Intraspecific variation in habitat availability among ectothermic animals near their climatic limits and their centres of range

1. In a modelling exercise, the quantity and distribution of habitat patches within a heathland biotope for four ectothermic heathland animals (silver-studded blue butterfly [ Plebejus argus ] , a red ant [ Myrmica sabuleti ] , heath grasshopper [ Chorthippus vagans ] and sand lizard [ Lacerta agilis ]) were compared in space and time assuming two climates: that experienced at the northern edge of the species' ranges and that 300–400 km further south, where mean summer temperatures are 2–3 °C warmer.
2. Habitats both at the northern edge of their ranges and 300–400 km further south for the four species were defined qualitatively from existing sources and then expressed quantitatively in terms of the attributes recorded in the Dorset Heathland Survey. The Survey was then used as a GIS to map the occurrence of the habitats of the four species under two climates and a decade apart.
3. The model predicts that an increase of 2–3 °C can result in a large increase in the area of habitat available to these north temperate species, that the length of time that individual patches of successional habitat may be occupied increases and that the distance between habitat patches within the biotope decreases.
4. The warmer conditions should result in a more stable metapopulation structure for P. argus , with fewer metapopulations existing in the landscape but each, on average, containing a greater number of larger and more stable constituent populations.
5. These predictions are of significance to ectothermic species which currently live at the northern limits of their ranges in the British Isles. The reverse effect is likely for species at the southern limits of their ranges. Conservationists who wish to maintain the status quo may be able to reduce some effects of these changes by appropriate habitat management.     

Changing dynamics of the pine beauty moth (Panolis flammea) in Britain: the loss of enemy free space?

1 The pine beauty moth Panolis flammea has two main host plants in Britain: Pinus sylvestris (Scots pine), which is the ancestral food plant where the insect is never abundant enough to cause tree mortality, and Pinus contorta (lodgepole pine), an introduced host tree that has experienced periodic widespread tree mortality due to this pest.
2 We review the recent literature, published mostly after the year 2000, regarding the impact of natural enemies on the population dynamics of P. flammea in Britain.
3 The natural enemies of P. flammea are more diverse and abundant in Scots pine habitat than in lodgepole pine habitat and some of them show differential selection for P. flammea larvae in Scots pine habitat over those located in lodgepole pine habitat.
4 It is concluded that the difference in the population dynamics of this insect in the two different habitats was probably the result of the P. flammea finding enemy-free space in lodgepole pine habitat.
5 Recent evidence on the diversity and impact of natural enemies on lodgepole pine has demonstrated that they currently have a much more significant impact on this pest than they did in the 1970s and 1980s, when outbreaks were frequent.     

Nitrogen deposition and the reduction of butterfly biodiversity quality in the Netherlands

Butterfly decline in Northern Europe is a cause of concern and it has been hypothesised that this is due to nitrogen deposition inducing excess early growth of plants. It has also been changing the quality of the food available to larvae. We tested these hypotheses by linking butterfly biodiversity quality indices (species richness, population, biomass, conservation value, evenness (Simpson's Index) and modelled species richness (Chao 1 and 2)) with nitrogen Critical Load Exceedence (nCLE) data. An index of butterfly sensitivity to nitrogen was also created (Species Nitrogen Value Index (SNVI)). Using PCA, datasets were tested for associations and relationships.The results included multiple biodiversity quality indices based on 17 years of data (aggregated into three periods of six, six and five years to give 287 datasets) in four habitat types (grassland, heathland, woodland and farmland). With the exception of heathland the analysis showed that nitrogen deposition and all other indices (except SNVI) were in decline.For heathland the last 11 years did not show any significant decline. Heathland also showed an anomalous biodiversity quality profile for these last 11 years, suggesting that the sensitivity of heathland to nitrogen deposition will require further considerable efforts to achieve a nitrogen deposition that is not in exceedence of the critical load.Habitat restoration will take time due to the multiple hindrances to colonisation, which in the case of heathland might prevent successful butterfly colonisation for the foreseeable future. These results indicate the efficacy of butterfly biodiversity quality and nCLE as indicators for the SEBI 2020 process (Streamlining European Biodiversity Indicators) by showing the relationship between them.     

Mosquitoes and other aquatic insects in fallow field biotopes and rice paddy fields

Fallow field biotopes that develop from abandoned rice fields are man‐made wetlands that provide new habitats for various aquatic animals. Although consideration of such biotopes generally focuses on their positive aspects, this study evaluated the negative aspects of establishing fallow field biotopes with regard to mosquito breeding sites. To determine whether fallow field biotopes become breeding habitats for vector mosquitoes, we evaluated mosquito fauna in fallow field biotopes and adjacent rice fields. We found larvae of Anopheles lesteri, Anopheles sinensis and Culex tritaeniorhynchus (all: Diptera: Culicidae) in the biotopes. Although abundances of mosquito larvae in the biotopes and rice fields were statistically similar, mosquito abundances in rice fields increased dramatically in August when the water level reduced after the rainy season. The abundance and variety of the mosquitoes' natural predators were greater in biotopes than in rice fields because the former are a permanent and stable aquatic environment. A generalized linear mixed model showed a negative effect of predator diversity on mosquito larvae abundance in both habitats. Although fallow field biotopes become breeding habitats for vector mosquitoes, establishing biotopes from fallow fields in order to protect various aquatic animals, including mosquito insect predators, may help to control mosquito breeding.     

The influence of wild boar <Emphasis Type="Italic">(Sus scrofa)</Emphasis> on microhabitat quality for the endangered butterfly <Emphasis Type="Italic">Pyrgus malvae</Emphasis> in the Netherlands

The decline of open habitats in Europe, such as semi-natural grasslands and heathlands, has caused a general decline in biodiversity, which has been well documented for butterflies. Current conservation practices often involve grazing by domestic livestock to maintain suitable butterfly habitats. The extent to which wild ungulates may play a similar role remains largely unknown. Through their rooting activity, wild boar could be effective to reduce grass encroachment and restore pioneer microhabitats that are vital to many grassland insects in temperate climates. Here, we assessed the microhabitat requirements of Pyrgus malvae, an endangered butterfly of heathland and grassland habitats in the Netherlands, with special attention for the influence of wild boar rooting. To date, oviposition site selection of this species has concentrated on calcareous grasslands, whereas we also include heathlands. Overall, larval occupancy was higher in warm, open and sparsely vegetated microhabitats, which supports earlier findings. In heathland, microhabitat occupancy was positively affected by bryophyte and litter cover. In heath-grassland mosaic, microhabitat occupancy was also influenced by bryophyte and litter cover, but in addition low grass cover increased occupancy by favouring host plants. In grassland, only low grass cover and host plant cover determined microhabitat quality. Across all habitats, occupied microhabitats were characterized by lower vegetation as well as higher average daytime temperatures than unoccupied microhabitats. We discovered that wild boar play an important role in reducing grass cover by shallow rooting in grass patches, thereby increasing host plant availability. Hence, wild boar may have an added value in maintaining and restoring P. malvae microhabitats in grassland habitats in addition to grazing by domestic livestock.     

Inconsistent detection of extinction debts using different methods

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species–area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data‐intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under‐reported.     

The phenological window of opportunity for early-season birch sawflies

Abstract 1. The phenological window of opportunity hypothesis posits that an insect herbivore cannot survive outside a specific leaf age interval of its host plant.
2. Previous studies have shown that mountain birch displays extensive seasonal changes in leaf biochemical characteristics. Young, expanding leaves are full of water and amino acids but they also contain large amounts of protein-precipitating gallotannins as well as flavonoid-glycosides. Concentrations of these compounds decline during leaf growth whereas concentrations of sugars and proanthocyanidins and leaf toughness increase.
3. Adult birch sawflies hatch in early summer and oviposit on growing leaves, which compels larvae to feed on leaves that are already well developed. The purpose of this study was to test whether leaves that were younger and biochemically different from the leaves available under natural conditions are within the phenological window of opportunity for the larvae of two early-season birch sawfly species, Amauronematus amplus Konow and Pristiphora alpestris (Konow).
4. Amauronematus amplus larvae survived better and developed faster, and P. alpestris larvae developed faster and became bigger, on atypically young leaves compared with larvae reared on leaves encountered normally. Therefore, these species can exploit the putative nutritional superiority of very young leaves, which probably outweighs the potential impact of the new set of secondary metabolites offered to them. In conclusion, young leaves that are consumed rarely by sawfly larvae are within their phenological window of opportunity, even though the timing of sawfly life cycles constrains their utilisation.     

Influence of larvae of Gastrophysa viridula on the distribution of conspecific adults in the field

1. Previous laboratory bioassays indicate that exocrine glandular secretions of larvae of Gastrophysa viridula repel conspecific adults and deter them from feeding and oviposition. The present study was conducted to investigate the influence of larvae of G. viridula on conspecific adults in the field.
2. Within the G. viridula population studied, two generations were observed in a year. Occurrence of the different developmental stages overlapped temporally.
3. Some individual plants of Rumex obtusifolius , the host of the G. viridula population studied, grew so close to each other that they were considered as a plant group. When investigating the spatial distribution of larvae and adults within such plant groups, larvae were rarely found on plant groups on which adults were feeding.
4. A field experiment revealed that adults avoided plants of R. obtusifolius infested by conspecific larvae of the second and third instar. Adults still avoided these damaged plants when larvae had left them for pupation.
5. Gastrophysa viridula avoided oviposition on leaves and plants infested by conspecific larvae. Larvae of the second instar significantly deterred oviposition when present at a high density (33.3 larvae/dm²), whereas larvae of the first instar did not deter oviposition of conspecifics at either density tested. The oviposition deterring effect was also observed when just exocrine glandular secretion of larvae of the second instar was applied to the leaves in amounts equivalent to a density of 33.3 larvae/dm².
6. Availability of food ( R. obtusifolius ) largely exceeded its exploitation in each group of plants examined. This may be due to either the spatial separation of adults and larvae or the low population density observed on these plant groups.     

A comparative study of the seedbanks of heathland and successional habitats in Dorset, Southern England

1 Many areas of lowland heaths are being lost due to invasion by Betula spp., Pinus sylvestris, Pteridium aquilinum, Rhododendron ponticum and Ulex europaeus. One of the factors influencing the success of restoration of heathland on such sites will be the content of their viable seedbanks.
2 Ten heathland areas in the Poole Basin area of Dorset, where succession to one or more of the above species had occurred were studied. The viable seedbanks of the successional sites were compared with those of nearby heathland using Canonical Discriminant Analysis.
3 The seedbanks of all the successional stages were significantly different from the seedbank of the heath.
4 The seedbanks from the Pinus sylvestris and Pteridium aquilinum successional stages contained significantly lower numbers of heathland species than did the heathland seedbank, although few non heathland species were present.
5 The seedbanks from the Betula spp., Rhododendron ponticum and Ulex europaeus successional sites contained both significantly lower numbers of heathland species and significantly higher numbers of non heathland species than the heathland seedbank.
6 The results are discussed in relation to the restoration of heathland on successional sites and the use of the seedbank as a source of propagales for the establishment of heathland species.     

A plant surface mutation mediates predator interference among ladybird larvae

Abstract.  1. We investigated mechanisms causing predator–predator interference between fourth instar Hippodamia convergens larvae foraging for pea aphids on pea plants, Pisum sativum , with a wild-type wax bloom, and the lack of such interference between larvae foraging on pea plants with a reduced-wax bloom caused by the single gene mutation wel .
2. Observations showed that behavioural interactions between larvae were not affected by wax phenotype. Specifically, larvae did not encounter one another more frequently on normal-wax peas as may have been predicted because reduced ability by coccinellids to attach to normal-wax plant surfaces could restrict them to foraging on only some parts of these plants.
3. In a controlled bioassay on normal-wax peas, H. convergens larvae avoided leaflets previously exposed to another larva. On reduced-wax peas, this effect was not detected.
4. In microcosm experiments, inter-predator interference in terms of prey consumption occurred on normal-wax peas, but not on reduced-wax peas. The interference on normal-wax peas occurred whether two H. convergens larvae were placed on a pea aphid-infested, normal-wax plant simultaneously or sequentially.
5. We conclude that the observed inter-predator interference is not as a result of direct physical contact, but rather arises because of (i) inhibition of foraging by chemical trails left by other larvae, (ii) the inability of larvae to access portions of the normal-wax plants creating aphid refugia, or (iii) a combination of these factors.     

Avoidance mechanisms of three Pieris butterfly species against the parasitoid wasp Apanteles glomerulus

Abstract. 1. Experimental studies have shown that larvae of three Pieris butterflies, P.rapae L., P.melete Mènètriés and P.napi L., are attacked by a parasitoid wasp, Apanteles glomeratus L. Although P.rapae larvae are parasitized heavily in the field, P.melete and P.napi are infrequently parasitized successfully because they possess mechanisms for encapsulating parasitoid larvae and for avoiding parasitism.
2. This study examines spatial and temporal variation in rates of parasitism of the three Pieris species by A.glomeratus in the field. We attempted to determine whether P.rapae possesses any means of avoiding parasitism by this wasp, and then to deduce why both P.melete and P.napi have more distinctive avoidance mechanisms than P.rapae.
3. Our results indicate that in temporary habitats, which are the main habitats of P.rapae, P.rapae is able to escape A.glomeratus in time and space by colonizing new habitats before the parasitoid arrives. In permanent habitats, however, such escape is not possible. P.rapae larvae lack physiological or behavioural avoidance mechanisms of reducing parasitism rates in permanent habitats. P.melete and P.napi , in contrast, live only in permanent habitats, where the parasitic pressure is potentially high, and have evolved active avoidance mechanisms.     

Natural pupation sites of swallowtail butterflies (Lepidoptera: Papilioninae): Papilio polyxenes Fabr., P.glaucus L. and Battus philenor (L.)

Abstract. 1. Natural pupation sites have been found in Papilio polyxenes and P.glaucus by releasing prepupal larvae marked with UV-fluorescent paint and locating them at night with a UV lamp, and in Battus philenor by searching a forest habitat where the larval foodplant is abundant.
2. P.polyxenes , a species of weedy habitats, pupates off the ground on a variety of substrates including grasses, weed stalks, posts, etc. The pupae may be green or brown, resembling the substrate.
3. P.glaucus , a species of forest habitats, pupates very close to the ground in the litter and has monomorphic brown pupae.
4. B.philenor , also a forest species, pupates on exposed surfaces (chiefly tree-trunks or cliffs) well off the ground. Its pupae may be brown or green, but the latter were found only on the slenderest twigs.
5. The results for polyxenes and glaucus support the generalization of Clarke & Sheppard (1972) that species of stable habitats are likely to have monomorphic pupae, while those of habitats in which available sites may not be so similar from one generation to the next will be dimorphic.
6. B.philenor is more problematical, but its tendency towards pupal monomorphism (brown) is logical in relation to its common pupation sites.     

Effects of specialist parasitoids on oviposition preference of phytophagous insects: encounter–dilution effects in a tritrophic interaction

Abstract.  1. Host plant preferences of the female diamondback moth Plutella xylostella were studied.
2. Female moths preferred conspecific-damaged cabbage plants over undamaged cabbage plants. The performance of P. xylostella larvae on conspecific-infested plants did not differ significantly from that of larvae on undamaged plants.
3.  Cotesia plutellae , the specialist parasitoid wasp of P. xylostella larvae, displayed equal preference for plants with differing levels of host-larvae damage, and the wasp attacked only one or two hosts on average before leaving an infested plant, irrespective of the number of hosts on the plant. It is hypothesised that the oviposition preferences of P. xylostella females for host plants already damaged by conspecific larvae demonstrate an encounter–dilution effect against C. plutellae .     

Defensive larval secretions of leaf beetles attract a specialist predator Parasyrphus nigritarsis

1. Noxious larval secretions of leaf beetles, which repel generalist predators, do not deter specialist syrphid fly predators (genus Parasyrphus ). These flies cause considerable mortality to the beetles, but little is known about their foraging behaviour.
2. Larvae of Parasyrphus nigritarsis were attracted to the volatile larval secretions produced by two prey species Phratora vitellinae and Linaeidea aenea. Parasyrphus nigritarsis feeds on both beetles in nature. Phratora vitellinae feeds on willows and utilizes host plant compounds for secretion production, while the alder-feeding L. aenea produces an autogenous secretion.
3. Fly larvae were strongly attracted to pieces of filter paper treated with larval secretion of the beetles. They attempted to feed on them for up to 7 min, and were equally attracted to the secretions of Ph. vitellinae and L. aenea . Fly larvae were also attracted to pure salicyl aldehyde, the main component of the secretion of Ph. vitellinae .
4. Fly larvae searched extensively for prey on leaves that had been damaged by beetle larvae. They also followed trails made with solutions containing faecal matter of prey larvae. They showed no differential preference for Ph. vitellinae or L. aenea , but always rejected larvae of the non-prey leaf beetle Agelastica alni .
5. Beetle secretions thus play an important, but unexpected, role in the feeding behaviour of P. nigritarsis . This predator uses the beetle secretion to locate its prey. The implications of these results for three trophic level interactions are discussed.     

Salt spray differentially affects water status, necrosis, and growth in coastal sandplain heathland species

Sandplain heathlands are disturbance-dependent plant communities that occur infrequently in coastal areas of the northeastern United States. We hypothesize that salt spray plays a role in maintaining the composition of the heathland community by excluding salt-intolerant species close to the ocean. We examined the distributions of Solidago nemoralis, Myrica pensylvanica, Pinus rigida, and Quercus spp. in heathlands and conducted greenhouse studies to determine whether different levels of salt spray tolerance explain patterns found in the field. We found that common heathland forb and shrub species grow closer to the ocean than successional woody species. In greenhouse experiments, these species differ in their water status, necrosis, and growth responses to salt spray. The tree species P. rigida and Q. rubra are more susceptible to salt spray than the common heathland species M. pensylvanica. Our results suggest that salt spray may prevent tree species in heathlands from growing close to the ocean and therefore might be an important factor in maintaining the characteristic community composition of these dwarf shrublands in coastal habitats.     

Species identification of spiny lobster phyllosome larvae via ribosomal DNA analysis.

Within the tropical northwestern Atlantic, Panulirus argus, P. guttatus, and P. laevicauda (Palinuridae family), are sympatric. Numerous studies have examined the distribution and abundance of planktonic phyllosome larvae with respect to recruitment of spiny lobsters to the benthic population, but the data are of limited use because larvae of these species cannot yet be distinguished from one another by morphological characteristics. A simple molecular method that unambiguously differentiates adults or larvae of P. argus, P. guttatus, and P. laevicauda is described: a 5' region of 28s ribosomal DNA is amplified in vitro and then cut with a diagnostic restriction enzyme to identify each species. Data are also presented from the application of this method to representative plankton tows.     

Effects of Habitat Quality and Isolation on the Colonization of Restored Heathlands by Butterflies

In areas with intensive land use, such as the Netherlands, habitat fragmentation and loss of habitat quality due to eutrophication and drainage are major threats to the preservation of species‐rich communities of heathland and acid grassland. Restoration of such nutrient‐poor habitats may be carried out by removing the topsoil from ex‐arable land, in order to lower the nutrient levels. However, the establishment of target plant communities is known to be fragmentary. The current study shows that this also applies to butterflies. Ten years after topsoil removal in eight study areas, on average, only 3.5 of 10 characteristic heathland species were recorded on the sites. Species that did colonize had a significantly lower density than in the source populations. Our study indicates that although isolation effects were limiting colonization, poor habitat quality was the main limiting factor, mainly due to lack of host plants, hydrological conditions, and, to a lesser extent, lack of nectar plants and excessive residual nutrient levels. An experiment with the introduction of cut heather in one study area showed a significantly higher abundance of both target and nontarget butterflies in manipulated sites than in control sites. It can be concluded that habitat restoration by topsoil removal can be successful for butterflies of especially wet heathland habitats, provided that source populations are at close range and care is taken that complete plant communities are restored.     

Oviposition habitat selection by the mosquito, Culiseta longiareolata: effects of conspecifics, food and green toad tadpoles

Abstract. 1. Ovipositing females should choose egg-laying sites which maximize the growth and survivorship of their offspring. Accordingly, they should avoid habitats with high densities of competitors and predators and should choose sites with adequate food. We tested experimentally the oviposition response of the mosquito, Culiseta longiareolata Macquart, to extra food and two species which are both potential competitors and predators of its offspring: conspecific larvae and green toad ( Bufo viridis ) tadpoles.
2. We conducted the experiment in outdoor artificial pools where wild C. longiareolata females were presented with choices of all combinations of the presence or absence of: (1) C.longiareolata larvae, (2) B.viridis tadpoles, and (3) extra food (a mixture of ground fish food and mouse chow).
3. Oviposition rates averaged approximately 4 times greater in response to extra food.
4. The presence of the tadpoles, particularly later in the experiment when they were larger, caused a significant reduction (approximately 50%) in oviposition rate.
5. We did not detect a statistically significant effect of conspecific larvae on oviposition habitat selection.
6. Our results indicate that Culiseta females choose habitats for oviposition based on food availability. They also tend to avoid habitats containing Bufo tadpoles, presumably to avoid predation and/or competition.     

Conservation of Heathland Ground Beetles (Coleoptera,Carabidae): The Value of Lowland Coniferous Plantations

The value of conifer-dominated plantation forestry for heathland carabids is examined, in Breckland, eastern England. Historically dominated by heathland habitats supporting scarce species of restricted distribution within the UK, approximately 47% of Breckland was afforested in the early 20th century. The carabid fauna of this forest, previously little known, was sampled by intensive pitfall trapping, and results compared to samples from heathland and arable. The results affirm that Breckland supports many range-restricted carabid beetles, including scarce species restricted to lowland heathland or sandy habitats. Approximately half of the heathland-associated species, and also half of the Nationally Scarce carabid species recorded in Breckland, were found within the forest landscape. Carabid species composition differed between closed-canopy forest, heathland and arable, while open forest habitats (clear-felled stands, young restocked stands, and track margins) contained a diverse fauna with elements from all habitats. The proportion of heathland-associated species was significantly greater in open forest habitats than in closed-canopy forest and was similar to that of heathland assemblages. Wing morphology did not differ between heathland species solely recorded from heathland and those also recorded in the forest. Nationally Scarce species from the forest had a wider UK distribution than those recorded only from heathland. Our study shows that, within conifer forest planted in lowland heathland areas, open habitats provided by trackway networks and clear-felling management can have significant value for the conservation of open-ground carabids, including species of conservation importance.     

Adaptations to low temperature in high altitude insects from Mount Kenya

Abstract. 1. The strategies for low temperature survival in insects on Mount Kenya were investigated. The insects were collected from their natural habitats and their supercooling points and low temperature tolerances determined.
2. Most insects showed no special adaptations to low temperature survival and seem to depend on spending the cold nights in protected habitats, such as beneath stones and fallen trunks of plants, as well as within the wet frills of dead leaves of alpine plants, where they are protected by the heat released from freezing water.
3. Some insects, e.g. Collembola, aphids and a curculionid beetle, which live in relatively unprotected habitats, had low supercooling points, allowing them to remain unfrozen when exposed to low night temperatures. A nucleator free diet is apparently essential for the survival of such species.
4. Two species of curculionid beetles were found to withstand freezing down to -7C. These beetles had nucleating agents in their haemolymph and higher supercooling points than most of the other species studied.
5. A moderate freezing tolerance was found in larvae of a midge that lives in the watery liquid between the leaves of Senecio brassica .