The influence of predator regime on reproductive traits in <Emphasis Type="Italic">Gammarus</Emphasis> <Emphasis Type="Italic">pulex</Emphasis> populations

Selection on traits conferring reduced predation may be opposed by selection on other traits associated with reproduction. Here, we examined the hypothesis that traits associated with reproduction in Gammarus pulex are driven by predation. We studied G. pulex originating from ponds with two different kinds of predator regimes: (1) ponds with fish—often large, non-gap-limited predators and (2) ponds without fish where invertebrates are the dominant predators—often small, gap-limited predators with a much more restricted prey size range. We examined the body size of males and females in G. pulex amplexus pairs originating from fish and fishless ponds. We also examined, in the laboratory, their mating success, the number of offspring per female and offspring mortality under different rearing conditions, with or without fish cue. Mating success, defined as the percentage of amplexus pairs that produced live offspring, was higher for G. pulex from fishless ponds independent of rearing condition. Individuals from fish ponds were larger and they produced a higher number of offspring which tended to be related to female body size. Offspring mortality was higher in populations from fish ponds compared to populations from fishless ponds. Despite the higher offspring mortality, females from fish ponds had a higher number of offspring alive after 13 weeks, which is the approximate time it takes for G. pulex to reach maturity. Our data imply that no trade-off between reducing body size to reduce mortality caused by fish and maximising reproductive success exist in G. pulex from fish ponds. The strategy with many offspring may be the correct strategy in fishponds where predation pressure generally is higher than in fishless ponds.     

Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

Interspecific differences in drift behaviour between the native <Emphasis Type="Italic">Gammarus pulex</Emphasis> and the exotic <Emphasis Type="Italic">Gammarus roeseli</Emphasis> and possible implications for the invader’s success

“Drifting” is known to subject aquatic invertebrates to intense predation by drift feeding fish. Consequently, interspecific variations in drifting behaviour could lead to differences in predation pressure between coexisting prey species. Predation being an important factor determining the success of invaders, differences in drift patterns could advantage either native or exotic invertebrates through differential predation by native fish predators. The exotic freshwater amphipod (Gammarus roeseli) has now largely colonized Western Europe where it is often found in sympatry with a native species (Gammarus pulex). Here we documented interspecific differences in drifting behaviour that might have favored the invader’s success through differential predation. Benthic and drifting amphipods were sampled three times at the same site to compare the proportion of each species within and between sample types (benthos or drift) across time. Compared with the benthos, where the invader (G. roeseli) was significantly less abundant than the native (G. pulex), G. roeseli was proportionally overrepresented in the drift but displayed a very different drifting pattern. While G. pulex drift rates remained roughly constant over a 24 h period, G. roeseli showed a marked diel periodicity with low diurnal and high nocturnal drift rates. Such drifting behaviour could procure this species with a competitive advantage regarding predation as most drift feeding fish are diurnal. As a result, the native appears more disadvantaged with respect to drift. This may partly explain the ability of G. roeseli to coexist with G. pulex in a habitat more suitable to the native.     

Predatory disturbance and prey species diversity: the case of gray whale (<Emphasis Type="Italic">Eschrichtius robustus</Emphasis>) foraging on a multi-species mysid (family <Emphasis Type="Italic">Mysidae</Emphasis>) community

Why competitive exclusion does not limit the number of coexisting plankton species is a persistent question for community ecology. One explanation, the intermediate disturbance hypothesis (IDH), proposes that elevated species diversity is a product of moderate levels of disturbance that allow the subsequent invasion of less competitive species. Here, we assess the shifts in species diversity in a mysid (Mysidae Dana, 1850) zooplankton community, where at least 10 species have, over the last 15 years, have come to comprise the primary prey base of summer resident gray whales (Eschrictius robustus Lilljeborg, 1861) in Clayoquot Sound, British Columbia. We evaluate trends in the community structure of mysids (species dominance, diversity, and richness) across mysid habitat in the study area during the gray whale foraging season (May–September) for the period 1996 and 2008. Mysid species composition varies among years and diversity has increased as whales shifted their predatory focus from benthic amphipods (Ampeliscidae Costa, 1857) to mysids, near our starting point in 1996. Holmesimysis sculpta Tattersall, 1933 was the dominant species in early years; however, in 2007, the dominance shifted to Neomysis rayi Murdoch, 1885. The habitat restrictions and life history attributes of local populations of coastal mysids leave them vulnerable to the cumulative impacts of increased predation pressure by gray whales. This case study presents a unique examination implicating predation as an agent of disturbance capable of altering the species structure of a local prey community.     

The effects of predation risk and current velocity stress on growth,condition and swimming energetics of Japanese minnow (<Emphasis Type="Italic">Pseudorasbora parva</Emphasis>)

A Japanese minnow, Pseudorasbora parva, was exposed simultaneously to multiple dangers in experimental tanks. The study aimed to quantify to what extent the risk of predation coinciding with an adverse environmental factor, high flow velocity, affects prey in terms of growth and energy expenditure. In this experiment, two measures of growth (i.e., body weight and length), condition, feeding, swimming cost and behavioral responses were analyzed. The results showed that in such an environment, prey showed lowered growth and were in a poorer condition. As the prey shifted to the shallow area with high flow velocity, the prey consumed a lower ration and incurred multiple costs for swimming locomotion that might reduce the allocation of energy to biomass and energy storage. Reduction in activity might decrease the cost of locomotion, but it did not have a considerable effect on overall swimming energy expenditure. In stream ecosystems, the high swimming energy expenditure appears to magnify the effects of predation risk by causing lowered growth and a poorer condition and, hence, fitness. The present study shows that high flow velocity is one of the environmental factors that determine the energetic responses of a potential prey to the presence of predators.     

Foraging under the risk of predation in desert grassland whiptail lizards (<Emphasis Type="Italic">Aspidoscelis uniparens</Emphasis>)

The whiptail lizard Aspidoscelis uniparens searches for fossorial prey using a series of moves and pauses punctuated by bouts of digging. We examined the effect of predation risk on foraging A. uniparens in outdoor enclosures, observing their behavior in the presence and absence of the predatory lizard Gambelia wislizenii. Predator presence led to changes in activity patterns and foraging behavior. When predators were present, A. uniparens reduced both the proportion of time active and time moving, shifted activity periods, reduced their movement rate, and changed the duration of moves. There were no apparent changes in digging effort, but the likelihood of successfully digging for food decreased when a predator was present.     

Potential long-term storage of the predatory mite <Emphasis Type="Italic">Phytoseiulus</Emphasis><Emphasis Type="Italic">persimilis</Emphasis>

Increasing the ability to store mass-reared natural enemies during periods or seasons of low demand is a critical need of the biocontrol industry. We tested the hypothesis that chemicals can enhance long-term cold storage of a predatory mite Phytoseiulus persimilis Athias-Henriot. The research objective was to determine the effect of cryoprotectant and carbohydrate chemicals on in-storage survival of predators. In-storage survival at 8°C was greater for predators sprayed with glycerol (5%, v/v) or glucose (10% and 20%, v/v) than with water spray controls. After 74 days in the cryoprotectant experiment, predator survival declined to 11.5% in the 5% glycerol treatment and 7.8% in the water spray control. After 88 days in the carbohydrate experiment, predator survival declined to 22% in the 20% glucose treatment and 2% in the water spray control. Although many individuals expired within 50 days in both experiments, a few females survived more than 200 days. This research suggests that select cryoprotectants and carbohydrates have a limited capacity to facilitate long-term storage of P. persimilis.     

Ecology of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in natural habitats within its native range

Originally distributed in northeast Asia, Harmonia axyridis (Coleoptera: Coccinellidae) is now found throughout much of the temperate zone. In its native area, H. axyridis maintains stable populations in heterogeneous and temporary habitats because of its great ability to find prey and reproduce, coupled with density-dependent and self-regulatory population regulation. A negative correlation of H. axyridis on the biodiversity of the aphidophagous community has been observed in its native range. The decrease in biodiversity may be mainly caused by the wider range of habitats available to H. axyridis than to the coexisting species. From a theoretical perspective, density-dependent regulation of H. axyridis populations, e.g., cannibalism, may be more important in maintaining the H. axyridis-dominated system, probably than is intraguild predation. Habitat heterogeneity may also be important to the coexistence of H. axyridis and other predators in both native and invaded areas.     

Spatial avoidance of littoral and pelagic invertebrate predators by <Emphasis Type="Italic">Daphnia</Emphasis>

Studies on spatial avoidance behaviour of predators by prey often ignored the fact that prey typically face multiple predators which themselves interact and show a spatial pattern in abundance and predation rates (PRs). In a series of laboratory experiments, we investigated predation risk (PRI) and horizontal migration of the cladoceran Daphnia magna between open water and vegetation in response to two important invertebrate predators with a contrasting spatial distribution: pelagic Choaborus and vegetation-associated Ischnura. As expected, PRI by Chaoborus was higher in open water due to higher numbers and higher PRs of Chaoborus, while for Ischnura, PRI was highest in the vegetation due to higher densities, despite lower PRs of Ischnura. In accordance with this, Daphnia moved into the vegetation in the presence of the pelagic Chaoborus alone. In the presence of Ischnura alone, however, Daphnia showed no response. We hypothesize this may be the result of a constitutive behaviour of Daphnia to avoid pelagic fish, which impedes a response to the open water. In the combined predator treatment, Daphnia migrated to the open water zone. The increased risk of predation in the vegetation, due to a facilitating effect of Chaoborus on Ischnura PRs is believed to have caused this migration of the Daphnia. This response of Daphnia declined through time and Daphnia moved toward the vegetation. A decline in the activity of the Ischnura larvae through time may have switched the risk balance in favour of the vegetation environment.     

<Emphasis Type="Italic">Spizaetus</Emphasis> hawk-eagles as predators of arboreal colobines

The predation pressure put on primates by diurnal birds of prey differs greatly between continents. Africa and South America have specialist raptors (e.g. crowned hawk-eagle Stephanoaetus coronatus and harpy eagle Harpia harpyja) whereas in Asia the only such specialist’s (Philippine eagle Pithecophaga jefferyi) distribution is largely allopatric with primates. The almost universal absence of polyspecific groups in Asia (common in Africa and South America) may indicate reduced predation pressure. As such there is almost no information on predation pressures on primates in Asia by raptors. Here we report successful predation of a juvenile banded langur Presbytis femoralis (~2 kg) by a changeable hawk-eagle Spizaetus cirrhatus. The troop that was attacked displayed no signs of being alarmed, and no calls were made before the event. We argue that in insular Southeast Asia, especially, large Spizaetus hawk-eagles (~2 kg) are significant predators of arboreal colobines. Using data on the relative size of sympatric Spizaetus hawk-eagles and colobines we make predictions on where geographically we can expect the highest predation pressure (Thai–Malay Peninsula) and which colobines are least (Nasalis larvatus, Trachypithecus auratus, P. thomasi) and most (P. femoralis, T. cristatus) affected.     

Effects of Chemical Cues on Foraging in Damselfly Larvae, <Emphasis Type="Italic">Enallagma antennatum</Emphasis>

Animals experiencing a trade-off between predation risk and resource acquisition must accurately predict ambient levels of predation risk to maximize fitness. We measure this trade-off explicitly in larvae of the damselfly Enallagma antennatum, comparing consumption rates in the presence of chemical cues from predators and injured prey. Damselflies distinguished among types of chemical cues based on species of prey injured or eaten. Injured coexisting heterospecific and unknown heterospecific chemical cues did not reduce foraging relative to starved predator cues, while cues arising from predators eating a coexisting heterospecific did decrease foraging. This study shows a cost in terms of reduced foraging in response to chemical cues and further defines the ability of prey to respond discerningly to chemical cues.     

No diapause prolongation response of <Emphasis Type="Italic">Daphnia</Emphasis> in the presence of planktivorous mosquitofish (<Emphasis Type="Italic">Gambusia holbrooki</Emphasis>)

Recent research suggests that planktonic organisms in resting stages can perceive predators chemically and delay hatching to evade predation. Using a co-occurring predator–prey pair (Gambusia holbrooki Girard and Daphnia curvirostris Eylmann) from a Mediterranean floodplain wetland, this study tested for such predator-avoidance behaviour of Daphnia. Results show that hatching patterns of Daphnia were not reduced by the presence of different biomass levels of Gambusia. This could be due to the fact that the density of Gambusia in the wetland is high from late spring to late autumn, suggesting that delayed hatchlings would face increased mortality through consumptive predation.     

The invasive predator <Emphasis Type="Italic">Bythotrephes</Emphasis> induces changes in the vertical distribution of native copepods in Lake Michigan

Invasive predators can have large negative effects on native prey populations. The susceptibility of native prey to invasive predators may depend on their ability to respond behaviorally to the presence of these non-native predators. In a field survey conducted in Lake Michigan over several years, we found that high densities of the invasive predatory cladoceran Bythotrephes were correlated with lower vertical distributions of some species and age classes of native copepods; moving from inhabiting primarily the epiliminion at low Bythotrephes density to primarily the hypolimnion at high Bythotrephes density. Five groups showed this pattern; diaptomid copepodites, adult cyclopoids, Diacyclops thomasi, and the adult diaptomids Leptodiaptomus ashlandi and L. minutus. In contrast, Bythotrephes density was not correlated with the vertical distribution of copepod nauplii and adult L. sicilis. Laboratory experiments suggest that the changes in the vertical distribution in the field at high Bythothrephes are due to an inducible, plastic response to predation threat from Bythotrephes signaled by water-borne cues. Species that were lower in the field at high Bythotrephes densities responded behaviorally to water-borne cues from Bythotrephes by moving to lower levels of experimental water columns. These species included D. thomasi and L. minutus, with L. ashlandi displaying a non-significant trend in the same direction. In contrast, L. sicilis, which was not correlated with Bythotrephes density in the field, was unaffected by the water-borne cues. Differences in vertical distribution shifts among these native copepod species and life-history stages are likely due to species-specific differences in spatial overlap with Bythotrephes and their relative ability to migrate large distances or employ alternative avoidance strategies. The varied responses exhibited among the copepod groups likely alter their interactions with each other, their resources and other predators, thus revealing the complex effects Bythotrephes can have on invaded communities.     

Advantage of early seedling emergence in <Emphasis Type="Italic">Fagus crenata</Emphasis>: importance of cotyledon stage for predator escape and pathogen avoidance

To examine how early-emerging seedlings take advantage of establishment in a deciduous forest, we explored the relationships among the emergence date, growth stage and major causes of mortality (damping-off by fungi and predation by rodents) in seedlings of Fagus crenata. The emergence of current-year seedlings and their survival and damage were followed at short (3–7 days in early spring) intervals for a growing season. The growth stage was divided into two stages, with only cotyledons (CT stage) and with true leaves (TL stage). The survival rate was negatively correlated with the emergence date, indicating the advantage of early seedling emergence. This advantage was largely explained by the lower occurrence of damping-off. In contrast, seedling predation occurred independently of the emergence date, but depended strongly on the developmental stage. Rodents consumed a considerable number of seeds during the early period after emergence, and strongly preferred CT-stage over TL-stage seedlings throughout the growing season. Therefore, seedling predation was inferred to be concentrated in a relatively short period while the remaining seeds were depleted and CT-stage seedlings were abundant. The seedling stage synchronously shifted from the CT to TL stage. This growth-stage transition was independent of the emergence date and appeared to correspond with the timing of seed depletion. Delayed stage transition resulted in a disproportionately high risk of damping-off later in the season. Our results indicate that early seedling emergence is advantageous for F. crenata for resistance to pathogens and that the timing of growth-stage progression of seedlings appears important to escape predation because of the distinct food preference of predators.     

Examination of boldness traits in sexual and asexual mollies (<Emphasis Type="Italic">Poecilia latipinna</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Considering the high costs of sexual reproduction (e.g., the production of males), its maintenance and predominance throughout the Animal Kingdom remain elusive. Especially the mechanisms allowing for a stable coexistence of closely related sexual and asexual species are still subject to a lively debate. Asexuals should rapidly outnumber sexuals due to higher population growth rates, unless they face some disadvantages. Here, we investigate potential differences in feeding behavior in a system of sexual (sailfin mollies, Poecilia latipinna) and coexisting gynogenetic fishes (Amazon mollies, Poecilia formosa). In two different experiments, we tested for differences in behavioral traits associated with boldness. Bold individuals take higher risks for gains in resources, so shyer individuals should be less competitive. Our study was motivated by the recent finding that P. formosa are less likely to be preyed upon by piscine predators than P. latipinna. We asked whether this result is indicative of low boldness in P. formosa. However, no differences between the two species were detectible in our behavioral experiments measuring (a) time to emerge from shelter to explore a novel environment, (b) latency time until feeding in a novel environment, and (c) recovery time until feeding restarted after a simulated predator attack. Furthermore, different boldness measures were not correlated with each other within individuals.     

Wrap attack of the spider <Emphasis Type="Italic">Achaearanea tepidariorum</Emphasis> (Araneae: Theridiidae) by preying on mealybugs <Emphasis Type="Italic">Planococcus citri</Emphasis> (Homoptera: Pseudococcidae)

Predation by Achaearanea tepidariorum (Koch 1841) on mealybugs Planococcus citri (Risso 1813) is facilitated by the design of its web, which features a tangle of sticky gumfooted lines, and wrap attacks as well as the ability to handle the prey, whose body is covered with a waxy secretion, via silk. Crawling, i.e., wingless, mealybugs (in particular those in the nymphal stages and adult females and, to a lesser extent, winged males) are caught by means of the gumfooted lines, covered with globules of an adhesive secretion. The process of wrap attack and subsequent handling of the captured prey is a series of the following consecutive events: (1) confining and immobilising the mealybugs with sticky silk; (2) biting with chelicerae and paralyzing the prey with a toxin; (3) detaching the confined prey, attached to the tense threads, from the plant surface and catapulting it toward the central section of the web; (4) wrapping the catapulted prey in viscid silk emitted by the spinning apparatus; (5) transporting the wrapped prey to the central section of the web; (6) wrapping the prey in the central section of the web in nonsticky silk, whose tufts are present in this part of the web even before the attack; (7) filling the prey with digestive fluid; (8) sucking the prey empty; and (9) cleaning the chelicerae and mouth parts. The process of silk tuft wrapping was described for the first time. The described ability to hunt mealybugs implies the possibility of using A. tepidariorum spiders for biological control of these pests.     

<Emphasis Type="Italic">Utricularia</Emphasis>: a vegetarian carnivorous plant?

Aquatic carnivorous plants of the genus Utricularia capture and utilise a wide range of small aquatic organisms. Most of the literature focuses on animals as prey. In this study, we investigate the occurrence of algae inside the traps of four species of bladderwort. We observed that algae of 45 genera form up to 80% of the total prey; algae were found frequently in traps without animal prey. The majority are coccal and trichal algae of the families Desmidiaceae and Zygnemataceae. The percentage of algae increases significantly with decreasing electric conductivity of the water (r _S = −0.417; P = 0,000). Thus, algae are the most frequent prey in extremely soft waters. The percentage of algae did not differ significantly, not within the investigated Utricularia species or within the various study sites. However, the taxonomic composition of the algal prey showed highly significant differences between different sites. More than 90% of the trapped algae were killed and degraded by the bladders. The recent data allow for two alternative hypotheses: either algal prey supplements animal prey in oligotrophic waters, or the unprofitable trapping of algae is rather an additional stress factor for Utricularia and contributes to its limited distribution in some peat bogs.     

Density-mediated indirect effects of a common prey tadpole on interaction between two predatory bugs: <Emphasis Type="Italic">Kirkaldyia deyrolli</Emphasis> and <Emphasis Type="Italic">Laccotrephes japonensis</Emphasis>

This paper suggests that the nymphs of a specialist predator, Kirkaldyia (=Lethocerus) deyrolli (Belostomatidae: Heteroptera), are indirectly affected by their tadpoles’ prey. K. deyrolli nymphs and the predator Laccotrephes japonensis (Nepidae: Heteroptera) adults coexist in rice paddy fields. It was predicted that the difference in tadpole density may influence the K. deyrolli nymph survival rate. We first compared survival rates of the first instar nymphs of K. deyrolli in June (high tadpole density period) and July (low tadpole density period). Secondly, we investigated the survival rate of K. deyrolli nymphs at different tadpole densities and under the presence or absence of L. japonensis adults to examine whether higher tadpole density moderates predation pressure from L. japonensis adults to K. deyrolli nymphs, e.g., density-mediated indirect effects. As a result of the comparison, the survival rate of K. deyrolli nymphs in June was higher than that in July. For the field experiment, the slopes between the survival rate of K. deyrolli nymphs and tadpole density were positive under both predator presence and absence. However, the slope under the presence of a predator was steeper than that under absence of the predator (“predator-by-tadpole density interaction” was significant). These results suggest that a higher tadpole density in June provides an abundant food resource for K. deyrolli nymphs and also moderates predation pressure from L. japonensis.     

Influence of sprint speed and body size on predator avoidance in New Mexican spadefoot toads (<Emphasis Type="Italic">Spea multiplicata</Emphasis>)

Predator–prey interactions play an important role in community dynamics and may be important for promoting genetic diversification. Diversification may be especially important when prey species have multiple anti-predator strategies available, but these strategies conflict with each other. For example, rapid sprint speed and large size are both thought to decrease vulnerability to many predators. A physiological trade-off between swimming speed and growth rate has been documented in many aquatic species and, as a result, individual genotypes may employ one strategy or the other, but not both. Although rapid sprint speed is often assumed to decrease vulnerability to predators, this has only rarely been tested. Here I provide evidence that both rapid sprint speed and large size in tadpoles of the New Mexico spadefoot toad (Spea multiplicata) decreases predation risk from carnivore morphs of its congener the Great Plains spadefoot toad (Spea bombifrons). Such conflicts, coupled with spatio-temporal variation in predation pressure, may be important in maintaining genetic variation for trade-offs.     

Pungent odor of the adult skipper butterfly <Emphasis Type="Italic">Erynnis</Emphasis><Emphasis Type="Italic">montanus</Emphasis> (Lepidoptera: Hesperiidae)

Adults of some butterfly species have odors, several components of which are known to have pheromonal or defensive functions. Little is known about the odors of hesperiid butterflies, however. Erynnis montanus (Hesperiidae), especially male adults, emit a pungent odor that is detectable by the human nose. Chemical analysis has revealed that crude extracts of wild individuals contained 10 volatile substances, of which docosane and heneicosane were the main components. Because males contain a significantly larger amount of p-cresol than females, this aromatic compound is characterized as a strong male odor. At the end of the adult occurrence period, p-cresol decreased substantially in each male. Benzothiazole, identified for the first time in lepidopteran adults, was present in both sexes in almost equal amounts. Among the 10 volatile substances detected in males, biased distribution in the wings rather than the body was observed for benzothiazole, heptanal, and p-cresol. Male adults have androconial organs in the costal part of the forewing on which benzothiazole and p-cresol tended to concentrate. However, these compounds were detected not only in other parts of the forewing but also in the hindwing and body, suggesting the presence of undiscovered scent-producing organs.