The contribution of the BIOTA/FAPESP Program to the advancement of the knowledge on terrestrial invertebrates / A contribuição do Programa BIOTA/FAPESP para o avanço no conhecimento sobre os invertebrados terrestres

Abstract The variability of the organisms living in a given area constitute what is referred to as biodiversity, one of nature's fundamental properties, responsible for the balance and stability of ecosystems. The loss of biodiversity has been of great concern to scientists, especially because of the role played by human activities in this regard, able to lead to irreversible circumstances. The São Paulo Research Foundation (Fundação de Amparo à Pesquisa do Estado de São Paulo, FAPESP) plays a major role in supporting research efforts in the most diverse branches of science. In the late 1990´s, FAPESP launched a major program to promote research on biodiversity, named BIOTA/FAPESP. So far, this program has financed the conduction of 26 projects, involving research activities in most of Brazil, while focusing mainly the State of São Paulo. These projects have generated about 1140 publications in peer-reviewed journals of high standard, providing relevant information, including the original description of 1187 species and 76 genera, the complementary description of 350 species, as well as a number of inventory works, biological studies, etc. The program has also been instrumental in the establishment or adequacy of research facilities and training of new taxonomists. Most extensively studied groups of terrestrial invertebrates include Insecta of the orders Hymenoptera, Lepidoptera and Diptera, and Arachnida of the subclasses Araneae and Acari. Distinct projects have also contributed to the detection of organisms potentially useful as biological control agents and in the determination of maps of major interest for the establishment of public policies. In the future, priority groups for study should include the Annelida and the Nematoda, for the potential both have as beneficial organisms, or for the potential some Nematoda have as organisms harmful to plants and animals.

Resumo A variabilidade dos organismos em uma determinada área constitui o que se denomina biodiversidade, uma das propriedades fundamentais da natureza, responsável pelo equilíbrio e estabilidade dos ecossistemas. A perda da biodiversidade tem sido uma grande preocupação para os cientistas, principalmente pelo papel desempenhado pelas atividades humanas, com potencial para desencadear circunstâncias irreversíveis. A Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) desempenha um papel importante no apoio às pesquisas nos mais diversos ramos da ciência. No final da década de 1990, a FAPESP lançou um grande programa de fomento à pesquisa em biodiversidade, denominado BIOTA/FAPESP. Até o momento, este programa financiou a realização de 26 projetos, envolvendo atividades na maior parte do Brasil, embora tenham como foco principal o estado de São Paulo. Esses projetos geraram cerca de 1.140 publicações em periódicos de alto impacto, fornecendo informações relevantes que incluem a descrição original de 1.187 espécies e 76 gêneros e a descrição complementar de 350 espécies, além de diversos trabalhos de inventário, estudos biológicos etc. O programa também tem sido fundamental para o estabelecimento ou adequação de instalações de pesquisa científica e o treinamento de novos taxonomistas. Os grupos de invertebrados terrestres mais estudados incluem os Insecta das ordens Hymenoptera, Lepidoptera e Diptera, e os Arachnida das subclasses Araneae e Acari. Projetos distintos também têm contribuído para a detecção de organismos potencialmente úteis como agentes de controle biológico e na determinação de mapas de áreas preferenciais para o estabelecimento de políticas públicas. No futuro, os grupos prioritários de estudo devem incluir os Annelida e os Nematoda, pelo potencial que ambos têm como organismos benéficos, ou pelo potencial que alguns Nematoda têm como organismos prejudiciais a plantas e animais.

Marimbondos: systematics, biogeography, and evolution of social behaviour of neotropical swarm-founding wasps (Hymenoptera: Vespidae: Epiponini).

Neotropical swarm-founding wasps are divided into 19 genera in the tribe Epiponini (Vespidae, Polistinae). They display extensive variation in several colony-level traits that make them an attractive model system for reconstructing the evolution of social phenotypes, including caste dimorphism and nest architecture. Epiponini has been upheld as a solid monophyletic group in most phylogenetic analyses carried out so far, supported by molecular, morphological and behavioural data. Recent molecular studies, however, propose different relationships among the genera of swarm-founding wasps. This study is based on the most comprehensive epiponine sampling so far and was analyzed by combining morphological, nesting and molecular data. The resulting phylogenetic hypothesis shows many of the traditional clades but still impacts the way certain behavioural characters, such as nest structure and castes, evolved, and thus requires some re-interpretations. Angiopolybia as sister to the remaining Epiponini implies that nest envelopes and a casteless system are plesiomorphic in the tribe. Molecular dating points to an early tribal diversification during the Eocene (c. 55-38 Ma), with the major differentiation of current genera concentrated in the Oligocene/Miocene boundary.

Taxonomic revision and cladistic analysis of the species of Zethus mexicanus group (Linnaeus, 1758) sensu Bohart amp; Stange, 1965 (Hymenoptera, Vespidae, Eumeninae).

The Zethus mexicanus group sensu Bohart Stange (1965) consists of four species in the subgenus Zethusculus: Zethus brasiliensis de Saussure, 1852, Z. mexicanus (Linnaeus, 1758) (both subdivided into three subspecies each), Z. magnus de Saussure, 1852 and Z. waldoi Zavattari, 1912. Color patterns and distribution have been used to differentiate the subspecies until now. However, these patterns may vary among individuals, and the distribution of specimens examined in this study showed overlap between subspecies. The analysis of external morphology did not show significant differences, making the subspecific division unnecessary. Thus, the following synonyms are proposed: Zethus brasiliensis fuscatus Boharth Stange, 1965 n. syn. and Z. brasiliensis panamensis Boharth Stange, 1965 n. syn. are junior synonyms of Z. brasiliensis brasiliensis; and Z. mexicanus erythrogena Boharth Stange, 1965 n. syn. and Z. mexicanus lugubris Perty, 1833 n. syn. are junior synonyms of Zethus mexicanus mexicanus. Furthermore, the following status changed are proposed: Z. brasiliensis brasiliensis de Saussure, 1852 stat. rev. to Z. brasiliensis; and Z. mexicanus mexicanus (Linnaeus, 1758) stat. rev. to Z. mexicanus. In addition, a cladistic analysis, using terminals with representatives of the three subgenera of Zethus and one species of Ctenochilus, recovered the monophyly of the Zethus mexicanus group, as well as the subgenus Zethusculus, but rendered the arietis group paraphyletic. An identification key and redescriptions are given for the species in the group of Zethus mexicanus sensu Bohart Stange (1965).

Zethus (Zethus) wileyi (Hymenoptera, Vespidae, Eumeninae), a new species with its own species-group.

Zethus wileyi Stange sp. nov. (Panama) is described and a cladistic analysis is carried in order to determine the relationship between the new species and the other species in the hilarianus species-group. The new species is assigned to its own new species-group due to evident differences in the morphology.

Notes on the Neotropical Zethus Fabricius, 1804 (Hymenoptera, Vespidae, Eumeninae) with the description of two new species from Brazil.

Two new species, Zethus (Zethus) aliceae Lopes, sp. nov. (Brazil) and Z. (Zethus) sinuostylus Lopes, sp. nov. (Brazil) are described and figured. New synonymy is proposed for Zethus (Zethoides) biglumis Spinola, 1841 (=Zethus ferrugineus de Saussure, 1852, syn. nov.

A survey of DNA methylation across social insect species, life stages, and castes reveals abundant and caste-associated methylation in a primitively social wasp.

DNA methylation plays an important role in the epigenetic control of developmental and behavioral plasticity, with connections to the generation of striking phenotypic differences between castes (larger, reproductive queens and smaller, non-reproductive workers) in honeybees and ants. Here, we provide the first comparative investigation of caste- and life stage-associated DNA methylation in several species of bees and vespid wasps displaying different levels of social organization. Our results reveal moderate levels of DNA methylation in most bees and wasps, with no clear relationship to the level of sociality. Strikingly, primitively social Polistes dominula paper wasps show unusually high overall DNA methylation and caste-related differences in site-specific methylation. These results suggest DNA methylation may play a role in the regulation of behavioral and physiological differences in primitively social species with more flexible caste differences.

Well, what about intraspecific variation? Taxonomic and phylogenetic characters in the genus Synoeca de Saussure (Hymenoptera, Vespidae).

Cely and Sarmiento (2011) took issue with the cladistic analysis of relationships among species of the genus Synoeca by Andena et al. (2009a), and presented a reanalysis. They claimed that intraspecific variation in the genus is meaningful, and proper consideration yields a conclusion different from that of Andena et al. Both their critique and reanalysis are vitiated by numerous errors, as is shown in the present paper.

Castes in the neotropical social wasp Leipomeles dorsata (Fabricius) (Hymenoptera: Vespidae): a window for workers achieving a new status in the colony.

Morphometric studies performed in several species of Neotropical social wasps from the tribe Epiponini showed that in some species there are marked differences between castes, while other species present highly distinct castes with differences only in ovarian development. This work analyzed females from six colonies of the social wasp Leipomeles dorsata (Fabricius) in which queens (egglayers) and workers showed differences in ovarian development and coloration. We propose that wasps with developed ovaries (egglayers) and coloration similar to those of workers are possibly intermediates that obtained the status of queens in the colony.

Castes in the neotropical social wasp Leipomeles dorsata (Fabricius) (Hymenoptera: Vespidae): a window for workers achieving a new status in the colony

Morphometric studies performed in several species of Neotropical social wasps from the tribe Epiponini showed that in some species there are marked differences between castes, while other species present highly distinct castes with differences only in ovarian development. This work analyzed females from six colonies of the social wasp Leipomeles dorsata (Fabricius) in which queens (egglayers) and workers showed differences in ovarian development and coloration. We propose that wasps with developed ovaries (egglayers) and coloration similar to those of workers are possibly intermediates that obtained the status of queens in the colony.

An improved bait method for collecting Hymenoptera, especially social wasps (Vespidae: Polistinae).

In this work, an alternative method for the capture of neotropical social wasps is described, which may supplement or replace the procedures commonly used in wasp surveys. The method was tested in a seasonal semideciduous forest in the northwest of São Paulo State, collecting a larger number of species and a greater abundance of social wasps and of other Hymenoptera than other methods also used in the same area, namely Malaise traps, bottle traps and meat baits. The proposed method was adapted from Wille (1962) who used an attractive solution based on water and honey for bee attraction. In here, the attractive solution was based on water, salt and crystal sugar, and was effective in attracting bees and solitary wasps, but mainly social wasps, which is the group of interest for this report.

An improved bait method for collecting hymenoptera, especially social wasps (Vespidae: Polistinae) / Método de armadilha melhorado para a coleta de hymenoptera, especialmente de vespas sociais (Vespidae: Polistinae)

In this work, an alternative method for the capture of neotropical social wasps is described, which may supplement or replace the procedures commonly used in wasp surveys. The method was tested in a seasonal semideciduous forest in the northwest of São Paulo State, collecting a larger number of species and a greater abundance of social wasps and of other Hymenoptera than other methods also used in the same area, namely Malaise traps, bottle traps and meat baits. The proposed method was adapted from Wille (1962) who used an attractive solution based on water and honey for bee attraction. In here, the attractive solution was based on water, salt and crystal sugar, and was effective in attracting bees and solitary wasps, but mainly social wasps, which is the group of interest for this report.

Neste trabalho é proposto método para a captura de vespas sociais neotropicais nos mais diferentes habitats e que pode substituir os métodos comumente utilizados em levantamentos faunísticos de curto prazo. O método foi inicialmente testado em um fragmento de floresta estacional semidecidual no Noroeste do estado de São Paulo, obtendo sucesso na coleta de vespas sociais e também de outros grupos de Hymenoptera. Coletou um número maior de espécies e também maior abundância do que outros métodos utilizados na mesma área, como armadilhas Malaise, garrafas-armadilha e iscas de carne. O método desenvolvido baseou-se na proposta de Wille (1962), que utilizou uma solução atrativa, composta por água e mel, para a coleta de abelhas. Neste trabalho, substituiu-se o mel por açúcar e adicionou-se sal, obtendo sucesso na atração não apenas de vespas sociais, mas também de abelhas e vespas solitárias.

Diversity of social wasps (Hymenoptera, Vespidae, Polistinae) in three fragments of semideciduous seasonal forest in the northwest of São Paulo State, Brazil / Diversidade de vespas sociais (Hymenoptera, Vespidae, Polistinae) em três fragmentos de floresta estacional semidecidual no noroeste do Estado de São Paulo, Brasil

Human exploitation of the forest in the northwest of São Paulo State has generated enormous fragmentation of that forest. Such disturbance has reduced the populations of insects in general. This work was a survey of social wasps (Hymenoptera, Vespidae; Polistinae) in three areas in different stages of regeneration: Paulo de Faria - SP (435 ha), Pindorama - SP (128 ha) and Neves Paulista - SP (1 ha). These three areas were chosen for comparative purposes. To capture the wasps, it was used: active collecting with attractant liquid (solution of water, salt and sugar) with the aid of a dorsal spray bag. During the period from July to December 2006, 414 social wasps were collected in Paulo de Faria, constituting seven species belonging to four genera; 111 social wasps were collected in Pindorama, constituting six species belonging to four genera, and 129 social wasps were collected in Neves Paulista constituting 12 species belonging to seven genera. In order to compare these three areas ecological indexes were calculated. Neves Paulista had the greatest diversity, and Paulo de Faria presented greater abundance. These factors were probably caused by neighboring areas and ecological corridors, which were limited in Paulo de Faria and Pindorama.

A ação antrópica no noroeste do Estado de São Paulo tem gerado grande fragmentação florestal. Tais perturbações reduziram as populações de insetos em geral. Neste trabalho foi realizado um levantamento de vespas sociais (Hymenoptera, Vespidae; Polistinae) em três áreas em diferentes estágios de regeneração, em Paulo de Faria - SP (435 ha), Pindorama (128 ha) e em Neves Paulista (1 ha), escolhidas para fins comparativos. Para capturar as vespas, foi utilizada: coleta ativa com utilização de líquido atrativo (solução de água, sal e açúcar) com auxílio de um borrifador dorsal de alavanca. Durante o período de Julho a Dezembro de 2006 foram capturados 414 vespídeos sociais em Paulo de Faria, sendo estes distribuídos em sete espécies pertencentes a quatro gêneros; em Pindorama, foram coletadas 111 vespas sociais distribuídos em seis espécies pertencentes a quatro gêneros, e em Neves Paulista, um total de 129 vespídeos sociais distribuídos em 12 espécies pertencentes a sete gêneros. A fim de comparar as três áreas foram utilizados índices ecológicos. A área de Neves Paulista obteve a maior diversidade e Paulo de Faria a maior abundância de vespas. Tais fatores provavelmente foram causados pela influência das áreas vizinhas e pelos corredores ecológicos, que eram limitados na área de Paulo de Faria e Pindorama.

Morphological caste studies in the neotropical swarm-founding Polistinae wasp Angiopolybia pallens (Lepeletier) (Hymenoptera: Vespidae) / Estudos morfológicos das castas da vespa Polistinae enxameadora neotropical, Angiopolybia pallens (Lepeletier) (Hymenoptera: Vespidae)

Neotropical swarm-founding wasps present polygynic colonies and a great variation in relation to caste differentiation, ranging from species in which queens and workers are similar in shape and size to those where variation in shape and size is conspicuous. Canonical discriminant analysis and Bonferroni t-test analysis on morphometric data collected from eight body parts of specimens from nine colonies of Angiopolybia pallens (Lepeletier) were undertaken as a step towards to a better understanding on caste differenciation in Epiponini. All specimens were dissected to verify the ovary developmental stage, and the spermatheca was removed to check for the mating status. Females were then grouped as queens (mated with developed ovaries), intermediates (virgin with developed ovaries) and workers (virgin with undeveloped ovaries). Even though differences were found for some measurements in seven out of the nine colonies evaluated, multivariate analysis (Wilks' lambda values) indicated castes could not be reliably distinguished only by morphology. We conclude that morphological differences among A. pallens castes are low or absent, reflecting a possible groundplan for the Epiponini, i.e. a post-imaginal pattern of caste differentiation.

As vespas neotropicais enxameadoras possuem colônias poligínicas e uma variação muito grande em relação à diferenciação das castas, desde espécies em que rainhas e operárias possuem formas e tamanhos semelhantes até aquelas em que grande diferença na forma e no tamanho são observados. Estudos morfométricos utilizando oito medidas corporais em nove colônias de Angiopolybia pallens (Lepeletier) foram realizados a fim de aumentar o conhecimento sobre diferenciação de castas em Epiponini, empregando-se análise canônica discriminante e teste t de Bonferroni. Os indivíduos foram dissecados para determinar o nível de desenvolvimento ovariano e as espermatecas foram removidas para verificar a ocorrência de inseminação e assim definir os três grupos de fêmeas: rainhas (ovários desenvolvidos e espermateca fecundada), intermediárias (ovários desenvolvidos e espermateca não-fecundada) e operárias (ovários não-desenvolvidos). Apesar de serem encontradas diferenças significativas para algumas medidas em sete das nove colônias, a análise multivariada (lambda de Wilks) mostrou que as castas são de difícil distinção apenas pela morfologia. É possível concluir que diferenças morfológicas entre castas de A. pallens são pequenas ou ausentes, o que reflete o plano basal para os Epiponini, i.e. padrão pós-imaginal de diferenciação castas.

Morphological caste studies in the neotropical swarm-founding Polistinae wasp Angiopolybia pallens (Lepeletier) (Hymenoptera: Vespidae).

Neotropical swarm-founding wasps present polygynic colonies and a great variation in relation to caste differentiation, ranging from species in which queens and workers are similar in shape and size to those where variation in shape and size is conspicuous. Canonical discriminant analysis and Bonferroni t-test analysis on morphometric data collected from eight body parts of specimens from nine colonies of Angiopolybia pallens (Lepeletier) were undertaken as a step towards to a better understanding on caste differenciation in Epiponini. All specimens were dissected to verify the ovary developmental stage, and the spermatheca was removed to check for the mating status. Females were then grouped as queens (mated with developed ovaries), intermediates (virgin with developed ovaries) and workers (virgin with undeveloped ovaries). Even though differences were found for some measurements in seven out of the nine colonies evaluated, multivariate analysis (Wilks' lambda values) indicated castes could not be reliably distinguished only by morphology. We conclude that morphological differences among A. pallens castes are low or absent, reflecting a possible groundplan for the Epiponini, i.e. a post-imaginal pattern of caste differentiation.

Castes and asynchronous colony cycle in Polybia bistriata (Fabricius) (Hymenoptera: Vespidae)

Neotropical swarm-founding wasps, the Epiponini, are an outstanding group of social insects whose societies are polygynic and complex nest builders. Caste dimorphism in these wasps ranges from incipient to highly distinct. Morphometric analyses of nine body parts, ovarian status, relative age and development of the 5th gastral sternite gland (Richards´ gland) of Polybia bistriata Fabricius were undertaken in order to estimate caste differentiation in nine colonies. Morphological and physiological data were used in multivariate analyses in order to evaluate the level of discrimination between inseminated and non-inseminated females. Clear physiological differences were found: queens had highly developed ovaries and they were inseminated, and workers had totally undeveloped ovaries or they had few developed oocytes (only in two colonies), but in both cases insemination was not detected. ANOVA and discriminant function analysis detected slight, but significant differences between castes. In relation to colony cycle, colonies were considered to be in the following stages: one in pre-emergence, four in worker production and four in sexuals production. Richards´ gland analyses indicated that in small colonies (<100 females) queens had a less developed gland than in medium (100-200 females), and large colonies (>200 females). Taking the whole data, it was possible to conclude that caste differences were slight, but more evident in some phases of the colony cycle, a phenomenon previously described for other epiponines.

Os Epiponini, vespas sociais neotropicais, são um grupo de insetos sociais com sociedades poligínicas, construtoras de ninhos complexos. O dimorfismo das castas varia de incipiente até altamente distinto. Análises morfométricas de nove partes corporais, estágio de desenvolvimento ovariano, idade relativa e desenvolvimento da glândula do quinto esternito do gáster (glândula de Richards) de Polybia bistriata (Fabricius) foram utilizados para calcular o nível de diferenciação das castas de nove colônias. Dados morfológicos e fisiológicos foram usados em análises multivariadas para avaliar o nível de discriminação entre fêmeas inseminadas e não-inseminadas. Foram encontradas diferenças fisiológicas claras: rainhas tinham ovários altamente desenvolvidos e inseminados, e as operárias não estavam inseminadas e tinham ovários não desenvolvidos ou uns poucos oócitos desenvolvidos (em duas colônias). ANOVA e análise da função discriminante detectaram diferenças significantes entre as castas. Considerou-se, pelos dados obtidos, que as colônias estavam nas seguintes fases em relação ao ciclo colonial: uma em pré-emergência, quatro em produção de operárias ;.e quatro em produção de sexuados. Análises das glândulas de Richards indicaram o seu menor desenvolvimento em rainhas de colônias pequenas (< 100 fêmeas) do que aquelas de colônias médias (100-200 fêmeas), e grandes (> 200 fêmeas). Considerando-se todos os dados, é possível concluir que as diferenças entre as castas são pequenas, mas evidentes em algumas fases do ciclo colonial, um fenômeno previamente descrito para outros epiponíneos.

Castes and asynchronous colony cycle in Polybia bistriata (Fabricius) (Hymenoptera: Vespidae).

Neotropical swarm-founding wasps, the Epiponini, are an outstanding group of social insects whose societies are polygynic and complex nest builders. Caste dimorphism in these wasps ranges from incipient to highly distinct. Morphometric analyses of nine body parts, ovarian status, relative age and development of the 5th gastral sternite gland (Richards' gland) of Polybia bistriata Fabricius were undertaken in order to estimate caste differentiation in nine colonies. Morphological and physiological data were used in multivariate analyses in order to evaluate the level of discrimination between inseminated and non-inseminated females. Clear physiological differences were found: queens had highly developed ovaries and they were inseminated, and workers had totally undeveloped ovaries or they had few developed oocytes (only in two colonies), but in both cases insemination was not detected. ANOVA and discriminant function analysis detected slight, but significant differences between castes. In relation to colony cycle, colonies were considered to be in the following stages: one in pre-emergence, four in worker production and four in sexuals production. Richards' gland analyses indicated that in small colonies (<100 females) queens had a less developed gland than in medium (100-200 females), and large colonies (>200 females). Taking the whole data, it was possible to conclude that caste differences were slight, but more evident in some phases of the colony cycle, a phenomenon previously described for other epiponines.

"Winter" aggregations, colony cycle, and seasonal phenotypic change in the paper wasp Polistes versicolor in subtropical Brazil.

Social wasps from temperate zones have clear annual colony cycles, and the young queens hibernate during winter. In the subtropics, the only previously reported evidence for the existence of "hibernation" is the facultative winter aggregations of females during harsh climate conditions. As in temperate-zone species analyzed so far, we show in this study that in the paper wasp, Polistes versicolor, a subtropical species, body size increases as an unfavorable season approaches. Our morphological studies indicate that larger females come from winter aggregations--that is, they are new queens. Multivariate analyses indicate that size is the only variable analyzed that shows a relationship to the differences. Given the absence of a harsh climate, we suggest that the occurrence of winter aggregations in tropical P. versicolor functions to allow some females to wait for better environmental conditions to start a new nest, rather than all being obliged to start a new nest as soon as they emerge.

Dados comportamentais na era da genômica / Behavioral data in a genomics age

Muitos acreditam que com o seqüenciamento do DNA não será mais necessária qualquer outra informação, como a relativa ao comportamento, para inferir passos e origens evolutivas. Em nossa opinião, sempre precisaremos de caracteres mais complexos, como os caracteres comportamentais, para a realização de estudos evolutivos. Estudos evolutivos ou filogenias sem referência aos caracteres fenotípicos são necessariamente incompletos. Não há sentido em estudar-se o DNA sem referência ao fenótipo. Com o advento da genômica, o estudo do comportamento pode parecer estar em segundo plano. Entretanto, argumentos importantes em favor desse estudo podem ser levantados (AU)

Dados comportamentais na era da genômica / Behavioral data in a genomics age

Muitos acreditam que com o seqüenciamento do DNA não será mais necessária qualquer outra informação, como a relativa ao comportamento, para inferir passos e origens evolutivas. Em nossa opinião, sempre precisaremos de caracteres mais complexos, como os caracteres comportamentais, para a realização de estudos evolutivos. Estudos evolutivos ou filogenias sem referência aos caracteres fenotípicos são necessariamente incompletos. Não há sentido em estudar-se o DNA sem referência ao fenótipo. Com o advento da genômica, o estudo do comportamento pode parecer estar em segundo plano. Entretanto, argumentos importantes em favor desse estudo podem ser levantados

Predatory behavior in a necrophagous bee Trigona hypogea (Hymenoptera; Apidae, Meliponini).

Although most bees feed on nectar and pollen, several exceptions have been reported. The strangest of all is the habit found in some neotropical stingless bees, which have completely replaced pollen-eating by eating animal protein from corpses. For more than 20 years, it was believed that carrion was the only protein source for these bees. We report that these bees feed not only off dead animals, but on the living brood of social wasps and possibly other similar sources. Using well developed prey location and foraging behaviors, necrophagous bees discover recently abandoned wasps' nests and, within a few hours, prey upon all immatures found there.