How do dispersal costs and habitat selection influence realized population connectivity?

Despite the importance of dispersal for population connectivity, dispersal is often costly to the individual. A major impediment to understanding connectivity has been a lack of data combining the movement of individuals and their survival to reproduction in the new habitat (realized connectivity). Although mortality often occurs during dispersal (an immediate cost), in many organisms costs are paid after dispersal (deferred costs). It is unclear how such deferred costs influence the mismatch between dispersal and realized connectivity. Through a series of experiments in the field and laboratory, we estimated both direct and indirect deferred costs in a marine bryozoan (Bugula neritina). We then used the empirical data to parameterize a theoretical model in order to formalize predictions about how dispersal costs influence realized connectivity. Individuals were more likely to colonize poor-quality habitat after prolonged dispersal durations. Individuals that colonized poor-quality habitat performed poorly after colonization because of some property of the habitat (an indirect deferred cost) rather than from prolonged dispersal per se (a direct deferred cost). Our theoretical model predicted that indirect deferred costs could result in nonlinear mismatches between spatial patterns of potential and realized connectivity. The deferred costs of dispersal are likely to be crucial for determining how well patterns of dispersal reflect realized connectivity. Ignoring these deferred costs could lead to inaccurate predictions of spatial population dynamics.     

How does the spatial structure of habitat loss affect the eco-epidemic dynamics?

Habitat loss is considered as one of the primary causes of species extinction, especially for a species that also suffers from an epidemic disease. Little attention has been paid to the combined effect of habitat loss and epidemic transmission on the species spatiotemporal dynamics. Here, a spatial model of the parasite–host/prey–predator eco-epidemiological system with habitat loss was studied. Habitat patches in the model, instead of undergoing a random loss, were spatially clustered by different degrees. Not only the quantity of habitat loss but also its clustering degree was shown to affect the equilibrium of the system. The infection rate and the probability of successful predation were keys to determine the spatial patterns of species. The epidemic disease is more likely to break out if only a small amount of suitable patches were lost. Counter-intuitively, infected preys are more sensitive to habitat loss than predators if the lost patches are highly clustered. This result is new to eco-epidemiology and implies a possibility of using spatial arrangement of suitable (or unsuitable) patches to control the spread of epidemics in the ecological system.     

Microbial and environmental effects on avian egg viability: do tropical mechanisms act in a temperate environment?

The viability of freshly laid avian eggs declines after several days of exposure to ambient temperatures above physiological zero, and declines occur faster in tropical than temperate ecosystems. Microbial infection during preincubation exposure has recently been shown as a second cause of egg viability decline in the tropics, but whether microbial processes influence the viability of wild bird eggs in temperate ecosystems is unknown. We determined the microbial load on eggshells, the incidence of microbial penetration of egg contents, and changes in the viability of wild bird eggs (Sialia mexicana, Tachycineta bicolor, Tachycineta thalassina) experimentally exposed to temperate-zone ambient conditions in situ in a mediterranean climate in northern California. Initial microbial loads on eggshells were generally low, although they were significantly higher on eggs laid in old boxes than in new boxes. Eggshell microbial loads did not increase with exposure to ambient conditions, were not reduced by twice-daily disinfection with alcohol, and were unaffected by parental incubation. The rate of microbial penetration into egg contents was low and unaffected by the duration of exposure. Nevertheless, egg viability declined very gradually and significantly with exposure duration, and the rate of decline differed among species. In contrast to studies performed in the tropics, we found little evidence that temperature or microbial mechanisms of egg viability decline were important at our temperate-zone site; neither temperatures above physiological zero nor alcohol disinfection was significantly related to hatching success. Delaying the onset of incubation until the penultimate or last egg of a clutch at our study site may maintain hatching synchrony without a large trade-off in egg viability. These results provide insight into the environmental mechanisms that may be responsible for large-scale latitudinal patterns in avian clutch size and hatching asynchrony.     

The unified neutral theory of biodiversity: do the numbers add up?

Hubbell's unified neutral theory is a zero-sum ecological drift model in which population sizes change at random in a process resembling genetic drift, eventually leading to extinction. Diversity is maintained within the community by speciation. Hubbell's model makes predictions about the distribution of species abundances within communities and the turnover of species from place to place (beta diversity). However, ecological drift cannot be tested adequately against these predictions without independent estimates of speciation rates, population sizes, and dispersal distances. A more practical prediction from ecological drift is that time to extinction of a population of size N is approximately 2N generations. I test this prediction here using data for passerine birds (Passeriformes). Waiting times to speciation and extinction were estimated from genetic divergence between sister populations and a lineage-through-time plot for endemic South American suboscine passerines. Population sizes were estimated from local counts of birds in two large forest plots extrapolated to the area of wet tropical forest in South America and from atlas data on European passerines. Waiting times to extinction (ca. 2 Ma) are much less than twice the product of average population size (4.0 and 14.4 x 10(6) individuals in South America and Europe) and generation length (five and three years) for songbirds, that is, 40 and 86 Ma, respectively. Thus, drift is too slow to account for turnover in regional avifaunas. Presumably, other processes, involving external drivers, such as climate and physiographic change, and internal drivers, such as evolutionary change in antagonistic interactions, predominate. Hubbell's model is historical and geographic, and his perspective importantly links local and regional process and pattern. Ecological reality can be added to the mix while retaining Hubbell's concept of continuity of communities in space and time.     

How do nutrient conditions and species identity influence the impact of mesograzers in eelgrass-epiphyte systems?

Coastal eutrophication is thought to cause excessive growth of epiphytes in eelgrass beds, threatening the health and survival of these ecologically and economically valuable ecosystems worldwide. Mesograzers, small crustacean and gastropod grazers, have the potential to prevent seagrass loss by grazing preferentially and efficiently on epiphytes. We tested the impact of three mesograzers on epiphyte biomass and eelgrass productivity under threefold enriched nutrient concentrations in experimental indoor mesocosm systems under summer conditions. We compared the results with earlier identical experiments that were performed under ambient nutrient supply. The isopod Idotea baltica, the periwinkle Littorina littorea, and the small gastropod Rissoa membranacea significantly reduced epiphyte load under high nutrient supply with Rissoa being the most efficient grazer, but only high densities of Littorina and Rissoa had a significant positive effect on eelgrass productivity. Although all mesograzers increased epiphyte ingestion with higher nutrient load, most likely as a functional response to the quantitatively and qualitatively better food supply, the promotion of eelgrass growth by Idotea and Rissoa was diminished compared to the study performed under ambient nutrient supply. Littorina maintained the level of its positive impact on eelgrass productivity regardless of nutrient concentrations.     

Which forcing factors fit? Using ecosystem models to investigate the relative influence of fishing and changes in primary productivity on the dynamics of marine ecosystems

Fishing mortality and primary production (or proxy for) were used to drive the dynamics of fish assemblages in 9 trophodynamic models of contrasting marine ecosystems. Historical trends in abundance were reconstructed by fitting model predictions to observations from stock assessments and fisheries independent survey data. The model fitting exercise derives values for otherwise unknown parameters that specify the relative strength of trophic interactions and, in some instances, a time series anomaly for changes in primary production. We measured how much better or worse were model predictions when bottom-up forcing by primary production were added to top-down forcing by fishing. Searching for cross system patterns, the relative contribution of fishing and changes in primary production, mediated through trophic interactions, are evaluated for the ecosystems as a whole and for selected similar species in different ecosystems. The analysis provides a simple qualitative way to explain which forcing factors have most influence on modeled dynamics. Both fishing and primary production forcing were required to obtain the best model fits to data. Fishing effects more strongly influenced 6 of 9 of the ecosystems, but primary production was more often found to be the main factor influencing the selected pelagic and demersal fish stock trends. Examination of sensitivity to ecological and model parameters suggests that the results are the product of complex food-web interactions rather than simple deterministic responses of the models.     

Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity?

Predators and prey assort themselves relative to each other, the availability of resources and refuges, and the temporal and spatial scale of their interaction. Predictive models of predator distributions often rely on these relationships by incorporating data on environmental variability and prey availability to determine predator habitat selection patterns. This approach to predictive modeling holds true in marine systems where observations of predators are logistically difficult, emphasizing the need for accurate models. In this paper, we ask whether including prey distribution data in fine-scale predictive models of bottlenose dolphin (Tursiops truncatus) habitat selection in Florida Bay, Florida, U.S.A., improves predictive capacity. Environmental characteristics are often used as predictor variables in habitat models of top marine predators with the assumption that they act as proxies of prey distribution. We examine the validity of this assumption by comparing the response of dolphin distribution and fish catch rates to the same environmental variables. Next, the predictive capacities of four models, with and without prey distribution data, are tested to determine whether dolphin habitat selection can be predicted without recourse to describing the distribution of their prey. The final analysis determines the accuracy of predictive maps of dolphin distribution produced by modeling areas of high fish catch based on significant environmental characteristics. We use spatial analysis and independent data sets to train and test the models. Our results indicate that, due to high habitat heterogeneity and the spatial variability of prey patches, fine-scale models of dolphin habitat selection in coastal habitats will be more successful if environmental variables are used as predictor variables of predator distributions rather than relying on prey data as explanatory variables. However, predictive modeling of prey distribution as the response variable based on environmental variability did produce high predictive performance of dolphin habitat selection, particularly foraging habitat.     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Diversity and host range of foliar fungal endophytes: are tropical leaves biodiversity hotspots?

Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests.     

The rarity of twins: a result of an evolutionary battle between mothers and daughters—or do they agree?

The aim of this study was to test the hypothesis that parent–offspring conflict over parental investment might have led to the rarity of dizygotic twins in humans. We explored the theoretical prediction that twins maximize their inclusive fitness by the death of a co-twin, while mothers of twins maximize fitness by raising both twins to independence. We used life history data (1700–1900) from two parishes in Central Norway to compare differences in inclusive fitness (measured as number of children reared to the age of 16 years, using Hamilton's rule) between twins and mothers of twins. Our results show that twins maximize their inclusive fitness by the death of a co-twin, while mothers of twins raise more children by rearing both twins to adulthood. However, because twins growing up as singletons may produce higher or at least equal number of offspring than the sum of the two twins growing up together, mothers might gain more grandchildren by allowing twins to grow up as singletons. To conclude, both selfish twins and their mothers might benefit by the death of a co-twin, indicating that there is no parent–offspring conflict responsible for the rareness of twins in these human populations. Finally, we discuss the results in the light of “The Insurance Egg Hypothesis” and “The Natural Selection Hypothesis”.     

Does the composition of the demersal fish assemblages in temperate coastal waters change with depth and undergo consistent seasonal changes?

The aim of this study was to determine whether the composition of the demersal fish fauna in coastal marine waters in temperate Australia changes markedly with increasing water depth and distance from the shore and whether the composition of the fish fauna in water depths of 5 to 35 m undergoes cyclic, seasonal changes. Samples of demersal fishes were therefore collected by trawling over the predominantly sandy substrate at nine sites located in water depths of 5 to 15 m or 20 to 35 m and within 20 km of the shore in four regions along ∼200 km on the lower west coast of Australia. The sampling regime involved trawling for fishes at each site at night in seven consecutive seasons between the summer of 1990/1991 and winter of 1992. A total of 72 435 fishes, representing 77 families, 143 genera and 172 species was caught. The compositions of the fish faunas in offshore waters with depths of 5 to 35 m were shown to differ markedly from those previously recorded for nearshore marine waters in the same regions. However, as some species, such as Sillago burrus, S. vittata, S. bassensis and Rhabdosargus sarba, increase in size, they move out from their nursery areas in nearshore waters into deeper and more offshore waters, where spawning occurs. Ordination showed that, in each of the four regions, the composition of the fish fauna in depths of 5 to 15 m differs from that in depths of 20 to 35 m. This difference is attributable to the fact that some species, such as  S. burrus, S. vittata and Upeneichthys lineatus, are far more abundant in depths of 5 to 15 m, whereas other species, such as S. robusta, U. stotti and Lepidotrigla modesta, occur predominantly in depths of 20 to 35 m. However, the samples collected from the single site that was inshore but in deeper water demonstrate that the composition of the fish fauna is influenced by distance from shore as well as by water depth. The compositions of the fish faunas differed with latitude, largely due to the fact that some subtropical species, such as Polyspina piosae, S. burrus and  S. robusta, did not extend down into the more southern regions. Ordination also showed that the composition of the fish faunas at all but one of the nine sites underwent pronounced and consistent cyclic, seasonal changes. This seasonal cyclicity at the different sites was attributable to sequential patterns of immigrations and emigrations by a number of fish species during the course of the year. These seasonal migrations involved, inter alia (1) movements of certain species from their nursery areas into these deeper waters, e.g.  S. bassensis and Scobinichthys granulatus; (2) migrations into and off the sandy areas of the inner continental shelf, e.g. Arnoglossus muelleri; (3) migrations to spawning areas, e.g. Sillago robusta; and (4) movements into areas where detached macrophytes accumulate in winter, e.g. Cnidoglanis macrocephalus and Apogon rueppellii. Received: 21 August 1998 / Accepted: 9 February 1999     

The pursuit of extra-pair copulations by monogamous female razorbills: how do females benefit?

Summary Monogamous female razorbills Alca torda actively sought extra-pair copulations (EPCs) in mating arenas outside their nesting colony. Females showed marked variation in promiscuity, defined as the number of EPCs they accepted (0–7), and in receptivity, defined as the percentage of EPC opportunities accepted (0%–100%). The opportunity of females to encounter males for EPCs was measured by (a) the time spent in the mating arena, and (b) the guarding effectiveness of their mates. Females appeared to encounter males and accept EPCs independently of opportunity, which suggests that variation in promiscuity and receptivity is caused by differences in the degree to which individual females might benefit from EPCs. I tested predictions of four hypotheses which propose benefits which females could obtain from EPCs: good genes, genetic diversity of offspring, insurance against infertility of mate, and acquisition of a new mate. No evidence was found to support two predictions made exclusively by the good genes hypothesis: (a) paired males did not achieve greater EPC success than unpaired males, and (b) males who were cuckolded did not obtain fewer EPCs than males whose mates avoided EPCs, suggesting that a male's attractiveness to other females does not ensure his own mate's fidelity. A prediction of both the insurance and good genes hypotheses was supported: females (who in this species retain their mates between years) showed similar receptivity to EPCs between years. This finding contradicts the genetic diversity hypothesis which predicts that in species that lay one egg per year, females who refuse EPCs in one year will accept them in the next. The good genes and genetic diversity hypotheses both predict that females who accept EPCs will behave as if to maximize their chances of being fertilized by extra-pair males. Results contradicted this prediction: females who accepted EPCs were more, rather than less receptive to their mates' copulation attempts than were females who did not accept EPCs. This finding is compatible with the insurance hypothesis which only requires females to store the sperm of extra-pair males, without necessarily being fertilized by it. Insurance against male infertility was the only hypothesis whose predictions were not contradicted by any of the results.     

Open and closed seascapes: where does habitat patchiness create populations with high fractions of self-recruitment?

Which populations are replenished primarily by immigrants (open) and which by local production (closed) remains an important question for management with implications for response to exploitation, protection, and disturbance. However, we lack methods for predicting population openness. Here, we develop a model for openness and show that considering habitat isolation explains the existence of surprisingly closed populations in high-dispersal species, including many marine organisms. Relatively closed populations are expected when patch spacing is more than twice the standard deviation of a species'. dispersal kernel. In addition, natural scales of habitat patchiness on coral reefs are sufficient to create both largely open and largely closed populations. Contrary to some previous interpretations, largely closed marine populations do not require mean dispersal distances that are unusually short, even for species with relatively long pelagic larval durations. We predict that habitat patchiness has strong control over population openness for many marine and terrestrial species with a highly dispersive life stage and relatively sedentary adults. This information can be used to make initial predictions about where populations will be more or less resilient to local exploitation and disturbance.     

Evaluating animal personalities: do observer assessments and experimental tests measure the same thing?

The animal personality literature uses three approaches to assess personality. However, two of these methods, personality ratings and experimentation, have been little compared in captivity and never compared in the wild. We assessed the boldness of wild chacma baboons Papio ursinus using both ratings and experimental methods. Boldness was experimentally assessed when individuals were presented with a novel food item during natural foraging. The boldness of the same individuals was rated on a five-point scale by experienced observers. The ratings and experimental assessments of boldness were found to correlate positively and in a linear fashion. When considered categorically the two approaches showed variable agreement depending on the number of categories assigned and the cut-off criteria adopted. We suggest that the variation between approaches arises because each method captures different aspects of personality; ratings consider personality in absolute terms (using predefined criteria) and multiple contexts, while experimental assessments consider personality in relative terms (using experimental scores relative to the population average) and in limited contexts. We encourage animal personality researchers to consider adopting both methodologies in future studies. We also propose that future studies restrict their analyses to continuous data, since the greatest comparability between methods was found with these data. However, if individuals must be categorised, we suggest that researchers either (a) analyse only those individuals categorised as bold or shy by both ratings and experimental approaches or, if these methods cannot be employed simultaneously, (b) do not use approach-specific criteria but choose a cut-off that can be compared by both approaches.     

Interacting effects of habitat destruction and changing disturbance rates on biodiversity: Who is going to survive?

Changes in disturbance rates due to climate change may increase or decrease diversity, whereas permanent loss of habitat is generally believed to decrease diversity. It is, however, very likely that the effects of disturbances and habitat destruction interact. Understanding such combined effects is essential to predict the response of communities to global changes and in particular which functional types of species are most endangered. Using an individual-based spatially explicit community model, we investigate (1) whether diversity-disturbance curves alter when spatially uncorrelated or autocorrelated habitat destruction is added, and (2) which functional types of species are able to survive under these altered conditions. Model communities consisted of four functional types of species trading off between colonisation ability and competition strength. We found that habitat destruction may alter both height and shape of diversity-disturbance curves: maximum diversity at intermediate disturbance rates may shift to other disturbance rates or even split into two peaks giving rise to bimodal diversity-disturbance relationships with different sub-communities persisting at low and high disturbance rates. Diversity responded differentially depending on how the colonisation-competition trade-off was represented. Our results suggest that, for trade-offs in seed production rate, generally the best coloniser will better withstand the interacting effects of habitat destruction and changing disturbance rates; however, for trade-offs in mean dispersal distances, functional types characterized by intermediate abilities will perform best. We conclude that predictions of the impacts of changing disturbance rates on biodiversity depend on community structure and cannot be made without knowledge of concurrent permanent habitat destruction.     

How do simple energy activities comprise complex behavior of life systems?: A conceptual synthesis and decomposition of the energy structure of life systems

Reduction and synthesis has always been Science’s way to learn about complex systems. The problem is, of course, that after studying the behavior of the system components, the reconstruction of system behavior cannot be any simple summation of the behavior of the individual components. How should we go about with the reconstruction is an umbrella question addressed in the paper. In particular, one should ask what is the contribution of each component to the system, and what kind and quantitative relations do the connections among the components signify in the total system? Answering these questions is necessary to understand the internal mechanism of a life system, the regularity of its dynamics, and the nature of the control mechanisms.The general approach of the paper is based on the assumption that the life energy transmission structure of the system is the fundamental principle of reduction and synthesis.The paper generalizes the energy transmission in terms of two basic types, based upon the mode of linkage in energy activities, such as sequential or parallel. The basic mathematical concept is an eigenparameter analysis in a model called Life Energy System Model (LESM). Simple examples illustrate the models application. It is true that the energy transmission structure of a natural life system is usually very complex, in which case the analysis has to incorporate more details than shown in the paper. Yet, the analysis still follows the same fundamental principle as outlined.     

Male mating history and female fecundity in the Lepidoptera: do male virgins make better partners?

In insects, large ejaculate and associated materials, including spermatophores, appear to have evolved via sexual selection acting on males to either delay female remating or to increase the rate of egg-laying. It is also possible, however, that females use nutrients transferred during mating to increase their lifetime fecundity. If so, male ejaculate size may also have evolved under natural selection as a form of paternal investment. In Lepidoptera, males with a greater number of prior matings tend to produce smaller spermatophores. However, the reported effects of male mating history on female fecundity vary widely among species. We therefore performed a meta-analysis using data from 29 studies of 25 species. Overall, the reproductive output of females mated to virgin males was significantly higher than that of females mated to sexually experienced males (Hedges d=0.33, P<0.01). A sample size of around 145 females per male mating type is required to detect an effect of this size with 80% statistical power at =0.05 (two-tailed). There was no difference in mean effect size between butterflies/skippers and moths. After controlling for any effect of taxonomic group, however, the mean effect size for polyandrous species was significantly greater than that for monandrous species (Hedges d=0.45 vs 0.25, P=0.01). We then discuss possible reasons why male mating history, presumably acting through its effect on spermatophore size, might have a stronger effect in polyandrous than monandrous species.Communicated by A. Cockburn     

Making scents of defense: do fecal shields and herbivore-caused volatiles match host plant chemical profiles?

Many plant families have aromatic species that produce volatile compounds which they release when damaged, particularly after suffering herbivory. Monarda fistulosa (Lamiaceae) makes and stores volatile essential oils in peltate glandular trichomes on leaf and floral surfaces. This study examined the larvae of a specialist tortoise beetle, Physonota unipunctata, which feed on two M. fistulosa chemotypes and incorporate host compounds into fecal shields, structures related to defense. Comparisons of shield and host leaf chemistry showed differences between chemotypes and structures (leaves vs. shields). Thymol chemotype leaves and shields contained more of all compounds that differed than did carvacrol chemotypes, except for carvacrol. Shields had lower levels of most of the more volatile chemicals than leaves, but more than twice the amounts of the phenolic monoterpenes thymol and carvacrol and greater totals. Additional experiments measured the volatiles emitted from M. fistulosa in the absence and presence of P. unipunctata larvae and compared the flower and foliage chemistry of plants from these experiments. Flowers contained lower or equal amounts of most compounds and half the total amount, compared to leaves. Plants subjected to herbivory emitted higher levels of most volatiles and 12 times the total amount, versus controls with no larvae, including proportionally more of the low boiling point chemicals. Thus, chemical profiles of shields and volatile emissions are influenced by the amounts and volatilities of compounds present in the host plant. The implications of these results are explored for the chemical ecology of both the plant and the insect.