Summary 1.In patchily distributed plant species, seed production is likely to be influenced both by local abiotic factors affecting plant size and conditions for fruit maturation, and by population characteristics affecting the intensity of interactions with mutualists and antagonists. However, the relative importance of these effects is poorly known. 2. We used multiple regression and path models to examine the importance of abiotic factors (sun exposure, soil depth) and population characteristics (size, density and connectivity) for variation in flower and fruit production and intensity of seed predation among 39 populations of the long-lived herb Vincetoxicum hirundinaria in three consecutive years. In addition, we manipulated water availability in a field experiment and recorded short-term and long-term effects on fruit output, and conducted a supplemental hand-pollination experiment. 3. Flower production varied little, while fruit initiation, fruit abortion and fruit predation varied considerably among years. Sun exposure and soil depth affected fruit production per plant indirectly and positively through their effects on flower number. Population density affected fruit production negatively through its effect on flower number. Both fruit initiation and the proportion of fruits attacked by the tephritid fly Euphranta connexa were related positively to population size. 4. The number of full-size fruits per plant was related positively to sun exposure and population size in two years each, and related negatively to population density in one year. However, because of seed predation, the number of intact mature fruits was related significantly to population characteristics in only one of three years. 5. The field experiments showed that both shortage of water and insufficient pollination may limit fruit set in V. hirundinaria . 6. Synthesis . These results demonstrate that the relative importance of local abiotic conditions and population characteristics may vary considerably along the chain of events from flower formation to intact fruit, and also among years. They further show that, at least in species with a naturally patchy distribution, connectivity may be relatively unimportant for variation in reproductive output compared to effects of habitat quality, population size and density.
Vincetoxicum hirundinaria is a longlived perennial herb. Its pod production was measured during seven years in about 40 small isolated patches in two study areas in southeastern Sweden. Total pod production in the two areas varied synchronously and up to about 200 times between years. This variation was mainly due to drought conditions as determined by the interplay of weather (sunshine h) and site characteristics.Populations of a univoltine tephritid fly, euphranta connexa, whose larva feeds on V. hirundinaria seeds in the maturing pods, were monitored for five and seven years in the two study areas respectively. Larval fly populations in the two areas varied 30 and 50-fold between years, with highs in years when pods were abundant and lows when pods were scarce. The percentage of pods attacked by E. connexa, however, varied in the opposite direction, with very high attack rates (about 100%) when pods were scarce and low rates (down to 10-20%) when pods were common. Thus the temporal tracking of food resources by E. connexa was poor, resulting in yearly variations in the amount of unattacked pods (and seeds) being about 2,00-fold.Many host plant patches were small and totally without pods in some years, causing frequent local extinctions of E. connexa. Patches were, however, readily colonized in later years when pods appeared again. Low overall attack rates in certain years were thus only to a minor extent explainable by host plant patches being uncolonized by the fly. The efficient spatial tracking of resources by the fly population allows this population system to be analyzed largely in terms of its temporal dynamics.Although E. connexa populations often exploit only a minor part of their potential larval resources, the rate of population change was closely correlated with these resources expressed on a per capita basis. Natural enemies only have negligible effects on E. connexa population trends. In this population system the herbivore (seed predator) population is thus controlled by its food resources, but weather-imposed temporal variations in resource set are too large for the herbivore population to closely track its resource base. This temporal tracking inertia has important implications for the long-term production of healthy seeds.
1. Horvathiolus gibbicoflis (Costa), a ground-living seedfeeding bug of the mediterranean region, has two wing morphs. In macropterous bugs both pairs of wings are fully developed. In brachypterous ones forewings are reduced to about two-thirds and hindwings to less than a third of their length in macropters.2. Each morph bred true with regard to wing length when reared under variable density, food and temperature conditions for several generations.3. All F, offspring between crosses of the two morphs were brachypterous. In F, approximately 25% were macropters and 75% brachypters implying monogenic control of wingform.4. Flight muscles in macropters vary from fully developed to totally reduced. This variation is determined by environmental conditions during adult life. Most young adult bugs have flight muscles, and totally starved or unmated bugs retain their flight muscles. Fed and mated females histolyse flight muscles as they start laying eggs, while most males of the same group retain their muscles.5. Brachypterous bugs have a smaller thorax and larger abdomen than macropterous ones. 6. Brachypterous bugs reach adulthood slightly before macropterous ones, and they have a distinctly shorter adult preoviposition period.7 . Lifetime egg production does not differ significantly between the two wing morphs. However, the temporal pattern of egg laying is different in the two morphs. The m,-curve of macropters starts later, then attains a higher peak and finally decreases faster than that of brachypters.8. Initially, macropters lay smaller eggs than brachypters, but egg volume increases with age in macropters and eventually approaches that of brachypters.9. The initial increase in reproductive effort (egg volumexegg number) of macropters is concomitant with wing muscle histolysis and the mobilization of thorax space for reproduction.10. Adult survival rate does not differ between the morphs.
Summary 1. The role of food plant and aggregation on the defensive properties of two aposematic sympatrically occurring seed bugs, Tropidothorax leucopterus and Lygaeus equestris (Lygaeinae, Heteroptera), was investigated. Larvae reared on seeds either of their natural host plant Vincetoxicum hirundinaria (Asclepiadaceae) or of sunflower Helianthus annuus were subjected to predation by chicks. 2. The two species differ in their dependency on the host for their defence. Lygaeus equestris was better defended on its natural host plant than on the alternative food, as indicated by fewer attacks, lower mortality, and predator avoidance after experience. No such effect of food plant could be found for T. leucopterus, suggesting the existence of alternative defences in this species. 3. The number of attacks was lower when host plant‐fed larvae of both species were presented in groups. 4. The discussion concerns how major components of an aposematic syndrome, such as host plant chemistry, insect colouration, and aggregation, are integrated with other life‐history traits to form alternative lifestyles in L. equestris and T. leucopterus.
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