Optimal number of matings in two aphidophagous ladybirds

Abstract.  1. The present study was designed to identify the optimal number of matings required for maximum fecundity and egg viability in two aphidophagous ladybirds, Cheilomenes sexmaculata and Propylea dissecta .
2. For this purpose, ladybirds were subjected to different numbers of matings and the reproductive responses were recorded thereafter.
3. The Gompertz model was used to draw asymptotic graphs for fecundity and per cent egg viability in both ladybird species. Ninety-five per cent and 50% of maximum theoretical fecundity and per cent egg viability were predicted from the model.
4. Ninety-five per cent maximum theoretical fecundity was obtained after 13.25 and 12.95 matings in C. sexmaculata and P. dissecta , respectively; and 8.95 and 11.25 matings were required for 95% maximum theoretical per cent egg viability in C. sexmaculata and P. dissecta , respectively.
5. The results of these experiments clearly support the existence of an optimal number of matings in these two ladybird species leading to maximum adult fitness.     

Effect of age on reproductive attributes of an aphidophagous ladybird, Cheilomenes sexmaculata

The effect of both male and female age was investigated on certain reproductive attributes, viz. mating incidence, mating duration, fecundity, percent egg viability, ratio of reproductive and non‐reproductive periods and reproductive rate, of an aphidophagous ladybird, Cheilomenes sexmaculata (Fabricius). Females started mating at the age of 8 hours post‐emergence (PE) and males at the age of 2 days PE. Mating in the laboratory was a male‐dominated phenomenon. The mating duration and reproductive rate of 10‐day‐old females when mated with males of varying ages increased up to the male age of 60 days, and thereafter decreased, whereas, fecundity, egg viability and ratio of reproductive and non‐reproductive periods increased up to the male age of 50 days, and thereafter declined. However, when females of varying ages were mated with 10‐day‐old males, fecundity and reproductive rate increased up to 40 days of female age, respectively, then decreased. The ratio of reproductive and non‐reproductive periods increased with increasing age of females. Mating age for optimal reproductive output was 10J50‐day‐old males and NE to 40‐day‐old females. Reproductive cessation in males was recorded after 50 days PE, whereas in females at the age of 40 days PE. Higher mating durations lead to elevated reproductive rates. Delay in the reproductive phase was positively correlated with longevity. The results of this study may aid mass multiplication of this ladybird by identifying and promoting usage of adults of optimal age. Our results also enhance our understanding of the effect of age on reproductive attributes in ladybirds.     

Larval cannibalism in aphidophagous ladybirds: Influencing factors, benefits and costs

Spray retention on scentless chamomile and round-leaved mallow was characterized in relation to droplet size and travel speed. The effect of varying retention on biocontrol efficacy was studied using Colletotrichum truncatum and Colletotrichum gloeosporioides f. sp. malvae, the bioherbicide agents for the two weeds, respectively. In retention studies, a tracer dye solution was applied using a cabinet sprayer fitted with a fine, medium, or coarse nozzle at travel speeds of 0.5, 1.0, and 2.0 km/h that resulted in approximate application volumes of 500–2000 L/ha. Retention efficiency, a ratio between the volume retained on plants and actual volume applied, was calculated for each application. In general, finer sprays achieved higher retention efficiency on whole plants of both weeds. On an average, fine sprays resulted in 68% greater retention than coarse sprays on scentless chamomile and 22–59% on round-leaved mallow. Faster travel speeds increased spray retention only slightly. Applying the biocontrol agents at 500 L/ha with different droplet sizes showed different effects on weed control on scentless chamomile and round-leaved mallow. C. truncatum applied with fine droplets was more efficacious than treatments with coarser droplets on scentless chamomile. On round-leaved mallow, however, the efficacy of C. gloeosporioides f. sp. malvae did not appear to be substantially affected by the droplet size. This varying effect may be due to reduced retention on the vertical stems of round-leaved mallow, which is critical to biocontrol of this weed by C. gloeosporioides f.sp. malvae.     

Functional response of two aphidophagous ladybirds searching in tandem

The functional response of two ladybird species, Propylea dissecta (Mulsant) and Coccinella transversalis Fabricius in combination was studied in an effort to determine the impact of two predators on the increasing density of aphid, Aphis gossypii Glover infesting leaves of bottle gourd, Lagenaria vulgaris Seringe. The ladybirds were exposed to prey density in three combinations using adult females: (i) one P. dissecta and one C. transversalis together, (ii) two P. dissecta, and (iii) two C. transversalis, for 24 h. Prey consumption increased curvilinearly with prey density in all three combinations. It was maximum when heterospecifics searched together. The prey were handled and consumed significantly faster (9.51 min) by heterospecifics in combination, followed by two C. transversalis (11.32 min) and two P. dissecta (12.52 min). The attack rate was maximum (0.00274) by heterospecifics followed by the two C. transversalis (0.00166) and two P. dissecta (0.00092). The synergistic effect on the prey consumption may possibly be due to spatial distribution of P. dissecta and C. transversalis in terms of feeding sites, with the former being negatively, and the latter being positively geotactic. Field release of two predator species in tandem is recommended for biocontrol of A. gossypii because of the potential for synergistic action by these natural enemies.     

Influence of parental age on reproductive performance of an aphidophagous ladybird, Propylea dissecta (Mulsant)

Abstract:  In the present study the influence of maternal and paternal ages on reproductive performance of an aphidophagous ladybird, Propylea dissecta (Mulsant) were investigated. Maternal age strongly influenced fecundity and pre-oviposition period while paternal age strongly influenced the egg viability. Maternal age also influenced percent egg viability, however, at lower levels of significance. The results revealed that the fecundity and percent egg viability increased up to the respective maternal and paternal ages of 30 days and thereafter declined, indicating the onset of senescence. The results of the present study can help in better mass multiplication of the ladybird concerned and in laying foundation for further age-related studies.     

Survival, development and life tables of two congeneric ladybirds in aphidophagous guilds

Two congeneric aphidophagous ladybirds, Coccinella septempunctata and Coccinella transversalis, were reared on three aphid species, Lipaphis erysimi, Myzus persicae and Aphis nerii, to estimate the effect of prey quality and intra- and interspecific interactions on their survival and development of life stages. Mortality of first instar ladybirds of both species was highest feeding on A. nerii. Preimaginal mortafity was lowest when feeding on L. erysimi （C. septempunctata, 1.6% and C. transversalis, 3.2%）, and highest when feeding on A. nerii （ C. septempunctata, 6.2% and C. transversalis, 8.2%）. Comparatively higher weight and larger size of C. septempunctata along with the lower levels of mortality recorded suggested that it is more likely to have acted as an intraguild predator than C. transversalis. High recorded mortality of C. transversalis is attributed to probable intraguild predation on account of its smaller size. The major sources of mortality were probably cannibalism, intraguild predation and other unknown factors. Lower prey quality increased the incidence of cannibalism and intraguild predation, especially in C. transversalis. The investigation suggests an intrinsic competitive advantage for C. septempunctata over C. transversalis in guilds of three aphid species.     

Evolution of cannibalism and female's response to oviposition-deterring pheromone in aphidophagous predators

1.  Egg cannibalism by larvae is common in Coccinellidae and is known to be advantageous for the cannibals. Furthermore, larvae of aphidophagous ladybirds usually produce an oviposition-deterring pheromone (ODP), which inhibits oviposition by adult females. It has been proposed that the response to ODP has evolved because of the high costs of cannibalism. However, this has never been formally proved.
2.  In this paper, we study the theoretical evolution of this system. We first look at the conditions under which cannibalism and the response to ODP can evolve. Subsequently, we examine the occurrence of polymorphism both in the production of larval tracks and in the sensitivity of females to specific pheromones.
3.  The models predict that the amount of cannibalism should not depend on prey density and that evolution should lead to a continuous increase in cannibalism, and consequently larvae should always cannibalize eggs when possible. In response to the cost of cannibalism, ODP recognition can evolve, so that females avoid laying eggs in patches of prey already occupied by conspecific larvae. The result is an arms race between larvae and adult females, which favours a diversification of ODP pheromones. Our models show that: (i) females should be able to recognize mixtures of hydrocarbons rather than a single molecule; and (ii) females should be more sensitive to the tracks of their own offspring than those of non-related larvae.     

Influence of mating duration on fecundity and fertility in two aphidophagous ladybirds

Abstract:  The influence of different mating durations on fecundity and fertility has been studied in two aphidophagous ladybirds, Cheilomenes sexmaculata and Coelophora saucia . Pre-oviposition period decreased while fecundity and fertility increased with increase in mating duration in both the ladybird species. The increase in fecundity with longer mating durations is probably due to the female response because of cryptic female choice. Mating of 10-s duration resulted in oviposition but of unviable eggs probably because of lack of sperm transfer. A minimum duration of 1 min of mating was probably essential for fertilization of the eggs. Absence of spermatophore in the reproductive tract of females is indicative of direct sperm transfer in both the ladybird species. Trend of fertility (similar fertility at 1 and 5 min followed by increase at 60 min which was again similar to that at complete mating) indicates probable sperm transfer in spurts in C. sexmaculata while continuous transfer in C. saucia is indicated by continuous increase in fertility with increase in mating duration.     

Does temperature modify slow and fast development in two aphidophagous ladybirds?

Development within a population is known to vary with abiotic and biotic factors. However, instances of slow and fast development, i.e. different development rates within the same egg batch or cohort, have not yet been rigorously investigated in many organisms. The present study was undertaken to assess the influence of an exogenous cue, temperature (15, 20, 25, 30 and 35 °C), on slow and fast development and its effect on reproductive attributes in two ladybird species, Menochilus sexmaculatus (Fabricius) and Propylea dissecta (Mulsant) (Coleoptera: Coccinellidae). A clear bimodal pattern of distribution with two peaks was found at each temperature where the first peak represented the fast developers and the second peak slow developers. Variation in ratio and development of slow and fast developers within a cohort at different temperatures was observed to ascertain whether the phenomenon has a purely genetic basis or is environmentally influenced. The ratio of slow and fast developers in a cohort differed with temperature but not with species. Slow developers showed a female biased sex ratio and low body mass. Fast developing females laid higher numbers of eggs with higher egg viability than slow developing ones. More slow developers were found at low temperature (15 and 20 °C), equal numbers at medium temperature (25 °C) and less at higher temperature (30 and 35 °C). Results of the study are indicative of stability of the slow and fast development in the egg batch and the likelihood that the variation observed at emergence was owing to exogenous cues influenced differential rates of mortality.     

Vulnerability of larvae of two species of aphidophagous ladybirds, Adalia bipunctata Linnaeus and Harmonia axyridis Pallas, to cannibalism and intraguild predation

Vulnerability of larvae of two species of aphidophagous ladybirds, Adalia bipunctata Linnaeus and Harmonia axyridis Pallas, to cannibalism and intraguild predation was assessed in the laboratory. In the first experiment, a first instar of one of the two above species was kept with a fourth instar of the other species in a Petri dish. The number of times each first instar larva was encountered by the fourth instar larva and the fate of the first instar was determined over a period of 10 min. The fourth instar larvae captured and killed all the first instar larvae of their own species at the first encounter. However, when presented with fourth instar larvae of the other species the first instar larvae of A. bipunctata and H. axyridis were encountered 6.4 ± 1.3 ( n  = 10) and 19.4 ± 2.1 ( n  = 10), respectively. In this experiment no first instar larvae of H. axyridis , whereas all those of A. bipunctata , were killed.     

Consequences of diapause on post‐diapause development,reproductive physiology and population growth of Chilo partellus (Swinhoe)

We investigate the effects of diapause on post‐diapause development, reproductive physiology and population growth of Chilo partellus (Swinhoe) (Crambidae: Lepidoptera). Aestivating and hibernating larvae of C. partellus are exposed to diapause terminating conditions (consisting of an LD 12 : 12 h photocycle at 27 ± 1 °C and 65 ± 5% relative humidity with a fresh diet) to terminate the diapause and observations are made on percentage pupation, pupal duration and weight, adult reproductive performance and population growth parameters. We find that the diapause in C. partellus significantly reduces the percentage pupation and weights of pupae, ultimately lowering the weight and reproductive performance of adults. Reduced weights of adult females are found to be directly associated with a lower number of egg cells in ovaries. Nevertheless, the reproductive performance of C. partellus males is also found to be greatly affected in the diapause (hibernation and aestivation) experiencing population in terms of the deposition of a lower number of spermatophores and eupyrene sperm in the reproductive tracts of females compared with the nondiapausing population. The results of the present study clearly indicate that a reduction in longevity, fecundity and egg viability, as well as a reduced rate of deposition of spermatophores and eupyrene sperm, in a diapause experiencing population of C. partellus ultimately leads to a reduction in population growth parameters, thus having implications for bio‐ecology and population dynamics under a changing climatic scenario.     

Interactions between ants and aphidophagous and coccidophagous ladybirds

Aphidophagous and coccidophagous coccinellids come into conflict with homopteran-tending ants for access to food. Antagonistic interactions between coccinellids and ants may be competitive or non-competitive. Competitive interactions occur when coccinellids attack aphids or coccids that are being tended by ants for honeydew. Non-competitive interactions include all interactions away from ant-tended homopteran colonies. We here review observations and studies of such interactions. We note that most competitive interactions occur at times when untended aphids/coccids are scarce. We describe the chemical and physical defences that coccinellids use against ant aggression and consider whether these have evolved as general anti-predator deterrents or specifically in response to ants. Myrmecophilous coccinellids are then considered, with particular focus on the two most studied species, Coccinella magnifica and Platynaspis luteorubra. We note that the myrmecophily of the two species has the same adaptive rationale—to enable the ladybirds to prey on ant-tended aphids at times of aphid scarcity—but that it is based on different traits to facilitate life with ants. Finally, we consider the role of ants in the evolution of habitat specialisation in some coccinellids.     

Strong material benefits and no longevity costs of multiple mating in an extremely polyandrous leaf beetle, Chrysochus cobaltinus (Coleoptera: Chrysomelidae)

It is well established that females of many species exhibitpolyandry. Although such behavior often increases female fitnessby augmenting fecundity or enhancing the genetic diversity andvigor of their offspring, it often reduces female longevity.It has been argued that trade-offs between these costs and benefitsshould limit the degree to which females remate. However, theexistence of highly polyandrous species suggests substantialpolyandry benefits and/or minimal costs in some systems. Femalesof the leaf beetle, Chrysochus cobaltinus, are extremely polyandrous,providing an opportunity to examine the factors influencingthe evolution of such behaviors. We compared the fecundity andlongevity of singly mated females, females that mated multipletimes with the same male, and females that mated multiple timeswith different males. Compared with females in the single matingtreatment, females in both multiple mating treatments exhibiteda significant reduction in latency to oviposition and, due toan increase in daily egg production, significant increases inlifetime fecundity. This difference diminished as the time sincelast mating increased. There were no differences in fecunditybetween the 2 multiple mating treatments, indicating that mateidentity does not influence the material benefits of multiplemating. Surprisingly, female longevity did not differ amongtreatments. The pronounced fecundity benefits that females gainfrom multiple mating, coupled with a lack of longevity costs,apparently explains the extreme polyandry in this species. Inaddition, the existence of material fitness benefits via conspecificmatings raises the intriguing possibility that in a C. cobaltinus–Chrysochusauratus hybrid zone, heterospecific matings may confer similarbenefits to Chrysochus females.     

The costs and benefits of fast living

Growth rates play a fundamental role in many areas of biology (Q. Rev. Biol., 67, 1992, 283; Life History Invariants. Some Explorations of Symmetry in Evolutionary Biology, 1993; Philos. Trans. R. Soc. Lond. B Biol. Sci., 351, 1996, 1341; Plant Strategies, Vegetation Processes, and Ecosystem Properties, 2002; Trends Ecol. Evol., 18, 2003, 471; Q. Rev. Biol., 78, 2003, 23; J. Ecol., 95, 2007, 926.) but the cost and benefits of different growth rates are notoriously difficult to quantify (Q. Rev. Biol., 72, 1997, 149; Funct. Ecol., 17, 2003, 328). This is because (1) growth rate typically declines with size and yet the most widely used growth measure – relative growth rate or RGR (conventionally measured as the log of the ratio of successive sizes divided by the time interval) – is not size-corrected and so confounds growth and size, (2) organisms have access to different amounts of resource and (3) it is essential to allow for the long-term benefits of larger size. Here we experimentally demonstrate delayed costs and benefits of rapid growth in seven plant species using a novel method to calculate size-corrected RGR. In control treatments, fast-growing plants benefited from increased reproduction the following year; however, fast-growing plants subjected to an experimental stress treatment (defoliation) showed strongly reduced survival and reproduction the following year. Importantly, when growth was estimated using the classical RGR measure, no costs or benefits were found. These results support the idea that life-history trade-offs have a dominant role in life-history and ecological theory and that the widespread failure to detect them is partly due to methodological shortcomings.     

Mating frequency positively associates with fitness in Ophraella communa

1. Why animals mate multiple times, owing to the lack of immediate fitness benefits, presents an intriguing problem for evolutionary biologists. Yet, the profusion of this behaviour suggests it must be maintained by natural selection via increased performance. 2. The possible benefits of multiple mating using the leaf beetles Ophraella communa LeSage, the biological control agent of the invasive common ragweed Ambrosia artemisiifolia L., were studied and the fitness consequences of single, twice, three, four, and unrestricted mating events were assessed. 3. Overall, it was observed that the number of copulation events was positively associated with fitness parameters of the insects. Insects performed the best under unrestricted mating regimes, with average increases of 48% in longevity, 75% in fecundity, and 55% in egg hatch rate. In addition, females that experienced unrestricted access to mates maintained very high viability over their entire reproductive lives. Nevertheless, insects also performed better when allowed to mate four times compared with once or twice. 4. The present findings thus support the hypothesis that multiple‐mating behaviour is maintained owing to increased fitness benefits in the current and the next generation. Selection for re‐mating is, therefore, expected in field populations, which is likely to happen naturally owing to the aggregate lifestyle of O. communa.     

The cost of floral longevity in Clarkia tembloriensis: An experimental investigation

The hypothesis that flower maintenance requires resources that would be used to support other plant functions (i.e. a cost of floral maintenance) was tested by experimentally manipulating floral longevity. Plants of Clarkia tembloriensis, a species with pollination-induced flower senescence, received either early or late pollinations (long and short longevities, respectively). We examined the effect of this manipulation on (1) per-flower allocation to nectar production and (2) flower, fruit and seed production per plant under two levels of resource availability. The direct costs of floral longevity measured in terms of nectar sugar were high: flowers that were maintained 35% longer invested proportionately more in nectar sugar (30%). At the whole-plant level, a cost of floral longevity was manifested as reduced seed production, but the magnitude of this cost varied with resource level. While plants with longer-lived flowers showed a 12% reduction in seed production, those that experienced reduced resource levels via partial defoliation, showed a decrement in seed production that was almost three times larger (34%). These differences were not brought about by changes in the number of flowers and fruits, but by significant alterations in their sizes. A model that expresses the cost of flower maintenance as a trade-off between floral longevity and seed production shows that an optimal flower longevity is determined by both the rate of fitness accrual and the cost of floral maintenance.     

Mating behavior of an aphidophagous ladybird beetle,Propylea dissecta （Mulsant）

Abstract The mating behavior of Propylea dissecta (Mulsant) was investigated in detail. Protandry was evident, as males mature earlier than females. Predicted age of sexual maturation of male and female ladybirds was 7.12 and 9.33 days, respectively. Males performed courtship in six steps, viz. approach, watch, examine, embrace, mount and attempt. Embrace is possibly an appeasement act, while examine serves to recognize mate. Chemical signals initiate male attraction toward females. Visual and tactile cues appear to secondarily help in mate recognition. Mating lasted longest (275.40±12.23 min) when it occurred between unmated individuals. It was much shorter (176.60±5.60 min) when prior mated individuals copulated. There was a significant decrease in mating durations when adults were subjected to five successive matings. Fecundity and percent egg viability increased significantly with increase in the number of matings.     

大猿叶虫卵的颜色与其后代滞育率和繁殖力的关系

大猿叶虫Colaphellus bowringi Baly卵的颜色有浅黄色和深红色2种类型。调查卵的颜色与其后代滞育的发生和繁殖力的关系,结果表明,在25℃、28℃,不同光周期L∶D(11∶13,12∶12,13∶11和15∶9)下,浅黄色卵和深红色卵的后代滞育率没有显著差异;但在L14∶D10下,浅黄色卵的后代滞育率(25℃为58.6%;28℃为33.9%)显著高于深色卵的后代(25℃为45.4%;28℃为11.2%)。卵的颜色与其后代成虫的产卵前期和成虫寿命没有显著相关,但浅黄色卵后代的平均产卵量和平均日产卵量(1663粒和58.6粒)均显著高于深红色卵的后代(1176粒和45.1粒)。     

Role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis

Abstract:  The role of surface chemicals in egg cannibalism and intraguild predation by neonates of two aphidophagous ladybirds, Propylea dissecta and Coccinella transversalis were examined. Neonates of both species prefer to eat non-sibling conspecific eggs than heterospecific eggs, with higher preference in P. dissecta . Surface chemicals appear to play a major role for the preference of conspecific eggs, as the ladybird behaviour was reversed when these chemicals were interchanged. The surface chemicals present on the eggs possibly act as feeding stimulants to conspecific neonates but not for heterospecific neonates. Egg clustering enhances the effect of surface chemical and stimulate non-sibling egg cannibalism and appears to reduce intraguild predation. Egg clustering is advantageous to ladybirds, as it can stimulate non-sibling cannibalism by neonates. It appears that aggregation of chemicals present on the conspecific egg surface possibly attracts the hungry conspecific neonate, providing the first meal for its survival, while protecting against heterospecific predators. Neonates of both ladybirds were reluctant to eat heterospecific eggs.     

SEXUAL SIZE DIMORPHISM AND SELECTION IN THE WILD IN THE WATERSTRIDER AQUARIUS REMIGIS: LIFETIME FECUNDITY SELECTION ON FEMALE TOTAL LENGTH AND ITS COMPONENTS

Darwin's fecundity advantage model is often cited as the cause of female biased size dimorphism, however, the empirical studies of lifetime selection on male and female body size that would be required to demonstrate this are few. As a component of a study relating sexual size dimorphism to lifetime selection in natural populations of the female size-biased waterstrider Aquarius remigis (Hemiptera: Gerridae), we estimated coefficients for daily fecundity selection, longevity selection, and lifetime fecundity selection acting on female body size and components of body size for two consecutive generations. Daily fecundity was estimated using females confined in field enclosures and reproductive survival was estimated by twice-weekly recaptures. We found that daily fecundity selection favored females with longer total length through direct selection acting on abdomen length. Longevity selection favored females with smaller total length. When daily fecundity and reproductive longevity were combined to estimate lifetime fecundity we found significant balancing selection acting on total length in both years. The relationship between daily fecundity and reproductive longevity also reveals a significant cost of reproduction in one of two years. We relate these selection estimates to previous estimates of sexual selection on male body size and consider the relationship between contemporary selection and sexual size dimorphism.