Helena C. Morais

Host specificity of Lepidoptera in tropical and temperate forests

For numerous taxa, species richness is much higher in tropical than in temperate zone habitats. A major challenge in community ecology and evolutionary biogeography is to reveal the mechanisms underlying these differences. For herbivorous insects, one such mechanism leading to an increased number of species in a given locale could be increased ecological specialization, resulting in a greater proportion of insect species occupying narrow niches within a community. We tested this hypothesis by comparing host specialization in larval Lepidoptera (moths and butterflies) at eight different New World forest sites ranging in latitude from 15° S to 55° N. Here we show that larval diets of tropical Lepidoptera are more specialized than those of their temperate forest counterparts: tropical species on average feed on fewer plant species, genera and families than do temperate caterpillars. This result holds true whether calculated per lepidopteran family or for a caterpillar assemblage as a whole. As a result, there is greater turnover in caterpillar species composition (greater β diversity) between tree species in tropical faunas than in temperate faunas. We suggest that greater specialization in tropical faunas is the result of differences in trophic interactions; for example, there are more distinct plant secondary chemical profiles from one tree species to the next in tropical forests than in temperate forests as well as more diverse and chronic pressures from natural enemy communities.

The global distribution of diet breadth in insect herbivores

Significance Dietary specialization determines an organism’s resource base as well as impacts on host or prey species. There are important basic and applied reasons to ask why some animals have narrow diets and others are more generalized, and if different regions of the Earth support more specialized interactions. We investigated site-specific host records for more than 7,500 species of insect herbivores. Although host specialists predominate, the proportion of specialists is affected by the diversity of hosts and shifts globally, supporting predictions of more exclusive tropical interactions. These results not only affect our understanding of the ecology of food webs, but also have implications for how they respond to environmental change, as well as for ecosystem management and restoration. Understanding variation in resource specialization is important for progress on issues that include coevolution, community assembly, ecosystem processes, and the latitudinal gradient of species richness. Herbivorous insects are useful models for studying resource specialization, and the interaction between plants and herbivorous insects is one of the most common and consequential ecological associations on the planet. However, uncertainty persists regarding fundamental features of herbivore diet breadth, including its relationship to latitude and plant species richness. Here, we use a global dataset to investigate host range for over 7,500 insect herbivore species covering a wide taxonomic breadth and interacting with more than 2,000 species of plants in 165 families. We ask whether relatively specialized and generalized herbivores represent a dichotomy rather than a continuum from few to many host families and species attacked and whether diet breadth changes with increasing plant species richness toward the tropics. Across geographic regions and taxonomic subsets of the data, we find that the distribution of diet breadth is fit well by a discrete, truncated Pareto power law characterized by the predominance of specialized herbivores and a long, thin tail of more generalized species. Both the taxonomic and phylogenetic distributions of diet breadth shift globally with latitude, consistent with a higher frequency of specialized insects in tropical regions. We also find that more diverse lineages of plants support assemblages of relatively more specialized herbivores and that the global distribution of plant diversity contributes to but does not fully explain the latitudinal gradient in insect herbivore specialization.

The Abundance of Insect Herbivore Species in the Tropics: The High Local Richness of Rare Species

Local sampling of larval lepidopterans on Erythroxylum host plant species in tropical savanna (cerrado) revealed a high species richness with low abundance per species. Cumulative numbers of morphospecies with increasing sampling effort yielded no asymptotic level of richness in sampling periods lasting 6 mo, 7 mo, and 23 mo. Peak richness was reached at 31 species in 1992 and 19 species in 1993, on the three Erythroxylum species sampled: E. deciduum, E. suberosum, and E. tortuosum. Less than one larva was found per plant during all sampling times, with a mean of 0.28 species per plant in 1992 and 0.12 in 1993. The number of specimens of all species combined discovered per plant was very low at 0.10 per plant. Many plants remained unattacked on any sampling date with 12 percent, 8 percent, and 16 percent of plants attacked in the species listed in order above. In general, there was an increase in species found during the late dry season when new leaves were produced, but plant phenology seemed to exert only a small influence. Comparisons with temperate samples of a similar kind, in savanna vegetation at the same altitude, indicate a very different assemblage. Comparison was based on four criteria: richness was from two to over three times higher in the tropics, even though sampling had not produced an asymptotic accumulation of species; the number of morphospecies per plant individual was similar at the sites, although total richness was lower in the temperate savanna; the number of total individuals per plant was 11-fold higher in the temperate samples; and the percent of plants with larvae present was over four times higher in the temperate zone (mean of 49%) than in the cerrado (12%). The high richness of relatively rare species in the cerrado site poses challenges in understanding the reasons for such rarity, the organization of such assemblages, the gradient of species richness from low to high latitudes, the estimation of biodiversity, and conservation management.

Patterns and correlates of interspecific variation in foliar insect herbivory and pathogen attack in Brazilian cerrado

Patterns of insect herbivore and leaf pathogen attack are described for 25 plant species (10 trees, 10 shrubs and five herbs) at a Brazilian savanna (cerrado) site. Plant and leaf traits were correlated with interspecific variation in attack by herbivores and pathogens in order to account for differences among plant species. Across all species, pathogen damage was 1.5 times higher than insect damage (17.3% vs. 6.8%, respectively). Most insect damage occurred to young leaves while they were expanding (end of the dry season). In contrast, pathogen attack was low on young expanding leaves at the end of the dry season, increased as those leaves matured in the wet season, but continued to increase through the next dry season. Protein-binding capacity was negatively associated with interspecific differences in insect damage to mature leaves. Protein availability and plant height were positive predictors of pathogen attack among plant species, while leaf expansion rate was a significant negative predictor. Interspecific differences in leaf phenology had little effect on the amount of damage caused by either insects or pathogens. However, new leaves produced during the wet season suffered less insect damage than leaves produced during the dry season, the time of greatest leaf production. Timing of young leaf production affected pathogen attack but the season of escape depended on plant species. In contrast, there was no evidence for escape in space as common species were less likely to suffer high pathogen attack than rare species. New and mature leaf toughness, and time for a leaf to reach full expansion all increased from herbs to shrub to trees, while mature leaf nitrogen decreased in that order.

Lepidopteran caterpillar fauna of cerrado host plants

The cerrado biome is rich in caterpillar species although the general biology and ecology of most tropical species remains unknown. Three host plant genera (Byrsonima, Erythroxylum and Qualea) were examined for caterpillars in four cerrado sensu stricto areas near Brasi´lia (DF, Brazil), from 1991 to 1995. Altogether, more than 16000 host plants were sampled and less than 20% of them possessed caterpillars. All the caterpillars found were reared under laboratory conditions. We successfully reared 137 species of 24 lepidopteran families. The average number of caterpillar species per host species was 28.3. The faunal similarity among plant genera, as well as among study sites, was low, and not related to the distances between them. The low proportion of host plants with caterpillars and the high incidence of rare species appears to be a general pattern for the cerrado. A large proportion (74%) of the caterpillar species occurred on only one host plant family. The most frequent species were either specialists (restricted to one host plant genus) or generalists. We provide a brief outline of some taxonomic problems, some observations on general biology, and temporal and local patterns of some specific caterpillars species. The methodology used, which included weekly data collection, in restricted study areas of the same habitat, with the same sampling method for recording the caterpillar species on the same host plant species, can be used as a tool to explore biodiversity and to discuss some aspects of the conservation of cerrado insects.

Effect of plant diversification on abundance of South American tomato pinworm and predators in two cropping systems

Several studies have demonstrated that the diversification of the agroecosystem may reduce insect pest infestation using the strategy of companion planting. We studied the effect of the presence of coriander and Gallant soldier as companion plants in the tomato crop under organic and conventional management on abundance of the tomato pests and predators. The experiment was carried out under field conditions, at Distrito Federal, Brazil, from July to October, 2004. The experimental design was of randomized blocks with three replicates in a 3 x 2 factorial arrangement [Three crop diversification levels: tomato in monoculture, tomato intercropped with coriander, and tomato intercropped with coriander and Gallant soldier; and two cropping systems: organic and conventional]. Each plot had 80 plants of the tomato and the pest-insects and predaceous arthropods were collected every week in five tomato plants/plot. Tuta absoluta (Meyr.) was the commonest pest species in both cropping systems. The tomato intercropped with coriander and Gallant soldier in organic cropping system had lower abundance and higher diversity of pests, and higher abundance and diversity of predaceous arthropods, mainly at 80 days after the tomato transplanting, coinciding with the population peak of T. absoluta (eggs and adults). The more abundant predators were spiders, lady beetles and ants. The spiders were abundant and uniformly distributed in both systems. The lady beetles appeared lately in the organic system, at the flowering season of the coriander. The ants were more abundant in the conventional systems.

Fatores de mortalidade na fase de ovo de Tuta absoluta em sistemas de produção orgânica e convencional de tomate

A traca-do-tomateiro Tuta absoluta (Meyrick) (Lep.: Gelechiidae) esta sujeita a fatores de mortalidade bioticos como predadores e parasitoides e fatores abioticos como agua da irrigacao e da chuva que influenciam sua dinâmica populacional. O objetivo deste trabalho foi quantificar a importância desses fatores na sobrevivencia de ovos da traca-do-tomateiro. Os experimentos foram desenvolvidos na Embrapa Hortalicas, de 1/8/2004 a 7/11/2005, em plantio de tomate e tomate consorciado em sistema orgânico e tomate em sistema convencional. As avaliacoes consistiram em: 1) coleta de ovos no campo em diferentes epocas do ciclo da cultura e sua incubacao em laboratorio ate a emergencia do parasitoide ou eclosao da lagarta; 2) exposicao de ovos sentinelas no campo por 72 horas; 3) ovos marcados e acompanhados a cada 24 horas ate eclosao das lagartas em campo. Ovos coletados no campo e incubados em laboratorio mostraram que a mortalidade por Trichogramma sp. foi de 32%. A mortalidade dos ovos que permaneceram no campo (metodologia 2 e 3), evidenciaram que o parasitismo por Trichogramma sp. foi de 12% e que os predadores e o efeito mecânico da agua produziram 36% de mortalidade dos ovos. Essa diferenca revela que a permanencia dos ovos no campo os expoe a competicao com predadores e a mortalidade causada pelo efeito mecânico da agua reduzindo o recrutamento de Trichogramma sp. Parasitoides de ovos, predadores generalistas associados a acao da agua acarretaram maior mortalidade de T. absoluta em sistemas de producao orgânica de tomate.

Abundance and temporal distribution of Gonioterma exquisita Duckworth (Lepidoptera, Elachistidae, Stenomatinae) on Byrsonima pachyphylla Griseb. (Malpighiaceae) in the Brazilian Cerrado

Abundance and temporal distribution of Gonioterma exquisita Duckworth (Lepidoptera, Elachistidae, Stenomatinae) on Byrsonima pachyphylla Griseb. (Malpighiaceae) in the Brazilian Cerrado. The seasonal characteristics of the Cerrado region strongly influence food resource predictability and vegetal tissue nutritional content. The aims of this work were to record the abundance and temporal distribution of Gonioterma exquisita Duckworth, 1964 (Lepidoptera, Elachistidae) and its relation with phenological, physical, and chemical traits of the host plant Byrsonima pachyphylla Griseb. (Malpighiaceae). Four nutritional quality parameters were determined for new and mature leaves: gross protein and nitrogen content, dry matter, and in vitro digestibility. We inspected 200 plants per month, searching for G. exquisita caterpillars. About 35.8% of the 2,400 plants inspected presented caterpillars, with an abundance peak in the wet season. Caterpillar abundance was positively correlated with mature leaf availability, their food resource. Although mature leaves presented lower gross protein and nitrogen contents than new leaves, this difference was small during the abundance peak of G. exquisita.

Lepidópteros folívoros em Roupala montana Aubl. (Proteaceae) no Cerrado Sensu Stricto

Richness of lepidopteran species in the cerrado is high. The confirmation of this richness is based on adult records but knowledge of larvae-host plants interactions is still incipient. The main purpose of this work was to identify folivorous caterpillar species on Roupala montana Aubl. in areas of a Brazilian cerrado sensu stricto, their dietary range, and the relative age of the consumed leaves richness. Three data sets, collected from 1991 to 2004 were used in this study. All caterpillars found on the host plants in the field were collected and reared in the laboratory. Sixty-two lepidopteran species from 22 families were recorded; they were found feeding on new (8%) and mature (92%) leaves of R. montana. Cerconota sciaphilina (Zeller) (Elachistidae), two unidentified Gracillariidae species and, one species of Gelechiidae fed only on new leaves. These species showed a short life cycle, which was very syncronized with the host plant foliar phenology. Among the species feeding on mature leaves, 36 were considered polyphagous and, these showed a large variation in the use of the food resource. Twenty-six species were found only on R. montana, 58% were very rare, a character that difficults the analysis of their dietary range. The monophagous species on R. montana were: Chlamydastisplatyspora (Meyrick) (Elachistidae), Stenoma cathosiota Meyrick (Elachistidae), Idalus prop. sublineata (Rothschild) (Arctiidae), and Eomichla sp. (Oecophoridae).

Larval Biology of Anthophagous Eumaeini (Lepidoptera: Lycaenidae, Theclinae) in the Cerrado of Central Brazil

ABSTRACT. The biology and morphology of the early stages of 22 species of Eumaeini (Lepidoptera: Lycaenidae, Theclinae) are presented. Observations were collected through the inspection of inflorescences in the field and the rearing of 214 larvae in laboratory. Allosmaitia strophius (Godart) associated with Malpighiaceae species and the polyphagous Strymon mulucha (Hewitson) were the most frequently collected species. Detritivory was observed in two species, Electrostrymon endymion (F.) and Kisutam syllis (Godman & Salvin), and myrmecophily in four other species, A. strophius, Ministrymon azia (Hewitson), Parrhasius polibetes (Stoll), and S. mulucha. Cannibalism was observed in A. strophius; in addition, the pupa of this and of three other species produced audible sounds. Paiwarria aphaca (Hewitson) was highlighted because of the great difference observed between its first and last instars, as well as the marked difference between that species and the larvae of Paiwarria umbratus (Geyer) documented in Costa Rica. Larvae of Calycopis mimas (Godman & Salvin) displayed “bungee jumping” behavior when stimulated. Parasitoids (Diptera, Hymenoptera) attacked 21 larvae of eight species, A. strophius, K. syllis, M. azia, Pai. aphaca, P. polibetes, Rekoa marius (Lucas), S. mulucha, and Tmolus venustus (H.H. Druce). Illustrations of immatures and parasitoids are provided.