The coadaptation of female morphology and offspring size: a comparative analysis in crickets

Few studies of invertebrates have considered combinations of morphological and life history traits in the context of the evolution of reproductive strategies. Cricket species that exploit habitats harsh with respect to egg survival have evolved a long ovipositor, presumably because laying deep in the soil reduces egg mortality. Yet hatchling mortality increases with laying depth, and the ability of hatchlings to climb through the soil increases with egg size. Thus a conflict may exist between survival of the egg and that of the hatchling, inducing a positive covariation between egg size and ovipositor length across species evolving under contrasting selective habitats. We used the phylogenetic autocorrelation method and a path analysis to assess whether egg size coevolved with ovipositor length across 40 species of crickets, and whether egg size was affected by body size or ecological factors that influence egg mortality. Body size and ovipositor length were affected by taxonomic association, whereas common ancestry had no significant effect on egg size, diapausing strategy, and oviposition preference for soil types. The path model indicated that 29.11% of the variance in egg size was explained by independent evolution. As expected, ovipositor length was positively correlated with egg size, and species diapausing in the egg stage produced larger eggs than crickets diapausing in the nymphal stage or with no diapause. Ovipositor length and diapausing strategy were the first and second most important traits, respectively, in term of the proportion of variance in egg size explained by specific values. These results support the hypothesis that the ability of hatchlings to climb through the soil, and variation in diapause strategies, are general selective factors affecting the evolution of egg size in crickets. Phylogeny explained 51.01% of the variance in egg size. Egg size in a current cricket species, however, was not directly determined by egg size in its ancestor. Instead, it was strongly related to the phylogenetic values of body size and ovipositor length. Such indirect phylogenetic effects of body size and ovipositor length may have arisen because clades originating from ancestors with different ovipositor lengths experienced different selective pressures on egg size. Recelived: 13 October 1995 / Accepted: 30 September 1996     

Quantitative genetics of oviposition behaviour and interactions among oviposition traits in the sand cricket

We performed a quantitative genetic study of oviposition behaviours and oviposition traits in the sand cricket Gryllus firmus. Egg survival in crickets depends on the depth at which they are inserted into the soil with the ovipositor. We examined whether egg depth depends on ovipositor length alone, or on both morphological and behavioural traits associated with oviposition. Heritability estimates were high (h² >0.5) for ovipositor length and small (h²=0.2) for oviposition behaviours. Negative genetic correlations between ovipositor length and some behavioural traits (digging depth and the behavioural component of egg depth) indicated compensation between oviposition traits on egg depth. Because of behavioural compensation, females with different ovipositor lengths subject to stabilizing selection on egg depth could have equal fitnesses. Females laid their eggs deeper, and their eggs were marginally more evenly distributed in dry than in wet sand. This suggests adaptive phenotypic plasticity in laying behaviour, but may also result from physical constraints of the substrate on the insertion of the ovipositor. The absence of significant between-family variation in oviposition traits in response to sand moisture indicates low evolutionary potential for phenotypic plasticity in oviposition traits according to soil moisture. In highly unpredictable environments, females could spread the risk of desiccation by laying eggs at different depths independently of environmental conditions (bet hedging). Our results show significant additive genetic influences on the ability of a female to spread risks as measured by genetic variation in egg distribution, suggesting that a bet-hedging strategy of egg laying has the potential to evolve in this population. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Climatic adaptation and species status in the lawn ground cricket

Summary The cricket tentatively identified as Pteronemobius taprobanensis shows a stepwise pattern of latitudinal variation in ovipositor length. Abrupt elongation of ovipositor at about 28° N marks the replacement of the subtropical form by the temperate form. The latter maintains almost a constant ovipositor length up to about 35° N within the bivoltine area. Further north in the univoltine area, an ascending cline extends to about 39° N, beyond which no further increase occurs. The ovipositor length adjusted for body size shows a northeastward increase in each of the univoltine and bivoltine areas, though this tendency is less clear in the latter. The optimum length of ovipositor would vary with the relative amounts of gain (due to the protection of eggs in the soil) and loss (due to the metabolic cost and the difficulty to emerge from the soil) in fitness. If so, it may be predicted that a longer ovipositor would be selected for, when the metabolic cost and the risk at hatching are smaller, the environmental pressure in the egg stage is stronger, and the egg stage lasts longer. The last parameter seems to be mainly responsible for the observed association of the type of life cycle with the geographic pattern of variation in ovipositor length.Contribution No. 76 from the Laboratory of Entomology, Hirosaki University     

Seasonal change in the spatial utilization of host mussels in relation to ovipositor length by female rosy bitterling Rhodeus ocellatus kurumeus

The seasonal changes in ovipositor length and utilization patterns of mussels for oviposition in the rosy bitterling Rhodeus ocellatus kurumeus (Cyprinidae) were investigated in a field experiment and field surveys during the breeding period (April to August). The mean length of ovipositors at oviposition was short at the start (early April) and end (July) of the breeding period. Females with long ovipositor at oviposition were collected between mid‐April and June. Mark‐and‐recapture data showed that ovipositor length at oviposition changed rhythmically throughout the breeding period, shortening and lengthening as the female entered the spawning and resting phase. The density of rosy bitterling embryos in mussels increased between April and June, peaking in May, but decreasing in July. The position of eggs on mussel gills varied from close to the exhalant siphon to deeper inside the gill during April, and periodically thereafter. There was a positive correlation between ovipositor length at oviposition and the distance from exhalant siphon of mussels to eggs deposited by females, suggesting that ovipositor length at oviposition determined the position of eggs deposited on a mussel gill. Because dissolved oxygen in mussel gills decreased with the density of bitterling embryos, suitable positions for embryo survival in gills changed with embryo density. By changing ovipositor length at oviposition, females might be able to spawn their eggs in a position that maximizes embryo survival. Thus, plasticity in ovipositor length at oviposition may play an important role as an adaptation of rosy bitterling in utilizing mussels when their quality as a spawning substratum fluctuates seasonally.     

高体鳑鲏的性选择问题研究

以高体鳑鲏为研究对象,分别设置个体大小不同的雌鱼和雄鱼进行性选择实验并验证性选择理论,对雌鱼的怀卵量和雄鱼的空间学习记忆能力也进行了检测。雄鱼对雌鱼的性选择实验结果表明,雄鱼对大个体雌鱼有明显的选择偏好;雄鱼选择雌鱼的次数及停留时间与雌鱼的体长、体高、产卵管长度等特征呈正相关。推测雄鱼是基于直接收益的角度,即最大限度的获得较多的后代及较高的后代存活率对雌鱼进行选择,因为大个体雌鱼拥有长的产卵管及绝对多的处于Ⅴ时相的成熟卵粒,可以提供较多的后代及较高的后代存活率。雌鱼对雄鱼的性选择结果表明,雌性高体鳑鲏对大个体雄鱼也表现出了明显的偏好;雌鱼选择雄鱼的次数及停留时间与雄鱼的体长、体高呈正相关;大个体雄鱼的空间学习记忆能力比小个体强,能更快捷地找到并占领质量好的河蚌而成为领域鱼。从直接收益的观点看,雌鱼选择大个体的雄鱼可能是因为大个体雄鱼可以更加容易、迅速和便捷地寻找到质量好的河蚌引领雌鱼进行产卵,从而保证子代的存活率。       

Neural mechanism for generating and switching motor patterns of rhythmic movements of ovipositor valves in the cricket

In adult female crickets (Gryllus bimaculatus), rhythmic movements of ovipositor valves are produced by contractions of a set of ovipositor muscles that mediate egg-laying behavior. Recordings from implanted wire electrodes in the ovipositor muscles of freely moving crickets revealed sequential changes in the temporal pattern of motor activity that corresponded to shifts between behavioral steps: penetration of the ovipositor into a substrate, deposition of eggs, and withdrawal of the ovipositor from the substrate. We aimed in this study to illustrate the neuronal organization producing these motor patterns and the pattern-switching mechanism during the behavioral sequence. Firstly, we obtained intracellular recordings in tethered preparations, and identified 12 types of interneurons that were involved in the rhythmic activity of the ovipositor muscles. These interneurons fell into two classes: ‘initiator interneurons’ in which excitation preceded the rhythmic contractions of ovipositor muscles, and ‘oscillator interneurons’ in which the rhythmic oscillation and spike bursting occurred in sync with the oviposition motor rhythm. One of the oscillator interneurons exhibited different depolarization patterns in the penetration and deposition motor rhythms. It is likely that some of the oscillator interneurons are involved in producing different oviposition motor patterns. Secondly, we analyzed oviposition motor patterns when the mecahnosensory hairs located on the inside surface of the dorsal ovipositor valves were removed. In deafferented preparations, the sequential change from deposition to withdrawal did not occur. Therefore, the switching from deposition pattern to withdrawal pattern is signaled by the hair sensilla that detect the passage of an egg just before it is expelled.     

The reproductive ecology of two subspecies of the bitterling Rhodeus atremius (Cyprinidae, Acheilognathinae)

The reproductive ecology of two endangered subspecies of the bitterling Rhodeus atremius was investigated in two rivers in Okayama and Fukuoka Prefectures, Japan. Several other bitterling species, R. ocellatus, Tanakia limbata, T. lanceolata, Acheilognathus tabira and A. rhombeus, are also found in these rivers. Spawning of both R. atremius subspecies was recorded between late March and August 2007, peaking between April and July. Females of R. atremius began to mature at small body size (minimum 25 mm). They had small clutch sizes (maximum 16 eggs) and developed a short ovipositor (mean 13.6 mm), which was used to deposit a bulb-shaped egg of comparatively large size (3.2 mm³, major and minor axes 2.8 and 1.5 mm). Clutch size, ovipositor length and egg size appear to be a function of female body size. Both the body size and ovipositor length of R. atremius with ripe ova and the size and shape of the eggs varied seasonally. Rhodeus a. suigensis achieved maturity at a smaller body size, had a longer ovipositor and produced smaller eggs, with the minor axis being wider relative to the major axis than those of R. a. atremius. Rhodeus atremius may be adapted for depositing small numbers of eggs in a much shallower position inside the gills of smaller host mussels than other bitterling species.     

Comparative effect of various powder diets on growth of big-eyed bug Geocoris ochropterus (Fieber) (Hemiptera: Geocoridae)

Application of powder diets may promote development of the predator Geocoris ochropterus (Fieber) when densities are low in greenhouses or fields. Development and nymphal survival ratio of G. ochropterus fed on different powder diets including silkworm pupae (A), crickets (B), silkworm pupae and sunflower seed (C) and crickets and sunflower seed (D) were examined. Total average development period was 30.14 days for instars fed on silkworm pupae, 34.43 days for instars fed on crickets, 26.28 days for instars fed on silkworm pupae and sunflower seed and 29.67 days for instars fed on crickets and sunflower seed, and they were significantly different. Head width, body length, forewing length, dry body weight of adults and sex ratio were not affected by the different diets. There were significantly different found in survival rates among the diets when the nymph reached 5th instar. Results indicate that the combination powder of silkworm pupae and sunflower seed will promise as an alternative diet in the greenhouses to maintain the population of G. ochropterus.     

Can there be an escalating arms race without coevolution? Implications from a host-parasitoid simulation

Summary Under a restricted set of conditions, predator-prey or parasite-host systems may exhibit an escalating arms race over several generations that is not coevolutionary. Preconditions for such a process include high correlation between prey/host quality and defensive capability, and phenotypic plasticity in predator/parasite-counter defenses that responds to quality. We present simulation models based on the parasitoid waspEurytoma gigantea, which lays its egg in the goldenrod gall induced by the flyEurosta solidaginis. For the parasitoid to successfully lay an egg, the gall walls must be thinner than the parasitoid's ovipositor is long. Wall thickness is highly correlated with gall size, so probability of successful attack declines with gall size. However, since the parasitoid eats the gall tissue, individuals developing in small galls have little food and mature with shorter ovipositors than those which develop in large galls. The simulation showed that the population mean parasitoid size is set by mean gall size. Since small galls are more frequently parasitized, there is a selection pressure on the gallmaker to induce larger galls. But, an additional simulation showed that since parasitoid ovipositor length depends on gall size, an evolutionary increase in gall size will also result in a non-evolutionary increase in parasitoid body size and ovipositor length over several generations.     

Effects of the Host Species and the Number of Parasitoids per Host on the Size of Some Trichogramma Species (Hymenoptera: Trichogrammatidae)

The size of some Trichogramma spp. adults and especially the ovipositor length depends on the species, but is also related to the host species and to the number of parasitoids per host. The length is greater in T. evanescens than in T. pretiosum itself greater than in T. exiguum, but the width is similar in the three species. For T. evanescens, the size obtained in Mamestra brassicae host when three or four insects emerged is similar to that obtained in Ephestia kuehniella host when singly parasitized. The size of the ovipositor is important because it may influence the possibility of in vitro egg laying in artificial host eggs. A shorter or a narrower ovipositor could cause difficulties in egg-laying into artificial host eggs composed of a membrane of unsuitable thickness.     

Does natural selection alter genetic architecture? An evaluation of quantitative genetic variation among populations of Allonemobiussocius and A. fasciatus

To make long-term predictions using present quantitative genetic theory it is necessary to assume that the genetic variance–covariance matrix ( G ) remains constant or at least changes by a constant fraction. In this paper we examine the stability of the genetic architecture of two traits known to be subject to natural selection; femur length and ovipositor length in two species of the cricket Allonemobius. Previous studies have shown that in A. fasciatus and A. socius natural selection favours an increased body size southwards but a decreased ovipositor length. Such countergradient selection should tend to favour a change in G . In the total sample of eight populations of A. socius and one of A. fasciatus we show that there is significant variation in all genetic covariance components, i.e. V_A for body size, V_A for ovipositor length, and Cov_A. This variation results entirely from an increase in the covariances of A. fasciatus. However, although larger, these components are approximately proportionally increased, thereby leading to no statistically significant change in the genetic correlation. A proportional increase in the covariance components is consistent with changes resulting from genetic drift. On the other hand, the genetic covariance components are significantly correlated with the length of the growing season suggesting that the change in the genetic architecture is the result of selection and drift.     

Sex differences in survival selection in the serin,Serinus serinus

Natural selection is demonstrated in most natural populations which suggests that populations are dispatched from their adaptive peaks as a result of selection on correlated characters, or conflicting selection between the sexes. We analysed patterns of survival selection in a population of serins (Serinus serinus) outside Barcelona over a period of 13 years. There was directional selection for increased wing length in males and females accompanied by strong disruptive selection on both tail and wing length in males and a selection against a positive correlation between the two characters in males. In females there was directional selection for increased bill width but decreased bill depth, which should be contrasted to the stabilizing selection acting on bill depth in males. There were conflicting selection on the characters within a sex and conflicting selection of the same characters between sexes, which constrain the rate of access to the nearest adaptive peak.     

Relationship Between Condition,Aggression, Signaling,Courtship, and Egg Laying in the Field Cricket,Gryllus assimilis

Sexual selection theory suggests males in good condition should be more successful than males in poor condition when competing with rivals for territories and mates. Understanding how condition influences the interplay between aggression, mate attraction, and courtship displays could help explain why variation is maintained in traits that confer fitness. Using laboratory‐reared Jamaican field crickets, Gryllus assimilis, we found that fine‐scale temporal components of mate attraction signals were positively correlated with body condition (residual body mass) and body size; signaling effort was positively correlated with both body condition and fine‐scale temporal signaling components; aggression was positively correlated with signaling effort; number of eggs laid was positively correlated with female body size, male body condition and aggression. Together our correlative study suggests that variation in body condition and size may drive some of the variation in cricket mate attraction signaling and aggression. Given condition and body size are influenced by foraging ability, nutrient availability and the organism’s ability to uptake and retain these essential nutrients could explain some of the persistent variation in fitness conferring traits.     

Viability selection on overwintering eggs in a field cricket mosaic hybrid zone

Habitat segregation allows recently diverged taxa to minimize competition and maximize fitness. Consequently, the overall distribution of a species in part will be determined by interactions with their close relatives. Two recently diverged but hybridizing field crickets, Gryllus firmus and G. pennsylvanicus , form a mosaic hybrid zone in Connecticut in which each species is associated with a specific soil type: G. firmus is found on sand soils and G. pennsylvanicus is found on loam soils. Both species produce diapausing eggs, which spend the winter in the ground. We investigate how viability selection on overwintering eggs in different soils influences the distributions of the two species, habitat partitioning between them, and the structure of the mosaic hybrid zone. Our results suggest that selection on eggs by soil types is not important in determining the success of crickets on different soils. However, winter climate has a strong effect on egg viability, and variation in egg survivorship over different winter climate regimes likely plays an important role in determining the distribution of these cricket species and the position of the hybrid zone.     

What is driving male mate preference evolution in Jamaican field crickets?

Male mating preferences are often a neglected aspect of studies on sexual selection. Male mating preferences may evolve if they provide males with direct‐fitness benefits such as increased opportunity to fertilize more eggs or indirect‐fitness benefits such as enhanced offspring survival. We tested these ideas using Jamaican field crickets, Gryllus assimilis, previously shown to exhibit male mating preferences. We randomly mated males to either their preferred or non‐preferred potential mates and then asked whether mating treatment influenced egg oviposition or offspring viability. Preferred females were not significantly more fecund and did not produce more viable eggs or offspring than non‐preferred females. Male mate preferences were therefore inconsistent with both the direct‐ and indirect‐fitness benefits hypotheses under the conditions of our experiment. Our null results leave us with an open question about what is driving the evolution of mating preferences in male crickets. Future research should explore the whether the offspring of preferred females are more attractive, have stronger immune systems, and/or experience higher adult longevity.     

Identity and Relative Importance of Egg Predators of Rice Leaffolders (Lepidoptera: Pyralidae)

Field andlaboratory studies on predation of rice leaffolder eggs (i.e., Cnaphalocrocis medinalis (Guenée) and Marasmia patnalis Bradley) were conducted to identify major predator species. Direct observations of predation on field-exposed eggs showed that in two seasons Metioche vittaticollis (Stål) and Anaxipha longipennis (Serville) were the major predators of leaffolder eggs. Together these crickets took the largest part of all eggs consumed during observation (92%) and had the highest ratio of visits with predation to their total observed visits to plants with leaffolder eggs. Furthermore, the activity pattern of the crickets matched best the daily pattern of egg disappearance, and the seasonal trends in their observed visits correlated best with the seasonal trends in egg disappearance. Minor predators feeding on field-exposed rice leaffolder eggs were Ophionea nigrofasciata Schmidt-Goebel, Micraspis sp., and Conocephalus longipennis (de Haan). The latter species was the most commonly observed egg predator, but had a negligible share in the total predation. In petri dish tests the consumption of leaffolder eggs by the predatory crickets M. vittaticollis and A. longipennis was far greater than that of four other predators. Female cricket adults consumed at least 80 eggs per day, and all individuals accepted leaffolder eggs as food. According to daily egg consumption and acceptance rates, the predators ranked as follows: M. vittaticollis, A. longipennis > Micraspis sp. > O. nigrofasciata > Paederus fuscipes Curtis, C. longipennis. Predator ranking according to the ratio of visits with predation to total visits in the field was identical to the ranking based on the egg consumption tests. Due to their large predation potential, predatory crickets will probably play an important role in leaffolder egg predation, even when their densities are low compared to those of other predator species.     

Initial preference for plant species and state during oviposition site selection by an odonate

Endophytic egg‐laying odonates use an ovipositor to insert their eggs inside plant tissues. Before egg deposition, oviposition site selection consists of two crucial steps: (i) the initial choice, typically decided in species that oviposit in tandem within vertical substrates by the male when landing and then by the female by staying on the substrate or flying to another; and (ii) the insertion site choice, made by the female who uses her ovipositor to palpate the substrate. Some odonates prefer to deposit their eggs within specific plant species. Some are able to discriminate between living and dead substrates during the initial choice. However, the extent to which odonates discriminate among distinct plant species during the initial choice is unknown. We studied the initial site preference in Lestes macrostigma (Odonata: Lestidae) to determine whether the males and/or females show a distinct preference among five types of shoots when landing on or when palpating the substrate, respectively. Male L. macrostigma preferred to land on Bolboschoenus maritimus and dead Juncus spp. When focusing on J. maritimus, females preferentially palpated the substrate when the male landed on dead shoots. We suggest that the male preference for these substrates is consistent with that of the female during insertion site choice but also during egg deposition. Such behavior should reduce the duration of oviposition, with benefits of reducing the predation risk and increasing available time for foraging. The advantage in preferring these substrates should be linked to a selection pressure acting on egg development and/or survival.     

Horn length is not correlated with calling efforts in the horn‐headed cricket Loxoblemmus doenitzi (Orthoptera: Gryllidae)

Field cricket species are ideal model organisms for the study of sexual selection because cricket calling songs, used to attract mating partners, are pronouncedly sexually dimorphic. However, few studies have focused on other sexually dimorphic traits of field crickets. The horn‐headed cricket, Loxoblemmus doenitzi, exhibits exaggerated sexual dimorphism in head shape: males have flat heads with triangular horns, while females lack horns. This study examines the relationship between horn length, male calling efforts and diet quality. Horn length was not found to be significantly correlated with calling efforts. When diet was manipulated for late‐stage nymphs, calling efforts in the group with poor‐quality diet treatment was significantly lower than that of crickets in the group with high‐quality diet treatment. However, horn length was not affected by diet quality. The implication of these results in the context of the evolution of multiple signals and sexual dimorphism is discussed.     

Divergence of ovipositor length and egg shape in a brood parasitic bitterling fish through the use of different mussel hosts

Bitterling fishes deposit their eggs on the gills of living mussels using a long ovipositor. We examined whether ovipositor length (OL) and egg shape correlated with differences in host mussel species in the family Unionidae among populations of the tabira bitterling (Acheilognathus tabira) in Japan. Bitterling populations that use mussels in the sub-family Anodontinae possessed longer ovipositors and more elongated eggs than those using mussels of Unioninae, as expected from the difference in host size between the sub-families (anodontine mussels are larger than unionine mussels). Based on a robust phylogeny of A. tabira populations, we demonstrated that the evolution of both OL and egg shape were correlated with host differences, but not with each other, suggesting that these traits have been selected for independently. Our study demonstrates how adaptive traits for brood parasitism may diverge with host shift due to different host availability and/or interspecific competition for hosts.     

SHIFT IN EGG‐LAYING STRATEGY TO AVOID PLANT DEFENSE LEADS TO REPRODUCTIVE ISOLATION IN MUTUALISTIC AND CHEATING YUCCA MOTHS

Through the process of ecological speciation, insect populations that adapt to new host plant species or to different plant tissues could speciate if such adaptations cause reproductive isolation. One of the key issues in this process is identifying the mechanisms by which adaptation in ecological traits leads directly to reproductive isolation. Here I show that within a radiation of specialist moths that pollinate and feed on yuccas, shifts in egg placement resulted in changes in female moth egg‐laying structures that led to concomitant changes in male reproductive morphology. As pollinator moths evolved to circumvent the ability of yuccas to selectively abscise flowers that contain pollinator eggs, ovipositor length became shorter. Because mating occurs through the ovipositor, shortening of the ovipositor also led to significantly shorter and wider male intromittent organs. In instances where two pollinator moth species occur in sympatry and on the same host plant species, there is one short and one long ovipositor species that are reproductively isolated. Given that many plant‐feeding insects lay eggs into plant tissues, changes in ovipositor morphology that lead to correlated changes in reproductive morphology may be a mechanism that maintains reproductive isolation among closely related species using the same host plant species.