Effects of female body size on lifetime fecundity of <Emphasis Type="Italic">Monochamus urussovii</Emphasis> (Coleoptera: Cerambycidae)

To analyze the relationship between female body size and lifetime fecundity (LF) in Monochamus urussovii (Fischer), 17 pairs of adults were reared on Picea jezoensis (Sieb. et Zucc.) Carrière twig sections and were allowed to oviposit on Abies sachalinensis (Fr. Schmidt) Masters bolts at 25 °C and 16:8 h light:dark. The means of life span and LF were 42.5 days and 28.2 eggs for fertile and sterile females combined, respectively. The mean lengths of preoviposition and oviposition periods (OPs) of fertile females were 12.1 and 28.1 days, respectively. The potential fecundity was estimated to be 72.1. Path analysis showed that female body size, i.e., mass and elytral length, had positive effects on LF through the oviposition rate but not through the OP. Ovariole number was 24 and did not vary according to body size, indicating a higher egg productivity per ovariole in large females than in small ones. Higher egg productivity per ovariole in large females is true for each of three other synovigenic lamiine beetles with an almost fixed number of ovarioles. The body size-dependent egg productivity per ovariole is considered to be one of the reasons for the body size–LF correlation in synovigenic species with a fixed ovariole number.     

Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

Relatively large males lower reproductive success in female zebrafish

Females can adjust their reproductive effort in relation to their partner’s perceived fitness value. In zebrafish (Danio rerio), large males are typically preferred mating partners. However, females have been observed to reduce their reproductive output with exceptionally large males but it remains unknown whether it is due to sexual harassment or aggressive behavior to establish and maintain dominance. Here, we study the association between relative male size, sexual harassment and dominance behavior, female stress status (stress behaviors and whole-body cortisol concentration), and reproductive success during a 4 day spawning trial. We found female cortisol to correlate negatively with female body size and positively with female dominance behavior. However, male and female behavior as well as female cortisol level were not related to relative male size. Females mating with relatively large males produced more and most of their eggs during the first spawning day, while females with smaller males produced few eggs during the first day but then increased egg production. Despite females produced more eggs when mating with relatively larger males, their eggs had substantially lower fertilization rates compared to females mating with relatively smaller males. Hence, overall, the reproductive fitness was lowest when females mated with a relatively large male. These findings could help to explain the maintenance of male size variation under natural conditions.     

Variation of clutch size and trophic egg proportion in a ladybird with and without male-killing bacterial infection

Maternally inherited bacterial endosymbionts can kill male embryos of their arthropod hosts to enhance the transmission efficiency of the endosymbionts. The resources from killed male eggs can be reallocated to infected female hatchlings as additional maternal investment. As a result, the number of offspring per patch and the maternal investment per offspring are expected to differ from the original optimal values for the host mother. Thus, in response to infection, these trait values should be adjusted to maximize the lifetime reproductive success of host females and the fitness of inherited endosymbionts as well. Here, we examined clutch size, egg size, and the proportion of trophic eggs (i.e., production of unhatched eggs, a maternal phenotype) per clutch of host mothers infected with male-killing bacteria. First, we developed a mathematical model to predict the optimal clutch size and trophic egg proportion in uninfected and infected females. Next, we experimentally compared these life-history traits in a ladybird, Harmonia yedoensis, between females infected or uninfected with male-killing Spiroplasma bacteria. Consistent with our predictions, clutch size was larger, egg size was smaller, and trophic egg proportion was lower in infected H. yedoensis females, compared with uninfected females. To our knowledge, this is the first empirical demonstration of variation in these life-history traits depending on infection with bacterial endosymbionts.     

Effects of male apparent length on female preference for absolute body size in <Emphasis Type="Italic">Xiphophorus helleri</Emphasis>

Female green swordtails (Xiphophorus helleri) prefer males of longer body and sword lengths, which may reflect perceptual constraints. Larger males project a larger image onto the female’s retina at a given viewing distance, which may elicit greater stimulation of the visual system and a stronger behavioral response. This study investigates the effects of male apparent size, as affected by proximity and sword length, on female preference for absolute body size in X. helleri. Females were presented pairs of dummy males in ten treatments organized into six categories: (1) the Actual Length Control where identically sized males were held at different viewing distances from the female; (2) the Sword Compensation Treatment in which viewing distance and apparent length (body + sword) were held constant, pairing sworded with swordless dummies of equal total length; (3) the Body Length Preference Treatment where male distance from the female was held constant, while varying body size; (4) the Apparent (Sword) Length Preference Treatments where male distance from the female was held constant, while varying sword length; (5) the Apparent Length Controls where body length, sword length, and viewing distances varied while holding apparent length constant; and (6) the Experimental Treatment in which the smaller-sworded male of the pair appeared larger. In each of the Actual Length Control, Body, and Sword Length Treatments, females preferred the closer, larger-bodied, and longer-sworded males, respectively. However, females exhibited preferences in neither any of the Apparent Size Control treatments nor any of the Sword Compensation Treatment. Additionally, females in the Experimental Treatment preferred the apparently longer-sworded dummy to the male whose sword was actually longer. These results suggest that female preference for male body and sword length is based on the male’s apparent rather than actual size.     

Costs and benefits of maternal behaviour in the green lynx spider (Oxyopidae,Peucetia viridans)

I examined the adaptiveness of maternal behaviour in the green lynx spider Peucetia viridans (Hentz), by measuring the costs and benefits to the female of egg-sac tending. P. viridans guards her egg sac for 6–8 weeks until the young have emerged and dispersed. I removed females from egg sacs in the field either immediately after oviposition or after they had guarded the egg sacs for 2 weeks, and compared the fate of these sacs with that of control sacs which had females present throughout. Survival rates of egg sacs were significantly improved by the presence of a female. The major sources of mortality were ant predation and sac dislodgement, both of which were significantly reduced by a guarding female, and mantispid brood parasitism, which was not affected by the female. To measure the spiders' lifetime reproductive output, I allowed caged females to construct egg sacs. Half had their first egg sac removed shortly after oviposition, while half were allowed to guard. Significantly more non-guarding than guarding females produced second egg sacs. Thus for the green lynx spider, egg sac guarding decreases lifetime egg production, but increases offspring survival. Under north Florida conditions, the benefits of guarding outweight the costs, and the females have a higher lifetime reproductive success than if they did not guard.     

Response of the predatory mite <Emphasis Type="Italic">Cydnoseius negevi</Emphasis> (Acari: Phytoseiidae) to webbing of the date palm mite, <Emphasis Type="Italic">Oligonychus afrasiaticus</Emphasis> (Acari: Tetranychidae), on date palm fruits and leaves

The behavioral response of a generalist phytoseiid predator, Cydnoseius negevi (Swirskii & Amitai) (Acari: Phytoseiidae) to the complicated webbing of the date palm mite (DPM) Oligonychus afrasiaticus (McGregor) (Acari: Tetranychidae) was evaluated for the first time on date palm leaves and fruits, both in the laboratory and on field-collected samples. Cydnoseius negevi used its first pair of legs in a swimming-like to-and-fro movement to penetrate the complicated webbing of DPM and demonstrated three attack behaviors, i.e., patrolling (PG), web invasion (WI), and web penetration (WP), against the webbing of DPM on date fruits and leaves. The time spent by the predator on attack behaviors was significantly longer in the laboratory for treatments where either more prey females or immature stages along with females were present. The time spent by C. negevi on response behaviors such as searching, grooming, and resting increased with increased number of DPM females and with the addition of immature stages along with DPM females. Cydnoseius negevi, even though being a generalist phytoseiid predator, showed its potential towards penetrating the complex dense webs of DPM on date palm fruits and leaves. If released in suitable numbers prior to establishment of DPM colonies, C. negevi may prove to be an effective biological control agent.     

The relationship between female body size and egg size in pipefishes

Comparing five species of pipefish, egg size was significantly larger in species with brood pouches (Syngnathus typhle, Syngnathus acus and Syngnathus rostellatus) than in species without brood pouches (Entelurus aequoreus and Nerophis ophidion). Egg size correlated positively with female body size in species with brood pouches, but was similar across female sizes in the species lacking pouches. These results may reflect differences in offspring competition as a consequence of variable offspring relatedness within a brood, due to the mating systems adopted by the different species and the presence or absence of a brood pouch.     

Attack Behavior of Two Wasp Species of the <Emphasis Type="Italic">Polysphincta</Emphasis> Genus Group (Hymenoptera,Ichneumonidae) on their Orb-Weaver Spider Hosts (Araneae,Araneidae)

Species in the Polysphincta genus group, as far as is known, are exclusively koinobiont ectoparasitoids of spiders. These wasps attack their hosts, inflicting a temporary paralysis, and then lay one egg on the host’s abdomen or prosoma. Parasitoid attack behavior is highly variable among species, including occasions where the wasp darts directly and holds the spider, as well as instances involving complex behavioral sequences. In the present study, we describe the attack behavior of Polysphincta sp. nr. purcelli and P. janzeni on Cyclosa fililineata and C. morretes, respectively. All attacks occurred at night. Initially, the female wasp landed on the web hub at the position occupied by the spider, with the spider always escaping from this initial attack. Subsequently, the wasp waited for up to 14 h at the web hub for the spider’s return. The wasp then inserted its ovipositor into the mouth of the spider, after which the spider became paralyzed and remained motionless for at least 30 min. The wasp laid one egg on the surface of the host’s abdomen and remained on the web for at least 1 h thereafter. The lie-in-wait and attack only after the return of the host to the web hub, as well as the permanence of the wasp on the web after the attack are not frequent behaviors described for polysphinctines. Behavioral idiosyncrasies, such as those observed here, are common among polysphinctines, suggesting a high level of specific adaptive matching of polysphinctine parasitoid behavior to their hosts’ biological characteristics.     

Female size and fitness in the leaf-cutter bee Megachile apicalis

1. The effects of female size on fitness of the leaf-cutter bee Megachile apicalis Spinola (Hymenoptera: Megachilidae) were examined using artificial nesting sites in the field.
2. Although female size had no significant effect on female longevity or sex ratio of progeny, it did have a significant effect on fecundity; larger females attained greater realized fecundity than did smaller females.
3. This significant effect of female size on fecundity occurred because larger females produced cells faster than did smaller females.
4. Female size also had a significant effect on egg size; larger females laid larger eggs than did smaller females.
5. Female size had a significant effect on the size of investment in each progeny. The size of investment estimated by brood cell weight was greater for larger females than for smaller females. This pattern was largely absent, however, when the size of investment was estimated by adult progeny weight.
6. Female size had a significant effect on nest usurpation behaviour; larger females had a higher capacity to usurp nests than did smaller females.     

Male-male interactions and their influence on the mating behavior and success in the fountain darter, <Emphasis Type="Italic">Etheostoma fonticola</Emphasis>

Behavioral interactions between individuals of the same sex can affect the outcome of intersexual selection. For this reason, intrasexual selection is oftentimes examined independently from intersexual selection in studies aimed at understanding mating behaviors. However, a more complete understanding of sexual selection effects within a population can only come from exploring both intrasexual and intersexual selection and the potential interactions between the levels of selection. Association preferences in male and female Etheostoma fonticola do not reveal a size preference for same sex or opposite-sex individuals. However, male-male interactions may influence female choice in E. fonticola. We examined whether male E. fonticola that differ in body size exhibit differences in agonistic behaviors and mating success. Larger males do exhibit higher rates of aggressive behaviors and smaller males, in turn, exhibit more defensive behaviors. However, differences between larger and smaller males in male-male interactions did not translate into differences in spawning success. These results suggest that male size influences dominance relationships in E. fonticola but not mating success. There were also no differences between large and small males in mating attempts, which could be an outcome of the year-round breeding season found in this species, females laying eggs singly, or males fertilizing eggs individually.     

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.     

Potential for interspecific hybridization between <Emphasis Type="Italic">Zizina emelina</Emphasis> and <Emphasis Type="Italic">Zizina otis</Emphasis> (Lepidoptera: Lycaenidae)

Environmental changes such as global warming and biological invasion caused by human activities raise the possibility of secondary contact between the endangered butterfly species Zizina emelina and its sibling species Zizina otis in Japan. To assess the possible risks from their habitats overlapping, we investigated the potential for hybridization and the development of F1 individuals. We observed successful mating of the two sibling species under artificial conditions. The presence of a postzygotic hybridization barrier was supported by the delay of larval development only in females; a delay did not occur in males. Existence of the barrier was also supported by a decreased egg hatching rate in one brood; this was likely associated with infection with Wolbachia, a bacterium manipulating the reproductive capability of its host. The size and wing markings of F1 hybrid individuals were intermediate between those of the two species. These results suggest that, if Z. emelina and Z. otis are distributed sympatrically in the future, there is a possibility of introgression and reproductive interference between the two species, which would increase the risk of decline of each species.     

High brood patch temperature of less colourful,less pheomelanic female Barn Swallows Hirundo rustica

In birds, even a minor difference in egg temperature (1–1.5 °C) has been shown to affect the fitness of offspring by changing hatching success, incubation period and nestling quality. Female, but not male, passerines develop brood patches. Thus if there are traits, such as plumage ornamentation, that indicate optimal egg temperature, males should pair with females that exhibit those traits. However, no study has yet investigated the relationship between female brood patch temperature, which would directly affect egg temperature, and female plumage ornamentation. In this study, we examined the surface temperature of female brood patches during nocturnal incubation and examined its relationship with female plumage ornaments in Asian Barn Swallows Hirundo rustica gutturalis. After controlling for ambient air temperature, brood patch temperature was negatively associated with colour saturation of the female throat patch. No other female ornaments, such as tail‐length, white tail spots or throat patch size, predicted brood patch temperature. When oral (mouth) temperature was statistically controlled, females with less colourful throats and longer tails showed higher brood patch temperature, indicating that these females had hotter brood patches in relation to the temperature of other body parts. Furthermore, we found a negative relationship between pheomelanin pigmentation and brood patch temperature after controlling for ambient air temperature or oral temperature. To our knowledge, this is the first study to show that female ornaments can predict the absolute/relative thermal investment in brood patches. This relationship, together with other aspects of female quality, may affect male mate preference and female ornamentation.     

Egg size and reproductive adaptations among Arctic deep-sea copepods (Calanoida,<Emphasis Type="Italic"> Paraeuchaeta</Emphasis>)

Reproductive strategies of the four congeneric and sympatric calanoid copepods Paraeuchaeta glacialis, P. norvegica, P. barbata, and P. polaris were studied in the Arctic Greenland Sea. Females of all species produce egg sacs and carry their brood attached to the genital opening until the offspring hatch. However, egg size and lipid content as well as clutch size and the fraction of females carrying egg masses show characteristic differences among the four species. P. glacialis and P. norvegica produce large numbers (37 to more than 50) of relatively small eggs, whereas P. barbata and P. polaris rely on small numbers (10 to 19 and 4 to 6, respectively) of large eggs with a high energy content. There is no correlation between female body size and egg size or clutch size, respectively. Females of the smallest species, P. polaris, produce relatively large eggs and show the highest energetic investment per egg. In contrast, energetic investment per clutch is highest in P. glacialis. Reproductive adaptations appear to be strongly related to the depth range inhabited by the respective species. In the central Greenland Sea P. glacialis and P. norvegica occur in the epipelagial and upper mesopelagial, whereas the other two species inhabit lower mesopelagic to bathypelagic depths. Thus, egg size increases with depth of occurrence, whereas clutch size is inversely correlated with depth. This observation leads to the hypothesis that the evolution of large eggs in deep-sea copepods may enable hatchlings to rely on a lecithotrophic development and thus represents a successful adaptation to cope with the limited food supply at great depths, whereas high offspring numbers in epipelagic species compensate for higher predation risks in the euphotic zone.Communicated by H.-D. Franke     

Paternity analysis in an egg‐carrying aquatic insect Appasus major (Hemiptera: Belostomatidae) using microsatellite DNA markers

The giant water bug Appasus major exhibits a peculiar reproductive behavior where females lay eggs on the backs of males. A male and female pair performs frequent repeat copulations during the oviposition behavior, and the male carries the deposited eggs until hatching. Such characteristic behaviors predict that the eggs are fertilized by the brooding males. If males carry eggs of other unrelated males, the egg carrying will drastically decrease the fitness of the carriers. In this study, we examined genetic relationships between the 489 eggs and nine males carrying them collected from the field, using microsatellite DNA markers. We revealed that in total, 28.4% of the eggs were of other male origin. This is the first report of frequent brood parasitism in an aquatic egg‐carrying hemipteran insect. The brood parasitism is adaptive for the females probably because it enables them to increase the chance of oviposition, or it can equalize motility risk of the eggs of each mate.     

Spiderling sex ratio and maternal investment in the bolas spider Mastophora cornigera (Araneae,Araneidae)

Male Mastophora cornigera exit egg sacs as adults, which allowed us to determine spiderling sex ratios and patterns of maternal investment in this species. We collected 15 egg sacs produced by seven mothers, which yielded 1945 emergent spiderlings which were sexed, 1850 of which were weighed. Two emergent broods were significantly male and female biased and were unaffected by pre-emergence mortality. The weights of male and female spiderlings differed in eight broods, with males and females being heavier in four cases each. Five of these broods were derived from multiple egg sac sets produced by one mother, and in each case, the total mean male and female spiderling weights for all broods in a set were biased in the same direction as the biased brood(s) within that set. Mean emergent spiderling weight was independent of brood size and sex ratio for both males and females. Despite such independence, sex allocation in M. cornigera can favor sons, daughters, or both equally, and by numbers, by weight, or both at once. The proximate mechanisms and adaptive significance of such variability is unknown. We also review evidence for gender-biased allocations in arachnid offspring and suggested mechanisms for their applicability to M. cornigera.     

A lengthy solution to the optimal propagule size problem in the large-bodied South American freshwater turtle, <Emphasis Type="Italic">Podocnemis unifilis</Emphasis>

In oviparous vertebrates lacking parental care, resource allocation during reproduction is a major maternal effect that may enhance female fitness. In general, resource allocation strategies are expected to follow optimality models to solve the energy trade-offs between egg size and number. Such models predict that natural selection should optimize egg size while egg number is expected to vary with female size, thus maximizing offspring fitness and consequently, maternal fitness. Deviations from optimality predictions are commonly attributed to morphological constraints imposed by female size, such as reported for small-bodied turtle species. However, whether such anatomical constraints exist in smaller-bodied females within large-bodied clades remains unstudied. Here we tested whether resource allocation of the river turtle Podocnemis unifilis (a relatively smaller member of the large-bodied Podocnemididae) follows optimality theory, and found a pattern of egg elongation in smaller females that provides evidence of morphological constraints and of a reproductive trade-off with clutch size, whereas egg width supports the existence of an optimal egg size and no trade-off. Moreover, larger females laid larger clutches composed of rounder eggs, while smaller females laid fewer and relatively more elongated eggs. Elongated eggs from smaller females have larger volume relative to female size and to round eggs of equal width. We propose that elongated eggs represent a solution to a potential morphological constraint suffered by small females. Our results suggest that larger females may optimize fitness by increasing the number of eggs, while smaller females do so by producing larger eggs. Our data supports the notion that morphological constraints are likely more widespread than previously anticipated, such that they may not be exclusive of small-bodied lineages but may also exist in large-bodied lineages.     

Description of a new species of <Emphasis Type="Italic">Ixodes</Emphasis> Latreille, 1795 (Acari: Ixodidae) and redescription of <Emphasis Type="Italic">I. priscicollaris</Emphasis> Schulze, 1932, parasites of New Guinea rodents (Rodentia: Muridae)

Ixodes goliath n. sp. (Acari: Ixodidae), is described based on females collected from the eastern hyomys, Hyomys goliath (Milne-Edwards) (Rodentia: Muridae) from Papua New Guinea. Females of I. goliath n. sp. are similar to those of I. priscicollaris Schulze, 1932 but can be distinguished by the overall size, porose areas sculpture and shape of palpal segment I ventrally. For comparative purposes, the female of I. priscicollaris is redescribed. Studied females of I. priscicollaris were found on murid rodents and Phascogale sp. (Dasyuromorphia: Dasyuridae) from Indonesia (Papua Province) and Papua New Guinea.     

Experimentally altering pre‐breeding sex steroids reduces extra‐pair paternity in female tree swallows

Extra‐pair paternity commonly occurs in many socially monogamous animal species, yet the reasons why females mate with males outside the social pair bond remain poorly understood. Because sex steroids mediate aggressive and sexual behaviors that peak prior to egg laying in female birds, we tested whether they also influence the rate of extra‐pair paternity in nests of female tree swallows (Tachycineta bicolor). In one treatment, we experimentally elevated testosterone (T) using implants containing exogenous T, and in another treatment, we blocked the estrogenic and androgenic actions of T using implants containing 1,4,6‐androstatrien‐3,17‐dione in combination with flutamide (ATD+F). Females with empty implants served as controls. Nests of females treated with T and those with ATD+F were less likely to contain extra‐pair offspring and had a lower proportion of extra‐pair offspring, compared to control females. Treating females with T also delayed clutch initiation dates and disrupted incubation behavior, as found in previous studies, but clutch sizes of T females did not differ from controls. Females treated with ATD+F tended to lay larger clutches than control and T‐treated females, which may be due to their clutch sizes not declining with later initiation dates as found in the T and control treatments; however, after controlling for brood size, ATD+F females produced nestlings that were lighter at day 16 than control females. Although the effect of T on extra‐pair paternity that we observed in tree swallows is consistent with previous studies in female birds, our results demonstrate that blocking the estrogenic and/or androgenic actions of T during pre‐breeding also lowers extra‐pair paternity. Overall, our study suggests that extra‐pair paternity in tree swallows is mediated by sex steroids of females.