Phenolic substances and cyanogenesis in galled and non-galled tissue of the fern species Microgramma vacciniifolia / Substâncias fenólicas e cianogênese em tecidos galhados e não galhados da samambaia Microgramma vacciniifolia

Galls, neo-formed plant structures that can occur in different organs, are generated by species-specific interaction with an inducing organism. Inducers manipulate the metabolism of its host. Microgramma vacciniifolia (Langsd. & Fisch.) Copel. is a Neotropical epiphytic fern that hosted two stem galls, one induced by a midge species (Diptera) and other by a micromoth species (Lepidoptera). The aim of this study was to evaluate the impact of these two gall-inducing insects on the biochemistry of phenolic acids and the cyanogenesis in galls, stems and leaves of M. vacciniifolia. High performance liquid chromatography (HPLC) indicated a total of 14 phenol derivatives, including caffeic and coumaric acid. Principal Coordinates Analysis (PCoA) of the phenolic substances indicated three groups consisting (1) non-galled stems and micromoth-induced galls; (2) midge-induced galls; (3) midge induced galls with parasitoids. Regarding the frequency of cyanogenesis assessed by the picrate paper test, the chi-squared test showed significant difference between fertile leaves (8.3%), sterile leaves (27.7%), non-galled stems (0%) and galls. Among galls, only the midge-induced galls analyzed were cyanogenic (15%). Our results indicated that the different gall-inducers (midge and micromoth) promote species-specific alterations to the phenolic substance composition of the host fern.

Galhas são estruturas vegetais neo-formadas que ocorrem em diferentes órgãos. Elas são geradas por uma interação espécie-específica com um organismo indutor. Os indutores manipulam o metabolismo do hospedeiro. Microgramma vacciniifolia (Langsd. & Fisch.) Copel. é uma samambaia epífita neotropical que hospeda duas galhas caulinares, uma induzida por uma espécie de mosquito (Diptera) e outra por uma micromariposa (Lepidoptera). O objetivo deste estudo foi avaliar o impacto desses dois insetos indutores de galhas na bioquímica dos ácidos fenólicos e da cianogênese em galhas, caules e folhas de M. vacciniifolia. Em análise de cromatografia líquida de alta eficiência (CLAE) foi possível indicar a presença de um total de 14 derivados fenólicos, incluindo ácido cafeico e ácido cumárico. Análise das Coordenadas Principais (ACoP) indicou três grupos (1) caules não galhados e galhas induzidas pela micromariposa; (2) galhas induzidas pelo mosquito; (3) galhas induzidas pelo mosquito com parasitoides. Em relação a frequência da cianogênese analisada com o teste do papel picrato, o teste do qui-quadrado apresentou diferença significativa entre as folhas férteis (8,3%), folhas estéreis (27,7%), caules não galhados (0%) e galhas. Entre as galhas, somente aquelas induzidas pelo mosquito foram cianogênicas (15%). Os resultados encontrados indicam, ao menos para as substâncias fenólicas, que os insetos indutores de galha (mosquito e micromariposa) promovem alterações espécie-específica na composição química da samambaia hospedeira.

A Review of Galls on Ferns and Lycophytes.

Fern-insect interactions have not received the same attention as angiosperm-insect interactions have. It has even been stated that ferns may have very few interactions with animals because of their lack of flowers, fruits, and seeds. Consequently, for many decades fern-insect interactions have been overlooked and underestimated, especially for highly developed interactions such as those with gall-formers. The present work aims to review the galls of ferns and lycophytes worldwide, to provide an updated checklist including unpublished data and to estimate the global gall diversity of ferns and lycophytes. We recorded 93 host species, belonging to 41 genera. Galls were found in 20 fern families and one lycophyte family (Selaginellaceae). Most galls occur within the more derived ferns of the order Polypodiales, especially the fern families Polypodiaceae (21 host species), Dryopteridaceae (14 host species) and Athyriaceae (11 host species). Thirty-eight of the 133 gall morphotypes were induced by mites and 95 by insects of six orders (Coleoptera, Diptera, Hymenoptera, Lepidoptera, Thysanoptera, and Hemiptera). Among the insects, Cecidomyiidae (Diptera) caused most of the galls (35 morphotypes). So far, most galls have been reported from the Neotropical region (40 spp.) and Oriental region (28 spp.).

Glycoside Hydrolase (GH) 45 and 5 Candidate Cellulases in Aphelenchoides besseyi Isolated from Bird's-Nest Fern.

Five Aphelenchoides besseyi isolates collected from bird's-nest ferns or rice possess different parasitic capacities in bird's-nest fern. Two different glycoside hydrolase (GH) 45 genes were identified in the fern isolates, and only one was found in the rice isolates. A Abe GH5-1 gene containing an SCP-like family domain was found only in the fern isolates. Abe GH5-1 gene has five introns suggesting a eukaryotic origin. A maximum likelihood phylogeny revealed that Abe GH5-1 is part of the nematode monophyletic group that can be clearly distinguished from those of other eukaryotic and bacterial GH5 sequences with high bootstrap support values. The fern A. besseyi isolates were the first parasitic plant nematode found to possess both GH5 and GH45 genes. Surveying the genome of the five A. besseyi isolates by Southern blotting using an 834 bp probe targeting the GH5 domain suggests the presence of at least two copies in the fern-origin isolates but none in the rice-origin isolates. The in situ hybridization shows that the Abe GH5-1 gene is expressed in the nematode ovary and testis. Our study provides insights into the diversity of GH in isolates of plant parasitic nematodes of different host origins.

Tri-trophic effects of seasonally variable induced plant defenses vary across the development of a shelter building moth larva and its parasitoid.

Plant chemical defenses can negatively affect insect herbivore fitness, but they can also decrease herbivore palatability to predators or decrease parasitoid fitness, potentially changing selective pressures on both plant investment in production of chemical defenses and host feeding behavior. Larvae of the fern moth Herpetogramma theseusalis live in and feed upon leaf shelters of their own construction, and their most abundant parasitoid Alabagrus texanus oviposits in early instar larvae, where parasitoid larvae lay dormant for most of host development before rapidly developing and emerging just prior to host pupation. As such, both might be expected to live in a relatively constant chemical environment. Instead, we find that a correlated set of phenolic compounds shows strong seasonal variation both within shelters and in undamaged fern tissue, and the relative level of these compounds in these two different fern tissue types switches across the summer. Using experimental feeding treatments, in which we exposed fern moth larvae to different chemical trajectories across their development, we show that exposure to this set of phenolic compounds reduces the survival of larvae in early development. However, exposure to this set of compounds just before the beginning of explosive parasitoid growth increased parasitoid survival. Exposure during the period of rapid parasitoid growth and feeding decreased parasitoid survival. These results highlight the spatial and temporal complexity of leaf shelter chemistry, and demonstrate the developmental contingency of associated effects on both host and parasitoid, implying the existence of complex selective pressures on plant investment in chemical defenses, host feeding behavior, and parasitoid life history.

A new genus and species of myrmecophilous brentid beetle (Coleoptera: Brentidae) inhabiting the myrmecophytic epiphytes in the Bornean rainforest canopy.

Pycnotarsobrentus inuiae Maruyama & Bartolozzi, gen. nov. and sp. nov. (Brentinae: Eremoxenini) is described from the Lambir Hills National Park, Borneo (Sarawak, Malaysia) based on specimens collected from Crematogaster difformis F. Smith, 1857 ant nests in the myrmecophytic epiphytic ferns Platycerium crustacea Copel. and Lecanopteris ridleyi H. Christ. A second species of Pycnotarsobrentus is known from Malaysia but is represented by only one female and consequently not yet described pending discovery of a male. Pycnotarsobrentus belongs to the tribe Eremoxenini and shares some character states with the African genus Pericordus Kolbe, 1883. No species of Eremoxenini with similar morphological modifications are known from the Oriental region.

Sexual compatibility among different host-originated isolates of Aphelenchoides besseyi and the inheritance of the parasitism.

Nine isolates of Aphelenchoides besseyi from two different hosts were studied. The isolates were identified at the species level according to morphometrics and fine structures observed under a scanning electron microscope. Two fern-originated isolates, Fu, and Fm, one rice-originated isolate, Rl, were not able to reproduce from a single juvenile, based on at least 50 replicates. The other six isolates were able to develop into a small population when inoculated with a single juvenile, demonstrating parthenogenesis. Crosses between isolates were conducted. In a compatibility cross experiment, three fern-originated isolates were selfed and crossed reciprocally, and all nine crossings had viable offspring. When fern isolates were used as paternal lines, the only two successful crosses were with the Rd line, and as maternal lines, only the Ff x Re and Fu x Rn crosses had viable offspring. Rl was used as the maternal line and Fm as the paternal line to study the inheritance of the bird's-nest fern parasitism. Twenty of the 80 attempted crosses resulted in viable offspring and among these; six lines had the ability to parasitize on the bird's-nest fern. When the F(1) lines were back-crossed to the Rl maternal line, 20 viable offspring lines were obtained and among them 4 were able to parasitize bird's-nest fern. These results indicate that bird's-nest fern parasitism can be transferred to new generations by cross fertilization.

Soral crypsis: protective mimicry of a coccid on an Indian fern.

Herbivory with crypsis is not well documented in ferns. The present record of cryptic coloration of coccid Saissetia filicum Boisduval (Homoptera: Coccidae) to the sori of a fern species Asplenium nidus L. (Aspleniaceae) is unique. Predatory beetles (Jauravia sp., Coleoptera: Coccinellidae) that feed on the coccids, are suggested to be selective pressure for the development of the present homopteran soral crypsis. A higher rate of effective predation is noticed in the vegetative leaves than the fertile leaves. Aggressive ants were found harvesting honeydew secretions from the coccids and defending the trophobionts as well as the host fern from their natural enemies. In addition, a possible three-way mutualistic relationship among the coccids, its host fern and the tending ant is suggested. Differential numbers of coccids on vegetative and fertile leaves is correlated with their phenol content and degree of predation by beetles. Such coloration mimicry by the coccids may enable them to obtain the necessary blend of sorus of the host fern needed to evade beetle detection and attack.

Life history and systematics of Albusambia elaphoglossumae (Lepidoptera:Crambidae): A new genus and species of musotimine with leaf-mining biology from Costa Rica

Albusambia elaphoglossumae Solis & Davis, a new genus and species,is described.It was discov- ered mining the fronds of the fern Elaphoglossum conspersum in Costa Rica (San José and Cartago Provinces, at elevations of 2300-3100 m).The type series was obtained by rearing of the immature stages in laboratory. The adult is defined by unique genital characters,and the pupa with a medial depression on the vertex and with two anterolateral horn-like structures on the prothorax. The larva is a gregarious leaf miner with its body flat- tened dorsoventrally and head prognathous; morphological adaptations to its leaf-mining habit are new to the Musotiminae.Fern-feeding musotimines are important to the discovery of new biological control agents for invasive ferns

Se describen un género y especie nuevos de Musotiminae,Albusambia elaphoglossumae Solis and Davis,de Costa Rica.Esta especie mina las frondas del helecho Elaphoglossum conspersum .La serie tipo se obtuvo mediante la recolección de las minas y mediante crianza en laboratorio.El adulto se define por caracteres específicos en el aparato genital;la pupa se caracteriza por una depresión media en el vértex y por dos estructuras anterolaterales en forma de cuernos en el protórax.La larva es un minador gregario de la fronda,con su cuerpo aplanado dorsoventralmente y su cabeza prognata;ambas adaptaciones morfológicas para minar frondas (vistas por primera vez en Musotiminae). Actualmente,Musotiminae es de alta prioridad en la búsqueda de nuevos agentes para el control biológico de helechos invasores

Biology and phenology of the eriophyid mite, Floracarus perrepae, on its native host in Australia, Old World climbing fern, Lygodium microphyllum.

The biology and phenology of the eriophyid mite, Floracarus perrepae Knihinicki and Boczek, a potential biological control agent of Lygoditon microphyllun (Cav.) R. Br., was studied in its native range--Queensland, Australia. F. perrepae forms leaf roll galls on the subpinnae of L. mnicrophyllum. It has a simple biology, with females and males produced throughout the year. The population was female biased at 10.5 to 1. The immature development time was 8.9 +/- 0.1 and 7.0 +/- 0.1 days; adult longevity was 30.6 +/- 1.6 and 19.4 +/- 1.2 days and mean fecundity per female was 54.5 +/- 3.2 and 38.5 +/- 1.6 eggs at 21 and 26 degrees C, all respectively. Field studies showed that the mite was active year round, with populations peaking when temperatures were cool and soil moisture levels were highest. Two species of predatory mites, Tarsonemus sp. and a species of Tydeidae, along with the pathogen Hirsutella thompsonii, had significant effects on all life stages of F. perrepae. Despite high levels of predators and the pathogen, F. perrepae caused consistent damage to L. microphyllum at all the field sites over the entire 2 years of the study.

Host plant selection of two Mansonia blanchard species (Diptera: Culicidae) in a heterogeneous habitat of Buenos Aires City, Argentina.

Larvae and pupae of the genus Mansonia Blanchard attach to the roots of aquatic plants by means of modified structures to obtain oxygen. A study of the association of larval Ma. indubitans and Ma. titillans with floating macrophytes was conducted at Macies Pond, Argentina. Fifty-four sampling units were taken from January to May 2003. Three genera of host plants were considered: Pistia, Limnobium, and Salvinia. A total of 402 immatures of Ma. indubitans and 217 of Ma. titillans were captured and associations between Mansonia immatures and roots of each genera were assesed. Significant association was noted between Ma. indubitans and certain host plant species (K-W H=42.74, df=2, p<0.001). The same result was observed for Ma. titillans (K-W H=23.42, df=2, p<0.001). Both Mansonia species utilized roots of P. stratiotes in significantly higher proportions than expected by random selection. Both species showed significant negative association with Salvinia spp., while no clear relationships were detected with L. laevigatum.

Life history and systematics of Albusambia elaphoglossumae (Lepidoptera: Crambidae): a new genus and species of musotimine with leaf-mining biology from Costa Rica.

Albusambia elaphoglossumae Solis & Davis, a new genus and species, is described. It was discovered mining the fronds of the fern Elaphoglossum conspersum in Costa Rica (San José and Cartago Provinces, at elevations of 2300-3100 m). The type series was obtained by rearing of the immature stages in laboratory. The adult is defined by unique genital characters, and the pupa with a medial depression on the vertex and with two anterolateral horn-like structures on the prothorax. The larva is a gregarious leaf miner with its body flattened dorsoventrally and head prognathous; morphological adaptations to its leaf-mining habit are new to the Musotiminae. Fern-feeding musotimines are important to the discovery of new biological control agents for invasive ferns.

Doubling the estimate of invertebrate biomass in a rainforest canopy.

Forest canopies represent the functional interface between 90% of the Earth's terrestrial biomass and the atmosphere and include some of the most threatened of all terrestrial ecosystems. However, we lack even a basic understanding of how the biomass of plants and animals is distributed throughout forest canopies, even though this information is vital for estimating energy flow, carbon cycling, resource use and the transfer of materials within this ecosystem. Here we measure the biomass of invertebrates living in a common rainforest epiphyte, describe a striking relationship between fern size and the biomass of animals within the ferns, and reveal that one large epiphyte may contain an invertebrate biomass similar to that found in the whole of the rest of the tree crown on which it is growing. Using these data, we show that including the fauna of these epiphytes--a neglected component in rainforest ecosystems--can more than double our estimate of the total invertebrate biomass in an entire rainforest canopy.