Spatial relationships between mate acquisition probability and aggregation size in a gregarious coreid bug, (Colpula lativentris): A case of the ideal free distribution under perceptual constraints

Summary Spatial relationships of mate acquisition probability for individuals of both sexes of a gregariously-mating coreid bug,Colpula lativentris, were studied in relation to aggregation size. Operational sex ratio was always strongly male biased. Mate acquisition probability of females was rather constant and independent of aggregation size, as predicted by an ideal free distribution. Moreover laboratory experiments showed that both multiple mating and rearing density little affected female fecundity, suggesting ideal free distribution of females in terms of reproductive success. On the other hand, mate acquisition probability of males was higher in larger aggregations, where more receptive females were available. This male discrepancy from an ideal free distribution was similar to the patterns predicted by an ideal free distribution under perceptual constraints (Abrahams, 1986), but not by that under unequal competitive ability.     

Comparative population studies of threePieris butterflies,P. rapae, P. melete andP. napi, living in the same area

Summary Utilization of patchy habitats by adult populations of threePieris butterflies,P. rapae, P. melete andP. napi was studied throughout the flight season in an area of their coexistence, about 3×1.5 km, in a farm village in the mountains in Inabu, Aichi Prefecture. Field study was by the mark-recapture method. Results were analyzed by dispersal distances and recapture duration decay curves for adults of different age-classes estimated on the basis of physical condition of their wings, together with supplementary information of daliy egg-laying rate of females, obtained in field cages. Sexually immature, mated femals ofP. rapae after teneral stage showed a migratory flight. On the other hand, reproductive females and all males ofP. rapae were strongly resident within suitable habitats, and reproductive females begun to lay eggs abundantly at sunny places of newly suitable areas within a short period.P. melete seemed to disperse gradually from emerged stites and females of this species continued to lay some constant numbers of eggs for more than ten days over a wider area.P. napi appeared more likeP. melete thanP. rapae. The habitats of the three species can be characterized as follows:P. rapae, temporary, continued for pre-reproductive females but localized for reproductive females and all males, and unstable;P. melete, permanent, widespread, and stable;P. napi, permanent, localized, and stable. The numbers of generations ofP. rapae, P. melete andP. napi were estimated to be about six, three and three, respectively. Seasonal fluctuations in the number of adults were influenced by the stability of their habitats, i. e., the population size fluctuated sharply inP. rapae, but it was much more stable inP. melete andP. napi. In view of these results, it can be said thatP. rapae fits the general characteristics of a r-strategist whereasP. melete andP. napi are more K-strategic thanP. rapae.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

The influence of urine from asymptotic laboratory populations on sexual maturation in prairie deermice

Summary Male and female prairie deermice housed singly from weaning (21 days) until 40 days of age had urine from one of the following sources applied daily to their nostrils: sexually mature laboratory population males or females, and sexually mature breeder males or females from our laboratory colony. Other males and females were treated with water. Records were obtained of the weights of the tests, seminal vesicles and bacula of males and the ovaries and uteri of females at 40 days of age. The major results of this study were as follow: Female urine from sexually mature and inhibited population or isolated mice stimulated the development of the male reproductive organs significantly more than water or male urine. Male urine applied to young males did not differentially affect the weights of their reproductive organs compared with water and thus showed no inhibition effects. The reproductive organs of females treated with male urine did not differ significantly in weight from those of females treated with female urine. Females treated with water developed significantly larger uteri than those treated with urine from sexually mature population females (P≤0.02), inhibited population females (P<0.001) and inhibited population males (P<0.007).     

A male fitness advantage to wing reduction in the oriental chinch bug,Cavelerius saccharivorus okajima (heteroptera: Lygaeidae)

Summary A male fitness advantage to wing reduction was investigated for the oriental chinch bug,Cavelerius saccharivorus, which is wing dimorphic. Field surveys for the frequency of matings between wing morphs showed that the mating probability was much higher in brachypterous males than in macropterous males in the early breeding season. Brachypterous males copulated with not only brachypterous females but also macropterous females in this season. This led to a considerable rate (30%) of insemination of macropterous females just before emigration even in the early breeding season. A rearing experiment for the pre-reproductive period of females revealed that females mated with brachypterous males copulated and oviposited earlier than females mated with macropterous males. These results indicate that there is a fitness advantage to wing reduction in the males ofC. saccharivorus in terms of the earlier sexual maturation.     

Kin-related social organization in a winter population of the voleClethrionomys rufocanus

Kinship amongClethrionomys rufocanus was investigated during the winter of 1992/93 in a 3-ha enclosure using both molecular and catch-mark-release techniques. Forty-six adult voles (22 males and 24 females) having high heterozygosities, which were collected from several natural populations, were released into the enclosure on 29 September 1992. Most fall-born individuals of both sexes stayed in their natal site during the non-breeding period (December–March), although reproductively active females dispersed during the fall breeding season (October–November). These philopatric individuals aggregated and formed an maternal family in the winter. Several females which failed to reproduce were solitary during this season. Some individuals which were derived from several families also aggregated into a mixed lineage group. Survival rate of fall-born voles from earlier litters was higher than that from later ones. Maternal families broke up soon after the onset of spring reproduction. Most females established a territory near the wintering site and made a kincluster, in which close relatives neighbored each other. Maternal families in winter bring about female kin-clusters in spring, which may influence reproductive output in the breeding season.     

Spatio-temporal variation of natural and sexual selection in breeding populations of the coreid bug,Colpula lativentris (Heteroptera: Coreidae)

Spatio-temporal variations of lifetime reproductive succes (LRS) of both male and female individuals of a coreid bugColpula lativentris were measured and analyzed using the multiple regression method of Arnold and Wade (1984a, b). The standardized variance of LRS was larger in males than that in females as males often to secure mates for a long period whereas females could easily find mates and oviposit simply dependent on ovarial maturation. LRS was partitioned into 4 consecutive fitness components: (1) reproductive lifespan, (2) copulating efficiency, (3) guarding efficiency (for males) or oviposition efficiency (for females), and (4) number of eggs per clutch. In males copulating efficiency was the largest determining factor of LRS, whereas in females reproductive lifespan was the most important factor. Such tendencies were stable on both a yearly and local basis. Patterns of relative contribution of natural selection (reproductive lifespan and number of eggs per clutch) and sexual selection (copulating efficiency and guarding or oviposition efficiency) to LRS were clearly different between males and females. This sexual difference is, at least to some extent, thought to be brought about by sexual selection among males for mating opportunity, though no physical fight was observed among males. Directional selection on body length was found only in relation to the clutch size of females because large females tended to lay larger clutches. No significant directional selection was found in other fitness components.     

Adult populations of the sugi bark borer,Semanotus japonicus Lacordaire (Coleoptera: Cerambycidae), in Japanese cedar stands: Population parameters, dispersal, and spatial distribution

Summary Adult populations of the sugi bark borer,Semanotus japonicus Lacordaire, which attacks the trunks of Japanese cedar and cypress, were studied in two cedar stands in 1981 and 1984, using a band-trapping method developed by the author. Adults began to emerge and fly earlier in 1981 than in 1984 because of the higher temperature after hibernation in 1981. Population size wasestimated by a markrelease-recapture method. The daily survival rate ranged from 0.86 to 0.92 and the mean longevity from 7.0 to 11.9 days. Females were fewer than males throughout the season, especially early in the flight period, because males consistently emerged earlier. The mean distances traversed among the cedar trees were 9.2 m for males and 16.3 m for females; maximum dispersal was 79.5 m for males, 149.5 m for females. There are two different patterns in their dispersal movement: the small-scale movement which consists of walking at cool temperatures and large-scale flight at warm temperatures. Adults of this species are somewhat sluggish and between-tree movement is infrequent. This lack of movement resulted in a contagious distribution pattern of adults.     

Competitive habitat utilization in the damselfly,Mnais nawai (Zygoptera: Calopterygidae) coexisting with a related species,Mnais pruinosa

Reproductive behaviors related to habitat utilization were studied in males of the damsefly,Mnais nawai, which has two male forms, territorial orange-winged males (nawai) and non-territorial pale-orange-winged males (sahoi), at the upper part of a mountain stream where they partiallycoexist with a related species,Mnais pruinosa, which also has two male forms, territorial orange-winged males (esakii) and non-territorial hyaline-winged males (strigata). These two species showed parapatric distribution; the lower part of the stream was occupied byM. nawai, and the upper part byM. pruinosa. In the present study, cross-matings occurred between bothMnais species, although normal intraspecific matings occurred more frequently than cross-matings. Territorial males of both species copulated with conspecific females that entered their territory and guarded the ovipositing females, probably to avoid sperm displacement resulting from subsequent copulations. Severe competition for oviposition sites by territorial males even occurred between the two species. On the other hand, non-territorial males of both species have alternative mating strategies (including several tactics such as sneaking, takeover and interception). The possible benefits from conflict among territorial males of both species is discussed.     

Scramble competition polygyny in the coreid bug (Colpula lativentris)

Summary The mating system of the coreid bug,Colpula lativentris, was studied, and compared with those in other hemipteran bugs. Copulating pairs formed a compact aggregation on a shoot of the Japanese knotweed,Polygonum cuspidatum, and there continued copulation and feeding for more than one day. This aggregation was defined as “a shoot aggregation”. Other than this aggregation composed of copulating pairs, there were loose aggregations on the ground composed of males and females not in copula. This loose aggregation was defined as “a ground aggregation”. Males searched for receptive females mostly in a ground aggregation. Sex ratio was constant and variance of sex ratio was also very small in any shoot aggregations and in large ground aggregations. However, in small ground aggregations, sex ratio was strongly male biased and variance of sex ratio was fairly large. This was because receptive females tended to leave a small ground aggregation and join a large one. These results suggest that a male can attain higher mating success in a larger ground aggregation.     

Survival rate and sex ratio in habu,Trimeresurus flavoviridis (Viperidae), on the subtropical Okinawa Island, Japan

We examined the age structure and sex ratio of a viperid snake,Trimeresurus flavoviridis, based on samples collected by hand by local inhabitants, habu hunters and by baited traps in the middle and south of Okinawa Island, Japan. Small individuals had a low possibility of capture in all the collecting methods. The age structure was estimated through summing up the age frequency in each snout-vent length (SVL) class in samples with a similar SVL structure. From the age structures, annual survival rates of adult females and males were estimated to be 0.7 and 0.8, respectively. In most samples the sex ratio was biased towards males, especially in March and August and in large individuals. However, the female proportion increased in June.     

Spatial pattern indices based on distances between individuals on a line-segment with finite length

Summary Let us consider a strip-wise habitat of line-segment, like a corridor, to simplify the subject mathematically, and assume that the length of the habitat is γ and there aren individuals. Here, we assume that the spatial pattern of the individuals is random if then distances from the left end of the habitat to each individual follow a uniform distribution on the strip. Under such an assumption, the variance of the distances between any two neighbors is represented by the formulanθ ²(n+1)⁻²(n+2)⁻¹ and the variance betweenn+1 distances betweenn individuals from the left end to the right end to the strip, is represented by the formulanθ ²(n+1)⁻²(n+2)⁻¹. These two kinds of variances can be used for determining (1) the spatial pattern of a population on the strip and (2) the spatial structure within the population, by comparison with the variances calculated from the data. Two examples cited from the literature, a cattle population on a pasture and an aphid population on a sycamore leaf, are presented.     

Mating and diapause in hibernating adults ofMenida scotti puton (Heteroptera: Pentatomidae)

InMenida scotti aduts which mate during hibernation, we examined the effect of photoperiod on diapause induction in terms of the developmental degree of reproductive organs and corpora allata. In a hibernating population, mating season and physiological changes related to dipause were investigated. When newly emerged adults were reared under long- or short-day conditions, reservoir of the ectodermal accessory gland and corpora allata in males developed, while the developments of ovary and corpora allata in females were suppressed under both conditions. This suggests that diapause of this species is induced only in females independently photoperiodic conditions during adult stage. In a hibernating population, mating was observed from early to late November and from late March to early May. Observations of the development of reproductive organs and corpora allata suggest that diapause is induced only in females in the former period, while both sexes are not in diapause in the latter period.     

A population study of the japanese wood mouse,Apodemus speciosus (Mammalia: Muridae), with reference to its social behavior

Summary The role of social behavior on the population regulation of the Japanese wood mouse,Apodemus speciosus, was studied in the Ashu Experimental Forest of Kyoto University by monthly trapping and direct observation on the social behavior at the artificial feeding site. The census was carried out from May to October in 1974 and 1975, and the direct observation in June, August and October, 1975. In the study area, the Japanese wood mouse has two breeding seasons in a year: in spring and in autumn. Minimum survival rate was high in the non-breeding season and low in the breeding season. It was negatively correlated to the number of sexually active adults in both sexes. Home ranges of females tended to be mutually exclusive, and female residents prevented the settlement of female immigrants. On the other hand, home ranges of males overlapped largely and males had a straight social hierarchy. Subordinate males tended to disappear more often than dominant males did, but male residents did not hinder the settlement of male immigrants. Males may regulate their number by driving out the subordinate males, while females by spacing behavior. Contribution from the Laboratory of Animal Ecology, Department of Zoology, Kyoto University, No. 445.     

Decrease in respiratory rate in a wolf spider,Pardosa astrigera (L. Koch), under starvation

Summary Effects of starvation on the suryival period and the respiratory rate in adults of a wolf spider,Pardosa astrigera (L. Koch), were investigated. The spiders used were divided into four groups: well-fed, starved and two limited food groups; in the latter two, each spider was supplied with one leafhopper every second or third day. Adult males and females ofP. astrigera could survive for a long time; 28.8±2.7 days and 54.4±18.9 days, respectively, without any food. The longevities shown here were 73.8% for males and 78.6% for females of those of well-fed spiders, indicating thatP. astrigera adults have a strong tolerance to starvation. The respiratory rate of well-fed adults showed no tendency to increase or decrease with their aging; the mean respiratory rates were 4.86×10⁻⁴ mg CO₂/mg f.w. (fresh body weight)/hr for males and 3.80×10⁻⁴ mg CO₂/mg f.w./hr for females. The respiratory rates of starved spiders increased during the first two days of starvation but decreased markedly from the third to the twelfth day, and thereafter retained an almost constant level for each sex. The mean respiratory rates after the twelfth day of starvation were 2.49×10⁻⁴ mg CO₂/mg f.w./hr for males and 2.76×10⁻⁴ mg CO₂/mg f.w./hr for females; these values were respectively 48.4% and 63.0% of those prior to starvation. The fresh body weight of starved spiders decreased linearly with time but the rate was small. The respiratory rates of the limited food groups tended to decline with time and thereby their weight losses were minimized. The decrease in the respiratory rate under starvation was considered not to be due to spider exhaustion or senescence but due to an intrinsic change in behaviour and/or metabolism, because when the spiders were supplied with ample food for five days after starvation, the respiratory rate and the body weight rapidly recovered to near the levels prior to starvation. It is suggested that starved spiders use a higher ratio of fat as catabolic substrate than normally fed or satiated ones. Feeding strategies of poikilo-therm predators are discussed. This work was partially supported by the Nippon Life Insurance Foundation Research Fund and Grant-in-Aid (No. 56480039) from the Ministry of Education, Science and Culture, Japan.     

Factors affecting female-biased sex ratio in a trap-nesting wasp, Trypoxylon malaisei

We examined the female-biased sex ratio of a trap-nesting wasp Trypoxylon malaisei considering the following factors: (1) local mate competition (LMC), (2) resource quality, (3) partial bivoltinism, and (4) presence of constrained females. The sex ratio (expressed as male ratio) at emergence was strongly female biased, i.e., 0.30 and 0.19, in terms of the number and investment, respectively. To evaluate the primary sex ratio, we analyzed the data from nests where all the offspring successfully emerged, excluding nests composed of single-sex offspring. The primary sex ratio was also female biased, at 0.33 and 0.21, in terms of the number and investment, respectively. LMC was highly responsible for the female-biased sex ratio because both the nonrandom oviposition sequence [females at inner cells and male(s) at outer cells] and earlier emergence of males allowed sib-matings to occur. In contrast, the other three factors little affected the female-biased sex ratio: the sex ratio was fairly constant when resource quality (nest size) varied, partial bivoltinism was extremely rare or absent, and constrained females were absent or did not reproduce at all. Received: June 19, 1998 / Accepted: January 18, 1999     

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

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

Effects of grouping and fighting on the reproductive tracts of male white-footed mice (Peromyscus leucopus)

Adult, male white-footed mice (Peromyscus leucopus) were subjected to a variety of social situations ranging from isolation during the 20 day experimental period to constant contact with both females and other adult males. Contacts included grouping (three or four males per cage) and exposure to fighters (once daily for 20 minutes). The following measurements were recorded: weights of the body, testes, epididymides, vesicular glands, vesicular gland tissue (wet and dry), seminal fluid of the vesicular gland, adrenal glands, and baculum; spermatozoan reserves of the testes and epididymides. Grouping significantly affected both the weight and spermatozoan reserves of the testes and epididymides, as well as both the tissue and seminal fluid weight of the vesicular glands. The results suggested a graded effect of all treatments on the reproductive tract. In order of magnitude of the associated response, from none to greatest, the treatments may be ranked as follows: pairing with females, isolating, handling, fighting, and grouping. All reproductive parameters measured showed this general ranking, suggesting that the response to the various treatments was similar and differed only quantitatively. The results further suggested decreased secretion of LH and testosterone, although measurements of testosterone did not substantiate this conclusion. The lack of significant effects of grouping on adrenal gland weights strengthened the argument that adrenal involvement is not a necessary adjunct to the suppression of the reproductive tract in groupedPeromyscus, but the adrenal may be involved if contacts between males result in overt fighting.     

Lifetime mating success of males in a natural population of the papilionid butterfly,Atrophaneura alcinous (Lepidoptera: Papilionidae)

Summary Lifetime mating success of males in a natural population of the papilionid butterfly,Atrophaneura alcinous, was investigated and causes of the variation were examined. The most successful males mated with 5 females, whereas about 73% of the males failed to mate. Body size of males was not correlated with their eclosion date, longevity and lifetime mating success. There was no trade-off between mating success and longevity, and long-lived males had a disproportionately high mating success. Although number of available females per male per day was not variable among males with different longevities, long-lived males had higher mating efficiency. Time interval between matings by non-virgin males was shorter than that from eclosion to the first mating. High lifetime mating success of long-lived males was strongly related to their mating experience, not to their ageper se.     

Ecological studies on a dragonfly,Nannophya pygmaea ramber (Odonata: Libellulidae) I. seasonal changes of adult population and its distribution in a habitat

Summary Adult population of a dragonflyNannophya pygmaea Ramber inhabited in a damp ground was investigated with mark-and-recapture method in 1975. The following results about the seasonal changes of the population size and distribution in the habitat were obtained. Adults emerged from late May to mid August. The number of the adults was most abundant in early June, but that of matured males in early July. From the recapture data, the estimate of daily survival rate was 0.82, and the length of immature stage in males was estimated as 5 days or so. Total number of post-teneral adults emerged in the habitat was estimated as about 9,000. The mean crowding-mean density regression method was applied for the analysis of the distribution pattern of the adults. Matured males showed a spaced-out distribution, while females and immatured males distributed themselves rather aggregatively. Such a distribution pattern of the matured male would be attributed to their territorial behaviour. The territorial behaviour was considered to force the matured males to extend the distribution area in July when they were most abundant. From the above-mentioned results and some observations, the meaning of the territoriality in this species was discussed.