Costs and benefits of reproduction in predaceous ladybird: Effect of multiple matings on reproduction and offspring development

A few matings are sufficient for females to maximize their reproductive success, while male fitness usually increases with an increase in the number of matings. However, females of a majority of insects mate multiple times. This presents an evolutionary puzzle and brings an understanding that some benefits are associated with it. Therefore, to understand the costs and benefits of multiple matings, we performed an experimental study in a ladybird beetle, Anegleis cardoni and observed reproductive performance and longevity of adults as direct benefits and offspring development and survival as indirect benefits. This is the first time that the effect of multiple matings is being evaluated on offspring development and survival in a ladybird beetle. Results clearly reveal that females directly benefit from multiple matings in terms of increased lifetime fecundity and egg viability, but their longevity decreases with increased number of matings. Best-fit curves on lifetime fecundity and percent egg viability revealed that maximum fecundity and egg viability were both attained after 17 matings. Developmental duration of offspring decreased and their survival increased with an increase in number of matings. Developmental duration was shortest after 20 matings and longest after a single mating.     

Prolonged copulation, sperm reserves and sperm competition in the Aquatic Warbler Acrocephalus paludicola

The Aquatic Warbler Acrocephalus paludicola has a quasi-promiscuous mating system with high levels of multiple paternity and, thus, intense sperm competition. The duration of copulation in the Aquatic Warbler is unusually long. In hand-reared, captive birds the duration of mounting was 23.7 ± 11.8 min (mean ± s.d.). On average, six cloacal contacts (inseminations) occurred during each copulation. Between inseminations the male remained in contact with the female, either on top of or directly behind her. Protracted copulation may be a form of contact mate guarding, a behaviour not previously recorded in birds. Copulation was most frequent during the evening and early morning. Male Aquatic Warblers also exhibit extreme morphological adaptations of their reproductive system compared with other Acrocephalus species and other birds: their testes, cloacal protuberance and seminal glomera (which contain 198 × 10⁶ spermatozoa) are all relatively large. Frequent insemination during protracted copulation and contact mate guarding may be alternatives to paternity guarding found in other birds.     

Strategic mate‐guarding behaviour in ladybirds

Mate‐guarding behaviour is regarded as a means of increasing paternity share by reducing sperm competition. It is known to be a plastic response which varies with operational sex ratios and competitor presence in the vicinity. In a recent study, prolonged mating duration in Menochilus sexmaculatus (Fabricius) (Coleoptera: Coccinellidae) has been found to incorporate mate‐guarding behaviour. The present investigation was conducted to assess its plasticity in the presence of competitors. The physical and chemical presence of competitors of both sexes at varying densities was provided to a pair of ladybirds, and their time to commence mating, latent period and mate‐guarding duration was observed. These were compared to a control treatment where other partners were absent. All treatments were conducted with sibling as well as non‐sibling competitors. It was our hypothesis that mate guarding would be increased in the presence of male competitors and would be reduced by female presence. The results revealed that while mate‐guarding duration was increased by the chemical presence of males it was decreased by their physical presence. The latter result was attributed to interference by other males who dislodge the mating male in order to access the female. Female chemical presence had no effect on mate guarding, while physical presence increased the duration of mate guarding. The reasons for the latter behaviour require further investigation. Responses were not significantly affected by the relationship between the focal pair and the competitor. The authenticity of the mate guarding in this ladybird is strongly affirmed by our results.     

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.     

Investment in Mate Guarding May Compensate for Constraints on Ejaculate Production in the Cricket Gryllodes sigillatus

Although there is a corpus of evidence that females of many taxa are choosy about males, there is less information on how males may react to females of different 'quality' (i.e. potential fecundity). The cricket Gryllodes sigillatus shows distinct mate guarding behaviour. We examined how long males mate guard females of different sizes (reflecting egg load and potential fecundity). We also examined the sperm number in ampullae donated to females of different sizes to see if males make a concomitant difference in investment in ejaculate. We also examined mate-guarding behaviour and ejaculate size of males mated to virgin and nonvirgin females of the same size to see if males equate size with increased age and increased likelihood of mating (increased sperm competition). The results showed that males mate guard larger females for longer but make no difference in ejaculate investment between sizes of female. Males make no significant difference in mate guarding investment or ejaculate investment between virgins and nonvirgins of the same size. There is evidence that other species of crickets do alter their ejaculate depending on the female size and mating history, but have less distinct guarding behaviour. We suggest that mate-guarding investment in G. sigillatus may serve a similar function to that of ejaculate investment in other crickets.     

Leg impairment affects the reproductive attributes of a ladybird,Menochilus sexmaculatus (Fabricius)

Natural injuries and their causes have recently garnered attention amongst researchers, however the effects of these injuries on life history traits are still sparsely investigated. In the few studies that have been conducted injuries are known to have a debilitating effect on male virility and reproductive performance. Thus the effect of leg injuries in males on mating behaviour and reproductive output in the ladybird beetle, Menochilus sexmaculatus was investigated. This is the first study on the effect of injury on any life history trait in Coccinellidae. The legs of the male were amputated at three different joints on the same side and opposite sides for one leg or two legs. These amputated males were paired with uninjured females and the mating behaviour and subsequent reproductive output was observed. Time to commence mating, latent period and mating duration in impaired males were observed and were all found to be higher than that of uninjured males. Fecundity and percent egg viability were reduced in all impaired male pairs. Males with physical disability were found to copulate for an extended duration, perhaps to ensure maximum paternity.     

Cost of mating in male Menochilus sexmaculatus (Fabricius) (Coleoptera: Coccinellidae)

Mating and reproduction not only provide benefits but can also incur substantial costs. An increasing number of recent studies have indicated that mate searching, mating, and ejaculate production requires large amount of energy which can result in high cost to males. Here, we examine the effect of number of matings of males and mating duration on mating and reproductive success in Menochilus sexmaculatus (Coleoptera: Coccinellidae) by subjecting them to 3 consecutive matings of 30, 60, 90 min, and complete mating (a) with no interval and (b) with interval of 24 hr between the matings. The results reveal a reduction in reproductive performance for both the setup and reduction is more when mating is consecutive without gap. This shows that between subsequent matings males require time to recuperate the sperm/ejaculate. As a male engages in a greater number of matings, the time it takes to begin mating when paired with a female increases and the overall duration of mating decreases. However, frequent mating attempts made by males result in reduction in their body weight. Thus, in this ladybird beetle mating and related physiological process consume lots of energy and the body weight of males tend to decrease which further modifies their reproductive performance. The study helps us to identify the cost of mating from male's perspective and indicates that males need to rejuvenate between the matings.     

Effects of hexane extract of Lantana camara leaves on reproductive bioactivities of Dysdercus koenigii Fabricius (Heteroptera: Pyrrhocoreidae)

Effects of hexane extract of Lantana camara leaves were investigated on reproductive bioactivities of Dysdercus koenigii by assessing mating behavior, oviposition behavior and fertility of the adults emerged from treated fifth instar nymphs. Leaves of L. camara were extracted in hexane by ‘cold extraction method’. The fifth instar nymphs were treated with the extract by ‘dry film residual method’ for 24 h and the adults emerged from these nymphs were used for the experiments. The results indicated that treated males showed altered courtship behaviour, lesser number of mounting attempts and took relatively more time to mate than the normal males. The treated females, very often, exhibited mating refusal and non-receptive behaviour towards the courting males. This led to decrease in percent successful mating. Also, the mating in the treated insects frequently got disrupted and terminated prior to insemination. The treatment of the females with the extract resulted in alteration of their oviposition behaviour. Consequently, the treated females laid lesser number of egg batches and total number of eggs in their life span. Although the eggs laid by the treated females were fertile, the percent hatchability was lesser than normal. The results signify that hexane leaf extract of L. camara possesses phytochemicals, which adversely impaired the reproductive bioactivities of D. koenigii. Therefore, some of these compounds individually or synergistically can be employed in integrated pest management of D. koenigii by hampering its reproductive potential.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

Flexible Mate Guarding Tactics in the Dragonfly Sympelrum Internum (Odonata: Libellulidae)

Mate guarding–a behaviour prevalent in odonates–is a post copulatory association during which males prevent females from re-mating. Some species use two forms of guarding: contact mate guarding, which is energetically costly but highly effective and non-contact mate guarding, which is less costly but less effective. This study aimed to determine if male Sympetrum internum (Odonata:Libellulidae) adjust the duration of contact mate guarding according to environmental, temporal and physiological factors. There was a significant interaction between male density and season on duration of contact mate guarding. Early in the season males increased the duration of contact guarding as the density of rivals increased. Later in the season males guarded mates longer irrespective of male density. Wind and temperature did not detectabiy alter the duration of contact mate guarding, suggesting that the trade-off between current and future reproductive success was more important than were physiological costs.     

Effects of multiple mating on the fecundity of an invasive pest (Octodonta nipae): the existence of an intermediate optimal female mating rate

Multiple mating is found in many insect taxa where both of the sexes can mate more than once. For males, this leads to the advantage of increasing their paternity by fertilizing more females. However, there is a trade‐off of resource allocation between reproduction and other life‐history characters. In the present study, the impact of increased mating rate on reproductive fitness of the invasive nipa palm hispid beetle Octodonta nipae Maulik (Coleoptera: Chrysomelidae) is investigated. A series of mating frequencies (i.e. 1, 5, 10, 15, 20 times) is selected from video frame playback, ranking from the minimum to maximum mating rate observed under laboratory conditions over a given time period. Fecundity parameters such as lifetime egg production, egg‐hatching rate, effective oviposition period and longevity are investigated for the evaluation of reproductive efficiency. For female O. nipae, increased fecundity is correlated with the mating frequency. Females mating 15 times lay the largest number of eggs (138.82 ± 6.87) and have a hatching rate of 47.43 ± 4.08%. After mating 20 times, females suffer significant declines in oviposition (90.31 ± 8.38 eggs) and egg‐hatching rate (34.16 ± 4.93%). Moreover, the population growth rate reaches a maximum in the females that mate 15 times. The results show that multiple matings in O. nipae have an intermediate optimal range within which female reproductive success is enhanced, providing empirical evidence for the existence of a trade‐off between costs and benefits during copulation based on resource allocation.     

Effects of Single Versus Multiple Mates: Monogamy Results in Increased Fecundity for the Beetle <Emphasis Type="Italic">Phoracantha semipunctata</Emphasis>

Effects of polyandry versus monogamy were assessed for the beetle Phoracantha semipunctata (Coleoptera: Cerambycidae) by comparing measures of female reproductive success, including fecundity, egg viability, time until eclosion, and clutch size. The effect of intermittent presence or absence of a male was also evaluated. Polyandry was detrimental to female reproductive success in comparison to monogamy. Fecundity, egg viability, and clutch size were lower, and time to eclosion were increased for eggs from females with multiple mates compared with monogamous females. Intermittent presence of males had no effect on female reproductive success. Possible explanations for the decreased fecundity experienced by females with multiple mates include sperm competition intensity, costs of male harassment, and exceeding the optimal mating frequency. Females may reduce costs associated with polyandry by spending less time on host logs where mating occurs.     

Effect of egg cannibalism on mating preferences and reproductive fitness of Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae)

Cannibalism has been reported in a large proportion of coccinellids in fields as well as in laboratories but studies involving mate preferences and potential benefits of cannibalism on reproduction in Menochilus sexmaculatus Fabricius (Coleoptera: Coccinellidae) have yet not been done. Thus, we assessed the effect of conspecific egg cannibalism on mate preferences and reproductive outputs including offspring development. Higher mate preferences were recorded for non-cannibalistic mates (fed on A. craccivora) than cannibalistic ones (fed on conspecific eggs). Mating parameters were significantly influenced by cannibalism. Time to commence mating lasted less for homogeneous diet pairs than heterogeneous diet pairs. Longer copulation duration and higher fecundity were recorded when one of the individuals in mating pair or both was a non-cannibal. Egg viability did not differ significantly in all the reciprocal crosses. In addition, the total developmental durations of offspring were similar for all mating pairs.     

Mate guarding and male mate choice in the chameleon grasshopper <Emphasis Type="Italic">Kosciuscola tristis</Emphasis> (Orthoptera: Acrididae)

In the wild, male chameleon grasshoppers (Kosciuscola tristis) are frequently observed mounted on the back of females even when not in copula, and will fight off other usurping males. If this behaviour is mate guarding and reflects investment in male mate choice, then we expect males to preferably guard females based on reliable cues of quality. Cues for female quality likely include female size and egg development that together may indicate fecundity. We investigated male mate choice in the field expressed as mate-guarding preference, by comparing size and egg development in guarded and unguarded females. We found no difference between guarded and unguarded females in measures of fecundity or body size. The majority of females sampled did not contain any viable eggs. This finding suggests that male K. tristis indiscriminately guard females in a scramble mating system.     

Benefits of polyandry in Parthenium beetle,Zygogramma bicolorata Pallister (Coleoptera: Chrysomelidae)

The influence of polyandry on the reproductive performance of females and on offspring fitness in Zygogramma bicolorata Pallister was investigated using four experimental treatments, viz. (A) monandrous, limited mating, (B) monandrous, unlimited mating, (C) polyandrous, no-choice limited mating, and (D) polyandrous, mate choice unlimited mating. Polyandrous females had higher reproductive performance than monandrous ones. Monandrous females subjected to unlimited matings had higher egg viability than those subjected to limited matings, but fecundity did not differ significantly. In polyandrous females, the freedom to choose mates did not affect reproductive performance. However, offspring of polyandrous females allowed mate choice developed fastest and had the highest survival at 25, 27, and 30 °C. Thus, polyandry in Z. bicolorata appears to provide both direct (material) and indirect (genetic) benefits resulting in better reproductive performance and increased adaptability of the offspring to counter environmental stresses. The present study not only adds to the knowledge of reproductive biology of Z. bicolorata but it could also be of economic value as it may help in the mass rearing of Z. bicolorata and in the management of Parthenium hysterophorus.     

Multiple mating and repeated copulations: effects on male reproductive success in red flour beetles

In many insects, both sexes mate multiple times and females use stored sperm for fertilizations. While males frequently engage in two distinct behaviours, multiple mating (with different females) and repeated copulations (with the same female), the reproductive consequences of these behaviours for males have been quantified for only a few species. In this study, males of the red flour beetle, Tribolium castaneum, were found to be capable of mating with as many as seven different virgin females within 15 min. Across sequential copulations with virgin females, there was no decline in either male insemination success or average female progeny production over 48 h. However, when males copulated with previously mated females, there was a significant decline in male paternity success across sequential copulations, possibly due to male sperm depletion. In separate experiments, T. castaneum males were found to engage in two to six repeated copulations with the same, individually marked female. These repeated copulations did not increase male insemination success, short-term female fecundity, or male paternity success. Repeated copulations may possibly play a role in sperm defence. This study indicates that males may frequently engage in multiple matings, but these additional matings may lead to diminishing male reproductive returns.     

The role of female dominance hierarchies in the mating behaviour of mosquitofish

While studies of sexual selection focus primarily on female choice and male-male competition, males should also exert mate choice in order to maximize their reproductive success. We examined male mate choice in mosquitofish, Gambusia holbrooki, with respect to female size and female dominance. We found that the number of mating attempts made by a male was predicted by the dominance rank of females in a group, with dominant females attracting more mating attempts than subordinates. The number of mating attempts made by males was independent of the female size. The observed bias in the number of mating attempts towards dominant females may be driven either by straightforward male mate choice, since dominance and female fecundity are often closely related, or via the dominant females mediating male mating behaviour by restricting their access to subordinate females.     

EVOLUTION OF MONOGAMY IN TERMITES

Two hypotheses have been proposed to explain the origin of lifetime monogamy in the Isoptera. The classic explanation is that (1) the male must be present to continually provide sperm for the vast number of eggs produced by the queen (Snyder, 1924: Brian, 1983). Thornhill & Alcock (1983) proposed that (2) synchrony in the availability of receptive females necessitates mate guarding; males subsequently gain if they improve the relative reproductive success of their sole partner.
Our review of the literature on termite flight behaviour, courtship behaviour, and incipient colony development indicates that neither of these two hypotheses satisfactorily explains the evolution of monogamy in termites. Because incipient colonies of lower termites exhibit a very low fecundity, it is doubtful that the continued presence of the male initially was due to the need for a continuous supply of spermatozoa. It is possible, however, that sperm requirements for the fertilization of numerous eggs over an extended period of time may be a factor in the persistence of the termites' monogamous mating system. Female alates are much more dispersed in time than implied by Thornhill & Alcock (1983) and there is no evidence of mate guarding. The importance of mate assistance is, however, supported by the literature. We propose a third hypothesis that incorporates the mate assistance element of the Thornhill & Alcock hypothesis: (3) the monogamous mating system of termites was structured by ecological constraints, namely, the low quality and scattered nature of their food/nesting material and the high costs of searching for a mate.     

Female Gryllus vocalis Field Crickets Gain Diminishing Returns from Increasing Numbers of Matings

Although female insects generally gain reproductive benefits from mating frequently, females do not mate unlimited numbers of times. This study asks whether the limit on female mating rate is imposed by trade‐offs between reproduction and survival. Female Gryllus vocalis were given the opportunity to mate 5, 10, or 15 times with novel males, and the effects on daily fecundity (egg production), fertility (proportion of eggs that were fertilized), and female post‐experimental longevity were measured. Females that mated 10 times laid more eggs and had a higher proportion of fertile eggs than females that mated 5 times. However, females that mated 15 times did not lay significantly more eggs or have a higher proportion of fertile eggs than females that mated 10 times. Although number of matings did not affect the date that females laid their last egg, mating more times was associated with a prolonged period of laying fertile eggs. Number of matings did not affect female post‐experimental longevity. Thus, there was no trade‐off between female reproductive effort and survival, even when females mated very large numbers of times. When females were allowed to mate ad libitum, the average number of times that females mated was greater than the number of times that confers maximal fitness. The lack of cost to mating explains why females might be willing to mate beyond the point of diminishing reproductive returns.     

Mating and fecundity of Dermatophagoides farinae

Studies of the life cycle of cultured Dermatophagoides farinae found that after an initial mating D. farinae females lived for 63.3 ± 64.6 (SD) dafter their egg production period ended .The long period after cessation of egg production for D. farinae suggested D. farinae females could mate multiple times and produce eggs continuously for a longer period. The purpose of this study was to determine if female D. farinae could mate at least two times, and subsequently increase the production of viable eggs over a longer period of time compared to a single mating. Female D. farinae were randomly selected from thriving cultures and isolated in cages. When the females had ceased to lay eggs a male was added to the cage. Fifty-seven percent of the isolated females mated again and produced a second batch of viable eggs. In natural or culture populations, females have continuous availability of males. Therefore, in another experiment, females that emerged from the tritonymphal stages were continuously exposed to fresh males and fecundity and lengths of the reproductive and post reproductive periods were determined. These females had a 11 d longer reproductive period and produced 30.7% more eggs compared to females that only mated one time after they emerged from the tritonymphal stage. However, the post reproductive period was still long (58.6 ± 11.4 [SE] d) the significance of which is not clear. In conclusion, this study revealed that D. farinae females are capable of more than one successful mating that results in increased egg production compared to that of a single mating. It is likely that females mate multiple times in natural and culture populations. It was observed that females actively attract males during the reproductive period but not afterward even though she continues to live a long time. This revised version was published online in July 2006 with corrections to the Cover Date.