Temporal and spatial variability in the interaction between the checkerspot butterfly,Euphydryas chalcedona and its principal food source,the Californian shrub,Diplacus aurantiacus

Summary The phenology of the Californian shrub, Diplacus aurantiacus has been shown to be closely tied to habitat water availability, and the life cycle of the checkerspot butterfly, Euphydryas chalcedona, is in turn tied to the phenological development of Diplacus. Here this relationship is further documented by showing how the activity patterns of both the shrub and the butterfly vary from year to year, but in synchrony, dependent on the breaking of the annual drought.The end of the feeding period for the post-diapause larvae coincides with flowering of Diplacus. At this time the quality of leaves as a food source for the larvae declines as nitrogen evidently moves from the leaves into plant reproductive parts.The abundance of leaves available as food for the larvae varies greatly with season. The few leaves present during the drought period, when the larvae are inactive, are low in nitrogen and high in resin content. Even during this period, though, these leaves contribute to the carbon economy of the plant. Of the leaves produced during the principal growing season, it is the youngest leaves with the highest nitrogen contents, and hence greatest potential carbon-gaining capacity, that have the highest resin contents. Larvae feed preferentially on the older, lower-resin-content leaves. It can be surmised thus that resin is a feeding deterrent and that its distribution within the plant results in the greatest protection of the plant's carbon-gaining capacity.Dedicated to Dr. Konrad Springer who has been a strong force in the development of ecology this past decade through his prolific publication series which have set the highest standards in the profession     

Coevolution of the checkerspot butterfly Euphydryas chalcedona and its larval food plant Diplacus aurantiacus: larval response to protein and leaf resin

Summary Prediapause larvae of the checkerspot butterfly Euphydryas chalcedona were raised from hatch until entrance into diapause on artificial diets. The proportions of protein and host plant leaf resin differed among the diets. Larval size growth rates and mortality were monitored and overall rates and efficiencies of food use were computed.Larval survivorship, growth rate and size of larvae at idapause were significantly enhanced by increasing dietary protein content, particularly over the range found in leaves of the host plant. In contrast, an increasing dietary content of Diplacus aurantiacus leaf resin significantly depressed larval surviviorship, growth rates and size of larvae at diapause. A simple dosedependent interaction was observed between the effects of dietary leaf resin and protein on larval success. Dietary content of leaf resin and protein significantly influenced some measures of food utilization efficiency (ECI and ECD), but not others (AD and NUE).The negative interaction between the effects of dietary leaf resin and protein content suggests the leaf resin phenolic compounds reduce the availability of protein to the larvae. The results for efficiency indices of larval food use are potentially in conflict with this interpretation.The influence of host plant leaf resin and protein on larval success, coupled with the relation between photosynthesis and leaf nitrogen content, are consistent with the hypothesis that productivity can be enhanced by herbivore deterrence resulting from leaf resin production.     

Ecological determinants of food plant choice in the checkerspot butterfly Euphydryas editha in Colorado

Summary The larvae of Euphydryas editha in Gunnison County, Colorado, feed on Castilleja linariifolia. Two related plants, C. chromosa and Penstemon strictus, occur in the same area and are equally nutritious, but are not eaten. Restriction to C. linariifolia appears to be a case of ecological monophagy — survival on the other two species is less likely not because of their biochemical make-up but because of their ecological characteristics, primarily their phenologies.     

Extinction,reduction, stability and increase: The responses of checkerspot butterfly (Euphydryas) populations to the California drought

Summary The California drought of 1975–77 has been correlated with unusual size changes in populations of two species of Euphydryas butterflies. Several populations became extinct, some were dramatically reduced, others remained stable and at least one increased. These differences in population dynamic response are not concordant with predictions made earlier that populations with heavy density-dependent mortality should be more stable in the face of drought than unregulated populations. The different responses are related to the fine details of the relationships between the insects and their host plants, relationships which are variable between populations. Revised predictions are given in the light of better knowledge of the variability and complexity of these insect-host relationships. The diversity of responses underlines the dangers of generalizing about the ecology of a taxonomic species. The extinctions support the view that such events are frequent and significant in the biology of populations.     

The role of adult feeding in egg production and population dynamics of the checkerspot butterfly Euphydryas editha

Carbohydrate intake increases longevity, body weight maintenance and egg production in female Euphydryas editha. Amino acid intake leads to heavier eggs, larvae from which are more likely to survive. Females fed nectar produce more eggs in later masses than females which are not fed. During years of normal and below normal precipitation, larvae emerging from these later eggs are unlikely to reach obligatory size for diapause before their food dries up. On Jasper Ridge, where mortality is density-independent, nectar plays an important role increasing production of late egg masses during years of greater than normal rainfall when larvae from these masses are likely to reach diapause. The resulting large population increases, though infrequent, are probably important in maintaining population sizes large enough to reduce the chances of extinction during dry years.     

Population differences in larval hostplant use in the checkerspot butterfly, Euphydryas chalcedona

Larvae from two populations of Euphydryas chalcedona Doubleday & Hewitson (Nymphalidae) were reared on their own hostplant and that of the other population, in both pre-diapause and post-diapause instars. One population, Chico, uses Penstemon breviflorus Lindl. (Scrophulariaceae), and the other, Echo Lake, uses P. newberryi Gray. Growth rate and survival were determined for pre-diapause and post-diapause larvae from both populations on both plant species; and digestive efficiencies were calculated during the prediapause instars. The results showed that larvae from the two populations differed in their responses to the two plant species. Pre-diapause larvae from Chico performed equally well on both plant species—survival and digestive indices were not significantly different for two Penstemon species. In contrast, pre-diapause larvae from Echo Lake performed significantly worse on the non-hostplant—growth and survival were significantly lower on the non-host, P. breviflorus. In addition, comparison of digestive efficiencies for the two plants showed that larvae from Echo Lake digested P. breviflorus better than P. newberryi, but were significantly less able to convert P. breviflorus to body mass. In the post-diapause instars, larvae from Chico grew faster on the host than on the non-host. Larvae from Echo Lake grew quite slowly on both plant species and significantly more of the Echo Lake larvae returned to diapause instead of completing development.

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Résumé Des chenilles de deux populations d'E. chalcedona ont été élevées sur leur propre plante-hôte et sur celle de l'autre population, aux stades avant et après diapause. Les deux populations s'alimentent sur différentes espèces de Penstemon (Scrophulariaceae), et une population—Echo Lake—est monophage sur P. newberry, tandis que l'autre—Chico—utilise d'abord P. breviflorus, mais les chenilles après diapause sont trouvées sur au moins deux autres espèces de plantes. Les taux de croissance et de survie ont été déterminés pour des chenilles avant et après diapause pour les deux populations sur les deux plantes; les efficacités digestives ont été calculées sur les chenilles avant diapause.Les résultats ont montré que les chenilles des deux populations différaient par leur degré de spécialisation digestive sur leur plante hôte normale: les chenilles de Chico ont utilisé aussi bien les deux plantes, tandis que celles d'Echo Lake le faisaient significativement moins bien sur la plante non-hôte, par suite de l'inaptitude à la digérer. Ainsi la population oligophage est alimentairement moins spécialisée et plus capable de se débrouiller avec une plante non-hôte. Après diapause, les chenilles de Chico s'alimentaient significativement mieux sur plante hôte que non-hôte, ce qui était le cas aussi pour la population monophage. Dans l'ensemble, les chenilles de la population monophage semblaient moins capables de se débrouiller dans des conditions défavorables ou moins avantageuses.

     

The structure and genetics of a montane population of the checkerspot butterfly,Chlosyne palla

Summary The population structure, genetics and ecology of the checkerspot butterfly, Chlosyne palla, in an area of Gunnison County, Colorado were investigated. The population structure was found to be quite different from that of most butterflies and from all of those aspects known for its thoroughly studied relative, Euphydryas editha. The population unit of Chlosyne palla may cover an area some five to eight times the size of the largest known Euphydryas population and twice the size of an Erebia epipsodea population in the same county of Colorado.Genetic variation at eleven loci of Chlosyne palla was examined by electrophoresis. Three samples of Chlosyne palla separated by 1.6, 4.7, and 12.0 km were not significantly different. Comparison with Euphydryas editha yields a genetic similarity of 0.186, about the same level as found by Ayala (1975) for different genera of Drosophila. Euphydryas editha from the same Colorado location were more similar to California E. editha than to C. palla, showing concordance with the phenetic classification. Decreased heterozygosity was observed for the Colorado E. editha and C. palla compared to California populations of E. editha and E. chalcedona.     

Nectar source distribution as a determinant of oviposition host species in Euphydryas chalcedona

The distribution of nectar sources is shown to affect both the distribution of adult Euphydryas chalcedona and their offspring. We suggest that nectar sources thereby influence the selection of oviposition host plant species in habitats where those species are spacially separated.     

Thermal influences on oviposition in the montane butterfly Euphydryas gillettii

Summary Euphydryas gillettii in a montane meadow in Wyoming oviposits in clusters on the highest large leaves of its larval host, the shrub Lonicera involucrata. Egg masses are found on the underside of leaves which predominantly face the southeast and thereby intercept morning sun. Measurements of leaf temperatures show that, during the cool, clear mornings, these leaves are warmer than leaves of the opposite orientation. Manipulative experiments demonstrate that eggs on these slightly warmer microsites hatch significantly faster than eggs on leaves of other orientations. Because of the rapid onset of winter, an increased developmental rate is likely to correlate directly with increased fitness. The proximal mechanism yielding such a result may be that females choose oviposition sites in the shade.     

Effect of group size on parasitism in a natural population of the Baltimore checkerspot Euphydryas phaeton

Summary The effect of group size of early instars on parasitism of Euphydryas phaeton (Nymphalidae) was examined. Different numbers of larvae were stocked per web to determine the effect of group size on parasitism. Larval aggregations of moderate size (the size occurring naturally) had the least parasitism. Larger larval groups had a disproportionately high rate of parasitism. The major larval parasitoids located vulnerable larvae within webs, instead of attacking larvae available on the outside of webs. Parasitism rates were similar for larvae of damaged and undamaged webs, a consequence of the behavior and location of larvae in the webs. Lower limit to group size was a function of facilitation of larval numbers in reaching the first feeding site, the top of the host plant. Feeding facilitation by larval aggregations was not a factor in larval survival or growth.     

Sites chosen by diapausing or quiescent stage quino checkerspot butterfly, <Emphasis Type=Italic>Euphydryas editha quino</Emphasis>, (Lepidoptera: Nymphalidae) larvae

This study examines whether in nature endangered quino checkerspot (Euphydryas editha quino) larvae will return to diapause and if so where they choose to hide. Multiple years of diapause probably help larvae survive drought years and sites chosen have high survival value to the species. Ninety square meters of habitat were created by removing non native plants and replacing them with natives found at checkerspot occupied sites. During the 2005–2006 winter 1,000 post-diapause larvae were released. From these larvae 31 adults (20 males and 11 females) developed over a 2.5 month period (March 20–June 6) from 41 pupae. One chrysalis was parasitized by a parasitic wasp Pteromalus puparum (L.) in the family Pteromalidae, one was partially eaten by an animal, while the remaining eight pupae died of unknown causes. Thirty quadrats (1 square meter each) were cleared of vegetation, leaf and branch litter, rocks, and checkerspot larvae from July 5 to August 1, 2006. Forty-nine larvae were found that returned to diapause. Most larvae (31) chose to make shelters on California buckwheat, which is not a checkerspot food plant, two to five cm above the ground. One shelter had 22, another had seven, and two others had single larvae. Five of 10 larvae found in leaf litter below California buckwheat were crawling and not associated with shelters suggesting they had been dislodged from shelters. California buckwheat may be important in habitat restoration for the checkerspot, particularly at sites below 900 meters elevation where summer conditions are hot and dry. No additional larvae were found the following spring, when they should have exited diapause. Therefore 910 (91%) larvae were lost to some undocumented form of mortality.     

Habitat use of the endangered butterfly Euphydryas maturna and forestry in Central Europe

The knowledge of ecological requirements of declining butterflies of European woodlands remains limited, which hinders conservation management of their localities. This also applies for continentally threatened scarce fritillary Euphydryas maturna . On the basis of the largest data set on its habitat use ever collected in Central Europe, we analyse habitat requirements of its populations in Austria (A), the Czech Republic (Cz) and Germany (D). All studied populations inhabit open-canopy sites within woodlands, but larval survival decreases under full sun and preferred sites are relatively humid and sheltered. Nests of pre-hibernation larvae occur at terminal branches of Fraxinus excelsior , 1.5–3 m above the ground. Pre-hibernation mortality reaches 70% (Cz, D). Another limiting factor is quality of woodland vegetation: post-hibernation larvae consume a wide range of herbs and shrubs, and adult distribution is linked to nectar availability. The butterfly thus depends on highly heterogeneous early successional stages of deciduous woods, historically maintained by coppicing (Cz, D) and forest pasture (A). Restoration of these traditional methods offers the only chance for survival of E. maturna in Central Europe, and the butterfly may become a flagship for other threatened organisms of open-canopy woodlands.     

The colony structure and reproductive biology of the afrotropical Mashona mole-rat, Cryptomys darlingi

Cryptomys darlingi occurs in the mesic Miombo woodland of north-eastem Zimbabwe. It occurs in colonies of up to nine individuals, in which reproduction is limited to one of the largest males and the largest female in the colony.
Reproduction and details of colony size and number of breeding animals in a colony are described for five complete field-captured colonies.
In captivity, mating is not confined to a particular season, and up to three litters of pups are orn per annum. The reproductive female initiates the pre-copulatory behaviour. The gestation lengti is 56–61 days ( n = 2 ). The new-born pups are altricial and litter size is small = 1.7 ± 0.5 ( n = 6). In this case, the pups first left the nest 10 days after birth, began to eat solids when 14 days old, and were fully weaned at five weeks. They began to spar with each other when 36–40 days old, but did not disperse and were incorporated into the colony. This suggests that the Mashona mole-rat colonies are composed of a founding pair and at least three successive litters of pups.     

Securing viable metapopulations of the Marsh Fritillary butterfly, Euphydryas aurinia, (Lepidoptera: Nymphalidae) in Northern England

The Marsh Fritillary butterfly, Euphydryas aurinia (Rott.), is regarded as the classic species that requires landscape-scale management in order to maintain viable populations. This paper describes the background to an extinction of this species in Northern England, United Kingdom, and the action taken to reintroduce and monitor the progress of sub-populations between 2007 and 2009. A captive breeding stock was secured using the last 155 known wild caterpillars in 2004 and supplemented by 95 caterpillars collected under licence from the nearest extant populations in western Scotland. Through captive breeding the source population for subsequent reintroduction was increased dramatically and 42,400 caterpillars were used in 2007 to reintroduce the butterfly to four locations in North and West Cumbria. Each of the release sites is within known historical network areas where the butterfly had thrived in past decades. The intention was to restore multiple habitat patches across each network area and eventually establish viable metapopulations across these landscapes. Adults and larval webs have been monitored since 2007 and show that introduction has been successful at three out of the four locations. The failure at the fourth site appears to be unrelated to habitat condition and the causes are currently being investigated. Work is continuing to secure appropriate management of suitable habitat within each landscape area and the number of sub-populations in one area will be increased in the 2010 season.     

Effects of maternal age and adult diet on egg weight in the butterfly Euphydryas editha

ABSTRACT. 1. Euphydryas editha (Boisduval) (Nymphalidae) butterflies exhibited no consistent pattern of change in egg weight over their lifetimes. This was the case for captive butterflies fed diets containing sugars and amino acids, for butterflies fed diets containing only sugar, and for those left completely at liberty in their natural habitats between ovipositions.
2. Two aspects of these results conflict with the findings of a previous study on the same subspecies. First, the previous study found that egg weight decreased with increasing maternal age. Second, this decrease was mitigated by addition of amino acids to the adult diet.
3. We accept the findings of these previous workers that egg weight decreased with maternal age under their laboratory conditions. The discrepancy between their data and ours is likely to stem from the more natural conditions under which our insects were kept. Our own data from captive insects are not distinguishable from our field data.
4. These results suggest that declines of egg weight with maternal age found in studies of other insects may also result from the artificial nature of laboratory conditions. We question whether such declines are general.
5. We suggest that the second discrepancy between the two sets of work (effects of amino acids) stems from a design problem in the previous study.     

Rainfall and the interaction of microclimate with larval resources in the population dynamics of checkerspot butterflies (Euphydryas editha) inhabiting serpentine grassland

Summary The interaction of host plant phenology and microclimatic heterogeneity was examined to determine its role in the population dynamics of checkerspot butterflies, Euphydryas editha, inhabiting serpentine grassland in California's outer Coast Range.Within the 2–3 hectares inhabited by a population of E. editha (Jasper Ridge Area H), microclimatic differences resulting from topographic heterogeneity largely determine the temporal and spatial pattern of senescence of the larval host plants, Plantago erecta and Orthocarpus densiflorus. Survival of larvae from hatching to diapause is extremely low as a result of unpredictable variation in the timing of larval development relative to the timing of host plant senescence, both of which are mediated by microclimatic patterns. During this study, population H declined to near extinction as a result of two consecutive years of record rainfall that apparently disrupted the tenuous temporal relationship between larval development and plant senescence. Retarded development of post-diapause larvae led to a late and extended flight season and delayed egg production; this in turn resulted in massive mortality of pre-diapause larvae due to starvation because host plant senescence occurred before larvae became large enough to enter diapause. Adult population size the following spring was the smallest in 25 years of study. This work emphasizes the importance of microclimatic heterogeneity for understanding population-level processes in small ectothermic animals and underlines the potential importance of such heterogeneity in the establishment of reserves designed to protect such animals     

Population structure and colony composition of two Zootermopsis nevadensis subspecies

Understanding the origin and maintenance of eusociality in termites has proved problematic, in part, due to a lack of knowledge concerning the variability and evolutionary changes in termite breeding structure. One way to address this is to compare the population genetics of a broad range of termite species. However, few studies have investigated the population genetics of basal termite taxa. We used 12 polymorphic microsatellite loci to characterize and compare the colony genetic structure of 18 colonies of two basal termite subspecies, Zootermopsis nevadensis nevadensis and Zootermopsis nevadensis nuttingi. The average relatedness (r) among individuals within a colony was high (0.59) and similar to values reported for other termite species. Average relatedness between colony founders was lower (0.21) suggesting the alates outbreed. Genotypes of workers and soldiers in 4 out of the 18 colonies were consistent with reproduction by a single pair of primary reproductives and the remaining colonies were inferred to have been derived from more than two reproductives. Eleven colonies with three or more reproductives were consistent with replacement reproductives (neotenics) and the remaining three colonies included genetic contribution from three or more primary reproductives. Comparisons between the subspecies revealed significant differences in breeding structure, specifically in the number and types of reproductives (that is, primaries or neotenics). Furthermore, we observed a larger proportion of colonies with greater than three primary reproductives compared to more derived termite lineages. Thus, our results suggest that breeding structure can vary significantly among termite taxa.     

The social biology of domiciliary cockroaches: colony structure, kin recognition and collective decisions

A substantial body of research on eusocial insects seen in the last decades has gone hand-in-hand with the development of social evolution theory. In contrast, little attention has been given to the non-eusocial insect species that nevertheless exhibit a rich spectrum of social behaviours, thus effectively skewing our vision of insect sociality. Recent studies on the behaviour, ecology and genetic of “gregarious” cockroaches (Blattodea) have revealed a diversity of social structures and group dynamics unique among insects, providing an important comparative model for the broader understanding of insect social evolution. Here, we present an overview of the social biology of the domiciliary cockroaches (ca. 25 species adapted to human habitats) based on research on two model species, Blattella germanica and Periplaneta americana. We discuss the evolution of these domiciliary cockroaches, considering them in the context of “social herds” within the insect sociality framework.