Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.     

Species richness and community composition of mat-forming ectomycorrhizal fungi in old- and second-growth Douglas-fir forests of the HJ Andrews Experimental Forest,Oregon, USA

We investigated the species identity of mat-forming ectomycorrhizal (EM) fungi associated with old- and second-growth Douglas-fir stands. Using molecular analyses of rhizomorphs and EM root tips, we characterized 28 unique internal transcribed spacer sequences and considered them proxies for mat-forming EM species. In both stand age classes, one Athelioid species in the genus Piloderma dominated our sample of the mat-forming fungal community. In second-growth stands, the second most frequently encountered mat-forming EM species belonged to the genus Hysterangium. In old-growth stands, several Ramaria species were associated with a frequently encountered mat morphology but no species dominated the community. After using rarefaction analysis to standardize sampling effort, the total species richness did not differ statistically between old- and second-growth habitats. Both an abundance of infrequently encountered species and incomplete sampling of the mat-forming EM community may have limited our ability to detect potential differences in species richness. Several frequently encountered Piloderma species appear to have broad (holarctic) distributions and diverse host associations and their potential importance in forest ecosystems warrants further study.     

Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis

Summary In the humid tropics of SE Asia there are some 14 myrmecophytic species of the pioneer tree genus Macaranga (Euphorbiaceae). In Peninsular Malaysia a close association exists between the trees and the small, non-stinging myrmicine Crematogaster borneensis. These ants feed mainly on food bodies provided by the plants and have their colonies inside the hollow internodes. In a ten months field study we were able to demonstrate for four Macaranga species (M. triloba, M. hypoleuca, M. hosei, M. hulletti) that host plants also benefit considerably from ant-occupation. Ants do not contribute to the nutrient demands of their host plant, they do, however, protect it against herbivores and plant competition. Cleaning behaviour of the ants results in the removal of potential hervivores already in their earliest developmental stages. Strong aggressiveness and a mass recruiting system enable the ants to defend the host plant against many herbivorous insects. This results in a significant decrease in leaf damage due to herbivores on ant-occupied compared to ant-free myrmecophytes as well as compared to non-myrmecophytic Macaranga species. Most important is the ants' defense of the host plant against plant competitors, especially vines, which are abundant in the well-lit pioneer habitats where Macaranga grows. Ants bite off any foreign plant part coming into contact with their host plant. Both ant-free myrmecophytes and non-myrmecophytic Macaranga species had a significantly higher incidence of vine growth than specimens with active ant colonies. This may be a factor of considerable importance allowing Macaranga plants to grow at sites of strongest competition.     

Evolution of egg dumping in a subsocial insect

Egg dumping, or abandonment of eggs and young to the care of other conspecifics, frees individuals from costs of maternal care while potentially imposing energetic and ecological costs on egg recipients. It is not clear, however, that egg dumping necessarily represents selfish manipulation of egg recipients, and in some ecological contexts, recipients may benefit from enlarged broods. Thus, egg dumping may either be mutually beneficial for dumpers and recipients or entail costs for dumpers that are compensated by other means, such as improving reproduction of genetically related egg recipients. Here I use field experiments to test the relative importance of manipulation (i.e., "parasitism"), mutualism, and kin selection in the evolution of egg dumping in the tingid lace bug Gargaphia solani. In support of mutualism and kin selection, I found that reproduction of egg recipient G. solani benefits from brood enlargement, most likely because eggs and gregarious nymphs find safety in greater numbers. But contrary to both parasitism and mutualism, egg dumper reproduction was not improved by offspring abandonment. Indeed, dumpers laid smaller clutches than recipients, and dumpers did not convert a survival advantage into greater future reproduction. Genetic analyses of a natural G. solani population revealed, however, that dumpers are related to their egg recipients. Moreover, Hamilton's rule showed that egg-dumping G. solani earn sufficient indirect genetic benefits for kin selection to favor the behavior. Thus, egg dumping in some species may be kin-selected cooperation rather than parasitism or mutualism.     

Influence of Host Plant Structure and Microclimate on the Abundance and Behavior of a Tephritid Fly

Microclimate and host plant architecture significantly influence the abundance and behavior of insects. However, most research in this field has focused at the invertebrate assemblage level, with few studies at the single-species level. Using wild Solanum mauritianum plants, we evaluated the influence of plant structure (number of leaves and branches and height of plant) and microclimate (temperature, relative humidity, and light intensity) on the abundance and behavior of a single insect species, the monophagous tephritid fly Bactrocera cacuminata (Hering). Abundance and oviposition behavior were signficantly influenced by the host structure (density of foliage) and associated microclimate. Resting behavior of both sexes was influenced positively by foliage density, while temperature positively influenced the numbers of resting females. The number of ovipositing females was positively influenced by temperature and negatively by relative humidity. Feeding behavior was rare on the host plant, as was mating. The relatively low explanatory power of the measured variables suggests that, in addition to host plant architecture and associated microclimate, other cues (e.g., olfactory or visual) could affect visitation and use of the larval host plant by adult fruit flies. For 12 plants observed at dusk (the time of fly mating), mating pairs were observed on only one tree. Principal component analyses of the plant and microclimate factors associated with these plants revealed that the plant on which mating was observed had specific characteristics (intermediate light intensity, greater height, and greater quantity of fruit) that may have influenced its selection as a mating site.     

Biomass allocation patterns in terrestrial, epiphytic and aquatic species of Utricularia (Lentibulariaceae)

Utricularia forms the largest genus of carnivorous plants and is characterized by the possession of typical traps (“bladders”). Total biomass allocation was examined in three aquatic, six terrestrial and one epiphytic species of Utricularia from natural habitats in West Africa and from the Botanical Gardens, Bonn. Total biomass of aquatic species was considerably higher than that of terrestrial or epiphytic species. Epiphytic Utricularia accumulate about 35% of their biomass in green leaves, in contrast to 65% of nearly chlorophylless reproductive structures and traps. Aquatic species allocated more than 85% of their total biomass to stolons, leaves and traps, but only 10–13% to reproductive structures. This is in stark contrast to the allocation patterns of terrestrial bladderworts. These species allocate nearly 90% of their total biomass in reproductive structures, and only about 10% to stolons, leaves and traps. This reduction of photosynthetically active plant tissue strongly suggests that as a consequence of the alternative resource of chemical energy, the carnivorous habit might have partly replaced autotrophy in certain terrestrial Utricularia species, especially in some smaller ones.     

Response of predatory mites with different rearing histories to volatiles of uninfested plants

The behavioural response of the predatory mite Phytoseiulus persimilis to volatiles from several host plants of its prey, spider mites in the genus Tetranychus, was investigated in a Y-tube olfactometer. A positive response to volatiles from tomato leaves and Lima bean leaves was recorded, whereas no response was observed to volatiles from cucumber leaves, or leaves of Solanum luteum and Solanum dulcamara.Different results were obtained for predators that differed in rearing history. Predators that were reared on spider mites (Tetranychus urticae) on Lima bean leaves did respond to volatiles from Lima bean leaves, while predators that had been reared on the same spider mite species but with cucumber as host plant did not respond to Lima bean leaf volatiles. This effect is compared with the effect of rearing history on the response of P. persimilis to volatile allelochemicals of prey-infested plant leaves.     

Effectiveness of three Glomus species in protecting pepper (Capsicum annuum L.) against verticillium wilt

The objective of this work was to study the influence of three Glomus species—Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe, Glomus intraradices (Schenck and Smith) and Glomus deserticola (Trappe, Bloss, and Menge)—on the development of Verticillium-induced wilt in Capsicum annuum cv. Piquillo. Results showed that the effectiveness of arbuscular mycorrhizal fungi (AMF) as biocontrol agents varied among different Glomus species. In pepper colonized by G. intraradices the severity of the disease was even higher than that observed in non-mycorrhizal plants in terms of plant growth and pepper yield. On the other hand, the high effectiveness exhibited by G. mosseae in improving plant growth and the early beginning of the reproductive stage in these plants was not associated with great plant protection and high pepper yield in diseased plants. Only plants associated with G. deserticola had greater yield than non-mycorrhizal ones despite the lower P fertilization applied to the mycorrhizal treatment and this fact was observed in both healthy and diseased plants. It is suggested that the higher specific phosphorus uptake in Verticillium-inoculated plants associated with G. deserticola could contribute to diminish the deleterious effect of pathogen on yield. On the other hand, the possible influence of endogenous phenolics in roots on the tolerance or resistance of pepper against wilt induced by Verticillium dahliae remains unclear.     

Dryinid (Hym.: Dryinidae) parasitoids ofDalbulus leafhopper (Hom.: Cicadellidae) in Mexico

Little is known about the natural enemies of the leafhopperDalbulus spp. (Homoptera: Cicadellidae). Searches for its dryinid (wasps) parasitoids were made in Jalisco, Mexico. Jalisco contains the greatest number ofDalbulus species, and is considered to be near to the center of origin of this leafhopper genus and its host plants: maize, teosintes (Zea spp.) and gamagrasses (Tripsacum spp.). The dryinidGonatopus bartletti was found parasitizingD. maidis on maize and on annual teosinteZea mays spp.parviglumis. G. flavipes was found parasitizingD. elimatus on perennial teosinteZ. perennis; and a new speciesG. moyaraygozai andAnteon ciudadi parasitizingD. quinquenotatus onTripsacum pilosum andT. dactyloides. Parasitism by dryinids was found at altitudes of 680–2,000 m.Dalbulus maidis, the leafhopper species which causes the greatest losses in maize in Latin America, was found to be parasitized from 680–1,760 m. TheDalbulus species associated with annual host plants (maize andZ. mays spp.parviglumis) were parasitized by dryinids during the rainy season, while theDalbulus species associated with perennial host plants (Z. perennis andTripsacum) were parasitized by dryinids during both the rainy and dry season. The greatest diversity of dryinid parasitoids ofDalbulus spp. and the highest levels of parasitism were recorded from perennial plants, indicating that such species are reservoirs of natural enemies ofDalbulus spp.     

Hormonal Control of Egg Dumping and Guarding in the Lace Bug, Gargaphia solani (Hemiptera: Tingidae)

This study examines the effect of juvenile hormone (JH) or a similarly acting juvenoid on the expression of maternal egg guarding and its life history alternative, egg dumping, in the lace bug, Gargaphia solani Heidemann. JH manipulations were indirect: methoprene, a synthetic JH analog, and precocene II, an allatocidal phytochemical commonly used to reduce JH synthesis, were applied exogenously to test the hypothesis that high JH titers promote egg production and egg dumping behavior, while low titers terminate egg production and initiate maternal care. As predicted, egg dumpers treated with precocene II ignored dumping opportunities and became egg guarders. Similarly, egg guarders that were treated with methoprene became gravid within 2 days and abandoned their eggs to become egg dumpers. These manipulations suggest that hormones can trigger the expression of both egg dumping and egg guarding in G. solani even when environmental conditions are inappropriate.     

Egg-dumping lace bugs preferentially oviposit with kin

Egg dumping, or abandoning eggs and young to the care of other conspecifics, results in an extreme form of alloparental care. It is unclear, however, if egg dumpers discriminate among kin and nonkin egg recipients. In the lace bug Gargaphia solani (Heteroptera: Tingidae), some females with eggs (guards) also accept and defend eggs of conspecifics. Other females (egg dumpers) abandon their offspring after oviposition, leaving a single guard as the caregiver. We asked if egg dumpers preferentially dump their eggs among unguarded eggs of kin or nonkin. When given a choice between dumping among eggs of full siblings and eggs of nonsiblings, most eggs (67%) were dumped with full siblings' eggs. Furthermore, egg dumpers were just as likely to oviposit among eggs of kin with whom they had interacted on a shared host plant during juvenile development as they were to oviposit with kin reared on different host plants. Thus, egg dumpers discriminate kin by using cues associated with eggs, and such cues are not likely to be acquired through interaction on a common host plant environment. Copyright 2000 The Association for the Study of Animal Behaviour.     

A three-trophic-level analysis of the effects of plant hybridization on a leaf-mining moth

We investigated relationships between host plant hybridization in the Quercus grisea x Q. gambelii species complex and the distribution and performance of a leaf-mining moth in the genus Phyllonorycter. In 2 years at two sites Phyllonorycter densities were lowest on Q. grisea and increased through the categories of hybrid host plants to Q. gambelii. Direct host plant effects on Phyllonorycter performance were consistent with Phyllonorycter distribution; unexplained mortality of larvae in the mines, which is often associated with direct plant effects, decreased from Q. grisea through the hybrids to Q. gambelii. Plant hybridization influenced parasitism of Phyllonorycter. Parasitism was density dependent, and across all densities was higher on Q. grisea and Q. gambelii than on hybrid host plants.     

Granitic and gneissic outcrops (inselbergs) as centers of diversity for desiccation-tolerant vascular plants

Although desiccation tolerance is common in non-vascular plants, this adaptive trait is very rare in vascular plants. Desiccation-tolerant vascular plants occur particularly on rock outcrops in the tropics and to a lesser extent in temperate zones. They are found from sea level up to 2800 m. The diversity of desiccation-tolerant species as measured by number of species is highest in East Africa, Madagascar and Brazil, where granitic and gneissic outcrops, or inselbergs, are their main habitat. Inselbergs frequently occur as isolated monoliths characterized by extreme environmental conditions (i.e., edaphic dryness, high degrees of insolation). On tropical inselbergs, desiccation-tolerant monocotyledons (i.e., Cyperaceae and Velloziaceae) dominate in mat-like communities which cover even steep slopes. Mat-forming desiccation-tolerant species may attain considerable age (hundreds of years) and size (several m in height, for pseudostemmed species). Both homoiochlorophyllous and poikilochlorophyllous species occur. In their natural habitats, both groups survive dry periods of several months and regain their photosynthetic activity within a few days after rainfall. Other desiccation-tolerant species colonize shallow depressions, crevices and even temporarily water-filled rock pools on inselbergs. Desiccation-tolerant vascular plants occur in 13 families and are best represented within the monocotyledons and ferns. Only a few desiccation-tolerant dicots exist, in the Gesneriaceae, Myrothamnaceae and Scrophulariaceae. In total, about 330 species of vascular desiccation-tolerant plants are known, of which nearly 90% occur on inselbergs. With regard to morphological adaptations, the mat-forming monocotyledons are particularly remarkable due to the possession of roots with a velamen radicum, which is reported here in the genus Borya for the first time.     

Larval performance of two leafroller species on known and potential hosts

Two univoltine leafroller species,Archips argyrospila Walker andArchips rosana L., were reared from eclosion to adulthood on known or potential host plants. Both species were able to complete development on eight of the ten plants tested, the exceptions being apricot and peach. Rates of development differed among hosts, and indicated a range of host suitability. Differences in female pupal weights were apparent following the different rearing treatments, but these did not correlate with subsequent female fecundity. When development was compared between the two leafroller species,A. rosana had higher survivorship, more rapid development and higher female fecundity than didA. argyrospila, regardless of host. These findings indicate thatA. rosana may successfully exploit a wider host range than doesA. argyrospila. Comparisons of survivorship and developmental rates among hosts for each species of leafroller suggest that both species may have the potential for host-range expansion.     

Development and reproduction of the foxglove aphid (Hemiptera: Aphididae) on invasive swallow-worts (Vincetoxicum spp.)

The foxglove aphid, Aulacorthum solani (Kaltenbach), recently was documented using the invasive species pale and black swallow-wort (Vincetoxicum rossicum (Kleopow) Barbar. and V. nigrum (L.) Moench, respectively) as host plants. Because these are new host plant records for this polyphagous species, we investigated foxglove aphid development and reproduction on pale and black swallow-wort relative to a known crop host, potato (Solanum tuberosum L.), at a 25:20°C thermophase and a photoperiod of 16:8 (L:D) h. Almost no such data have been previously reported for a noneconomic host plant. Larval development and survival, as well as adult reproductive development and fecundity, were similar between the two swallow-wort species and potato. Adult aphids lived significantly longer on pale swallow-wort than the other two host plants, but this extended longevity encompassed the postreproductive stage. Foxglove aphid population parameters were therefore similar among the three plant species as well as most previous reports on crop plants. Pale and black swallow-wort appear to be suitable secondary hosts for foxglove aphid; other factors possibly limit aphid abundance on these two plant species in the field.     

Artificial algae as habitats for mobile epifauna: factors affecting colonization in a Japanese Sargassum bed

The utility of artificial habitats as tools in benthic studies was assessed by comparing the faunas associated with natural and artificial plants in a Japanese Sargassum bed. The 35 most common epifaunal species collected were all found to associate with both artificial and natural habitats, however, the abundances of species varied considerably between habitats. Colonization of artificial habitats was most rapid between 8 and 24 days after being placed in the field. This increased rate of colonization at intermediate time periods possibly reflected increasing attractiveness of habitats as a periphytic coating developed after the first week. Artificial habitats preconditioned in a sunlit environment to possess a layer of epiphytic algae attracted much higher numbers of epifaunal animals than habitats placed in a darkened but otherwise similar preconditioning environment. Epifaunal species appear to be attracted to artificial substrata, and presumably also natural habitats, largely because of associated diatoms and epiphytic plants.     

Pollination Ecology and Seed Production of Rhododendron Ponticum in Native and Exotic Habitats

Alien plants may be reproductively limited in exotic habitats because of a lack of mutualistic pollinators. However, if plants are adequately served by generalist pollinators, successful reproduction, naturalisation and expansion into exotic habitats may occur. Rhododendron ponticum is very successful, ecologically damaging invasive plant in Britain and Ireland, but is in decline in its native Iberian habitat. It spreads locally by sending out lateral branches, but for longer distance dispersal it relies on sexually produced seeds. Little is known about R. ponticum's pollination ecology and breeding biology in invaded habitats. We examined the flower-visiting communities and maternal reproductive success of R. ponticum in native populations in southern Spain and in exotic ones in Ireland. R. ponticum in flowers are visited by various generalist (polylectic) pollinator species in both native and exotic habitats. Although different species visited flowers in Ireland and Spain, the flower visitation rate was not significantly different. Insects foraging on R. ponticum in Spain carried less R. ponticum pollen than their Irish counterparts, and carried fewer pollen types. Fruit production per inflorescence varied greatly within all populations but was significantly correlated with visitation at the population level. Nectar was significantly depleted by insects in some exotic populations, suggesting that this invasive species is providing a floral resource for native insects in some parts of Ireland. The generality of the pollination system may be factor contributing to R.ponticum's success in exotic habitats.     

A mowing experiment to evaluate the influence of management on the activity of host ants of <Emphasis Type="Italic">Maculinea</Emphasis> butterflies

Maculinea butterflies obligatory parasitize certain species of Myrmica ants. Thus, the presence of the host ant species is a limiting factor for the survival of a Maculinea population. Here, we analyse the influence of vegetation structure and ground temperature on ant diversity and abundance on Maculinea habitats, with the final aim of identifying the environmental variables determining patterns of variation in species composition in order to recommend a mowing regime that will promote our three target species: Maculinea teleius, M. nausithous and M. alcon. Experimental plots with different mowing regimes were established at eight sites in South-Eastern Germany, a region which still contains a number of relatively large, stable populations of these threatened butterfly species. Among the seven different ant species recorded, four belong to the genus Myrmica (M. scabrinodis, M. rubra, M. ruginodis and M. vandeli). Among these, M. scabrinodis results most abundant at all sites. In a CCA analysis of environmental variables recorded at the studied plots, ant species diversity appears largely determined by litter cover, mean temperature, and mean grass cover. Mowing once a year, in the second half of September, after the larvae have left their host plants, enhances the abundance of Myrmica ants in the meadows, and would be the best management compromise for all three species.     

When genes meet nomenclature: tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone

We used a five-gene data set (mtDNA: 12S rRNA, 16S rRNA, cyt-b; nDNA: Cmos, Rag2) comprising approximately two-thirds of all extant testudinid species and, for the first time, including all five Testudo species to investigate the question of whether all western Palaearctic testudinids are monophyletic. Further, we examined whether the recently suggested allocation of the African Geochelone pardalis in the otherwise exclusively South African genus Psammobates and of the Malagasy G. yniphora in the monotypic genus Angonoka is justified in the face of considerable morphological evidence against such placements. Our phylogenetic analyses do not support the paraphyly and generic break-up of Testudo, as suggested by previous papers using a smaller taxon sampling and mtDNA data only. We propose a continued usage of the generic name Testudo for all five western Palaearctic tortoise species. Within Testudo, two monophyletic subclades are present, one containing T. hermanni+T. horsfieldii, and the other comprising (T. kleinmanni+T. marginata)+T. graeca. Nomenclaturally, we demonstrate that Eurotestudo Lapparent de Broin et al., 2006, which was recently erected with the type species T. hermanni, is an objective junior synonym of Chersine Merrem, 1820 and Medaestia Wussow, 1916. Recognition of a monotypic genus Angonoka for G. yniphora is unwarranted according to both our re-analysis of sequence data and morphological data. Acknowledging the strong morphological similarity between G. yniphora and G. radiata, we suggest placing both species into the genus Astrochelys. Although sequence data for only one of the three Psammobates species was available for analysis, there is currently no cause to challenge the monophyly of this genus as established on the basis of morphological evidence. Thus, we hypothesize that G. pardalis is sister to a monophyletic Psammobates. In light of the clear morphological gap between G. pardalis and Psammobates species, the recognition of a distinct genus Stigmochelys for the former seems justified.     

The host range and selectivity of a parasitic plant: Rhinanthus minor L.

Summary Rhinanthus minor (Yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Britain. Association analysis of the dune vegetation at Holme-next-the-Sea in eastern England revealed only two potential host plants through positive association. In contrast direct examination of the root systems revealed haustorial connections with 20 host species. The number of species parasitized by one plant ranged from one to seven. Data from another four sites in Britain and one in central Europe indicate that the natural host range of R. minor encompasses at least 50 species from 18 families with 22% in the Leguminosae and 30% in the Gramineae. Comparison of the number of haustorial connections made to each species with the abundance of roots in the soil shows that R. minor is a highly selective parasite, but that the selectivity is not consistent between populations or between plants from different parts of the same population. The reasons for host selectivity are discussed.