盐长蝽亚科外部形态及其系统学意义的研究：半翅目：长蝽科

本文研究了半翅目长蝽科盐长蝽亚科（Ｈｅｎｅｓｔａｒｉｎａｅ）的外部形态,希图为长蝽总科复合体的系统发育研究提供资料。材料选用盐长蝽ＨｅｎｅｓｔａｒｉｓｏｓｃｈａｎｉｎｉＢｅｒｇｒｏｔｈ１９１９和卤长蝽ＥｎｇｉｓｔｕｓｓａｌｉｎｕｓＪａｋｏｖｌｅｖ１８７４,分别作为该亚科３个成员属中两个属（即盐长蝽属ＨｅｎｅｓｔａｒｉｓＳｐｉｎｏｌａ１８３９和卤长蝽属ＥｎｇｉｓｔｕｓＦｉｅｂｅｒ１８６４）的代表。文章还初步提出了该亚科的一些共有新征,并作了有关讨论,认为此分类单元为一单系群。     

The extent and costs of reproductive interference among four species of true bug

Reproductive interference arises when individuals of one species engage in reproductive activities with individuals of another, leading to fitness costs in one or both species. Reproductive interference (RI) therefore has two components. First, there must be mis-directed mating interactions. Second, there must be costs associated with these mis-directed interactions. Here we consider RI between four species of true bug in the family Lygaeidae, focusing in particular on the fitness consequences to Lygaeus equestris. The species we consider vary in their relationships with each other, including species in the same or different genus, and with or without natural overlap in their geographic ranges. First we show that inter-specific mating interactions, although not a certain outcome, are common enough to perhaps influence mating behaviour in these species (arising in up to 10 % of inter-specific pairings). Second, we show that reproductive interference can seriously reduce female fitness in L. equestris. Importantly, different species impose different costs of RI on L. equestris, with interactions with male Spilostethus pandurus inflicting fitness costs of similar magnitude to the costs of mating with con-specifics. On the other hand, mating interactions with male Oncopeltus fasciatus appear to have no effect on female fitness. In a follow-up experiment, when we allowed competition amongst just females of S. pandurus and L. equestris, the fitness of the latter was not reduced, arguing more strongly for the role of reproductive interference. However, in our final experiments under mass mating conditions with extended ecological interactions (including scope for competition for resources and cannibalism), the costs of RI were less apparent. Our data therefore suggest that the costs of RI will be context-specific and may act in concert with, or be swamped by, other ecological effects. We suggest that comparative studies of this sort that both mimic naturally occurring reproductive interference events, and also artificially generate new ones, will be necessary if we are to better understand the ecological and evolutionary significance of reproductive interference.     

Are environmental factors responsible for geographic variation in the sex ratio of the Greenlandic seed‐bug Nysius groenlandicus?

Until recently nothing indicated an unequal sex ratio in the widespread Greenland seed‐bug Nysius groenlandicus (Zetterstedt) (Heteroptera: Lygaeidae). However, recently populations more or less devoid of males were discovered in high arctic Northeast Greenland. This initiated an inspection of the entire material of the species collected in Greenland and now preserved at the Zoological Museum, University of Copenhagen. It was found that the sex ratio varied significantly among different locations. In most cases females were most abundant, but males were either scarce or absent only in samples from Northeast Greenland, indicating that here the species reproduces asexually. This paper demonstrates that the differing sex distributions can be explained by climatic factors (temperature, precipitation) and that the degree of continentality (distance from the open sea) promotes female‐biased sex ratios.     

Females Remate for Sperm Replenishment in a Seed Bug: Evidence from Offspring Viability

Females of many insects mate multiply but why they do so remains controversial. Here we investigated the effects of multiple matings on female reproductive success of a New Zealand seed bug, Nysius huttoni. We found little evidence for females to gain material (nutritional) benefits through multiple matings because the number of matings did not have significant effect on female fecundity. Females remated to the same males or different males produced similar number of viable offspring, suggesting that females do not obtain genetic benefit from remating in terms of offspring viability. With the increase of the number of matings, however, overall fertility rate significantly increased and daily fertility rate declined significantly slower over time. These results suggest that females remate for the replenishment of sperm. Five matings are sufficient for females to maximize their reproductive success, and additional matings appear to be superfluous. However, the females of this bug mate as many as 68 times if males and females are paired for lifetime. This can be explained by the convenience hypothesis, i.e., females remate superfluously to minimize the costs of harassment by promiscuous males.     

Coexistence of bisexual and unisexual populations of Nysius groenlandicus in the Zackenberg Valley,Northeast Greenland

The seed‐feeding true bug Nysius groenlandicus (Zetterstedt) (Heteroptera: Lygaeidae) has temperature and humidity preferences similar to many desert‐inhabiting insects. Nevertheless, it is widely distributed in Greenland and occurs even in high arctic areas. This seeming paradox implies that the species has developed physiological, ecological, and behavioral mechanisms that enable it to cope with the extreme arctic environment. One of these adaptations is its ability to reproduce both sexually and asexually. Sexual reproduction is most common in coastal populations, whereas asexual reproduction (parthenogenesis) occurs mainly in inland populations. Preliminary data from the Zackenberg Valley in Northeast Greenland indicated that both reproductive strategies occurred within the same area. We therefore decided to focus on this area to find evidence of (1) coexistence of unisexual and bisexual populations, (2) factors that correlate with the distribution of the two reproductive types, and (3) morphological differences between females from sexual and asexual populations. We sampled N. groenlandicus at 51 sites within the Zackenberg area. Males and females were found in almost equal numbers in only four of the samples, and only in samples taken close to the coast line, whereas samples from the inner part of the valley consisted almost entirely of females. Thus, the distribution of uni‐ and bisexual populations at the local scale agreed with the pattern previously found for Greenland as a whole. Using data obtained from two climate stations operated in the area, we showed that average summer temperatures tend to be more favorable for N. groenlandicus in the inner part of the valley than at the coast. The inland populations inhabited small isolated patches, whereas the sexual populations were found in the more homogeneous areas along the coast. Data support the hypothesis that coexistence of the two reproductive types is maintained by a combination of environmental variability, which gives sexual reproduction an advantage, and metapopulation dynamics characterized by frequent extinctions and re‐colonizations, which favors asexual reproduction. Head width of females from unisexual populations was on average 7% larger than that of females from bisexual populations. The mechanism behind this morphological difference needs further investigation.     

The evolutionary ecology of the Lygaeidae

The Lygaeidae (sensu lato) are a highly successful family of true bugs found worldwide, yet many aspects of their ecology and evolution remain obscure or unknown. While a few species have attracted considerable attention as model species for the study of insect physiology, it is only relatively recently that biologists have begun to explore aspects of their behavior, life history evolution, and patterns of intra‐ and interspecific ecological interactions across more species. As a result though, a range of new phenotypes and opportunities for addressing current questions in evolutionary ecology has been uncovered. For example, researchers have revealed hitherto unexpectedly rich patterns of bacterial symbiosis, begun to explore the evolutionary function of the family's complex genitalia, and also found evidence of parthenogenesis. Here we review our current understanding of the biology and ecology of the group as a whole, focusing on several of the best‐studied characteristics of the group, including aposematism (i.e., the evolution of warning coloration), chemical communication, sexual selection (especially, postcopulatory sexual selection), sexual conflict, and patterns of host‐endosymbiont coevolution. Importantly, many of these aspects of lygaeid biology are likely to interact, offering new avenues for research, for instance into how the evolution of aposematism influences sexual selection. With the growing availability of genomic tools for previously “non‐model” organisms, combined with the relative ease of keeping many of the polyphagous species in the laboratory, we argue that these bugs offer many opportunities for behavioral and evolutionary ecologists.     

Semiochemistry of aposematic seed bugs

(E)-2,7-Octadienyl acetate and (E)-2-octenyl acetate (1:10 by volume) were identified as a pheromone attractive to both sexes of the lygaeid bug, Tropidothorax cruciger. In a parallel investigation of Neacoryphus bicrucis (Lygaeidae), (E,E)-2,4-hexadienyl acetate and phenethyl acetate (≈9:1) were identified from males, and found attractive to both sexes of adults in the field plus a tachinid fly parasitoid of the bugs. In N. bicrucis, the pheromone was clearly shown to come from the tubular accessory glands of the metathoracic scent gland; this evidence, plus earlier literature reports for other species, indicate that male lygaeids are the pheromone emitters. In another lygaeid, Oncopeltus fasciatus, 2-isobutyl-3-methoxypyrazine was identified in the cardiac glycoside-laden fluid sequestered from milkweed hosts and expelled by these bugs when they are attacked. Alkyl methoxypyrazines are warning odorants associated with poisonous insect secretions, and their presence in O. fasciatus indicates that the plant-derived chemical defense of lygaeines is more elaborate than previously appreciated.     

Precopulation Sexual Selection in Nysius huttoni White (Heteroptera: Lygaeidae) in Relation to Morphometric Traits

Nysius huttoni White is a polygamous bug, endemic to New Zealand, and an important pest of wheat and brassicas. This bug has a female-biased sexual size dimorphism but relative to body length, males have longer antennae, suggesting that the allometric scales of antennal–body relationships may be highly selective in sexual selection. Body weight and most morphometric traits measured have no effect on mating success of either sex. Males significantly preferred mating with females having thicker abdomens, more mature eggs, and longer ovipositors. This result suggests that males may select their mates on the basis of immediate reproductive benefit: fertilizing more eggs and ensuring better survival of these eggs. Males with large genital structures have mating advantages over those with small ones, suggesting that precopulation sexual selection in this species act on male genital traits rather than body weight and nonsexual traits. Finally, females significantly preferred males with greater slopes for the antennal-body relationship for mating. The allometry in the male antennal length may be an indicator of male reproductive fitness.