Integrative species delimitation in Nearctic ambush bugs (Heteroptera: Reduviidae: Phymatinae): insights from molecules,geometric morphometrics and ecological associations

Ambush bugs (Hemiptera: Reduviidae: Phymatinae) are sit-and-wait predators of flower-visiting insects including pollinators. Broad species distribution ranges, intraspecific polymorphism, sexual dimorphism and subtle interspecific differences all contribute to making species delimitation especially difficult in this group, which is used as a model in the study of interactions between sexual dimorphism and sexual selection. Species boundaries among Nearctic ambush bugs in the common and frequently collected erosa species group (11 species, nine subspecies) have therefore remained unclear, resulting in a complex and poorly justified taxonomy. Recent molecular phylogenetic research suggested that several widespread Nearctic species are para- or polyphyletic. We here build on this research, integrating geometric morphometrics, molecular species delimitation approaches and host plant association data to provide a comprehensive dataset with respect to both taxon and character sampling with the goal of teasing apart evolutionary lineages of Nearctic Phymata Latreille. Although molecular-based species delimitation analyses suggested a variety of species hypotheses, probably as a result of striking discordance between mitochondrial and nuclear ribosomal genes, the combination of these with geometric morphometric data enabled us to confidently delimit several of these problematic taxa. In addition, geometric morphometric analysis of pronotal shape revealed undocumented morphological patterns that appear to be useful in the diagnosis of many of the surveyed taxa. The results from this study provide an objective foundation for the much-needed taxonomic revision of the most ubiquitous ambush bugs in North America.     

Higher taxa and the identification of areas of endemism

Quantitative analyses of areas of endemism have rarely considered higher taxa. This paper discusses aspects related to the use of higher taxa in the analysis of areas of endemism, and computer implementations. An example of the application of the method is provided, with a data set for Nearctic mammals, showing that some of the areas recognized by species‐level taxa also adjust well to the distribution of other taxa of higher level (genera, monophyletic groups).     

Impact of Nosema (Microsporidia) infection and fumagillin treatment on Lygus lineolaris (Hemiptera: Miridae)

A consistent supply of healthy tarnished plant bug, Lygus lineolaris (Palisot de Beauvois) (Hemiptera: Miridae), is necessary for the development of novel management strategies targeting this pest. After being in culture for several years, a substantial portion of a tarnished plant bug colony was found to be infected with a Nosema (Microsporidia) species. Studies were subsequently undertaken to evaluate the impact of Nosema infection on tarnished plant bug productivity and to test the efficacy of fumagillin to treat this infection. Using buffalo black stain, infections could not be reliably detected in adult tarnished plant bugs until adults were 6-8days post eclosion. Nosema infections reduced adult longevity and fecundity. Maximum fecundity was restored using a concentration of 16.8ppm fumagillin while maximum longevity for females was at a concentration of 33.6ppm fumagillin incorporated into the tarnished plant bug diet. Minimum infection scores were obtained at 67.2ppm, the highest concentration tested. A field survey of tarnished plant bugs in Mississippi found Nosema infections in 3% of wild tarnished plant bugs.     

The potential of a jumping spider, Phidippus clarus, as a biocontrol agent

Spiders, particularly assemblages of species, have been shown to be effective in reducing pest insects and crop damage in field crops and orchards. We investigated the potential for a single jumping spider species to reduce pests in a greenhouse setting. We placed three treatments in large enclosures: 1) control treatment of only sweet basil, Ocimum basilicum L.; 2) sweet basil and a phytophagous pest, fourlined plant bug, Poecilocapsus lineatus (F.) (Heteroptera: Miridae); and 3) sweet basil, fourlined plant bug, and jumping spider Phidippus clarus (Keyserling 1884). After 1 wk, jumping spiders reduced the number of plant bugs. Plants exposed to plant bugs alone were significantly shorter than either control plants or plants exposed to plant bugs and spiders. Chlorophyll concentration did not significantly differ across treatments. We discuss the feasibility of using P. clarus and similar salticids in biocontrol.     

Molecular phylogeny of the plant bugs (Heteroptera: Miridae) and the evolution of feeding habits

The first comprehensive cladistic analysis of Miridae, the plant bugs, is presented based on analysis of 3935 base pairs of mitochondrial (16S, COI) and nuclear (18S, 28SD3) DNA for 91 taxa in seven subfamilies. Data were analysed using maximum likelihood (ML), parsimony and Bayesian inference (BI) phylogenetic frameworks. The phylogenetic results are compared with previous hypotheses of higher relationships in the family using alternative hypothesis tests. A Bayesian relaxed molecular clock is used to examine divergence times, and ancestral feeding habits are reconstructed using parsimony and a Bayesian approach. Clades recovered in all analyses are as follows: Cimicomorpha, Miroidea and Miridae; Bryocorinae: Bryocorini; Stenodemini; Mirinae; Deraeocorinae (Clevinemini + Deraeocorini); Cylapinae; Isometopinae; Bryocorinae: Dicyphini; Orthotylini; Phylinae (Phylini + Pilophorini), and Phylinae as sister group to all the remaining mirid taxa. These results are largely congruent with former hypotheses based on morphological data with respect to the monophyly of various subfamilies and tribes; however, our results indicate that the subfamily Bryocorinae is not monophyletic, as the two tribes, Dicypini and Bryocorini, were separated in the phylogenetic results. Divergence time estimates indicate that the radiation of the Miridae began in the Permian; most genus‐level radiations within subfamilies began in the late Cretaceous, probably in response to the angiosperm radiation. Ancestral feeding state reconstructions based on Bayesian and parsimony inference were largely congruent and both reconstructed phytophagy as the ancestral state of the Miridae. Furthermore, the feeding habits of the common ancestors of Mirinae + Deraeocorinae, Bryocorinae + Cylapinae + Isometopinae + Orthotylinae, and the remaining taxa excluding Phylinae, were inferred as phytophagous. Therefore, at least three shifts from phytophagy or polyphagy to predation occurred within the Miridae. Additionally, based on the mirid host‐plant records, we discovered several trends, such as a strong relationship between host‐plant ranges and a facultative feeding habit. © The Willi Hennig Society 2011.     

Changes in abundance of native and introduced parasites (Hymenoptera: Braconidae), and of the target and non-target plant bug species (Hemiptera: Miridae), during two classical biological control programs in alfalfa.

High numbers of tarnished plant bugs [Lygus lineolaris (Palisot)], were once common in alfalfa, as was a low level of parasitism (9%) by the native Peristenus pallipes (Curtis). After the bivoltine European parasite Peristenus digoneutis Loan became well established, average parasitism of the first and second generations increased to 64%, and tarnished plant bug numbers dropped by 65%. This reduced host density eventually caused a decline in total parasitism by both parasite species to 22%. A few P. digoneutis also attacked the alfalfa plant bug, Adelphocoris lineolatus (Goeze), but did not reduce this pest or increase its parasitism rate. At another location, where P. digoneutis is not established, parasitism of first generation alfalfa plant bugs, an adventive (accidently introduced) pest, was increased to 21% by the introduced univoltine parasite, Peristenus conradi Marsh, and a slight reduction in the pest may have resulted. P. digoneutis did not parasitize the meadow plant bug, Leptopterna dolabrata (L.), an adventive pest of forage grasses, so did not affect this mirid or its parasite. Neither introduced parasite eliminated the native parasites of the tarnished or alfalfa plant bugs. The narrow host ranges of the braconid parasites of mirid nymphs are contrasted with the broad host range of the native tachinid parasite [Phasia robertsoni (Towns.)] of adult mirids. The major changes in mirid abundance and their mortality by parasites that slowly occurred during this 19-year study demonstrate the need for long-term field research, to adequately document and understand these complex interactions.     

Sampling strategies for square and boll-feeding plant bugs (Hemiptera: Miridae) occurring on cotton

Sampling methods for square and boll-feeding plant bugs (Hemiptera: Miridae) occurring on cotton, Gossypium hirsutum L., were compared with the intent to assess if one approach was viable for two species occurring from early-season squaring to late bloom in 25 fields located along the coastal cotton growing region of south Texas. Cotton fleaphopper, Pseudatomoscelis seriatus (Reuter), damages squares early-season and dominated collections using five sampling methods (approximately 99% of insects collected). A major species composition shift occurred beginning at peak bloom in coastal fields, when verde plant bug, Creontiades signatus Distant, represented 55-65% of collections. Significantly more cotton fleahoppers were captured by experienced samplers with the beat bucket and sweep net than with the other methods (30-100% more). There were more than twice as many verde plant bugs captured by experienced and inexperienced samplers with the beat bucket and sweep net than captured with the KISS and visual methods. Using a beat bucket or sweep net reduced sampling time compared with the visual method for the experienced samplers. For both species, comparing regressions of beat bucket-based counts to counts from the traditional visual method across nine cultivar and water regime combinations resulted in only one combination differing from the rest, suggesting broad applicability and ability to translate established visual-based economic thresholds to beat bucket-based thresholds. In a first look at sample size considerations, 40 plants (four 10-plant samples) per field site was no more variable than variation associated with larger sample sizes. Overall, the beat bucket is much more effective in sampling for cotton fleahopper and verde plant bug than the traditional visual method, it is more suited to cotton fleahopper sampling early-season when plants are small, it transitions well to sample for verde plant bug during bloom, and it performs well under a variety of soil moisture conditions and cultivar selections.     

Herbivory by Strongylocoris leucocephalus (Hemiptera: Miridae) on a novel host plant Adenophora triphylla var. japonica in Japan

When monophagous and oligophagous insect herbivores colonize new areas, they sometimes use novel hosts. The availability of the new hosts can facilitate further expansion into regions beyond the geographic range of the original hosts, resulting in specialization on different host plant taxa in different parts of the herbivore's geographic range. Strongylocoris leucocephalus Linnaeus (Hemiptera: Miridae), which is a plant bug attacking Campanula spp., is widely distributed in the Eurasian continent, North Africa, and northern Japan. In the process of geographic range expansion, S. leucocephalus may incorporate novel host plants into their host range. I investigated the host plant species of S. leucocephalus in central Japan. Field observations revealed that the plant bug fed on the novel host plant Adenophora triphylla (Thunb.) A. DC. var. japonica (Regel) H. Hara (Campanulaceae), although feeding by S. leucocephalus was not associated with plant mortality. However, S. leucocephalus never fed on Campanula punctata Lam. var. hondoensis (Kitam.) Ohwi, the only Campanula species at the study site. The plant bug S. leucocephalus and the novel host plant A. triphylla var. japonica are not entirely distributed sympatrically, suggesting that the plant bug has expanded its geographic range by incorporating new hosts.     

Effect of seasonal variation in host-plant quality on the rice leaf bug, Trigonotylus caelestialium

The rice leaf bug, Trigonotylus caelestialium (Kirkaldy) (Heteroptera: Miridae), is an important insect pest, causing pecky-rice damage in Japan. This bug has a multivoltine life cycle and feeds on various species of the Poaceae. The phenology of the host-plants differs between species and the plant quality of a single host species may change with the seasons. The fitness of the bug may be affected by these seasonal changes in host-plant quality. In the current study, the effects of such quality changes on adult and nymphal performance were examined using a population of bugs collected in Joetsu, Niigata, Japan. Rearing of newly emerged adults collected from spring to autumn from Italian rye grass, Lolium multiflorum Lam., and southern crabgrass, Digitaria ciliaris (Retz.), showed that body size and performance of adults differed between collection seasons and plant species, indicating that nymphal conditions including host-plant quality have crucial effects on their fitness. Rearing nymphs from hatching to adult emergence on various field-collected grasses under laboratory conditions showed that the performance of nymphs depends on the plant species and varies greatly with the season, within a given species. In general, nymphs performed more poorly on most of the plants collected late in the season, but they performed well on some plants unavailable early in the season. Therefore, in T. caelestialium , seasonal and spatial variations are important factors that affect fitness of nymphs and adults, and this suggests that oviposition of diapause eggs in summer is an adaptation to deteriorating food conditions.     

Influence of corn on stink bugs (Heteroptera: Pentatomidae) in subsequent crops

In southeastern United States farmscapes, corn, Zea mays L., is often closely associated with peanut, (Arachis hypogaea L.), cotton, (Gossypium hirsutum L.), or both. The objective of this 3-yr on-farm study was to examine the influence of corn on stink bugs (Heteroptera: Pentatomidae), Nezara viridula (L.), and Euschistus servus (Say), in subsequent crops in these farmscapes. Adults of both stink bug species entered corn first, and seasonal occurrence of stink bug eggs, nymphs, and adults indicated that corn was a suitable host plant for adult survival and nymphal development to adults. Stink bug females generally oviposited on cotton or peanut near the interface, or common boundary, of the farmscape before senescence of corn, availability of a new food, or both. Adult stink bugs dispersed from crop to crop at the interface of a farmscape in response to senescence of corn, availability of new food, or both. In corn-cotton farmscapes, adult stink bugs dispersed from senescing corn into cotton to feed on bolls (fruit). In corn-peanut farmscapes, adult stink bugs dispersed from senescing corn into peanut, which apparently played a role in nymphal development in these farmscapes. In the corn-cotton-peanut farmscape, stink bug nymphs and adults dispersed from peanut into cotton in response to newly available food, not senescence of peanut. Stink bug dispersal into cotton resulted in severe boll damage. In conclusion, N. viridula and E. servus are generalist feeders that exhibit edge-mediated dispersal from corn into subsequent adjacent crops in corn-cotton, corn-peanut, and corn-peanut-cotton farmscapes to take advantage of suitable resources available in time and space for oviposition, nymphal development, and adult survival. Management strategies for crops in this region need to be designed to break the cycle of stink bug production, dispersal, and expansion by exploiting their edge-mediated movement and host plant preferences.     

New World biogeography and the evolution of polychromatism: evidence from the bee assassin genus Apiomerus (Heteroptera: Reduviidae: Harpactorinae)

The crassipes and pictipes species groups of Apiomerus Hahn together contain 12 species, many with high intraspecific chromatic variability, which represent the majority of Nearctic species in this New World assassin bug genus. Because of their geographical distribution and their varying degrees of polychromatism, these two species groups provide a unique opportunity to study the evolution of polychromatism and analyse relationships among areas of endemism in the Nearctic, as well as determine the boundary between the Nearctic and Neotropical regions. The results of a morphology based phylogenetic analysis allowed investigation of these questions while also determining relationships among the 12 species in the two species groups. The crassipes and pictipes species groups were each supported as monophyletic and as sister taxa. Apiomerus rufipennis (Fallou) was not included in the original concept of the crassipes species group, but is shown here to be a member of the group. Apiomerus barrocoloradoi Forero, Berniker & Szerlip, which had been hypothesized previously to belong to the pictipes species group, is excluded from this group. Intraspecific polychromatism for each species was identified as being present in one of three states: no polychromatism; limited polychromatism; or polychromatism as discrete colour morphs. Limited polychromatism was here found to be the ancestral state for Apiomerus, and species with discrete colour morphs are restricted to the crassipes and pictipes species groups. Polychromatism appears to be a greatly homoplastic character within the genus. A Brooks parsimony analysis recovered distinct Nearctic and Neotropical clades. The Nearctic clade is divided between areas in the central and eastern U.S.A. and areas in the Western U.S.A. and Mexico. The Nearctic–Neotropical boundary for the taxa included in the analysis is along the Isthmus of Tehuantepec in southern Mexico.     

Testing the resource concentration hypothesis with tarnished plant bug on strawberry: density of hosts and patch size influence the interaction between abundance of nymphs and incidence of damage

Abstract. 1. Attributes of patches with strawberry plants were manipulated experimentally to assess the impact of patch size and host density on the abundance of tarnished plant bug (Hemiptera: Miridae) and incidence of fruit damage.
2. The density of nymphs per inflorescence increased with patch size and host density for some but not all generations of plant bug, providing partial support for the resource concentration hypothesis, while emphasising the importance of replicating experiments across several generations of herbivore.
3. Increasing density of nymphs with increasing patch size and host density did not translate into a high incidence of damage, which may be due to the relatively low feeding impact of nymphs in patches with numerous fruits and to the relatively high perimeter-to-area ratio in small patches resulting in a relatively high proportion of damaged fruits.
4. The decreasing proportion of damaged fruits with increasing density of fruits per patch suggests that nymphs exhibit a saturating functional response to density of strawberry fruits. Functional responses of herbivores may be a critical yet overlooked component that influences interactions between insects and their host plant. In particular, concentrated plant resources may result in increased numbers of herbivores yet sill have a negligible influence on plant damage and/or fitness.     

Facilitation of a native pest of rice, Stenotus rubrovittatus (Hemiptera: Miridae), by the non-native Lolium multiflorum (Cyperales: Poaceae) in an agricultural landscape

Source populations of polyphagous pests often occur on host plants other than the economically damaged crop. We evaluated the contribution of patches of a non-native meadow grass, Lolium multiflorum Lam. (Poaceae), and other weeds growing in fallow fields or meadows as source hosts of an important native pest of rice, Stenotus rubrovittatus (Matsumura) (Hemiptera: Miridae), in an agricultural landscape of northern Japan. Periodical censuses of this mirid bug by using the sweeping method, vegetation surveys, and statistical analysis revealed that L. multiflorum was the only plant species that was positively correlated with the density of adult S. rubrovittatus through two generations and thus may be the most stable and important host of the mirid bug early in the season before the colonization of rice paddies. The risk and cost of such an indirect negative effect on a crop plant through facilitation of a native pest by a non-native plant in the agricultural landscape should not be overlooked.     

Differential induction of volatiles in rice plants by two stink bug species influence behaviour of conspecifics and their natural enemy Telenomus podisi

Tritrophic interactions mediated by semiochemicals have been intensively studied from the viewpoint of ecological relationships with Nearctic tritrophic organisms. However, there are few studies involving interactions with different herbivores on the same host plant in Neotropical systems. The objective of the current study was to investigate the effects of herbivory by two species of stink bugs (Heteroptera: Pentatomidae) with the same feeding habit – Tibraca limbativentris Stål and Glyphepomis spinosa Campos & Grazia – on indirect and direct defence strategies of rice plants. The responses of each stink bug species (virgin and mated females) and of their main natural enemy, the egg parasitoid Telenomus podisi Ashmead (Hymenoptera: Platygastridae; mated females), to volatiles from undamaged and herbivore-damaged rice plants were evaluated using a Y-tube olfactometer. The results showed that rice plants responded differently to T. limbativentris or G. spinosa herbivory, enhancing the production of a different blend of volatile compounds, which reduced the attraction for conspecific stink bugs and elicited the foraging behaviour of T. podisi.     

Biogeographic regions of North American mammals based on endemism

Since the 19th Century, two regions have been recognized for North American mammals, which overlap in Mexico. The Nearctic region corresponds to the northern areas and the Neotropical region corresponds to the southern ones. There are no recent regionalizations for these regions under the criterion of endemism. In the present study, we integrate two methods to regionalize North America, using species distribution models of mammals: endemicity analysis (EA) and parsimony analysis of endemicity (PAE). EA was used to obtain areas of endemism and PAE was used to hierarchize them. We found 76 consensus areas from 329 sets classified in 146 cladograms, and the strict consensus cladogram shows a basal polytomy with 14 areas and 16 clades. The final regionalization recognizes two regions (Nearctic and Neotropical) and a transition zone (Mexican Transition Zone), six subregions (Canadian, Alleghanian, Californian‐Rocky Mountain, Pacific Central America, Mexican Gulf‐Central America, and Central America), two dominions (Californian and Rocky Mountain), and 23 provinces. Our analysis show that North America is probably more complex than previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 485–499.     

THE EVOLUTION OF LOCAL ENDEMISM IN MADAGASCAR: WATERSHED VERSUS CLIMATIC GRADIENT HYPOTHESES EVALUATED BY NULL BIOGEOGRAPHIC MODELS

Substantial insular speciation has resulted in exceptionally high levels of endemism in Madagascar, creating locally restricted species' ranges that remain poorly understood. The contributions of alternative processes that could influence patterns of local endemism—including speciation by geographic isolation or adaptation to environmental gradients—are widely debated, both for Madagascar and elsewhere. A recently proposed hypothesis (the "watershed hypothesis") suggests that allopatric speciation driven by isolation in watersheds during Quaternary climate shifts provides a general explanation for patterns of local endemism across taxa in Madagascar. Here we tested coincidence between species' distributions and areas of endemism predicted by two contrasting biogeographic hypotheses: (1) the watershed hypothesis, and (2) an alternative hypothesis driven by climatic gradients (the "current climate hypothesis"). Statistical significance of coincidence was assessed by comparing against a null model. Surprisingly, we found that extant distributions of lemurs, geckos, and chameleons reveal species patterns that are significantly coincident with the watershed and current climate hypotheses. These results strongly support local endemism developing from multiple processes, even among closely related species. Our findings thus indicate that pluralistic approaches will offer the best option both for understanding processes that generate local endemism, and for incorporating endemism within conservation priority setting.     

Influence of genetic architecture on contemporary local evolution in the soapberry bug,Jadera haematoloma: artificial selection on beak length

Little is known about the influence of genetic architecture on local adaptation. We investigated the genetic architecture of the rapid contemporary evolution of mouthparts, the flight polymorphism and life history traits in the soapberry bug Jadera haematoloma (Hemiptera) using laboratory selection. The mouthparts of these seed‐feeding bugs have adapted in 40–50 years by decreasing in length following novel natural selection induced by a host switch to the seeds of an introduced tree with smaller fruits than those of the native host vine. Laboratory selection on beak length in both an ancestral population feeding on the native host and a derived population feeding on the introduced host reveals genetic variance allowing a rapid response (heritabilities of 0.51–0.87) to selection for either longer or shorter beaks. This selection resulted in reverse evolution by restoring long beaks in the derived population and forward evolution by re‐creating short beaks in the ancestral bugs. There were strong genetic correlations (0.68–0.84) in both populations between beak lengths and the frequency of flight morphs, with short beaks associated with short wings. The results reveal a genetically interrelated set of adaptive multivariate traits including both beak length and flight morph. This suite of traits reflects host plant patchiness and seeding phenology. Weaker evidence suggests that egg mass and early egg production may be elements of the same suite. Reversible or forward evolution thus may occur in a broad set of genetically correlated multivariate traits undergoing rapid contemporary adaptation to altered local environments.     

Nesidiocoris tenuis as a pest in Northwest Europe: Intervention threshold and influence of Pepino mosaic virus

The use of Nesidiocoris tenuis (Hemiptera: Miridae) as a biocontrol agent is controversial as it is considered a pest in Northwest European tomato greenhouses, due to its tendency to damage the plant and fruit. Necessary chemical plant protection products to control N. tenuis have toxic side effects on important beneficials like Macrolophus pygmaeus (Hemiptera: Miridae), which jeopardizes the whole IPM programme. In this study, several commercial tomato greenhouses were monitored for mirid populations. The relationship between the number of N. tenuis individuals and plant damage was assessed in function of availability of prey and interaction with M. pygmaeus. These greenhouse data were used to determine a practical density intervention threshold. Next, the hypothesis that a Pepino mosaic virus (PepMV) infection increases plant and fruit damage by N. tenuis (as has been shown for M. pygmaeus) was tested. Plant damage occurred when the average number of predatory bugs in the head of the plant exceeded 16 per ten plants. Plant damage increased in severity at increasing predatory bug densities, independent of the availability of prey and M. pygmaeus presence. Plant and fruit damage were not affected by the presence of PepMV, as was shown for fruit damage in previous studies for M. pygmaeus. Our study provides a practical density intervention threshold for growers in greenhouse crops. Simple monitoring of the number of predatory bugs in the head of the plant can be used to take specific biocontrol actions. It was also shown that only the predatory bug N. tenuis itself causes damage, and there is no interaction with PepMV.     

Global patterns in species richness of pelagic seabirds: the Procellariiformes

Quantitative information on large scale spatial patterns of biodiversity remains poor, especially for pelagic systems. In this paper the regional diversity of procellariiforms is mapped worldwide at the species level. These seabirds do not display a conventional latitudinal gradient of decreasing species richness towards high latitudes, but rather are most speciose between 37° and 59°S in all ocean basins. Based on data for foraging ranges, areas with the highest species richness and the most species with smaller range sizes are all found in the vicinity of New Zealand and its sub-Antarctic islands. In contrast, data for breeding ranges show islands in the southern Indian and Atlantic oceans to have the highest number of breeding species, while these islands and New Zealand have the most species with smaller range sizes. No northern hemisphere regions are amongst the top ten grid cells for foraging and breeding species richness, although Hawaii has the highest species richness of procellariilforms north of the equator. Northern Baja California. Madeira, the Canary islands, and the west coast of South America are all important sites of narrow endemism in the northern hemisphere. High species richness and narrow endemism coincide with areas of significant longline fishing activity in the southern hemisphere. Near-minimum sets based on one and three representations demonstrate that if all procellariiform species are to be retained, large areas of the ocean and almost all breeding sites require conservation.     

Potential for classical biological control of the potato bug Closterotomus norwegicus (Hemiptera: Miridae): description, parasitism and host specificity of Peristenus closterotomae sp. n. (Hymenoptera: Braconidae)

The potato bug, Closterotomus norwegicus (Gmelin) (Hemiptera: Miridae) is an introduced pest of lucerne, white clover and lotus seed crops in New Zealand and a key pest of pistachios in California, USA. Efforts were made to identify potential biological control agents of C. norwegicus in Europe. A total of eight parasitoids, including six primary parasitoids from the genus Peristenus (Hymenoptera: Braconidae) and two hyperparasitoids from the genus Mesochorus (Hymenoptera: Ichneumonidae), were reared from C. norwegicus nymphs collected in various habitats in northern Germany. With a proportion of more than 85% of all C. norwegicus parasitoids, Peristenus closterotomae (Hymenoptera: Braconidae), a new species, was the most dominant parasitoid, whereas other parasitoid species only occurred sporadically. Peristenus closterotomae did not fit in the keys to any described species and is described as new to science. Parasitism caused by P. closterotomae was on average 24% (maximum 77%). To assess the host specificity of parasitoids associated with C. norwegicus, the parasitoid complexes of various Miridae occurring simultaneously with C. norwegicus were studied. Peristenus closterotomae was frequently reared from Calocoris affinis (Herrich-Schaeffer), and a few specimens were reared from Calocoris roseomaculatus (De Geer) and the meadow plant bug, Leptopterna dolobrata (Linnaeus) (all Hemiptera: Miridae). The remaining primary parasitoids associated with C. norwegicus were found to be dominant in hosts other than C. norwegicus. Whether nymphal parasitoids may potentially be used in a classical biological control initiative against the potato bug in countries where it is introduced and considered to be a pest is discussed.