Ants associated with fronds of the tropical bracken fern Pteridium esculentum subsp. arachnoideum / Formigas associadas às frondes da samambaia Pteridium esculentum subsp. arachnoideum

Abstract: Pteridium is a cosmopolitan genus of ferns that possess nectaries on its fronds (fern leaves), thereby attracting ants. Foliar (or extrafloral) nectaries are nectar-producing glands that are not related to pollination, but rather attract ants and other arthropods. Foliar nectaries are found in 101 fern species, belonging to 11 genera and six families. The aim of the study is to characterize the community of ants that visit the fronds of Pteridium esculentum subsp. arachnoideum, as well as daily and seasonal ant abundance in different frond development stages. The study was conducted in the Atlantic Forest of Rio de Janeiro state, Brazil. Bimonthly collections were established, where 30 expanding fronds and 30 fully expanded fronds were randomly marked. In each 1-hour shift starting at 8:30 am and ending at 5:30 pm, the fronds were observed for the presence of ants. Thirty three ant species were recorded on the Pteridium esculentum subsp. arachnoideum fronds, distributed into six subfamilies and 13 genera. The most abundant species were Solenopsis sp.1 and Ectatomma tuberculatum. Eight ant species were observed foraging the nectaries of tropical bracken fern fronds. Ectatomma tuberculatum has been observed feeding on the nectaries and patrolling the fronds. Ant activity peak was on mid-day during the rainy season. The tropical bracken fern Pteridium esculentum subsp. arachnoideum has a rich (the highest recorded until now on Pteridium species) and diverse ant community on its fronds, mainly on the expanding fronds. The presence of generalist predatory ants (Ectatomma tuberculatum and Solenopsis sp.1) during the entire study period suggests a positive interaction between ants and Pteridium esculentum subsp. arachnoideum.

Resumo: Pteridium é um gênero cosmopolita de samambaias que possui nectários em suas frondes (folhas de samambaias), atraindo formigas. Nectários foliares (ou extraflorais) são glândulas produtoras de néctar que não estão relacionadas com a polinização, mas podem atrair formigas e outros artrópodes. Nectários foliares já foram registrados em 101 espécies de samambaias, pertencentes a 11 gêneros e seis famílias. O objetivo do estudo é caracterizar a comunidade de formigas que visitam as frondes de Pteridium esculentum subsp. arachnoideum, bem como a abundância diária e sazonal das formigas em diferentes estágios foliares. O estudo foi realizado na Mata Atlântica do estado do Rio de Janeiro, Brasil. Foram estabelecidas coletas bimestrais, onde foram marcadas aleatoriamente 30 frondes em expansão e 30 frondes totalmente expandidas. Em cada turno de 1 hora com início às 8h30 e término às 17h30, as frondes marcadas foram observadas quanto à presença de formigas. Trinta e três espécies de formigas foram registradas nas frondes do Pteridium esculentum subsp. arachnoideum, distribuídas em seis subfamílias e 13 gêneros. As espécies mais abundantes foram Solenopsis sp.1 e Ectatomma tuberculatum. Foram registradas oito espécies de formigas forrageando os nectários foliares da samambaia. Ectatomma tuberculatum foi observada se alimentando nos nectários e patrulhando as frondes. As formigas tiveram o pico de atividade ao meio-dia e na estação chuvosa. A samambaia tropical Pteridium esculentum subsp. arachnoideum tem uma elevada riqueza (a maior já registrada até o momento para espécies de Pteridium) e diversidade de formigas em suas frondes, com maior frequência nas frondes não totalmente expandidas. A presença de formigas predadoras generalistas (Ectatomma tuberculatum e Solenopsis sp.1) durante todo o período de estudo sugere uma interação positiva entre as formigas e Pteridium esculentum subsp. arachnoideum.

True bugs living on ant-defended acacias: evasion strategies and ant species preferences, in Costa Rica and Panama

Introduction: Herbivores specialized in consuming ant-defended plants evolve strategies to prevent the attack of ant workers. When the plant can associate with more than one ant species, the herbivore evasion strategies may either be species-specific, or flexible enough to successfully deter workers of different ant-plant species. Objectives: We studied the behavior of an herbivore bug (Piezogaster reclusus) on ant-defended acacia trees (Vachellia collinsii), which associates with one of three mutualistic Pseudomyrmex ant species, and report the geographical distribution of the acacia bug species of Costa Rica and Panama. Methods: We tested whether herbivore bugs (1) associate with a particular ant species; (2) use chemical or behavioral strategies to evade the ant workers; (3) adjust the evasion strategy to the ant species living on the acacia tree. We also compared collected acacia bugs with Museum specimens to clarify the identification from Costa Rica and Panama. Results: We found bugs more often on trees with ants, particularly Ps. spinicola, and never on trees with Ps. nigrocinctus. To avoid ant attacks, bugs use evasive behaviors to prevent encounters with the ant workers, that depended on the ant species. Also, indirect evidence of intra and interspecific transfer experiments suggest species-specific chemical camouflage or repellence. We also report an expansion of the Southern limit of Pi. reclusus distribution in Central Panama, and reduced the distribution of Pi. chontalesis to the Chiriquí region. Conclusions: Similar to herbivores specialized on chemically defended plants, herbivores on ant-defended trees could evolve specific mechanisms to deal with the plant defenses. However, plants associated with multiple partners are a challenge to herbivore specialization, and might require behavioral plasticity, as our evidence suggests.

Introducción: Herbívoros especializados en consumir plantas defendidas por hormigas evolucionaron estrategias para prevenir el ataque de las obreras, que pueden ser específicas o flexibles para repeler obreras de diferentes especies. Objetivos: Estudiamos el comportamiento del chinche herbívoro (Piezogaster reclusus), que consume la savia de árboles de acacia, que se pueden asociar con una de tres especies de hormigas mutualistas del género Pseudomyrmex y reportamos el rango geográfico de las especies de chinches de acacias en Costa Rica y Panamá. Métodos: Evaluamos si los chinches herbívoros (1) se asocian preferiblemente con una especie de hormiga; (2) usan estrategias de comportamiento para evadir a las obreras; (3) ajustan su estrategia a la especie de hormiga residente en la acacia. También, comparamos especímenes de los chinches con especímenes de museos, para clarificar la identificación en Costa Rica y Panamá. Resultados: Los chinches fueronmás frecuentes en árboles con hormigas, especialmente Ps. spinicola, y nunca estuvieron en árboles con Ps. nigrocinctus. Los chinches mostraron diferentes comportamientos evasivos dependiendo de la especie de hormiga para prevenir encuentros con las obreras. También, evidencia indirecta de experimentos de transferencia sugiere que hay camuflaje especie-específico o repelencia. Además, reportamos que el límite de distribución de Pi. reclusus llega al centro de Panamá, mientras que Pi. chontalensis solamente está en la región de Chiriquí. Conclusiones: Igual que los herbívoros se especializan en plantas con defensas químicas, herbívoros en plantas con hormigas pueden evolucionar mecanismos específicos para lidiar con las defensas de las plantas. Sin embargo, plantas que se asocian a múltiples especies de hormigas se vuelven un reto para la especialización del herbívoro, y pueden requerir plasticidad de comportamiento como sugieren nuestros datos.

Are ants attracted to herbivorized leaves of Caryocar brasiliense Camb. (Caryocaraceae)? / As formigas são atraídas por folhas herbivoradas de Caryocar brasiliense Camb. (Caryocaraceae)?

Abstract Different profiles of secondary compounds are released by plants after herbivore attack. Many of these compounds are used by predators and parasitoids to locate herbivores that are damaging leaves. Such an induced indirect defense was tested with the Caryocar brasiliense-ant system in the dry season, when C. brasiliense has old leaves, and in the rainy season, when C. brasiliense has new leaves. A total of 20 plants were analyzed per season. Two opposite leaves of the same branch were selected for each plant. Approximately 40% of the area of one leaf was removed (treatment leaf) while the other leaf remained intact (control). The number of ants that visited each leaf was counted simultaneously for a period of 15 minutes and the mean difference in ant number was tested by paired t-test. The mean number of ants differed significantly between treatment and control only in the rainy season (t = 3.004, df = 19, p = 0.007). This finding suggests the presence of induced defense in this system only when the leaves are young. The study supports the Optimal Defense Theory since young leaves of C. brasiliense with artificial damage attracted significantly more ants than leaves without damage and represents the first evidence of an induced defense mechanism in the C. brasiliense-ant system.

Resumo Diferentes compostos químicos são liberados pelas plantas após o ataque dos herbívoros. Muitos desses compostos são usados por predadores e parasitoides para localizar os herbívoros que estão injuriando as plantas. Esse tipo de defesa induzida indireta foi testada no sistema Caryocar brasiliense e formigas em duas estações: seca (quando C. brasiliense está com folhas maduras) e chuvosa (quando C. brasiliense está com folhas jovens). Nós analisamos 20 plantas por estação do ano. Em cada planta nós selecionamos duas folhas opostas de um mesmo ramo. Nós removemos cerca de 40% da área de uma das folhas, deixando a outra folha intacta. Nós mensuramos simultaneamente o número de formigas que visitaram cada tipo de folha por 15 min e analisamos a diferença entre o número de formigas em cada tipo de folha através de teste t pareado. Nós observamos diferença significativa no número de formigas que patrulham as folhas tratamento (com herbivoria artificial) e controle (sem hervivoria artificial) apenas na estação chuvosa (t = 3,004, df = 19, p = 0,007). Isso sugere que existe defesa induzida nesse sistema somente quando as folhas são jovens. Nosso estudo corrobora a Teoria de Defesa Ótima já que apenas as folhas jovens de C. brasiliense com dano artificial atraíram significativamente mais formigas do que as folhas sem danos. Esta é a primeira vez que mecanismos de defesa induzida são observados no sistema C. brasiliense-formigas.

Ant-diaspore interactions during secondary succession in the Atlantic forest of Brazil

Animal-plant interactions are important for the recovery of diversity and processes in secondary forests, which increasingly dominate the tropical landscape. We used a combination of observational and experimental approaches to study the interactions of ants with diaspores across a successional gradient of forests in Southern Brazil, from August 2007 to April 2008. In addition to diaspore removal rates, we assessed the species richness, diversity and behaviour of ants interacting with diaspores, in three replicated sites of four successional stages of forests. We recorded 22 ant species interacting with diaspores (an estimated 15% of the total species pool in the region). Species richness and diversity did not differ among successional stages but the behaviour of ants towards diaspores changed with the age of secondary forests. In old successional stages the removal of entire diaspores was more common than in young successional stages of forests. Concordantly, diaspore removal rates were lowest in the youngest successional stage of secondary forests and increased with the age of forests. These results indicate that ant-diaspore interactions in secondary forests are disturbed and lower removal rates in secondary forests are likely to constrain the recruitment of plant populations during secondary succession. Rev. Biol. Trop. 60 (2): 933-942. Epub 2012 June 01.

Las interacciones entre animales y plantas son importantes para la recuperación de la diversidad y los procesos en los bosques secundarios, los cuales cada vez más tienden a dominar el paisaje tropical. Nosotros utilizamos una combinación de métodos experimentales y observaciones para estudiar las interacciones entre hormigas y diásporas a través de un gradiente de sucesión en los bosques del sur de Brasil, entre agosto 2007 y abril 2008. Además de las tasas de eliminación de diásporas, evaluamos la riqueza de especies, la diversidad y el comportamiento de las hormigas que interactúan con las diásporas, con tres repeticiones por sitio de cuatro estadios de sucesión del bosque. Se registraron 22 especies de hormigas que interactúan con diásporas (se estima un 15% del total de especies en la región). La riqueza y diversidad de especies no varió entre las etapas de sucesión, pero el comportamiento de las hormigas hacia las diásporas cambio con la edad de los bosques secundarios. En las etapas sucesionales avanzadas del bosque la eliminación de diásporas fue más común que en las etapas tempranas. En concordancia, las tasas de eliminación de diásporas fueron menores en la etapa más temprana de la sucesión de bosques secundarios y aumenta con la edad de los mismos. Estos resultados indican que las interacciones hormigasdiásporas en los bosques secundarios son alteradas y las menores tasas de extracción en los bosques secundarios es probable que limiten el reclutamiento de las poblaciones de plantas durante la sucesión secundaria.

Glandular trichomes in Connarus suberosus (Connaraceae): distribution, structural organization and probable functions

Connarus suberosus is a typical species of the Brazilian Cerrado biome, and its inflorescences and young vegetative branches are densely covered by dendritic trichomes. The objective of this study was to report the occurrence of a previously undescribed glandular trichome of this species. The localization, origin and structure of these trichomes were investigated under light, transmission and scanning electron microscopy. Collections were made throughout the year, from five adult specimens of Connarus suberosus near Botucatu, São Paulo, Brazil, including vegetative and reproductive apices, leaves and fruits in different developmental stages, as well as floral buds and flowers at anthesis. Glandular trichomes (GTs) occurred on vegetative and reproductive organs during their juvenile stages. The GTs consisted of a uniseriate, multicellular peduncle, whose cells contain phenolic compounds, as well as a multicellular glandular portion that accumulates lipids. The glandular cell has thin wall, dense cytoplasm (with many mitochondria, plastids and dictyosomes), and a large nucleus with a visible nucleolus. The starch present in the plastids was hydrolyzed during the synthesis phase, reducing the density of the plastid stroma. Some plastids were fused to vacuoles, and some evidence suggested the conversion of plastids into vacuoles. During the final activity stages of the GTs, a darkening of the protoplasm was observed in some of the glandular cells, as a programmed cell death; afterwards, became caducous. The GTs in C. suberosus had a temporal restriction, being limited to the juvenile phase of the organs. Their presence on the exposed surfaces of developing organs and the chemical nature of the reserve products, suggest that these structures are food bodies. Field observations and detailed studies of plant-environment interactions, as well as chemical analysis of the reserve compounds, are still necessary to confirm the role of these GTs as feeding rewards.

Connarus suberosus es una especie típica del bioma Cerrado brasileño, y sus inflorescencias junto con los ramos vegetativos jóvenes están densamente cubiertos por tricomas dendríticas. El objetivo de este estudio es presentar la ocurrencia de un tricoma glandular sin descripción previa para esta especie. La localización, origen y estructura de estos tricomas se observó mediante microscopia óptica, así como electrónica de transmisión y de barrido. Los tricomas glandulares (TGs) se producen en los órganos vegetativos y reproductivos, en sus fases jóvenes. Los TGs consisten en un pedúnculo uniseriado y multicelular, cuyas células contienen compuestos fenólicos, así como un sitio en el cual las células glandulares acumulan compuestos pécticos, almidón y lípidos. La acumulación de aceite en el citosol es impresionante, pues ocupa una gran parte del volumen celular. La porción glandular presenta células con paredes delgadas, un denso citoplasma y un núcleo grande con nucleolo visible. El citoplasma contiene gran cantidad de mitocondrias, plástidos y dictiosomas. El almidón presente en los plástidos se hidroliza durante la fase de síntesis, de tal modo que reduce la densidad del estroma plastidial. Algunos plástidos se fusionan con las vacuolas, además hay pruebas de la conversión de los plástidos en vacuolas. Durante las etapas finales de la actividad de los TGs, se observó un oscurecimiento del protoplasma en algunas de las células glandulares lo que sugiere la existencia de la muerte celular programada; después de lo anterior, los TGs se convierten en caducas. Por lo tanto la aparición de TGs en las superficies expuestas de los órganos en desarrollo en C. suberosus y la naturaleza química de los productos de reserva sugieren que estas estructuras son cuerpos alimenticios. Hay evidencias sobre el papel de estos tricomas en las interacciones planta-hormiga, especialmente en el bioma Cerrado.

O bambu Merostachys fischeriana (Bambusoideae: Bambuseae) como habitat para formigas de Floresta Tropical Montana / The bamboo Merostachys fischeriana (Bambusoideae: Bambuseae) as a canopy habitat for ants of Neotropical Montane Forest

Although Merostachys fischeriana is very abundant in the Brazilian Atlantic Rainforest, little attention has been paid to the biological interactions with other animals. The present study describes some of the interactions between ants and this bamboo species. The experiment was carried out in a fragment of a montane tropical forest in the Parque Estadual do Itacolomi, near Ouro Preto, MG, Brazil. Thirty culms of bamboo were randomly collected. The ants were obtained by direct collection from nodes and internodes. Morphometric variables of the bamboo were recorded for characterization of potential ant habitat. Merostachys fischeriana grows in rosettes as a thin bamboo (average = 1,0 cm; se = 0,27; n = 20) and is tall enough to reach the upper canopy of this low forest (average = 9,1 m; se = 2,72; n = 20). Fifteen ant species were sampled. Brachymyrmex heeri Forel was the most abundant in the nodes, while Camponotus crassus Mayr (Hymenoptera: Formicidae) was the most abundant in the internodes. The composition of the species that inhabit the internodes was different from the composition in the node (Q-test: Q = 3,76; P = 0,05). The level of occupation was defined by the number of holes (F = 10,33; P < 0,01), the number of internodes in the canopy (F = 6,84; P = 0,01) and the length of the culm (F = 7,52; P = 0,01). The plant's morphology allowed the occurrence of additional species of ants in the canopy and influenced the composition of the entire ant assemblage.

Nest structure and occurrence of three species of Azteca (Hymenoptera, Formicidae) in Cecropia pachystachya (Urticaceae) in non-floodable and floodable pantanal areas / Arquitetura de ninho e ocorrência de três espécies de Azteca (Hymenoptera, Formicidae) em Cecropia pachystachya (Urticaceae) em ambiente alagável e não alagável no Pantanal

Thirty Cecropia pachystachya trees were examined in non-floodable and floodable areas to investigate the association between C. pachystachya and Azteca ants in the Pantanal of Mato Grosso do Sul, Brazil. The species Azteca ovaticeps, Azteca isthmica, and Azteca alfari were found nesting inside domatia of C. pachystachya. A. ovaticeps was the most frequent species in the trees in the floodable area, while A. isthmica and A. alfari, in the non-floodable area. A. ovaticeps and A. isthmica maintained more entrance/exit holes in comparison to A. alfari. All Azteca species maintained entrance/exit holes in the closest domatia to the apical area of the branch, due to proximity to Müllerian and pearl bodies, suggesting that these species of Azteca were influenced by their environment during evolution and became specialized. All internodal septa of each examined branch were perforated by ants, indicating the branches were inhabited by a single colony.

Foram analisadas 30 plantas de Cecropia pachystachya em cada ambiente alagável e não alagável no Pantanal sul-mato-grossense, Brasil, com o objetivo de investigar a associação entre formigas Azteca e C. pachystachya. Foram encontradas as espécies Azteca ovaticeps, Azteca isthmica e Azteca alfari nidificando nas domáceas da planta. A. ovaticeps foi mais frequente em plantas de área alagável, enquanto A. isthmica e A. alfari em plantas em área não alagável. A. ovaticeps e A. isthmica apresentaram maior quantidade de orifícios de entrada/saída em relação à A. alfari e todas as espécies mantêm próximo da região apical do ramo, orifícios de entrada/saída nas domáceas, devido à proximidade com os corpúsculos müellerianos e pérola. Isto é, essas espécies de Azteca especializaram-se ao longo da evolução influenciadas pelo ambientes. Todos os septos internodais de cada ramo analisados apresentaram-se perfurados pelas formigas, sugerindo que os mesmos são habitados por uma única colônia.

Spiders associated with Psychotria carthagenensis Jacquin. (Rubiaceae): vegetative branches versus inflorescences, and the influence of Crematogaster sp. (Hymenoptera, Formicidae), in South-Pantanal, Brazil / Aranhas associadas à Psychotria carthagenensis Jacq. (Rubiaceae): ramos vegetativos versus inflorescências, e a influência de Crematogaster sp. (Hymenoptera, Formicidae), no Pantanal Sul, Brasil

The aim of this study was to analyze: i) the spider community in vegetative and reproductive branches of Psychotria carthagenensis concerning relative abundance, guild composition and body size distribution; ii) ant abundance in diferent types of branches and iii) the spider behavior when experimentally put in contact with inflorescences covered with ants. There was no difference between vegetative and reproductive branches in relation to spider abundance, composition of guilds and body size distribution of spiders. However, there was a significant difference in ant abundance. In the behavioral experiment, 90 percent of the spiders were expelled from inflorescences by ants; in control treatment, 100 percent remained in the inflorescences. The ant density in different parts of the plant may explain the spider distribution.

Aranhas associadas à Psychotria carthagenensis Jacq. (Rubiaceae). O objetivo deste estudo foi analisar: i) a comunidade de aranhas em ramos vegetativos e reprodutivos de P. carthagenensis quanto à abundancia relativa, composição das guildas e distribuição de tamanhos; ii) a abundância de formigas nos diferentes tipos de ramos; e iii) o comportamento das aranhas quando colocadas em inflorescências cobertas por formigas. Não houve diferença significativa entre ramos vegetativos e reprodutivos em relação à composição de guildas, abundância relativa e distribuição de tamanhos de aranhas. Mas quando comparado à abundância de formigas, houve diferença. No experimento comportamental, 90 por cento das aranhas foram expulsas das inflorescências pelas formigas; no tratamento controle; 100 por cento das aranhas permaneceram nas inflorescências. Para P. carthagenensis, talvez a densidade de formigas em diferentes partes da planta explique melhor a distribuição das aranhas.

Protection of Vochysia elliptica (Vochysiaceae) by a nectar-thieving ant

Vochysia elliptica (Vochysiaceae) is a shrubby plant, which does not have EFNs. Camponotus ants thieve nectar, and can decrease plant fitness by making flowers less attractive to pollinators. However, ants remove herbivores, wich can be beneficial. Results show that plants from which ants were excluded had lower rates of termite (simulated herbivore) removal than did plants visited by ants. Plants accessible to ants showed higher rates of termite removal in the base of leaves and in the inflorescence, than in the tip of leaves. This occurs because ants must pass through the principal axis to reach the inflorescence. Conclusive results of this cost/benefit analysis of the Camponotus sp. presence for V. elliptica can be obtained, with experimental manipulations.

Vochysia elliptica (Vochysiaceae) é uma planta arbustiva que não porta NEFs. Formigas Camponotus roubam néctar e podem diminuir a aptidão da planta, tornando as flores menos atrativas para polinizadores. O efeito das formigas na remoção dos herbívoros, no entanto, pode ser benéfico. Resultados evidenciaram que plantas cujas formigas foram excluídas tiveram menor taxa de remoção de cupins (herbívoros simulados) do que plantas visitadas por formigas. Plantas com acesso às formigas apresentaram maior taxa de cupins removidos na base das folhas e na inflorescência do que na ponta das folhas. Isso ocorre porque formigas devem passar através do eixo principal para alcançar a inflorescência. Resultados conclusivos dessa análise de custo/benefício da presença de Camponotus sp. para V. elliptica deverão ser obtidos no futuro, com manipulações experimentais.

Formigas (Hymenoptera: Formicidae) visitantes de Ilex paraguariensis (Aquifoliaceae) / Visitor ants (Hymenoptera: Formicidae) of Ilex paraguariensis (Aquifoliaceae)

A cultura de erva-mate, Ilex paraguariensis, base econômica de vários municípios do Rio Grande do Sul, possui mais de uma centena de insetos associados, no entanto, sem registros da família Formicidae. Assim, visando identificar as espécies de formicídeos que visitam I. paraguariensis, foram realizadas coletas em duas áreas de uma monocultura, uma com plantas sadias e outra com plantas atacadas por Hedypathes betulinus, e em um pequeno plantio dessa espécie. Paralelamente, foram conduzidos testes com térmitas, simulando possíveis herbívoros, para avaliar a atividade predatória das formigas que forrageiam em I. paraguariensis. Considerando as três áreas avaliadas, foram encontradas dezesseis espécies de formigas na erva-mate. Não foi observada interação significativa entre formigas e nectários extraflorais em I. paraguariensis, não tendo ocorrido uma proteção diferencial das folhas (com nectários extraflorais) em relação ao caule (sem nectários extraflorais). A maior ocorrência de formigas na erva-mate com homópteros, bem como nidificando nas galerias deixadas pelo cerambicídeo H. betulinus sugere que as formigas estão apenas explorando recursos proporcionados por outros insetos.

Maté crop, Ilex paraguariensis, is the economic base of several municipalities of Rio Grande do Sul. Despite the fact of presenting hundreds of associated insects, family Formicidae has not been registered yet. Therefore, aiming to identify ant species visiting I. paraguariensis, collections were made in two maté crop areas, one with healthy plants and the other with plants attacked by Hedypathes betulinus, and in a third small area. At the same time, tests with termites to simulate possible herbivores were conducted in order to evaluate predator activity of the ants which forage in I. paraguariensis. Considering the evaluated areas, sixteen ant species were found in maté. Significant interaction between ants and extrafloral nectaries in I. paraguariensis was not observed, and differential protection of leaves (with extrafloral nectaries) compared to stem (without extrafloral nectaries) did not occur. The greatest occurrence of ants in maté with Homoptera, as well as nesting in galleries left by H. betulinus (Cerambycidae) suggests that ants are only exploring resources provided by other insects.