Conservation and Management for Multiple Species: Integrating Field Research and Modeling into Management Decisions

Multiple-species reserves aim at supporting viable populations of selected species. Population viability analysis (PVA) is a group of methods for predicting such measures as extinction risk based on species-specific data. These methods include models that simulate the dynamics of a population or a metapopulation. A PVA model for the California gnatcatcher in Orange County was developed with landscape (GIS) data on the habitat characteristics and requirements and demographic data on population dynamics of the species. The potential applications of this model include sensitivity analysis that provides guidance for planning fieldwork, designing reserves, evaluating management options, and assessing human impact. The method can be extended to multiple species by combining habitat suitability maps for selected species with weights based on the threat faced by each species, and the contribution of habitat patches to the persistence of each species. These applications and extensions, together with the ability of the model to combine habitat and demographic data, make PVA a powerful tool for the design, conservation, and management of multiple species reserves.     

Effects of Population Size and Food Stress on Fitness-Related Characters in the Scarce Heath, a Rare Butterfly in Western Europe

Abstract: Knowledge about the effects of inbreeding in natural populations is scarce, especially in invertebrates. We analyzed to what extent fitness-related traits in the scarce heath (  Coenonympha hero ), a butterfly, are affected by population size and isolation and whether differences in food quality influence these effects. We categorized nine populations as either large or small and isolated. Full-sib groups of offspring from 27 females were followed under seminatural conditions. Because of increased zygote mortality, egg hatchability was significantly lower in the small and isolated populations than in the large ones. Population category had no effect on larval weight under optimal conditions, but weight was significantly lower in the small-isolated category with low food quality. The effects of inbreeding can thus be hidden when conditions are benign but can appear under stress. Survival also differed significantly between population categories, and larval developmental time tended to be longer in the small-isolated category, irrespective of food conditions. We suggest that the differences in fitness between offspring from large and small isolated populations are at least partly due to inbreeding. This adds a further threat to a species that is already suffering from decreasing population sizes and increasing isolation among populations.     

Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Protein content of spermatophores in relation to monandry/polyandry in butterflies

The evolution of a mating system, and specifically mating frequency, is dependent on the costs and benefits to both sexes of mating once or several times. In butterflies, males transfer a spermatophore that contains both sperm and accessory gland products. Accessory gland substances contain nutrients which, in some species, females use to increase their reproductive output and longevity. Nutrients contained in these packaged ejaculates represent investment by males in reproduction. Consequently, the nutritional composition of spermatophores may vary depending on the mating system. There are two lines of arguments concerning the evolution of the nutrient content of ejaculates. One hypothesis argues that male nuptial gifts evolved in the context of certainty of paternity and ease of finding mates; thus spermatophores of polyandrous males (with lower certainty of paternity and greater ease of finding mates) should contain less protein than those of monandrous males, since more spermatophores are produced on average. The other hypothesis argues that polyandry evolved in the context of maximization of male transfer of nutrients to females, and hence spermatophores of polyandrous males should contain more protein than those of monandrous males. In an attempt to distinguish between these two hypotheses, we determined how protein content of ejaculates varied with the degree of polyandry in nine species of pierid and two species of satyrid butterflies. We found that both relative ejaculate mass and protein content increased with the degree of polyandry. Hence our results are consistent with the view that polyandry has evolved in the context of male transfer of nutrients to females, and provides another example of a male adaptation to multiple mating in butterflies.     

Effects of size and nuptial gifts on butterfly reproduction: can females compensate for a smaller size through male-derived nutrients?

In butterflies and other insects, fecundity generally increases with female adult weight. Hence, most butterflies are essentially "capital breeders", because nutrients acquired during the larval stage are stored and subsequently used for egg production during the adult stage. However, in some species, males transfer a large nutritious ejaculate to the female at mating. These females can partly be characterized as "income breeders", and female mass can potentially be decoupled from fecundity to some extent. In the gift-giving green-veined white butterfly Pieris napi, it has been shown that female fecundity and longevity increase with number of matings and also that females mature at smaller size under poor food conditions compared to males. So it has been suggested that females can compensate for their smaller size through nuptial feeding. Here we test this hypothesis in P. napi by assessing female fecundity and longevity in relation to female mass and polyandry. The results showed no support for the hypothesis. Smaller females were not capable of increasing their mating rate to compensate for a low weight at eclosion. Instead, larger females remated sooner. Also, smaller females suffered from both a reduced daily and total fecundity compared to larger females and this decrease in fecundity was independent of female mating status, i.e. females allowed to mate only once and multiply mated females suffered to the same extent from their smaller size.     

Food information in the Black-headed Gull,Larus ridibundus

Summary The information-centre hypothesis suggests that bird colonies function as sources of information about good feeding sites, to which unsuccessful birds may follow foragers. One assumption of the hypothesis is that unusually successful foragers are followed by other colony members when returning to a newly found, rich food source. We tested this assumption in a colony of Black-headed Gulls (Larus ridibundus). Parents feeding their young from a rich, artificial food source were observed on their return trips to the feeding site. In none of 50 cases did other colony members follow them to the newly found food. However, the gulls were attracted to groups of foraging conspecifics. In experiments with paired food piles and a group of model gulls at one pile in each pair, Black-headed Gulls always alighted first at the piles with models. Hence the information-centre mechanism was refuted, but the gulls did acquire food information from each other in another way.     

“An eye for an eye?”—on the generality of the intimidating quality of eyespots in a butterfly and a hawkmoth

Large eyespots on the wings of butterflies and moths have been ascribed generally intimidating qualities by creating a frightening image of a bird or mammal much larger than the insect bearing the eyespots. However, evidence for this anti-predator adaptation has been largely anecdotal and only recently were peacock butterflies, Inachis io, shown to effectively thwart attacks from blue tits, Parus caeruleus. Here, we test whether large eyespots on lepidopterans are generally effective in preventing attacks from small passerines and whether the size of insect or bird can influence the outcome of interactions. We staged experiments between the larger eyed hawkmoths, Smerinthus ocellatus, and the smaller peacock butterflies, I. io, and the larger great tits, Parus major, and the smaller blue tits, P. caeruleus. Survival differed substantially between the insect species with 21 of 24 peacock butterflies, but only 6 of 27 eyed hawkmoths, surviving attacks from the birds. Thus, surprisingly, the smaller prey survived to a higher extent, suggesting that factors other than insect size may be important. However, great tits were less easily intimidated by the insects’ eyespots and deimatic behaviour and consumed 16 of 26, but the blue tits only 8 of 25, of the butterflies and hawkmoths. Our results demonstrate that eyespots per se do not guarantee survival and that these two insects bearing equally large eyespots are not equally well protected against predation.     

Mating system evolution in response to search costs in the speckled wood butterfly, Pararge aegeria

A general and intuitive prediction from models of mate preference is that when the cost of searching for mates increases, individuals should become less choosy. Here, we test this prediction by comparing the mating propensity of females in two populations of the butterfly Pararge aegeria. The populations originated from southern Sweden and Madeira and due to different adult emergence patterns throughout the year, the average density of males per female is likely to be lower on Madeira. Therefore, we expected that the cost of searching should be greater on Madeira and, consequently, that the Madeiran females should be less choosy. In line with predictions, the Madeiran females mated significantly sooner after the first interaction with males than did females from southern Sweden. This difference may reflect a weaker preference for territorial males over non-territorial patrollers in the Madeiran population, because of the greater costs of searching. The Madeiran females also showed a shorter time lag between mating and the start of oviposition. We discuss this unexpected result and propose that the same mechanism could also explain this population difference, i.e. different costs of searching for suitable host plants. Both search processes are fundamental for female reproductive success and we find it plausible that they can be generalised into the same theory of optimal search behaviour. Received: 14 May 1998 / Accepted after revision: 13 December 1998     

Seasonal polyphenism in life history traits: time costs of direct development in a butterfly

Insects with two or more generations per year will generally experience different selection regimes depending on the season, and accordingly show seasonal polyphenisms. In butterflies, seasonal polyphenism has been shown with respect to morphology, life history characteristics and behaviour. In temperate bivoltine species, the directly developing generation is more time-constrained than the diapause generation, and this may affect various life history traits such as mating propensity (time from eclosion to mating). Here, we test whether mating propensity differs between generations in Pieris napi, along with several physiological parameters, i.e. male sex pheromone synthesis, and female ovigeny index and fecundity. As predicted, individuals of the directly developing generation—who have shorter time for pupal development—are more immature at eclosion; males take longer to synthesise the male sex pheromone after eclosion and take longer to mate than diapause generation males. Females show the same physiological pattern; the directly developing females lay fewer eggs than diapausing females during the first days of their life. Nevertheless, the directly developing females mate faster after eclosion than diapausing females, indicating substantial adult time stress in this generation and possibly an adaptive value of shortening the pre-reproductive period. Our study highlights how time stress can be predictably different between generations, affecting both life history and behaviour. By analysing several life history traits simultaneously, we adopt a multi-trait approach to examining how adaptations and developmental constraints likely interplay to shape these seasonal polyphenisms.