Herbivore specificity and the chemical basis of plant-plant communication in Baccharis salicifolia (Asteraceae).

It is well known that plant damage by leaf-chewing herbivores can induce resistance in neighbouring plants. It is unknown whether such communication occurs in response to sap-feeding herbivores, whether communication is specific to herbivore identity, and the chemical basis of communication, including specificity. We carried out glasshouse experiments using the California-native shrub Baccharis salicifolia and two ecologically distinct aphid species (one a dietary generalist and the other a specialist) to test for specificity of plant-plant communication and to document the underlying volatile organic compounds (VOCs). We show specificity of plant-plant communication to herbivore identity, as each aphid-damaged plant only induced resistance in neighbours against the same aphid species. The amount and composition of induced VOCs were markedly different between plants attacked by the two aphid species, providing a putative chemical mechanism for this specificity. Furthermore, a synthetic blend of the five major aphid-induced VOCs (ethanone, limonene, methyl salicylate, myrcene, ocimene) triggered resistance in receiving plants of comparable magnitude to aphid damage of neighbours, and the effects of the blend exceeded those of individual compounds. This study significantly advances our understanding of plant-plant communication by demonstrating the importance of sap-feeding herbivores and herbivore identity, as well as the chemical basis for such effects.

Traits underlying community consequences of plant intra-specific diversity.

A plant's performance and interactions with other trophic levels are recorgnized to be contingent upon plant diversity and underlying associational dynamics, but far less is known about the plant traits driving such phenomena. We manipulated diversity in plant traits using pairs of plant and a substitutive design to elucidate the mechanisms underlying diversity effects operating at a fine spatial scale. Specifically, we measured the effects of diversity in sex (sexual monocultures vs. male and female genotypes together) and growth rate (growth rate monocultures vs. fast- and slow-growing genotypes together) on growth of the shrub Baccharis salicifolia and on above- and belowground consumers associated with this plant. We compared effects on associate abundance (# associates per plant) vs. density (# associates per kg plant biomass) to elucidate the mechanisms underlying diversity effects; effects on abundance but not density suggest diversity effects are mediated by resource abundance (i.e. plant biomass) alone, whereas effects on density suggest diversity effects are mediated by plant-based heterogeneity or quality. Sexual diversity increased root growth but reduced the density (but not abundance) of the dietary generalist aphid Aphis gossypii and its associated aphid-tending ants, suggesting sex mixtures were of lower quality to this herbivore (e.g. via reduced plant quality), and that this effect indirectly influenced ants. Sexual diversity had no effect on the abundance or density of parasitoids attacking A. gossypii, the dietary specialist aphid Uroleucon macolai, or mycorrhizae. In contrast, growth rate diversity did not influence plant growth or any associates except for the dietary specialist aphid U. macolai, which increased in both abundance and density at high diversity, suggesting growth rate mixtures were of higher quality to this herbivore. These results highlight that plant associational and diversity effects on consumers are contingent upon the source of plant trait variation, and that the nature of such dynamics may vary both within and among trophic levels.

The first record of Baccharis L. (Asteraceae) as a host-plant genus for Nepticulidae (Lepidoptera), with description of new Stigmella species from South America.

We record the first Nepticulidae species found to feed on Baccharis L. (Asteraceae). Despite the high species richness of Baccharis in the Western Hemisphere, no nepticulid has ever been recorded feeding on Baccharis. In this paper we describe six new Stigmella Schrank species feeding on Baccharis: S. emarginatae Diskus & Stonis, sp. nov., S. bipartita Diskus & Stonis, sp. nov., S. tripartita Diskus & Stonis, sp. nov., S. latifoliae Remeikis, Diskus & Stonis, sp. nov., S. baccharicola Diskus & Stonis, sp. nov., and S. confertae Diskus & Stonis, sp. nov. The remaining two taxa are left unnamed. All taxa are illustrated with photographs of adults, their genitalia, and their leaf-mines. Additionally, leaf-mines on Baccharis salicifolia are documented.

A new cecidogenous species of Eugnosta Hübner (Lepidoptera: Tortricidae) associated with Baccharis salicifolia (Asteraceae) in the northern Chilean Atacama Desert: Life-history description and phylogenetic inferences.

Eugnosta Hübner, 1825 (Lepidoptera, Tortricidae, Tortricinae, Cochylini, Cochylina) is reported for the first time in Chile. Male and female adults, the pupa, the last-instar larva, and galls of Eugnosta azapaensis Vargas & Moreira, sp. n., are described and illustrated from the Azapa Valley in the northern Atacama Desert. The larvae induce fusiform galls on shoots of the shrub Baccharis salicifolia (Ruiz & Pav.) Pers. (Asteraceae). An assessment of phylogenetic relationships of E. azapaensis with two congeneric species based on mitochondrial DNA is provided.

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation.

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and micro-habitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having a geographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system.

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and microhabitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having ageographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system. Rev. Biol. Trop. 59 (3): 1419-1432. Epub 2011 September 01.

La hipótesis de heterogeneidad espacial se ha utilizado para explicar el aumento en la diversidad de especies desde los polos a los trópicos: los trópicos pueden ser más diversos ya que están conformados por una mayor cantidad de hábitats y micro-hábitats. En este estudio, la hipótesis de heterogeneidad espacial se puso a prueba evaluando la variación en la riqueza de dos gremios de insectos herbívoros (formadores de agallas y de alimentación libre) asociados con B. dracunculifolia (Asteracea) a lo largo de un gradiente latitudinal en Brasil. Las diecisiete poblaciones de B. dracunculifolia seleccionadas para el muestreo de insectos herbívoros estaban en hábitats con una estructura similar, a lo largo del límite Norte-Sur de distribución de la planta hospedera, cerca de la costa brasileña. De cada población de planta hospedera, se muestrearon treinta arbustos y se obtuvo un total de 8 201 agallas y 864 insectos de alimentación libre pertenecientes a 28 familias y 88 especies. La mayoría de los insectos que se encontraron en B. dracunculifolia estaban restringidos a un sitio específico en lugar de tener una distribución geográfica similar a la de la planta hospedera. La riqueza de especies de insectos de alimentación libre no se vió afectada por el gradiente latitudinal, por lo que se corroboró la hipótesis de heterogeneidad espacial. Mientras que la riqueza de especies de insectos formadores de agallas se correlacionó positivamente con la latitud, probablemente debido a que los insectos asociados al género Baccharis se extendieron hacia el sur de Brasil. Otros índices de diversidad y equidad estimados no variaron con la latitud para ninguno de los dos grupos de insectos herbívoros evaluados, lo que sugirie una estructura general para diferentes conjuntos de herbívoros asociados con la planta hospedera B. dracunculifolia. Por lo tanto, es probable que, la muestra de insectos en cada sitio sea resultado de eventos a gran escala, como la especiación, migración y coevolución; mientras que a nivel local la población de estos insectos está regulada por fuerzas ecológicas que operan dentro del sistema.

Plant architecture and meristem dynamics as the mechanisms determining the diversity of gall-inducing insects.

Plant architecture is considered to affect herbivory intensity, but it is one of the least studied factors in plant-insect interactions, especially for gall-inducing insects. This study aimed to investigate the influence of plant architecture on the speciose fauna of gall-inducing insects associated with 17 species of Baccharis. Five architectural variables were evaluated: plant height, number of fourth-level shoots, biomass, average level and number of ramifications. The number of galling species associated with each host plant species was also determined. To test the effects of plant architecture on gall richness at the individual level, we used another data set where the number of fourth-level shoots and gall richness were determined for B. concinna, B. dracunculifolia, and B. ramosissima every 3 weeks during 1 year. The average similarity between host species based on gall fauna was low (9%), but plants with the same architectural pattern tended to support similar gall communities. The most important architectural trait influencing gall richness at the species level was the number of fourth-level shoots, which is indicative of the availability of plant meristems, a fundamental tissue for gall induction and development. This variable also showed a positive correlation with gall richness at the individual level. We propose that variations in gall richness among host species are driven by interspecific differences in plant architecture via availability of young, undifferentiated tissue, which is genetically controlled by the strength of the apical dominance. Plant architecture should have evolutionary consequences for gall communities, promoting insect radiation among architecturally similar plants through host shift and sympatric speciation. We also discuss the role of plant architecture in the global biogeography of gall-inducing insects.

Effect of microenvironment on development of a gall midge.

This study assessed the influence of microenvironment on the establishment and relative reproductive success of the gall-forming midge Rhopalomyia californica Felt on its host plant Baccharis pilularis De Candolle in Marin County, CA. Mesh cages were used to alter the microenvironment, which also allowed us to assess the validity of using these types of experimental manipulations in this system. Temperature, light intensity, wind speed, and stem growth were compared in caged and uncaged B. pilularis plots in two sites during three seasons. Cage presence significantly altered the microenvironment of R. californica but did not affect development. R. californica establishment was greater when growing on host plants with increased stem growth. Season had the largest impact on gall establishment and reproductive success, with the highest establishment and success rates in late winter to early spring, which correlated with the growing period of B. pilularis. These results suggest that the seasonality of R. californica reproductive success is linked to the phenology of its host plant. When the growing conditions for the plant are less than ideal, R. californica performance is stimulated by increased stem growth. Cage presence was not a significant driver of population dynamics because it did not change the environment in an ecologically meaningful way. We therefore assert that the use of cages for experimental manipulations in this study system does not alter R. californica performance.

Assessing trophic interactions in a guild of primary parasitoids and facultative hyperparasitoids: stable isotope analysis.

Facultative hyperparasitism is likely to be the most common form of intraguild predation among parasitoids. However, difficulties associated with studying facultative hyperparasitoids in the field have hampered a thorough understanding of their trophic ecology. In this study, we used a combination of stable isotope analysis and published natural history information to infer trophic interactions in a guild of field-collected primary parasitoids and facultative hyperparasitoids that attack a gall-making midge on Baccharis pilularis. Our three a priori hypotheses were: (1) stable isotope values should increase incrementally from the host plant to higher trophic levels; (2) the two species of ectoparasitoids should exhibit higher stable isotope signatures than the two endoparasitoids, and; (3) the two facultative hyperparasitoids should exhibit stable isotope signatures that fall between zero and one trophic level steps above that observed for the primary parasitoids. Food webs inferred from stable isotope data generally agreed with previously published accounts of community structure. As expected, both delta(13)C and delta(15)N were progressively enriched in the heavy isotope from the host plant to the herbivorous midge to the parasitic wasps. Multivariate analysis of stable isotope data revealed that the two primary ectoparasitoids occupied a similar trophic niche, but were significantly different from the primary endoparasitoids. We attribute this result to "coincidental intraguild predation" by ectoparasitoids that develop on already-parasitized midge larvae. One of the facultative hyperparasitoids, Zatropis capitis, exhibited a stable isotope signature approximately one trophic step above the primary parasitoids. Unexpectedly, the second facultative hyperparasitoid, Mesopolobus sp., appeared to be developing as a primary parasitoid at all sites. Coupled with independent assessments of community structure, stable isotope analysis validated trophic links constructed by previous researchers and identified potential taxon-specific differences in trophic interactions for two facultative hyperparasitoids in the B. pilularis gall community.

Morphogenesis of galls induced by Baccharopelma dracunculifoliae (Hemiptera: Psyllidae) on Baccharis dracunculifolia (Asteraceae) leaves

Morfogênese das galhas induzidas por Baccharopelma dracunculifoliae (Hemiptera, Psyllidae) em folhas de Baccharis dracunculifolia (Asteraceae). A galha entomógena mais comum em folhas de Baccharis dracunculifolia (Asteraceae) é induzida por Baccharopelma dracunculifoliae (Hemiptera, Psyllidae). O inseto galhador ataca tanto folhas jovens não distendidas quanto as completamente distendidas. Quatro fases de desenvolvimento foram observadas nesse tipo de galha: 1) Fase de dobramento, na qual o limbo dobra-se ao longo da nervura central e devido à aproximação das margens, uma fenda é formada na porção superior da folha. Uma câmara única está presente na face adaxial da lâmina foliar; 2) Fase de intumescimento, em que a folha dobrada torna-se espessada e suas margens aproximam-se, fazendo com que a fenda se estreite. Nesta fase, a galha está madura, suculenta e fusiforme, apresentando coloração verde claro. A câmara ninfal abriga de 1 a vários indutores e é recoberta por cera branca; 3) Fase de deiscência, caracterizada pela abertura da fenda, liberando os indutores; e 4) Fase de senescência, quando torna-se escura e seca. O sistema de revestimento da galha madura é formado por epiderme uniestratificada. O mesofilo apresentou-se intumescido, devido principalmente à hiperplasia do tecido parenquimático. Na nervura central, os tecidos vasculares mostraram-se conspícuos e as fibras perivasculares com aspecto semelhante ao de células parenquimáticas. Nas cavidades secretoras hipertrofiadas, o conteúdo lipofílico era escasso. Nessa galha não se formou tecido nutritivo, mas foram vistas bainhas salivares deixadas pelos indutores junto ao parênquima, aos feixes vasculares e às cavidades secretoras. Os estudos feitos complementam as informações sobre a biologia da galha e contribuem para a compreensão da plasticidade dos tecidos vegetais estimulados por fatores bióticos.

Morphogenesis of galls induced by Baccharopelma dracunculifoliae (Hemiptera: Psyllidae) on Baccharis dracunculifolia (Asteraceae) leaves.

The commonest insect gall on Baccharis dracunculifolia (Asteraceae) leaves is induced by Baccharopelma dracunculifoliae (Hemiptera, Psyllidae). The gall-inducing insect attacks young leaves in both the unfolded and the fully expanded stages. Four developmental phases were observed in this type of gall: 1) A folding phase, during which the leaf lamina folded upward alongside the midrib and the edges of the upper portion of the leaf approached each other, forming a longitudinal slit. A single chamber was formed on the adaxial surface of the leaf; 2) A swelling phase, in which the folded leaf tissues thickened and the edges of the leaf drew closer together, narrowing the slit. In this phase the gall matured, turning succulent, fusiform and pale green. The single nymphal chamber was lined with white wax and was able to house from one to several nymphs; 3) A dehiscence phase, characterized by the opening of the slit to release inducers; and 4) A senescence phase, when the gall turned dark and dry. The dermal system of the mature gall was composed of a single-layered epidermis. The mesophyll was swollen, and the swelling was due mainly to hyperplasia of the parenchyma. The vascular tissues along the midrib vein were conspicuous and the perivascular fibers resembled parenchymal cells. The hypertrophied secretory cavities contained low lipophylic content. This gall does not form nutritive tissue, but salivary sheaths left by the inducers were observed near the parenchyma, vascular bundles and secretory cavities. This study complements our current knowledge of gall biology and sheds further light on the plasticity of plant tissues stimulated by biotic factors.