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.     

Environmental cues or conspecific attraction as causes for egg mass aggregation in hydrobiosid caddisflies

The distribution of egg masses throughout the environment can strongly influence the population dynamics of aquatic insects. Many species lay eggs in aggregations and most eggs will subsequently hatch from only a few locations—knowing how and why these aggregations arise is therefore needed to understand the population dynamics of these species. The hydrobiosid caddisfly Ulmerochorema rubiconum lays eggs in large aggregations on the undersides of emergent rocks in streams. Our aim was to test whether females oviposit in response to conspecific egg masses or the environmental characteristics of rocks by manipulating the number and age of egg masses initially present on rocks and monitoring the accumulation of new masses. Our first experiment used rocks that had recently been used for oviposition and initial egg masses encompassed a range of ages; our second used rocks with no history of oviposition and initial masses of uniform age. Females did not respond to conspecific eggs of any age, as removing initial egg masses had no impact on the number of new masses laid in either experiment. There was a strong positive relationship between the number of initial and new masses, regardless of whether initial egg masses were removed, at the scale of whole rocks and for rock microhabitats. The results suggest that females select oviposition sites based primarily on physical characteristics of emergent rocks. We demonstrate for the first time that the spatial arrangement of egg masses may be consistent and predictable through time, with females showing a high degree of fidelity for particular rocks and microhabitats.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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Selective oviposition of the mayfly Baetis bicaudatus

Selective oviposition can have important consequences for recruitment limitation and population dynamics of organisms with complex life cycles. Temporal and spatial variation in oviposition may be driven by environmental or behavioral constraints. The goals of this study were to: (1) develop an empirical model of the substrate characteristics that best explain observed patterns of oviposition by Baetis bicaudatus (Ephemeroptera), whose females lay eggs under rocks protruding from high-elevation streams in western Colorado; and (2) test experimentally selective oviposition of mayfly females. We surveyed the number and physical characteristics of potential oviposition sites, and counted the number and density of egg masses in different streams of one watershed throughout two consecutive flight seasons. Results of surveys showed that variability in the proportion of protruding rocks with egg masses and the density of egg masses per rock were explained primarily by seasonal and annual variation in hydrology, and variation in geomorphology among streams. Moreover, surveys and experiments showed that females preferred to oviposit under relatively large rocks located in places with high splash associated with fast current, which may provide visual, mechanical or both cues to females. Experiments also showed that high densities of egg masses under certain rocks were caused by rock characteristics rather than behavioral aggregation of ovipositing females. While aggregations of egg masses provided no survival advantage, rocks selected by females had lower probabilities of desiccating during egg incubation. Our data suggest that even when protruding rocks are abundant, not all rocks are used as oviposition sites by females, due to female selectivity and to differences in rock availability within seasons, years, or streams depending on variation in climate and hydrogeomorphology. Therefore, specialized oviposition behavior combined with variation in availability of quality oviposition substrata has the potential to limit recruitment of this species.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Environmental constraints on oviposition limit egg supply of a stream insect at multiple scales

Species with complex life cycles pose challenges for understanding what processes regulate population densities, especially if some life stages disperse. Most studies of such animals that are thought to be recruitment limited focus on the idea that juvenile mortality limits the density of recruits (and hence population density), fewer consider the possibility that egg supply may be important. For species that oviposit on specific substrata, environmental constraints on oviposition sites may limit egg supply. Female mayflies in the genus Baetis lay egg masses on the underside of stream rocks that emerge above the water’s surface. We tested the hypothesis that egg mass densities are constrained by emergent rock densities within and between streams, by counting egg masses on emergent rocks. All emergent rocks were counted along 1-km lengths of four streams, revealing significant variation in emergent rock density within streams and a more than three-fold difference between streams. In each stream, egg mass density increased with the density of emergent rocks in 30-m stretches. We used regression equations describing these small-scale relationships, coupled with the large-scale spatial variation of emergent rocks, to estimate egg mass densities for each 1-km stream length, a scale relevant to population processes. Scaled estimates were positively associated with emergent rock density and provided better estimates than methods that ignored environmental variation. Egg mass crowding was inversely related to emergent rock density at the stream scale, a pattern consistent with the idea that oviposition substrata were in short supply in streams with few emergent rocks, but crowding did not compensate entirely for differences in emergent rock densities. The notion that egg supply, not larval mortality, may limit population density is an unusual perspective for stream insects. Environmental constraints on egg supply may be widespread among other species with specialised oviposition behaviours.     

A test of the preference–performance hypothesis with stream insects: selective oviposition affects the hatching success of caddisfly eggs

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Hydrologic and behavioral constraints on oviposition of stream insects: implications for adult dispersal

The supply of recruits plays an important role in plant and animal population dynamics, and may be governed by environmental and behavioral constraints on animals. Mated females of the mayfly genus Baetis alight on rocks protruding from streams, crawl under water and deposit a single egg mass under a rock. We surveyed oviposition and emergence of a bivoltine population of B. bicaudatus in multiple stream reaches in one high-altitude watershed in western Colorado over 3 years to establish qualitative patterns at a regional scale (entire watershed), and quantitative patterns over six generations at a local scale (one stream reach). We also measured characteristics of preferred oviposition substrates, performed experiments to test hypotheses about cues used by females to select oviposition sites, and measured mortality of egg masses in the field. Our goals were to determine whether: (1) hydrologic variation necessitated dispersal of females to find suitable oviposition sites; (2) the local supply of females could provide the supply of local recruits; and (3) local recruitment determined the local production of adults. The onset of oviposition corresponded with the decline of spring run-off, which differed dramatically among years and among sites within years. However, eggs appeared before any adults had emerged in 8 of 22 site-years, and adults emerged 2-3 weeks before any eggs were oviposited in 3 site-years. Furthermore, the size distribution of egg masses differed from that predicted by the size distribution of females that emerged from seven of nine stream reaches. Protruding rocks and eggs appeared earlier each summer in smaller tributaries than in larger mainstream reaches, suggesting that hydrologic and behavioral constraints on oviposition may force females to disperse away from their natal reach to oviposit, and possibly explain the predominantly upstream flight of Baetis females reported in other studies. Local oviposition rates in one third-order stream-reach increased rapidly as soon as substrates protruded from the water surface, and females preferred large rocks that became available early in the flight season. However, females oviposited on <10% of all available rocks, and <65% of preferred rocks as determined by an empirical model. These data indicated that the timing of appearance of suitable oviposition sites determined the phenology of local recruitment, but that preferred oviposition sites were not saturated. Thus, the magnitude of local recruitment was not limited by the absolute abundance of preferred oviposition sites. Only 22% of egg masses observed in the field suffered mortality during their embryonic development, and per capita Baetis egg mass mortality was significantly lower on rocks with higher densities of egg masses. Thus, we suspect that specialized oviposition behavior may reduce the probability of egg mortality, potentially compensating for the costs of dispersal necessary to locate suitable oviposition sites. Finally, the number of adults that emerged at one stream reach was independent of the number of egg masses oviposited over six generations of Baetis; and local recruitment was not a function of the number of adults of the previous generation that emerged locally. The patterns of oviposition and emergence of Baetis found in this study are consistent with the following hypotheses. Recruitment of eggs in a stream reach is not limited by the local supply of adults, but is a function of the regional supply of dispersing adults, which are constrained by the spatial and temporal distribution of preferred oviposition habitat. Furthermore, subsequent local production of adults is not a function of the supply of recruits, arguing for post-recruitment control of local populations by processes operating in the larval stage (e.g., predation, competition, dispersal, disturbance). Processes affecting larval and adult stages of Baetis act independently and at different scales, thereby decoupling local population dynamics of successive generations.     

Aggregative oviposition varies with density in processionary moths—Implications for insect outbreak propensity

1. In gregarious insects, groups commonly originate from females laying eggs in masses and feeding groups are established as soon as larvae hatch. Some group-living insect species may aggregate beyond the individual parent level, such that offspring from two or more egg masses develop within a common resource.
2. Here we show that aggregative oviposition can vary with population density at oviposition and possibly be an important factor in outbreak dynamics of phytophagous insects.
3. We analysed density data with respect to egg mass aggregation for two species of pine processionary moths, Thaumetopoea pinivora (in Sweden 2005–2019) and T. pityocampa (in Spain 1973–1991). Both species lay their eggs in egg masses and feed in groups. During the study periods, insect population density for both species varied by at least an order of magnitude.
4. The two species showed strikingly similar patterns of egg mass aggregation. Egg masses were overdispersed at high population density, with few trees showing a high load of egg masses.
5. Our data suggest that aggregative oviposition can be important in explaining the previously documented higher propensity for outbreaks in insects laying eggs in clusters, compared with those laying individual eggs.

     

Experimental evidence for physical cues involved in oviposition site selection of lotic hydrobiosid caddis flies

Cues involved in determining the distribution of invertebrate propagules within a stream landscape contribute greatly to our knowledge of the supply and arrangement of new recruits and thus an improved understanding of factors that might ultimately affect population parameters. Previous observations indicated that both current velocity and rock size were important determinants of the egg mass distribution of certain hydrobiosid caddis flies that lay their eggs in single masses beneath emergent rocks. These observations were tested experimentally in a temperate, upland Australian stream. Manipulations of current speed confirmed that females of Ulmerochorema sp. and the Taschorema complex deposited more eggs on rocks in elevated current speeds (>0.40 ms(-1)) whereas Apsilochorema sp. deposited more eggs on rocks in slow currents (<0.30 ms(-1)). This latter result did not coincide with previous observational data. The anomaly between observational and experimental data, however, was reconciled by the outcome of a further experiment that tested the influence of the emergent or 'landing pad' size of rocks as the abundance of Apsilochorema egg masses increased with landing pad size independent of the prevailing flow conditions. Landing pad size did not influence the abundance of egg masses of Taschorema or Ulmerochorema. Patterns of female visits to rocks indicated that taxa might distinguish between favoured egg-laying sites prior to landing on rocks. Large aggregations of adult male and female Ulmerochorema collected from rocks favoured for oviposition provide indirect evidence for mating induced swarming behaviour associated with oviposition site selection. This study provides a framework for more sophisticated questions relating to the influence of oviposition site selection on structuring populations of lotic macroinvertebrates.     

Recognition and avoidance of previously laid egg masses by the oblique-banded leafroller (Lepidoptera: Tortricidae)

Investigations were conducted on oviposition by laboratory-reared oblique-banded leafrollers, Choristoneura rosaceana Harris. In paperbag oviposition chambers, females deposited their egg masses in a significantly nonrandom, uniform manner in 16 of 20 replicates. Inpetri dish oviposition chambers, gravid females were exposed to egg masses recently deposited by other females. As determined by the magnitude of the mean direction vector of the second egg mass in relation to the first, females detected and avoided egg masses laid by other females. In addition, they avoided waxed paper from which a recently oviposited egg mass had been removed. This phenomenon is most likely due to a pheromone laid down with each egg mass.     

Seasonal decline in plant defence is associated with relaxed offensive oviposition behaviour in the viburnum leaf beetle Pyrrhalta viburni

1. Plant defence of Viburnum shrubs against oviposition by its specialist herbivore, the viburnum leaf beetle [VLB Pyrrhalta viburni(Paykull)], involves an egg‐crushing wound response in twigs. Although the response is variable among Viburnum species, it can have a strong impact on egg survivorship. Beetles typically aggregate egg masses with conspecifics along infested twigs, forming clusters that can overwhelm the twig response. It was investigated whether twig responses and beetle oviposition behaviour vary seasonally. 2. In a field experiment, twig defences decreased towards the end of the VLB oviposition period: wound response of the North American Viburnum dentatum L. and the European V. opulus L. was reduced by 100% and 54%, respectively, in September compared with the July to August period. 3. Oviposition trials demonstrated a corresponding behavioural change: VLB females displayed aggregative oviposition in August, but not in September. 4. Further tests revealed that late‐season VLB females reverted to aggregative oviposition after being kept on uninfested twigs, whereas females kept on heavily infested twigs did not. This behavioural change suggests that relaxation of aggregative oviposition originates from cues associated with high densities of egg masses. 5. Relaxation of aggregative oviposition may be adaptive (and beneficial for invasion) on shrubs with low levels of defences by reducing intra‐specific competition.     

Brochosome influence on parasitisation efficiency of Homalodisca coagulata (Say) (Hemiptera: Cicadellidae) egg masses by Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae)

Abstract. 1. Many cicadellid females in the tribe Proconiini (Hemiptera: Cicadellidae) cover their egg masses with specialised, usually rod‐shaped, brochosomes as the eggs are being laid. The brochosomes are produced in Golgi complexes in the Malpighian tubules of Cicadellidae. In contrast to the gravid females, adult males, pre‐reproductive adult females, and nymphal males and females produce specialised, usually spherically shaped brochosomes. Brochosomes are also used to cover the external surfaces of nymphs and newly moulted adult males and females. 2. The function of the brochosome covering the egg masses is unknown but various hypotheses have been suggested, including protecting the eggs against pathogens, predators, and parasitoids. Based on preliminary observations of Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae) parasitising the eggs of the cicadellid, Homalodisca coagulata (Say), it is speculated here that brochosomes covering an egg mass hinder parasitisation of eggs by G. ashmeadi. This hypothesis was tested by observing G. ashmeadi females foraging on leaves with H. coagulata egg masses heavily covered with rod‐shaped brochosomes vs. those lacking brochosomes. 3. Cox's proportional hazards model was used to evaluate the probability, per unit time, that a female G. ashmeadi displayed the sequence of behaviours that ended in successful oviposition as influenced by five variables: (a) presence or absence of brochosomes on an egg mass, (b) the leaf surface, upper or lower, being searched by the parasitoid (the egg masses are laid in the parenchyma on the lower leaf surface), (c) the parasitoid's previous ovipositional experience, (d) egg mass size, and (e) the parasitoid's age. 4. Brochosomes significantly decreased oviposition efficacy of G. ashmeadi females. Scanning electron microscopy showed that females exposed to brochosome‐covered egg masses had brochosomes adhering to their tarsi, legs, antennae, and eyes, all of which prompted extensive bouts of grooming.     

A quantified ethogram for oviposition behavior and oviposition preference in the hemipterophagous butterfly Spalgis epius (Westwood) (Lepidoptera: Lycaenidae)

In this study, we examined the oviposition behavior and preference of Spalgis epius, a potential predator of mealybug crop pests. An ethogram of oviposition behavior was constructed based on observations made in an oviposition cage. Ovipositional behavioral acts were catalogued and separated into two behavioral repertoires: searching and egg laying. Gravid females of S. epius oviposited similar numbers of eggs on three mealybug species. Females preferred eggs and adults to nymphs of mealybugs for oviposition. Among three species of mealybugs attended by ants, females laid fewer eggs in the mealybug mass attended by Oecophylla smaragdina than on mealybugs attended by Tapinoma melanocephalum and Camponotus variegatus. Females preferred mealybug masses already containing conspecific eggs to mealybug masses containing conspecific larvae or Cryptolaemus montrouzieri larvae for egg deposition. Gravid females laid larger numbers of eggs under bright sunlight than in diffused sunlight or shade. The results of this study showed that S. epius can effectively attack any species of mealybugs, avoid intra- and interspecific competition, and co-exist with some species of ants attending mealybugs. With the knowledge of these behaviors, this predator can be effectively utilized as a major biological control agent of mealybugs.     

Large‐scale manipulation of oviposition substrata affects egg supply to populations of some stream‐dwelling caddisflies

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Lasting effects of maternal behaviour on the distribution of a dispersive stream insect

1. Predicting population dynamics at large spatial scales requires integrating information about spatial distribution patterns, inter-patch movement rates and within-patch processes. Advective dispersal of aquatic species by water movement is considered paramount to understanding their population dynamics. Rivers are model advective systems, and the larvae of baetid mayflies are considered quintessential dispersers. Egg laying of baetids along channels is patchy and reflects the distribution of oviposition sites, but larvae are assumed to drift frequently and far, thereby erasing patterns created during oviposition. Dispersal kernels are often overestimated, however, and empirical tests of such assumptions are warranted because of the pivotal role distribution patterns can have on populations. 2. We tested empirically whether the egg distribution patterns arising from oviposition behaviours persisted and were reflected in the distribution patterns of larval Baetis rhodani. In field surveys, we tested for associations between egg mass and larval densities over 1 km lengths of four streams. A control species, the mayfly Ephemerella ignita, was employed to test for covarying environmental factors. We estimated drift rates directly to test whether larvae dispersed between riffles (patches of high egg mass density) and whether drift rates were density-dependent or density-related - expected outcomes if drift erases patterns established by maternal behaviours. 3. Positive associations between egg masses and larval benthic densities were found for neonate and mid-stage larvae of Baetis, but not the control species, suggesting persistence of the patchy distribution patterns established at oviposition. Drift rates were high, and riffles were net exporters of neonate and mid-stage larvae, but drift rates were unrelated to benthic densities and few drifters reached the next riffle. Riffles were sinks for large larvae, suggesting ontogenetic shifts in habitat use, but little long-distance dispersal. 4. Overall, the results suggest that most neonate and mid-stage larvae of B. rhodani remain close to the natal riffle, and late-stage larvae disperse shorter distances than routinely assumed. The persistence of maternal effects on distribution patterns well into juvenile life of an allegedly iconic disperser suggests that traditional models of how dispersal influences the population dynamics of many lotic invertebrates may be incorrect.     

Effects of fruit fly host, fruit species, and host egg to female parasitoid ratio on the laboratory rearing of Biosteres arisanus

The host suitability of the oriental fruit fly, Bactrocera dorsalis (Hendel), for development of Biosteres arisanus (Sonan), a braconid parasitoid, was compared with three other fruit fly species, namely, Mediterranean fruit fly, Ceratitis capitata Weidemann, melon fly, Bactrocera cucurbitae Coquilett, and Malaysian fruit fly, Bactrocera latifrons (Hendel). In addition, effects of five different fruit species, namely, Carica papaya L. (solo papaya), Musa sapientum (L.) O. Ktze. (apple banana), Mangifera indica (L.) (Haden mango), Terminalia catappa (L.) (false kamani), and Citrus aurantiifolia (Christman) Swingle (common lime), on the parasitization rate of B. dorsalis and sex ratio of parasitoid progenies were evaluated. Effects of host egg to female B. arisanus ratios on parasitoid progeny yields were likewise determined. The host suitability of fruit flies for development of B. arisanus was ranked as: B. dorsalis>C. capitata=B. latifrons=B. cucurbitae. Based on percent parasitization of B. dorsalis, preference of B. arisanus females for host eggs varied with fruit species, however, preferential oviposition displayed by female parasitoids did not influence sex ratios of subsequent parasitoid progenies. Increases in host egg to female parasitoid ratios of 5:1, 10:1, 20:1, 25:1, and 30:1 corresponded with increases in parasitoid progeny yield reaching a plateau at 20:1.     

Insect oviposition plasticity in response to host availability: the case of the tephritid fruit fly Bactrocera dorsalis

1. Insect oviposition behaviour is ecologically and physiologically plastic. For tephritid fruit flies, Bactrocera dorsalis Hendel, host availability varies spatially and temporally. Females are expected to adopt adaptive oviposition strategies to maximise lifetime reproductive fitness, including survival. Bactrocera dorsalis oviposition tactics in response to different host availabilities were investigated. 2. This study includes three treatments: (i) variable host densities (host density varied according to a fixed cycle from day to day over values of 1, 5, 10 and 20 hosts per cage), (ii) a fixed high host density (20 hosts per cage), and (iii) a fixed low host density (1 host per cage). 3. Daily egg‐laying number per female over the course of 27 days was entirely independent of host density and highly dependent on female age. As host availability increased, females accepted significantly more hosts, generally laid small egg clutches, and more broadly distributed the eggs. 4. Tephritid fruit flies adaptively adjusted egg clutches in ways that reflected the variability of host availability. Egg‐ and time‐limitation constraints appeared to drive these adjustments. Female egg maturation was triggered by oviposition activity and reflected marked lifetime trade‐offs. Such strategies involved specific time schedules for egg laying. 5.This study defined the oviposition plasticity of the tephritid fruit fly. These results have general implications for the behavioural ecology of insect herbivores and parasitoids.     

Active female courtship behavior and male nutritional contribution to female fecundity in Bruchidius dorsalis (Fahraeus) (Coleoptera: Bruchidae)

Mating behavior and the male's contribution to female fecundity were studied in the bean weevil Bruchidius dorsalis (Fahraeus) (Coleoptera: Bruchidae) in comparison with two other species, Callosobruchus chinensis (which infests stored beans) and Kytorhinus sharpianus (which feeds on wild legumes). Only females of B. dorsalis showed multiple mating and characteristic precopulatory behavior that appeared to solicit the male's nutritious secretion. In contrast, all females of the other two species did not copulate multiply and did not show such precopulatory behavior. In B. dorsalis, the decrement of male body weight just after copulation indicated that seminal fluid weighing as much as approximately 7% of the male's body weight was transferred to the female. Fecundity was more than eight times higher in females that had copulated ten times than in females that had copulated only once, indicating that males paid most of the nutritional cost of egg production. These facts suggest that the sex role is reversed in B. dorsalis. Received: May 22, 1998 / Accepted: July 19, 1999