream, stream subunit, substratum) to choose oviposition sites. Females of O. unicolor seemed to choose at only two different levels of spatial resolution (stream, substratum). 4. The formation of large aggregations of egg masses in N. rickeri and O. unicolor suggests that females actively choose oviposition sites where conspecific egg masses are already attached. The clustering of egg masses may minimise the risk of predation through the dilution effect, because egg masses of N. rickeri and O. unicolor are both attacked by dipteran predators. 5. Females of the three Trichoptera species studied are able to make a clear choice of oviposition site in the heterogeneous stream environment. In comparison with other stream microhabitats, these are characterised by stable and relatively predictable environmental conditions during the time of egg development, which is seen as the main selective pressure leading to the observed egg‐laying behaviour.     

Substrate selection for pit making and oviposition in an antlion, Myrmeleon bore Tjeder, in terms of sand particle size

The larvae of the pit‐making antlion Myrmeleon bore Tjeder live in open sand in riverbeds with a substratum consisting of various particle sizes. We analyzed the spatial distribution of their pits in a sandy floodplain to determine their larval and adult responses to the heterogeneous substrate. The spatial distribution pattern of their pits had an aggregated distribution, and there was a significant positive correlation between pit density and the ratio of medium‐size sand particles to total weight of sand. We examined the size of sand particles selected in the larval pit‐building behavior and the oviposition behavior of the adult. Both larvae and adults selected medium‐size sand particles. The larvae of M. bore are relatively sedentary predators and rarely move great distances. Thus, the present results suggest that habitat selection by adult females is a major factor causing the aggregative distribution of the pits.     

Schwankungen im Spektrum von Kriebelmückenarten und räumliche Konstanz der Eiablageplätze in einem Niederungsbach (Simuliidae,Diptera). Variation in Species Composition of Black Flies (Simuliidae,Diptera) and Spatial Stability of Oviposition Sites in a Lowland Stream

The species composition of black flies (Simuliidae, Diptera) in a lowland stream of Aller-Leineregion (northern Germany, FRG) is analysed in the course of several years. Normally, Odagmia ornata (MG .) is the dominating species. For a short sharply defined period Boophthora erythrocephala DE GEER appeared in a great number. Possible reasons and related research on spatial stability of oviposition sites are presented.     

The distribution patterns of dragonflies (Insecta: Odonata) along the Kiewa River,Australia, and their relevance in conservation assessment

Sampling of larval and adult Odonata from 16 sites along the Kiewa River, Victoria, yielded 34 species: 10 Zygoptera, 24 Anisoptera. Patterns of larval and adult incidence were appraised, and showed that most species were restricted in incidence to several consecutive sites along the river, and that there is clear distinction also between the faunas of the potamon, rhithron and eucrenon regions. Different species of some genera of Anisoptera displayed different zonational distributions, and patterns of incidence and relative abundance of larvae and adults confirmed zonational occupancy. For larvae, these distribution patterns transcended the mode of collection, although many species were found most abundantly in one microhabitat or by one of several sampling methods employed at each site. Sampling of the two stages separately showed considerable concurrence of distributional patterns, so that either stage alone may provide data of value in faunal and conservation assessment. This revised version was published online in July 2006 with corrections to the Cover Date.     

Microhabitat preferences of longear sunfish: low light intensity versus submerged cover

Both cover and light intensity are important factors in microhabitat selection of fishes, suggesting a possible hierarchical relationship. Many centrarchids have been shown to prefer low light intensities and heavy submerged cover, but adult nonbreeding longear sunfish are reported to occupy sparsely vegetated, clear, shallow water during the brightest part of the day, probably for foraging. To determine the relative importance of cover and light intensity to longear sunfish, we presented nonbreeding adults with choices between these two conditions in a series of three laboratory experiments. Longear sunfish preferred low light to high light intensity and preferred submerged cover to no cover. However, when given a choice, they preferred to occupy low light intensity conditions without cover rather than submerged cover under higher light intensity conditions. Thus, low light intensity, such as is found in greater pool depths, may be the preferred refuge from predation for longear sunfish.     

Differential Habitat Selection by Pygmy Grasshopper Color Morphs; Interactive Effects of Temperature and Predator Avoidance

Habitat selection behavior is affected by complex interplays between competing requirements. Here we combine field observations with laboratory experiments to examine how thermal benefits and predator avoidance influences habitat selection by different color morphs of the pygmy grasshopper Tetrix undulata. The composition of substrate types and surface temperatures in areas selected by free-ranging individuals did not reflect relative availability, and varied among morphs and sexes. Surface temperatures of selected habitats deviated less from the range of preferred body temperatures than would result from a random utilization of surface temperatures, suggesting that grasshoppers selected habitats with thermal properties which were suitable for maintaining preferred body temperatures. The thermal property of habitats occupied by different color morphs suggests that darker morphs (which absorb more solar radiation) selected cooler habitats to avoid overheating. Dissimilarities in substrate use among color morphs in the field and laboratory emphasize a role also of predator avoidance by background matching for habitat choice. The degree of habitat selectivity was lowest in the striped morph, supporting the notion that a disruptive color pattern may constitute a solution to the trade-off between relative crypsis in different visual backgrounds. Finally, individuals modified their habitat use when subjected to elevated risk of predation, showing that habitat choice is governed by conflicting priorities. Collectively, our findings suggest that, as a result of direct and indirect effects of coloration on performance, alternative color morphs use different solutions to the trade-off between competing requirements. Our results also lend support to the notion that relative fitness of alternative color morphs and sexes may be dependent on microhabitat selection, as predicted by the theory of multiple niche polymorphisms.     

Detection and Avoidance of Predators in White-Tailed Deer (Odocoileus virginianus) and Mule Deer (O. hemionus)

In this paper, we investigate the relationship between early detection of predators and predator avoidance in white-tailed deer ( Odocoileus virginianus ) and mule deer ( O. hemionus ), two closely related species that differ in their habitat preferences and in their anti-predator behavior. We used observations of coyotes ( Canis latrans ) hunting deer to test whether the distance at which white-tails and mule deer alerted to coyotes was related to their vulnerability to predation. Coyote encounters with both species were more likely to escalate when deer alerted at shorter distances. However, coyote encounters with mule deer progressed further than encounters with white-tails that alerted at the same distance, and this was not due to species differences in group size or habitat. We then conducted an experiment in which a person approached groups of deer to compare the detection abilities and the form of alert response for white-tails and mule deer, and for age groups within each species. Mule deer alerted to the approacher at longer distances than white-tails, even after controlling for variables that were potentially confounding. Adult females of both species alerted sooner than conspecific juveniles. Mule deer almost always looked directly at the approacher as their initial response, whereas white-tails were more likely to flee or to look in another direction with no indication that they pinpointed the approacher during the trial. Mule deer may have evolved the ability to detect predators earlier than white-tails as an adaptation to their more open habitats, or because they need more time to coordinate subsequent anti-predator defenses.     

Frequency Dependence of Host Plant Choice within and Between Patches: A Large Cage Experiment

Oviposition preference is considered to be one of the most important factors behind patterns of host use among herbivorous insects. However, preference is defined as host plant choice under equal host abundance and availability, and it is likely that frequency-dependent effects will alter the actual pattern of host use beyond what preference trials reveals. The effects of such alterations are poorly known but could be important for the understanding of specialization and host shifts. We investigated how changes in frequency of a preferred and a less preferred host affected movement patterns and egg deposition within and among patches in a polyphagous butterfly, Polygonia c-album. Two experiments were carried out in large (8 × 30 m) outdoor cages, artificially divided into distinct patches with different frequencies of the two hosts: one that allowed for limited movement between patches and one that did not. There was a clear effect of frequency on patch selection; females spent more time in and laid more eggs in patches with a high frequency of the preferred host, which will potentially have a large effect on host use by modifying encounter rates in favor of the preferred host. However, there was no significant frequency-dependent plant choice within patches in any experiment. Instead, results indicate that females are distributing their eggs among plants species according to specific likelihoods of oviposition, independent of encounter rates, which is compatible with a strategy of risk-spreading.Co-ordinating editor: N. Yamamura     

Predation Risk Avoidance by Terrestrial Amphibians: The Role of Prey Experience and Vulnerability to Native and Exotic Predators

We studied avoidance, by four amphibian prey species (Rana luteiventris, Ambystoma macrodactylum, Pseudacris regilla, Tarichia granulosa), of chemical cues associated with native garter snake (Thamnophis elegans) or exotic bullfrog (R. catesbeiana) predators. We predicted that avoidance of native predators would be most pronounced, and that prey species would differ in the intensity of their avoidance based on relative levels of vulnerability to predators in the wild. Adult R. luteiventris (presumably high vulnerability to predation) showed significant avoidance of chemical cues from both predators, A. macrodactylum (intermediate vulnerability to predation) avoided T. elegans only, while P. regilla (intermediate vulnerability to predation) and T. granulosa (low vulnerability to predation) showed no avoidance of either predator. We assessed if predator avoidance was innate and/or learned by testing responses of prey having disparate levels of prior exposure to predators. Wild‐caught (presumably predator‐exposed) post‐metamorphic juvenile R. luteiventris and P. regilla avoided T. elegans cues, while laboratory‐reared (predator‐naive) conspecifics did not; prior exposure to R. catesbeiana was not related to behavioural avoidance among adult or post‐metamorphic juvenile wild‐reared A. macrodactylum and P. regilla. These results imply that (i) some but not all species of amphibian prey avoid perceived risk from garter snake and bullfrog predators, (ii) the magnitude of this response probably differs according to prey vulnerability to predation in the wild, and (iii) avoidance tends to be largely learned rather than innate. Yet, the limited prevalence and intensity of amphibian responses to predation risk observed herein may be indicative of either a relatively weak predator–prey relationship and/or the limited importance of predator chemical cues in this particular system.