Auditory defence in the peacock butterfly (<Emphasis Type="Italic">Inachis io</Emphasis>) against mice (<Emphasis Type="Italic">Apodemus flavicollis</Emphasis> and <Emphasis Type="Italic">A. sylvaticus</Emphasis>)

Morphological and behavioural traits can serve as anti-predator defence either by reducing detection or recognition risks, or by thwarting initiated attacks. The latter defence is secondary and often involves a ‘startle display’ comprising a sudden release of signals targeting more than one sensory modality. A suggested candidate for employing a multimodal defence is the peacock butterfly, Inachis io, which, by wing-flicking suddenly, produces sonic and ultrasonic sounds and displays four large eyespots when attacked. The eyespots make small birds retreat, but whether the sounds produced thwart predator attacks is largely unknown. Peacocks hibernate as adults in dark wintering sites and employ their secondary defence upon encounter with small rodent predators during this period. In this study, we staged predator–prey encounters in complete darkness in the laboratory between wild mice, Apodemus flavicollis and Apodemus sylvaticus, and peacocks which had their sound production intact or disabled. Results show that mice were more likely to flee from sound-producing butterflies than from butterflies which had their sound production disabled. Our study presents experimental evidence that the peacock butterfly truly employs a multimodal defence with different traits targeting different predator groups; the eyespots target birds and the sound production targets small rodent predators.     

Fire Management,Managed Relocation,and Land Conservation Options for Long‐Lived Obligate Seeding Plants under Global Changes in Climate,Urbanization, and Fire Regime

Most species face multiple anthropogenic disruptions. Few studies have quantified the cumulative influence of multiple threats on species of conservation concern, and far fewer have quantified the potential relative value of multiple conservation interventions in light of these threats. We linked spatial distribution and population viability models to explore conservation interventions under projected climate change, urbanization, and changes in fire regime on a long‐lived obligate seeding plant species sensitive to high fire frequencies, a dominant plant functional type in many fire‐prone ecosystems, including the biodiversity hotspots of Mediterranean‐type ecosystems. First, we investigated the relative risk of population decline for plant populations in landscapes with and without land protection under an existing habitat conservation plan. Second, we modeled the effectiveness of relocating both seedlings and seeds from a large patch with predicted declines in habitat area to 2 unoccupied recipient patches with increasing habitat area under 2 projected climate change scenarios. Finally, we modeled 8 fire return intervals (FRIs) approximating the outcomes of different management strategies that effectively control fire frequency. Invariably, long‐lived obligate seeding populations remained viable only when FRIs were maintained at or above a minimum level. Land conservation and seedling relocation efforts lessened the impact of climate change and land‐use change on obligate seeding populations to differing degrees depending on the climate change scenario, but neither of these efforts was as generally effective as frequent translocation of seeds. While none of the modeled strategies fully compensated for the effects of land‐use and climate change, an integrative approach managing multiple threats may diminish population declines for species in complex landscapes. Conservation plans designed to mitigate the impacts of a single threat are likely to fail if additional threats are ignored. Manejo de Incendios, Reubicación Administrada y Opciones de Conservación de Suelo para Plantas de Vida Larga con Sembrado Obligado bajo los Cambios Globales en el Clima, la Urbanización y el Régimen de Incendios     

Male butterfly investment in successive ejaculates in relation to mating system

In Lepidoptera, female mating systems range from strict monandry to strong polyandry. Males transfer an ejaculate during copulation that contains both sperm and accessory gland substances. In butterflies the male ejaculate has at least three effects: it (1) contains sperm that can fertilize the eggs of the male’s mating partner, (2) influences the refractory period of the mated female, and (3) contains nutrients that can be used by the female to increase her reproductive output. A number of recent studies have shown that males in polyandrous mating systems are endowed with adaptations to increase mating capabilities. Relative to males in more monandrous species they transfer larger first ejaculates which contain relatively more protein, and allocate proportionally more resources to reproduction. The objective of this study was to compare male reproductive investment in successive ejaculates, in terms of both mass and quality, to determine if males of polyandrous species are capable of maintaining the production of larger and more nutritious ejaculates than males of more monandrous species. We used three species of pierid butterfly, with mating systems ranging from relative monandry to polyandry. The degree of polyandry had a substantial effect on the reproductive performance of males. The cumulative protein content of ejaculates increased with the degree of polyandry. Only males of the most polyandrous species, Pieris rapae, produced three successive ejaculates of similar mass and protein content. In the relatively monandrous species, Aporia crataegi and P. brassicae, males in subsequent matings never produced another ejaculate as large as that transferred by males mating for the first time, although the protein content (mg) of ejaculates did not differ in second and third ejaculates. Moreover, the ability to remate varied with the degree of polyandry. Given the opportunity to remate, the majority of P. rapae males mated three times, whereas relatively few A. crataegi and P. brassicae males were able to perform three matings. These results suggest that male capacity to produce large, nutritious ejaculates is limited in relatively monandrous species. In this study only males belonging to the most polyandrous species had the ability to recuperate quickly from a mating event and remate. Our results suggest that males in polyandrous systems are better adapted to mating more than once. Received: 3 November 1995/Accepted after revision: 13 July 1996     

Polyandry and its effect on female reproduction in the green-veined white butterfly (Pieris napi L.)

Summary In many insects nutrients transferred by the male to the female at mating are later incorporated into both the eggs and soma of the mated females. Accordingly, it has been suggested that female insects can use these male-derived nutrients both for somatic maintenance and to increase both the number and quality of their offspring. Moreover, much discussion is presently devoted to whether the male nuptial gift represents paternal investment, defined as any increase in given male's total surviving progeny by increasing the reproductive output by a given female, or mating effort which obtains if a male gains by increasing the proportion of eggs he fertilizes from a given female (Parker and Simmons 1989). If the male nuptial gift represents parental investment it should be expected to benefit predominantly the offspring sired by the donor, whereas the physiological fate of the male nuptial gift is somewhat irrelevant under the mating effort explanation. In this paper we test these issues by studying the lifetime fecundity, egg weights and longevity of two groups of females of the polyandrous green-veined white butterfly, Pieris napi, one group of which was allowed to mate only once and the other of which was allowed to mate at liberty, the latter group of females mating on average 2.28 times. Moreover, to test the incorporation rate of male-derived nutrients, we performed a second set of experiments where females were allowed to mate with radioactively labelled males. The results showed that polyandrous females had higher lifetime fecundity compared to monandrous females, laying on average 1.61 as many eggs, and that the difference in cumulative fecundity between the two groups was statistically significant from the 5th day of egg-laying onwards. Polyandrous females also lived longer and maintained egg weight at a high level for longer than monandrous females. Largely concomitant with egg-laying rate, incorporation rate of male-derived nutrients peaked 3–4 days after mating, subsequently tapering off to stabilize at about 40% of the maximum. Given the opportunity, female P. napi remated after 3–5 days, the duration of the refractory period being positively correlated with ejaculate mass. Hence, although the nutrient investment of the first male to mate with a female subsidizes the progeny of later-mating males, the male nuptial gift in P. napi clearly qualifies as both paternal investment and mating effort. Correspondence to: C. Wiklund     

Mass and production rate of ejaculates in relation to monandry/polyandry in butterflies

Summary The mating system maintained in a species has a strong effect on the degree of sperm competition, and certainty of paternity should accordingly influence the optimal sperm content, nutrient content, and mass of the ejaculate. We investigated how ejaculate mass relates to the degree of polyandry in 20 species of butterflies belonging to the families Pieridae and Satyridae. We found that the degree of polyandry has a substantial effect on the reproductive performance of males. The allometric line between ejaculate mass and male body mass has a higher elevation in the pierids compared to the satyrids. The mean number of matings performed by the pierid species is also higher compared to the mean of the satyrids. Thus, the relative ejaculate mass is larger in the family in which polyandry is more pronounced. A within family effect of degree of polyandry on relative ejaculate mass was also detected in the pierids. Since males of polyandrous species on average mate more often than males of monandrous species, they should be expected to have a higher capacity for producing many ejaculates. We investigated how this capacity was influenced by the degree of polyandry, by allowing males of seven different species (Danaus plexippus, Lasiommata megera, Papilio machaon, Pararge aegeria, Pieris napi, Pieris rapae, and Polygonia c-album) to mate twice, with different time intervals between matings. The results showed that not only is the mass of the ejaculate greater in more polyandrous species, but also the rate at which males are able to produce sperm and accessory substances is greater. Hence our data indicate that sperm competition is important for explaining variation in ejaculate mass in butterflies.     

New tools in local energy planning: experimenting with scenarios,public participation and environmental assessment

This article presents a model for local energy planning and its application in a full-scale experiment in a Swedish municipality. The model is based on legal requirement, research findings and standards of good practice and includes a combination of analytical and procedural tools intended to support rational decision-making: external scenarios, a citizens’ panel, life cycle analysis and qualitative environmental assessment (EA). The application of the model indicates that the decision-support tools selected can give several new and valuable inputs to local energy planning, such as local knowledge and values through citizen dialogue and comprehensive EAs. However, the experiment also shows that there are several challenges involved in applying the tools, for example, it is difficult to get citizens and the industry to participate and that it is complicated to combine several different tools for decision-making into a single planning process. Moreover, the experiences from the application suggest that the model for local energy planning show great potential but needs to be improved before it can be used as a standard of good practice.     

Quantification of Extinction Risk: IUCN's System for Classifying Threatened Species

Abstract: The International Union for Conservation of Nature (IUCN) Red List of Threatened Species was increasingly used during the 1980s to assess the conservation status of species for policy and planning purposes. This use stimulated the development of a new set of quantitative criteria for listing species in the categories of threat: critically endangered, endangered, and vulnerable. These criteria, which were intended to be applicable to all species except microorganisms, were part of a broader system for classifying threatened species and were fully implemented by IUCN in 2000. The system and the criteria have been widely used by conservation practitioners and scientists and now underpin one indicator being used to assess the Convention on Biological Diversity 2010 biodiversity target. We describe the process and the technical background to the IUCN Red List system. The criteria refer to fundamental biological processes underlying population decline and extinction. But given major differences between species, the threatening processes affecting them, and the paucity of knowledge relating to most species, the IUCN system had to be both broad and flexible to be applicable to the majority of described species. The system was designed to measure the symptoms of extinction risk, and uses 5 independent criteria relating to aspects of population loss and decline of range size. A species is assigned to a threat category if it meets the quantitative threshold for at least one criterion. The criteria and the accompanying rules and guidelines used by IUCN are intended to increase the consistency, transparency, and validity of its categorization system, but it necessitates some compromises that affect the applicability of the system and the species lists that result. In particular, choices were made over the assessment of uncertainty, poorly known species, depleted species, population decline, restricted ranges, and rarity; all of these affect the way red lists should be viewed and used. Processes related to priority setting and the development of national red lists need to take account of some assumptions in the formulation of the criteria.     

Fighting without weaponry: a review of male-male contest competition in butterflies

The males of many butterfly species compete via pair-wise intrasexual contests, yet lack any obvious morphological traits conventionally associated with animal aggression. The evolution of contest behavior in this group is therefore interesting because the means by which individuals are able to inflict costs upon each other during combat are unclear. Here we review the relevant empirical information within a game-theoretical context in order to analyze contest-related costs and mechanisms of dispute settlement. Territorial butterfly contests carry obvious role asymmetries, and individuals in the 'resident' role are more likely to win across all studied species. We use this phenomenon as a framework for gaining further insight into the evolution of contest settlement in this group. Four principal hypotheses are evaluated: (1) that residency is used as an arbitrary convention, (2) that residents stand to gain a greater payoff if successful, (3) that residency bestows higher resource-holding potential (RHP) on males in that role, and (4) that residents are males of intrinsically higher RHP. We found most support for the latter explanation (the 'superior-competitor' hypothesis), partly because the alternatives require a restrictive set of ecological conditions, but also due to positive support for the idea of intrinsic RHP. The strongest evidence is provided by species in which repeated interactions (between the same individuals) always have the same outcome, regardless of changes in residency status. However, we also found a consistent effect on contest duration due to apparent 'confusion over residency,' which suggests a conventional element to contest behavior. Although butterfly contests must be costly in some way, the literature provides limited insight into how costs accrue, and hence the determinants of contest RHP remain obscure. We suggest that careful experimentation, using 'natural' manipulations where possible, should prove most beneficial in identifying costs and exploring schedules of contest settlement in this group. Moreover, future investigations should be sensitive to the fact that butterfly territoriality and associated male-male interactions are seated within a broader ecological and life-historical context.     

Mating in the afternoon: Time-saving in courtship and remating by females of a polyandrous butterfly Pieris napi L.

Summary Among butterflies where females are polygamous, interactions with males after the first mating entail time costs associated with both courtship and mating. In the green-veined white butterfly, Pieris napi, spermatophore counts on wild-caught females revealed that the average number of matings that females had performed increased gradually from 1.1 during the first week of the flight period to 2.4 during the 4th and final week. By dissecting pairs found in copula in the field, we found that female rematings occurred solely in the afternoon, whereas virgin females mated most frequently earlier in the day. We suggest that non-virgin females minimize the time cost of rematings by allocating them to an hour when the egglaying frequency has decreased to a low level. Virgin females were very attractive to males, judging by their courtship persistence (several minutes), but when mated, females became so unattractive that courting males left them within a few seconds. However, females' attractiveness increased again with time elapsed since the last mating. These observations support the idea that females emit repellent pheromones after mating, which may be male derived. Since newly mated females were not harassed by males or hindered in their egglaying activities, it is conceivable that the time cost of remating may be outweighed by the benefit of the regained ability to curtail male courtship. This advantage to newly remated females may in itself be an explanation for polyandry in P. napi and is alternative or complementary to the idea that females that remate receive a material benefit that can be used to increase their realized fecundity.