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''Brachygastra lecheguana'' (Latreille 1824), formerly known as ''Nectarina lecheguana'' <ref name="Bequaert">Bequaert, J.Q. (1932). “The Nearctic social wasps of the subfamily polybiinae (Hymenoptera; Vespidae). Entomologica Americana. 13(3):87-150.</ref>, is a species of dark [[paper wasp]] <ref name="Alves-Silva 2013">Alves-Silva, E., Baronio, G.J., Torezan-Silingardi, H.M., Del-Claro, K. (2013). “Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on [[Banisteriopsis]] malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system.” Entomological Science. 16:162-169.</ref> found across Central and South America <ref name="Bequaert" />. It nests in underbrush in grassland-type environments, and produces [[honey]], characteristic of the [[genus]] ''Brachygastra'' <ref name="Bequaert" />. Common names include marimbondo-do-campo, marimbondo-do-pasto <ref name="Henrique-Simões 2011">Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)</ref>, marimbondo-exu, marimbondo-de-pote, marimbondo-de-purrão, and avispa-de-pote <ref name="Costa Neto 2004">Costa Neto, E.M. (2004). “La etnoentomología de las avispas (Hymenoptera, Vespoidea) en el poblado de Pedra Branca, estado de Bahia, nordeste de Brasil.” Boletín de la Sociedad Entomológica Aragonesa. 34:247-262.</ref>, among others depending on the location.
''Brachygastra lecheguana'' (Latreille 1824), formerly known as ''Nectarina lecheguana'' <ref name="Bequaert">Bequaert, J.Q. (1932). “The Nearctic social wasps of the subfamily polybiinae (Hymenoptera; Vespidae). Entomologica Americana. 13(3):87-150.</ref>, is a species of dark [[paper wasp]] <ref name="Alves-Silva 2013">Alves-Silva, E., Baronio, G.J., Torezan-Silingardi, H.M., Del-Claro, K. (2013). “Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on [[Banisteriopsis]] malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system.” Entomological Science. 16:162-169.</ref> found across Central and South America <ref name="Bequaert" />. It nests in underbrush in grassland-type environments, and produces [[honey]], characteristic of the [[genus]] ''[[Brachygastra]]'' <ref name="Bequaert" />. Common names include marimbondo-do-campo, marimbondo-do-pasto <ref name="Henrique-Simões 2011">Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)</ref>, marimbondo-exu, marimbondo-de-pote, marimbondo-de-purrão, and avispa-de-pote <ref name="Costa Neto 2004">Costa Neto, E.M. (2004). “La etnoentomología de las avispas (Hymenoptera, Vespoidea) en el poblado de Pedra Branca, estado de Bahia, nordeste de Brasil.” Boletín de la Sociedad Entomológica Aragonesa. 34:247-262.</ref>, among others depending on the location.
== Taxonomy ==
== Taxonomy ==
''B. lecheguana'' belongs to the [[Epiponini]] tribe of [[Polistinae]] wasps <ref name="Noll 2004">Noll, F.B., Wenzel, J.W., Zucchi, R. (2004). “Evolution of caste in Neotropical swarm-founding wasps (Hymenoptera: Vespidae; Epiponini).” American Museum Novitates. 3467:1-24.</ref>, sometimes referred to as Polybiinae wasps <ref name="Bequaert" /><ref name="Noll 2004" />. It is a Neotropical social wasp species that falls under the genus ''Brachygastra'' <ref name="Noll 2004" />.
''B. lecheguana'' belongs to the [[Epiponini]] tribe of [[Polistinae]] wasps <ref name="Noll 2004">Noll, F.B., Wenzel, J.W., Zucchi, R. (2004). “Evolution of caste in Neotropical swarm-founding wasps (Hymenoptera: Vespidae; Epiponini).” American Museum Novitates. 3467:1-24.</ref>, sometimes referred to as Polybiinae wasps <ref name="Bequaert" /><ref name="Noll 2004" />. It is a Neotropical social wasp species that falls under the genus ''Brachygastra'' <ref name="Noll 2004" />.
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''[[Brachygastra]]'' wasps are identified, as noted in Andena & Carpenter (2012), by an anatomical projection over their posterior. The [[Scutellum (insect anatomy)|scutellum]], [[metanotum]], and [[propodeum]], three of the [[hymenopteran]] back structures, combined form a flat, vertical surface in the middle of the body ([[mesosoma]]) <ref name="Andena 2012">Andena, S.R. (2012). “A phylogenetic analysis of the social wasp genus Brachygastra Perty, 1833, and Description of a new species (Hymenoptera: Vespidae: Epiponini).” American Museum Novitates. 3753:1-38</ref>.
''[[Brachygastra]]'' wasps are identified, as noted in Andena & Carpenter (2012), by an anatomical projection over their posterior. The [[Scutellum (insect anatomy)|scutellum]], [[metanotum]], and [[propodeum]], three of the [[hymenopteran]] back structures, combined form a flat, vertical surface in the middle of the body ([[mesosoma]]) <ref name="Andena 2012">Andena, S.R. (2012). “A phylogenetic analysis of the social wasp genus Brachygastra Perty, 1833, and Description of a new species (Hymenoptera: Vespidae: Epiponini).” American Museum Novitates. 3753:1-38</ref>.


''B. lecheguana'' is very closely related to ''[[Brachygastra mellifica|B. mellifica]]'' and ''B. borellii'', and indeed some authors suggest that ''B. lecheguana'' and ''B. mellifica'' can be differentiated based solely on location <ref name="Bequaert" />. However there are morphological and genetic differences between these species <ref name="name">details of the citation</ref><ref name="Andena 2012" /><ref name="Zhu 2000">Zhu, Y., Queller, D.C., Strassman, J.E. (1999). “A phylogenetic perspective on sequence evolution in microsatellite loci.” Journal of Molecular Evolution. 50:324-338.</ref>.
''B. lecheguana'' is very closely related to ''[[Brachygastra mellifica|B. mellifica]]'' and ''B. borellii'' <ref name="Andena 2012" />, and indeed some authors suggest that ''B. lecheguana'' and ''B. mellifica'' can only be differentiated based on where they are found with respect to their reported distribution throughout the Americas <ref name="Bequaert" />. However there are morphological and genetic differences between these species <ref name="name">details of the citation</ref><ref name="Andena 2012" /><ref name="Zhu 2000">Zhu, Y., Queller, D.C., Strassman, J.E. (1999). “A phylogenetic perspective on sequence evolution in microsatellite loci.” Journal of Molecular Evolution. 50:324-338.</ref>.
== History of Study ==
== History of Study ==
''Brachygastra lecheguana'' was first described by [[Pierre André Latreille|Latreille]] in 1824 <ref name="Bequaert" />. It was originally named ''Nectarina lecheguana'' <ref name="Bequaert" />, and the switch from ''Nectarina'' to ''Brachygastra'' as a name for this genus was not standard in the scientific literature until the 1940’s <ref name="Andena 2012" />. Between 1824 and 1943, this species was documented as a member of the following genera: ''[[Polistes]]'', ''[[Vespa]]'', ''[[Nectarinia]]'' (a misspelling of ''Nectarina'', and a misuse of this genus of birds), ''Brachygaster'' (another misspelling, and a misuse of this genus of parasitic wasp), ''Melissaia'', and ''Caba'' <ref name="Bequaert" /><ref name="Andena 2012" />.
''Brachygastra lecheguana'' was first described by [[Pierre André Latreille|Latreille]] in 1824 <ref name="Bequaert" />. It was originally named ''Nectarina lecheguana'' <ref name="Bequaert" />, and the switch from ''Nectarina'' to ''Brachygastra'' as a name for this genus was not standard in the scientific literature until the 1940’s <ref name="Andena 2012" />. Between 1824 and 1943, this species was documented as a member of the following genera: ''[[Polistes]]'', ''[[Vespa]]'', ''[[Nectarinia]]'' (a misspelling of ''Nectarina'', and a misuse of this genus of birds), ''Brachygaster'' (another misspelling, and a misuse of this genus of parasitic wasp), ''Melissaia'', and ''Caba'' <ref name="Bequaert" /><ref name="Andena 2012" />.


In 1923, Bequeart performed a survey of the literature on this species, compiling accounts and descriptions by such entomologists as [[Ferdinand de Saussure|de Saussure]], [[Hermann von Ihering|von Ihering]], and du Buysson among numerous others, creating a source of descriptive information <ref name="Bequaert" />. One study has focused on the morphological caste differentiation in this species <ref name="Shima">Shima, S.N., Noll, F.B., Zucchi, R. (2000). “Morphological caste differences in the Neotropical swarm-founding Polistine wasp, Brachygastra lecheguana (Hymenoptera: Vespidae, Polistinae, Epiponini).” Sociobiology. 36(1):41-52.</ref> and another directly studied the [[Foraging|foraging behavior]] of this wasp species <ref name="Alves-Silva 2013" />. While not the focus, ''B. lecheguana'' has been included in studies of groups of wasps known to be [[natural enemies]] of agricultural pests <ref name="Fernandes 2013">Fernandes, F.L., Da Silva, P.R., Gorri, J., Pucci, L.F., Da Silva, I.W. (2013). “Selectivity of old and new organophosphate insecticides and behavior of Vespidae predators in coffee crop.” Sociobiology. 60(4):471-476.</ref><ref name="Gusmão 2000">Gusmão, M.F., Picanço, M., Alfredo, H.R.G., Moura, M.F. (2000). “Selectividade fisiológica de insecticidas a vespidae predadores do bicho-mineiro-do-cafeeiro.” Pesquisa Agropecuária Brasileira. 35(4):681-686.</ref><ref name="Reis 2000">Reis Jr., R., DeSouza, O., Vilela, E.F. (2000). “Predators impairing the natural biological control of parasitoids.” Anais da Sociedade Entomológica do Brasil. 29(3):507-514.</ref>, and has been documented in a number of papers cataloging flower visitation by insects and birds <ref name="Santos 2006">Santos, G.M.D.M., Aguiar, C.M.L., Gobbi, N. (2006). “Characterization of the social wasp guild (Hymenoptera: Vespidae) visiting flowers in the Caatinga (Itatim, Bahia, Brazil).” Sociobiology. 47(2):1-12.</ref><ref name="Traveset 2013">Traveset, A., Heleno, R., Chamorro, S., Vargas, P., McMullen, C.K., Castro-Urgal, R., Nogales, M., Herrera, H.W., Olesen, J.M. (2013). “Invaders of pollination networks in the Galápagos Islands: emergence of novel communities.” Proceedings of the Royal Society B. 280:1-9.</ref>.
In 1923, Bequeart performed a survey of the literature on this species, compiling accounts and descriptions by entomologists [[Ferdinand de Saussure|de Saussure]], [[Hermann von Ihering|von Ihering]], and du Buysson among numerous others, creating a source of descriptive information on this species <ref name="Bequaert" />. One study has focused on the morphological caste differentiation of ''B. lecheguana'' <ref name="Shima">Shima, S.N., Noll, F.B., Zucchi, R. (2000). “Morphological caste differences in the Neotropical swarm-founding Polistine wasp, Brachygastra lecheguana (Hymenoptera: Vespidae, Polistinae, Epiponini).” Sociobiology. 36(1):41-52.</ref> and another directly studied its [[Foraging|foraging behavior]] <ref name="Alves-Silva 2013" />. While not the focus, ''B. lecheguana'' has been included in studies of groups of wasps known to be [[natural enemies]] of agricultural pests <ref name="Fernandes 2013">Fernandes, F.L., Da Silva, P.R., Gorri, J., Pucci, L.F., Da Silva, I.W. (2013). “Selectivity of old and new organophosphate insecticides and behavior of Vespidae predators in coffee crop.” Sociobiology. 60(4):471-476.</ref><ref name="Gusmão 2000">Gusmão, M.F., Picanço, M., Alfredo, H.R.G., Moura, M.F. (2000). “Selectividade fisiológica de insecticidas a vespidae predadores do bicho-mineiro-do-cafeeiro.” Pesquisa Agropecuária Brasileira. 35(4):681-686.</ref><ref name="Reis 2000">Reis Jr., R., DeSouza, O., Vilela, E.F. (2000). “Predators impairing the natural biological control of parasitoids.” Anais da Sociedade Entomológica do Brasil. 29(3):507-514.</ref>, and has been documented in a number of papers cataloging flower visitation by insects and birds <ref name="Santos 2006">Santos, G.M.D.M., Aguiar, C.M.L., Gobbi, N. (2006). “Characterization of the social wasp guild (Hymenoptera: Vespidae) visiting flowers in the Caatinga (Itatim, Bahia, Brazil).” Sociobiology. 47(2):1-12.</ref><ref name="Traveset 2013">Traveset, A., Heleno, R., Chamorro, S., Vargas, P., McMullen, C.K., Castro-Urgal, R., Nogales, M., Herrera, H.W., Olesen, J.M. (2013). “Invaders of pollination networks in the Galápagos Islands: emergence of novel communities.” Proceedings of the Royal Society B. 280:1-9.</ref>.
== Description and Identification ==
== Description and Identification ==
Bequaert provides a detailed description of the anatomy and features of ''B. lecheguana'' <ref name="Bequaert" />. This species is black, with an abdomen colored with yellow stripes, having patches on the [[thorax]] and head colored yellow as well <ref name="Henrique-Simões 2011" />. The abdomen is wide and truncate <ref name="Andena 2012" />, and nests are globular <ref name="Henrique-Simões 2011" />, grey, and close to the ground <ref name="Bequaert" />. Queens and workers were recorded to be ~7.5-9 mm long and males ranged from ~7.5-8 mm in length <ref name="Bequaert" />. The species ''B. lecheguana'' can be differentiated from ''[[Brachygastra mellifica|B. mellifica]]'' based on the male reproductive anatomy; in ''B. lecheguana'' the males have an apically wide digitus while the males of ''B. mellifica'' have a comparatively long digitus <ref name="Andena 2012" />.
Bequaert provides a detailed description of the anatomy and features of ''B. lecheguana'' <ref name="Bequaert" />. This species is black, with an abdomen colored with yellow stripes, having patches on the [[thorax]] and head colored yellow as well <ref name="Henrique-Simões 2011" />. The abdomen is wide and truncate <ref name="Andena 2012" />, and nests are globular <ref name="Henrique-Simões 2011" />, grey, and close to the ground <ref name="Bequaert" />. Queens and workers were recorded to be ~7.5-9 mm long and males ranged from ~7.5-8 mm in length <ref name="Bequaert" />. The species ''B. lecheguana'' can be differentiated from ''[[Brachygastra mellifica|B. mellifica]]'' based on the male reproductive anatomy; in ''B. lecheguana'' the males have an apically wide digitus while that of ''B. mellifica'' males is relatively narrow <ref name="Andena 2012" />.
[[File:Brachygastra.lecheguana2.jpg.png|thumb|''B. lecheguana'' individual (lateral view).
[[File:Brachygastra.lecheguana2.jpg.png|thumb|''B. lecheguana'' individual (lateral view).


Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)]]
Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)]]
== Distribution and Habitat ==
== Distribution and Habitat ==
B. lecheguana is found across [[Central and South America]], ranging from [[Mexico]] to [[Argentina]] <ref name="Bequaert" />. This species has been observed a couple times as far north as [[Texas]] and [[Arizona]], but tends to be rare north of Mexico <ref name="Bequaert" />.
''B. lecheguana'' is found across [[Central and South America]], ranging from [[Mexico]] to [[Argentina]] <ref name="Bequaert" />. This species has been observed a couple times as far north as [[Texas]] and [[Arizona]], but tends to be rare north of Mexico <ref name="Bequaert" />.


[[Nests]] are oval, about the size of a human head, made of gray paper-like material, close to the ground. Bequeart includes a lengthy description of the construction process for one of these nests in his 1923 publication on Polybiinae wasps <ref name="Bequaert" />. Nests are located in undergrowth <ref name="Henrique-Simões 2011" />, and it appears that these wasps prefer more [[temperate]] locations characterized by open, less humid, grassland environments <ref name="Hermes 2006">Hermes, M.G. & Köhler, A. (2006). “The flower-visiting social wasps (Hymenoptera, Vespidae, Polistinae) in two areas of Rio Grande do Sul State, southern Brazil.” Revista Brasileira de Entomologia. 50(2):268-274.</ref><ref name="Santos 2006" /><ref name="Souza 2014">De Souza, M.M., Pires, E.P., Prezoto, F. (2014). “Seasonal richness and composition of social wasps (Hymenoptera, Vespidae) in areas of cerrado biome in Barroso, Minas Gerais, Brazil.” Bioscience Journal. 30(2):539-545.</ref>.
[[Nests]] are oval, about the size of a human head, made of gray paper-like material, and close to the ground <ref name="Bequaert" />. Bequeart includes a lengthy description of the construction process for one of these nests in his 1932 publication on Polybiinae wasps <ref name="Bequaert" />. Nests are located in undergrowth <ref name="Henrique-Simões 2011" />, and it appears that these wasps prefer more [[temperate]] locations characterized by open, less humid, grassland environments <ref name="Hermes 2006">Hermes, M.G. & Köhler, A. (2006). “The flower-visiting social wasps (Hymenoptera, Vespidae, Polistinae) in two areas of Rio Grande do Sul State, southern Brazil.” Revista Brasileira de Entomologia. 50(2):268-274.</ref><ref name="Santos 2006" /><ref name="Souza 2014">De Souza, M.M., Pires, E.P., Prezoto, F. (2014). “Seasonal richness and composition of social wasps (Hymenoptera, Vespidae) in areas of cerrado biome in Barroso, Minas Gerais, Brazil.” Bioscience Journal. 30(2):539-545.</ref>.
[[File:Brachygastra.lecheguana3.jpg.png|thumb|''B. lecheguana'' nest.
[[File:Brachygastra.lecheguana3.jpg.png|thumb|''B. lecheguana'' nest.


Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)]]
Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)]]
== Colony Cycle ==
== Colony Cycle ==
As a member of the polistine wasp tribe [[Epiponini]], ''B. lecheguana'' are a [[Swarm behaviour|swarm-founding]] species <ref name="Noll 2004" />. Their nests are [[polygynous]] <ref name="Noll 2004" />, with reproductively active females numbering anywhere from 1% <ref name="Shima" /> to possibly even 17% of a colony <ref name="Sugden 1994">Sugden, E.A. & McAllen, R.L. (1994). “Observations on foraging and nest biology of the Mexican honey wasp, Brachygastra mellifica (Say) in Texas (Vespidae: Polybiinae).” Journal of the Kansas Entomological Society. 67(2):141-155.</ref>. Reports of the total number of individuals in a given colony have been recorded by a couple different sources as 13800 <ref name="Shima" /> and 15000 <ref name="Bequaert" />, although it appears that a limited number of colony counts are documented in the scientific literature. The male to female ratio was reported as 0 in a colony collected in May <ref name="Shima" />, 1:15 in a colony collected in January, and 1:1 in a colony observed in the fall <ref name="Bequaert" />.
As a member of the polistine wasp tribe [[Epiponini]], ''B. lecheguana'' is a [[Swarm behaviour|swarm-founding]] species <ref name="Noll 2004" />. Their nests are [[polygynous]] <ref name="Noll 2004" />, with reproductively active females numbering anywhere from 1% <ref name="Shima" /> to possibly even 17% of a colony <ref name="Sugden 1994">Sugden, E.A. & McAllen, R.L. (1994). “Observations on foraging and nest biology of the Mexican honey wasp, Brachygastra mellifica (Say) in Texas (Vespidae: Polybiinae).” Journal of the Kansas Entomological Society. 67(2):141-155.</ref>. Reports of the total number of individuals in a given colony have been recorded by a couple different sources as ranging from 13800 <ref name="Shima" /> to 15000 <ref name="Bequaert" />, although it appears that a limited number of colony counts are documented in the scientific literature. The male to female ratio was reported as 0 in a colony collected in May <ref name="Shima" />, 1:15 in a colony collected in January, and 1:1 in a colony observed in the fall <ref name="Bequaert" />.


Nests are recorded to be [[perennial]], often lasting several years <ref name="Bequaert" />. New [[colonies]] are formed in the spring when several fertile queens, accompanied by a few worker wasps in a swarm, establish new nest sites <ref name="Bequaert" />.
Nests are recorded to be [[perennial]], often lasting several years <ref name="Bequaert" />. New [[colonies]] are formed in the spring when several fertile queens, accompanied by a few worker wasps in a swarm, establish new nest sites <ref name="Bequaert" />.
== Caste Differences ==
== Caste Differences ==
In a study that characterized the morphological difference between different [[Caste (disambiguation)|castes]] in ''B. lecheguana'' used both [[ovary]] development to classify workers, queens, and an “intermediate” group <ref name="Shima" />. Workers had either undeveloped or slightly developed ovaries, and constituted around 39% of the studies’ pool of individuals, while queens, with well developed ovaries containing 6-12 [[oocytes]], accounted for around 0.7% of the individuals <ref name="Shima" />. A third group, that with moderately developed ovaries containing 1-5 oocytes was described as the intermediate caste, and accounted for around 60.3% of the colony <ref name="Shima" />. While the ovaries belonging to queen individuals had evidence of [[insemination]], there was no detectable [[sperm]] in the reproductive organs of the intermediate individuals <ref name="Shima" />.
In a study that characterized the morphological difference between different [[Caste (disambiguation)|castes]] in ''B. lecheguana'', researchers used [[ovary]] development to classify individuals as workers, queens, or “intermediates” <ref name="Shima" />. Workers had either undeveloped or slightly developed ovaries, and constituted around 39% of the study's pool of individuals, while queens, with well developed ovaries containing 6-12 [[oocytes]], accounted for around 0.7% of the individuals <ref name="Shima" />. A third group, with moderately developed ovaries containing 1-5 oocytes, was identified as an intermediate caste, and accounted for around 60.3% of the colony <ref name="Shima" />. While the ovaries belonging to queen individuals had evidence of [[insemination]], there was no detectable [[sperm]] in the reproductive organs of the intermediate individuals <ref name="Shima" />.


While anecdotal accounts of B. lecheguana suggest that queens are larger than workers <ref name="Bequaert" />, it appears through more careful statistical analysis that there is no overall size difference between queens, workers, or intermediates <ref name="Shima" />. There is no difference in overall size, but the queens have a set of anatomical proportions distinct from the intermediates and workers, smaller in some structures, and larger in others <ref name="Shima" />.
Anecdotal accounts of ''B. lecheguana'' nests suggest that queens are larger than workers <ref name="Bequaert" />. However it appears that a more careful statistical analysis indicates that there is no overall size difference between queens, workers, or intermediates <ref name="Shima" />. There is no difference in overall size, but the queens have a set of anatomical proportions distinct from the intermediates and workers, smaller in some structures, and larger in others <ref name="Shima" />.


While it appears that caste might be discernible through ovary development <ref name="Shima" />, the absence of morphological differences between individuals in ''B. lecheguana'' suggests that there is no meaningful morphological caste <ref name="Noll 2004" />. However it is clear that individuals can be designated reproductives and nonreproductives <ref name="Noll 2004" />.
While it appears that caste might be discernible through ovary development <ref name="Shima" />, the absence of morphological differences between individuals in ''B. lecheguana'' suggests that there is no meaningful morphological caste <ref name="Noll 2004" />. However it is clear that individuals can be designated reproductives and nonreproductives <ref name="Noll 2004" />.
== Mimicry ==
== Mimicry ==
Bequaert describes the extensive color and shape similarities between ''B. lecheguana'' and a number of other species, using the word “homeochromic” to name these similarities instead of “[[Mimicry|mimic]]” <ref name="Bequaert" />. He lists a number of [[Vespidae|vespid wasps]], [[Potter wasp|potter wasps]], belonging to the genera ''[[Pachodynerus]]'', ''[[Odynerus]]'', and ''[[Ancistrocerus]]'', along with a number of [[Crabronidae|crabronid wasps]] of the genera ''Gorytes'' and ''[[Cerceris]]'' <ref name="Bequaert" />. Along with wasps, there are several bee species that are homeochromic, including [[Mason bee|mason bees]] of the genera ''[[Megachile]]'', ''Anthidiellum'', ''Hypanthidium'', and ''Dianthidium'', [[Cuckoo bees]] belonging to the genus ''Epeolus'', stingless ''[[Trigona]]'' bees, and [[Sweat bee|sweat bees]] from the genus ''[[Halictus]]'' <ref name="Bequaert" />. He also lists a [[soldier fly]] of the genus ''[[Stratiomys]]'' as a homeochromic species <ref name="Bequaert" />. While it is unclear what species he is referring to, he cites ''[[Stelis]] costaricensis'' as a homeochromic species, and while ''Stelis'' is a genus of [[orchid]], he did not elaborate enough to rule out the possibility that he was simply using an outdated name for another other insect <ref name="Bequaert" />.
Bequaert describes the extensive color and shape similarities between ''B. lecheguana'' and a number of other species, using the word “homeochromic” to name these similarities instead of “[[Mimicry|mimic]]” <ref name="Bequaert" />. He lists a number of [[Vespidae|vespid wasps]], [[Potter wasp|potter wasps]] belonging to the genera ''[[Pachodynerus]]'', ''[[Odynerus]]'', and ''[[Ancistrocerus]]'', and several [[Crabronidae|crabronid wasps]] of the genera ''Gorytes'' and ''[[Cerceris]]'' <ref name="Bequaert" />. Along with wasps, there are several bee species that are homeochromic, including [[Mason bee|mason bees]] of the genera ''[[Megachile]]'', ''Anthidiellum'', ''Hypanthidium'', and ''Dianthidium'', [[Cuckoo bees]] belonging to the genus ''Epeolus'', stingless ''[[Trigona]]'' bees, and [[Sweat bee|sweat bees]] from the genus ''[[Halictus]]'' <ref name="Bequaert" />. He also lists a [[soldier fly]] of the genus ''[[Stratiomys]]'' as a homeochromic species <ref name="Bequaert" />. While it is unclear what species he is referring to, he cites ''[[Stelis]] costaricensis'' as a homeochromic species, and while ''Stelis'' is a genus of [[orchid]], he did not elaborate enough to rule out the possibility that he was simply using an outdated name for another insect <ref name="Bequaert" />.


While ''B. lecheguana'', Bequaert suggests, is the most abundant of this group of species that all share color and morphological similarities, he does not make the claim that these similarities arose from [[mimicry]] in the classic sense, but he suggests that further research would be helpful <ref name="Bequaert" />.
While ''B. lecheguana'', Bequaert suggests, is the most abundant in this group of homeochromic species, he does not make the claim that these similarities arose from [[mimicry]] in the classic sense, but he suggests that further research would be helpful <ref name="Bequaert" />.
== Diet ==
== Diet ==
=== Nectar ===
=== Nectar ===
B. lecheguana is known to visit a number of plants to obtain [[nectar]] resources <ref name="Alves-Silva 2013" />. This wasp has been documented visiting the flowers of ''[[Baccharis]] spp.'' <ref name="Freitas 2006">Freitas, L., & Sazima, M. (2006). “Pollination biology in a tropical high-altitude grassland in Brazil: interactions at the community level.” Annals of the Missouri Botanical Garden. 93(3):465-516.</ref>, ''[[Erythrina crista-galli|Erythrina crista-galli]]'' <ref name="Galetto 2000">Galetto, L., Bernardello, G., Isele, I.C., Vesprini, J., Speroni G., Berduc, A. (2000). “Reproductive biology of Erythrina crista-galli (Fabaceae).” Annals of the Missouri Botanical Garden. 87(2):127-145.</ref>, ''[[Ziziphus]] cotinifolia'', ''[[Solanum paniculatum]]'', ''Sidastrum paniculatum'', and ''Erythroxylon catingae'' <ref name="Santos 2006" /> and more <ref name="Bequaert" />. In addition to obtaining nectar from flowers, ''B. lecheguana'' has been reported as a visitor of extrafloral nectaries on such species as ''[[Banisteriopsis|Banisteriopsis]] malifolia'' <ref name="Alves-Silva 2013" />.
B. lecheguana is known to visit a number of plants to obtain [[nectar]] resources <ref name="Alves-Silva 2013" />. This wasp has been documented visiting the flowers of ''[[Baccharis]] spp.'' <ref name="Freitas 2006">Freitas, L., & Sazima, M. (2006). “Pollination biology in a tropical high-altitude grassland in Brazil: interactions at the community level.” Annals of the Missouri Botanical Garden. 93(3):465-516.</ref>, ''[[Erythrina crista-galli|Erythrina crista-galli]]'' <ref name="Galetto 2000">Galetto, L., Bernardello, G., Isele, I.C., Vesprini, J., Speroni G., Berduc, A. (2000). “Reproductive biology of Erythrina crista-galli (Fabaceae).” Annals of the Missouri Botanical Garden. 87(2):127-145.</ref>, ''[[Ziziphus]] cotinifolia'', ''[[Solanum paniculatum]]'', ''Sidastrum paniculatum'', and ''Erythroxylon catingae'' <ref name="Santos 2006" /> and more <ref name="Bequaert" />. In addition to obtaining nectar from flowers, ''B. lecheguana'' has been reported as a visitor of extrafloral nectaries on such species as ''[[Banisteriopsis|Banisteriopsis]] malifolia'' <ref name="Alves-Silva 2013" />.
=== Predation ===
=== Predation ===
This wasp does not rely solely on nectar for food, as it is a well-documented predator of a number of different [[arthropod]] species <ref name="Reyes-Roras 2011">Reyes-Rosas, M.A., López-Arroyo, J.I., Buck, M., Loera-Gallardo, J. (2011). “First report of a predaceous wasp attacking nymphs of Diaphorinacitri (Hemiptera: Psyllidae), Vector of Hlb.” Florida Entomologist. 94(4):1075-1077.</ref>. Among its prey is the [[beetle]] genus ''[[Anthonomus]]'' <ref name="Alves-Silva 2013" /><ref name="Reyes-Roras 2011" /> and the [[Leaf miner|coffee leaf miner]] ''[[Leucoptera|Leucoptera coffella]]'' <ref name="Gusmão 2000" />. It is known to forage on a number of plant species seeking out insects including ''[[Eugenia uniflora]]'' fruits <ref name="Souza 2013">Souza, G.K., Pikart, T.G., Jacques, G.C., Castro, A.A., DeSouza, M.M., Serrão, J.E., Zanuncio, J.C. (2013). “Social wasps on Eugenia uniflora Linnaeus (Myrtaceae) plants in an urban area.” Sociobiology. 60(2):204-209.</ref>, ''[[Banisteriopsis]] malifolia'' <ref name="Alves-Silva 2013" />, and a number of agriculturally relevant plants <ref name="Santos 2010">Santos, G.M.D.M. & Presley, S.J. (2010). “Niche overlap and temporal activity patterns of social wasps (Hymenoptera: Vespidae) in a Brazilian cashew orchard.” Sociobiology. 56(1):121-131.</ref><ref name="Souza 2011">De Souza, A.R., Venancio, D.F.A., Zanuncio, J.C., Prezoto, F. (2011). “Sampling methods for assessing social wasps species diversity in a Eucalyptus plantation.” Journal of Economic Entomology. 104(3):1120-1123.</ref><ref name="Reis 2000" />.
This wasp does not rely solely on nectar for food; it is a well-documented predator of a number of different [[arthropod]] species <ref name="Reyes-Roras 2011">Reyes-Rosas, M.A., López-Arroyo, J.I., Buck, M., Loera-Gallardo, J. (2011). “First report of a predaceous wasp attacking nymphs of Diaphorinacitri (Hemiptera: Psyllidae), Vector of Hlb.” Florida Entomologist. 94(4):1075-1077.</ref>. Among its prey are [[beetles]] of the genus ''[[Anthonomus]]'' <ref name="Alves-Silva 2013" /><ref name="Reyes-Roras 2011" /> and the [[Leaf miner|coffee leaf miner]] ''[[Leucoptera|Leucoptera coffella]]'' <ref name="Gusmão 2000" />. It is known to forage on a number of plant species including ''[[Eugenia uniflora]]'' fruits <ref name="Souza 2013">Souza, G.K., Pikart, T.G., Jacques, G.C., Castro, A.A., DeSouza, M.M., Serrão, J.E., Zanuncio, J.C. (2013). “Social wasps on Eugenia uniflora Linnaeus (Myrtaceae) plants in an urban area.” Sociobiology. 60(2):204-209.</ref>, ''[[Banisteriopsis]] malifolia'' <ref name="Alves-Silva 2013" />, and several agriculturally relevant plants <ref name="Santos 2010">Santos, G.M.D.M. & Presley, S.J. (2010). “Niche overlap and temporal activity patterns of social wasps (Hymenoptera: Vespidae) in a Brazilian cashew orchard.” Sociobiology. 56(1):121-131.</ref><ref name="Souza 2011">De Souza, A.R., Venancio, D.F.A., Zanuncio, J.C., Prezoto, F. (2011). “Sampling methods for assessing social wasps species diversity in a Eucalyptus plantation.” Journal of Economic Entomology. 104(3):1120-1123.</ref><ref name="Reis 2000" />.
== Species Interactions ==
== Species Interactions ==
=== Competition ===
=== Competition ===
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The limited information on this species documents very few instances of [[parasitism]], although it is unlikely that this wasp is free from parasites <ref name="Bequaert" />. One report suggests that ''B. lecheguana'' is subject to parasitism by [[Strepsiptera]] <ref name="Bequaert" />.
The limited information on this species documents very few instances of [[parasitism]], although it is unlikely that this wasp is free from parasites <ref name="Bequaert" />. One report suggests that ''B. lecheguana'' is subject to parasitism by [[Strepsiptera]] <ref name="Bequaert" />.
=== Predators ===
=== Predators ===
B. lecheguana is a prey target for a number of different animals <ref name="Jeanne 1975">Jeanne, R.L. (1975). “The adaptiveness of social wasp nest architecture.” The Quarterly Review of Biology. 50(3):267-287.</ref>. The [[lizard]] ''[[Tupinambis teguixin]]'' is known to prey on this wasp, having easy access to its nests since they are low to the ground and easily accessible <ref name="Bequaert" />. A number of [[birds]] are suspected to prey on B. lecheguana, although there are relatively few documentations of this in the literature <ref name="Bequaert" /><ref name="Costa Neto 2004" />. Also the army ant ''[[Eciton]] dulcius'' is reported to prey on this wasp <ref name="Jeanne 1975" />. Another source of predation is by [[Asilidae|asilids]], or robber flies <ref name="Bequaert" />.
''B. lecheguana'' is a prey target for a number of different animals <ref name="Jeanne 1975">Jeanne, R.L. (1975). “The adaptiveness of social wasp nest architecture.” The Quarterly Review of Biology. 50(3):267-287.</ref>. The [[lizard]] ''[[Tupinambis teguixin]]'' is known to prey on this wasp, having easy access to its nests since they are low to the ground and easily accessible <ref name="Bequaert" />. A number of [[birds]] are suspected to prey on B. lecheguana, although there are relatively few documentations of this in the literature <ref name="Bequaert" /><ref name="Costa Neto 2004" />. Also the army ant ''[[Eciton]] dulcius'' is reported to prey on this wasp <ref name="Jeanne 1975" />. Another source of predation is by [[Asilidae|asilids]], or robber flies <ref name="Bequaert" />.
== Human Interactions ==
== Human Interactions ==
=== Agriculture ===
=== Agriculture ===
''B. lecheguana'' is a known natural enemy of the coffee leaf miner ''[[Leucoptera]] coffeella'', along with several other species of wasp including ''[[Polistes]] versicolor'', ''[[Polybia]] paulista'', <ref name="Scalon 2011">Scalon, J.D., Avelar, M.B.L., Alves, G.F., Zacarias, M.S. (2011). “Spatial and temporal dynamics of coffee-leaf-miner and predatory wasps in organic coffee field in formation.” Ciencia Rural, Santa Maria. 41(4):646-652.</ref> ''[[Polybia occidentalis]]'' <ref name="Gusmão 2000" />, ''[[Polybia]] scutellaris'', and ''Protonectarina sylveirae'' <ref name="Reis 2000" />. They are often used as [[biological controls]] in [[coffee]] [[plantations]] in order to prevent the proliferation of the devastating leaf miner <ref name="Fernandes 2013" />. In order to test the effect [[pesticides]] have on the environment, and to test the efficacy of using pesticides in conjunction with natural predators, the effect of a number of pesticides was described for ''B. lecheguana'' and other wasps <ref name="Fernandes 2013" />. This wasp species is found to be highly susceptible to [[chlorpyrifos]] and other [[organophosphate]] pesticides <ref name="Fernandes 2013" />. Even at half the recommended dosage of a number of these pesticides, the wasp was found to have nearly 100% mortality <ref name="Gusmão 2000" />.
''B. lecheguana'' is a known natural enemy of the coffee leaf miner ''[[Leucoptera]] coffeella'', along with several other species of wasp including ''[[Polistes]] versicolor'', ''[[Polybia]] paulista'', <ref name="Scalon 2011">Scalon, J.D., Avelar, M.B.L., Alves, G.F., Zacarias, M.S. (2011). “Spatial and temporal dynamics of coffee-leaf-miner and predatory wasps in organic coffee field in formation.” Ciencia Rural, Santa Maria. 41(4):646-652.</ref> ''[[Polybia occidentalis]]'' <ref name="Gusmão 2000" />, ''[[Polybia]] scutellaris'', and ''Protonectarina sylveirae'' <ref name="Reis 2000" />. They are often used as [[biological controls]] in [[coffee]] [[plantations]] in order to prevent the proliferation of the devastating leaf miner <ref name="Fernandes 2013" />. In order to test the effect [[pesticides]] have on the environment, and to test the efficacy of using pesticides in conjunction with natural predators, the effect that several pesticides have on the health of ''B. lecheguana'' and other wasps was assayed <ref name="Fernandes 2013" />. This wasp species is found to be highly susceptible to [[chlorpyrifos]] and other [[organophosphate]] pesticides <ref name="Fernandes 2013" />. Even at half the recommended dosage of a number of these pesticides, the wasp was found to have nearly 100% mortality <ref name="Gusmão 2000" />.


The wasp has also been documented foraging in [[cashew]] farms <ref name="Santos 2010" /> and inhabiting [[eucalyptus]] plantations <ref name="Souza 2011" />.
In addition to coffee plantations wasp has also been documented foraging in [[cashew]] farms <ref name="Santos 2010" /> and inhabiting [[eucalyptus]] plantations <ref name="Souza 2011" /> where it may play a role in pest control.
=== Stings and Aggressiveness ===
=== Stings and Aggressiveness ===
The [[aggressiveness]] of this wasp is disputed <ref name="Bequaert" /><ref name="Henrique-Simões 2011" /><ref name="Costa Neto 2004" />. While some report that the wasp is rather unaggressive even when disturbed <ref name="Bequaert" />, others warn of its aggressive behavior towards human victims <ref name="Henrique-Simões 2011" /><ref name="Costa Neto 2004" />. Having a moderately painful sting <ref name="Starr 1985">Starr, C.K. (1985). “A simple pain scale for field comparison of Hymenopteran Stings.” Journal of Entomological Science. 20(2):225-232.</ref>, ''B. lecheguana'' uses an [[Autotomy|autotomous]] stinging strategy, employing a barbed stinger specialized for [[venom]] delivery and release of the stinger in the sting site <ref name="Shorter 2012">Shorter, J.R. & Rueppell, O. (2012). “A review on self-destructive defense behaviors in social insects.” Insectes Sociaux. 59:1-10.</ref>.
The [[aggressiveness]] of this wasp is disputed <ref name="Bequaert" /><ref name="Henrique-Simões 2011" /><ref name="Costa Neto 2004" />. While some report that the wasp is rather unaggressive even when disturbed <ref name="Bequaert" />, others warn of its aggressive behavior towards human victims <ref name="Henrique-Simões 2011" /><ref name="Costa Neto 2004" />. Having a moderately painful sting <ref name="Starr 1985">Starr, C.K. (1985). “A simple pain scale for field comparison of Hymenopteran Stings.” Journal of Entomological Science. 20(2):225-232.</ref>, ''B. lecheguana'' uses an [[Autotomy|autotomous]] stinging strategy, employing a barbed stinger specialized for [[venom]] delivery and release of the stinger in the sting site <ref name="Shorter 2012">Shorter, J.R. & Rueppell, O. (2012). “A review on self-destructive defense behaviors in social insects.” Insectes Sociaux. 59:1-10.</ref>.
Line 81: Line 81:
Some sources indicate that the [[honey]] from the ''B. lecheguana'' hives is harvested regularly and consumed <ref name="Bequaert" />. In some places in Mexico it appears that ''B. lecheguana'' is maintained in a state of semi-[[domestication]] <ref name="Costa Neto 2004" />. It is warned, however, that the honey can be [[poisonous]] at times <ref name="Costa Neto 2004" />. The toxicity of the honey is a result of the wasp collecting nectar during the ''[[Datura]]'' blooming season <ref name="Bequaert" />.
Some sources indicate that the [[honey]] from the ''B. lecheguana'' hives is harvested regularly and consumed <ref name="Bequaert" />. In some places in Mexico it appears that ''B. lecheguana'' is maintained in a state of semi-[[domestication]] <ref name="Costa Neto 2004" />. It is warned, however, that the honey can be [[poisonous]] at times <ref name="Costa Neto 2004" />. The toxicity of the honey is a result of the wasp collecting nectar during the ''[[Datura]]'' blooming season <ref name="Bequaert" />.
=== Medicine ===
=== Medicine ===
Ethnoentomologists and [[anthropologists]] have documented use of ''B. lecheguana'' as a medical treatment in certain communities <ref name="Costa Neto 2000">Costa Neto, E.M. (2002). “The use of insects in folk medicine in the state of Bahia, Northeastern Brazil, with notes on insects reported elsewhere in Brazilian folk medicine.” Human Ecology. 30(2):245-263</ref><ref name="Costa Neto 2000" />. Some eat both the honey and the larvae of the wasp <ref name="Costa Neto 2004" />, while others use the adult wasps themselves <ref name="Costa Neto 2000" />. It appears that the honey has been used to treat [[coughing]] or [[asthma]] in the Pankararé and [[Matinha]] communities in [[Brazil]] <ref name="Costa Neto 2000" />. The wasp itself has been used in the [[Serrinha]] area as a pain relief for stings <ref name="Costa Neto 2000" />.
Ethnoentomologists and [[anthropologists]] have documented use of ''B. lecheguana'' as a medical treatment in certain communities <ref name="Costa Neto 2000">Costa Neto, E.M. (2002). “The use of insects in folk medicine in the state of Bahia, Northeastern Brazil, with notes on insects reported elsewhere in Brazilian folk medicine.” Human Ecology. 30(2):245-263</ref><ref name="Costa Neto 2000" />. Some eat both the honey and the larvae of the wasp <ref name="Costa Neto 2004" />, and others use the adult wasps themselves <ref name="Costa Neto 2000" />. It appears that the honey has been used to treat [[coughing]] or [[asthma]] in the Pankararé and [[Matinha]] communities in [[Brazil]] <ref name="Costa Neto 2000" />. The wasp itself has been used in the [[Serrinha]] area as a pain relief for stings <ref name="Costa Neto 2000" />.


== References ==
== References ==

Revision as of 04:46, 26 September 2014

Brachygastra lecheguana
File:Brachygastra.lecheguana1.jpg.png
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Hymenoptera
Suborder:
Apocrita
Superfamily:
Vespoidea
Family:
Vespidae
Subfamily:
Polistinae
Tribe:
Epiponini
Genus:
Brachygastra

Perty, 1833
Species:
lecheguana

Latreille, 1824

Brachygastra lecheguana (Latreille 1824), formerly known as Nectarina lecheguana [1], is a species of dark paper wasp [2] found across Central and South America [1]. It nests in underbrush in grassland-type environments, and produces honey, characteristic of the genus Brachygastra [1]. Common names include marimbondo-do-campo, marimbondo-do-pasto [3], marimbondo-exu, marimbondo-de-pote, marimbondo-de-purrão, and avispa-de-pote [4], among others depending on the location.

Taxonomy

B. lecheguana belongs to the Epiponini tribe of Polistinae wasps [5], sometimes referred to as Polybiinae wasps [1][5]. It is a Neotropical social wasp species that falls under the genus Brachygastra [5].

Brachygastra wasps are identified, as noted in Andena & Carpenter (2012), by an anatomical projection over their posterior. The scutellum, metanotum, and propodeum, three of the hymenopteran back structures, combined form a flat, vertical surface in the middle of the body (mesosoma) [6].

B. lecheguana is very closely related to B. mellifica and B. borellii [6], and indeed some authors suggest that B. lecheguana and B. mellifica can only be differentiated based on where they are found with respect to their reported distribution throughout the Americas [1]. However there are morphological and genetic differences between these species [7][6][8].

History of Study

Brachygastra lecheguana was first described by Latreille in 1824 [1]. It was originally named Nectarina lecheguana [1], and the switch from Nectarina to Brachygastra as a name for this genus was not standard in the scientific literature until the 1940’s [6]. Between 1824 and 1943, this species was documented as a member of the following genera: Polistes, Vespa, Nectarinia (a misspelling of Nectarina, and a misuse of this genus of birds), Brachygaster (another misspelling, and a misuse of this genus of parasitic wasp), Melissaia, and Caba [1][6].

In 1923, Bequeart performed a survey of the literature on this species, compiling accounts and descriptions by entomologists de Saussure, von Ihering, and du Buysson among numerous others, creating a source of descriptive information on this species [1]. One study has focused on the morphological caste differentiation of B. lecheguana [9] and another directly studied its foraging behavior [2]. While not the focus, B. lecheguana has been included in studies of groups of wasps known to be natural enemies of agricultural pests [10][11][12], and has been documented in a number of papers cataloging flower visitation by insects and birds [13][14].

Description and Identification

Bequaert provides a detailed description of the anatomy and features of B. lecheguana [1]. This species is black, with an abdomen colored with yellow stripes, having patches on the thorax and head colored yellow as well [3]. The abdomen is wide and truncate [6], and nests are globular [3], grey, and close to the ground [1]. Queens and workers were recorded to be ~7.5-9 mm long and males ranged from ~7.5-8 mm in length [1]. The species B. lecheguana can be differentiated from B. mellifica based on the male reproductive anatomy; in B. lecheguana the males have an apically wide digitus while that of B. mellifica males is relatively narrow [6].

File:Brachygastra.lecheguana2.jpg.png
B. lecheguana individual (lateral view). Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)

Distribution and Habitat

B. lecheguana is found across Central and South America, ranging from Mexico to Argentina [1]. This species has been observed a couple times as far north as Texas and Arizona, but tends to be rare north of Mexico [1].

Nests are oval, about the size of a human head, made of gray paper-like material, and close to the ground [1]. Bequeart includes a lengthy description of the construction process for one of these nests in his 1932 publication on Polybiinae wasps [1]. Nests are located in undergrowth [3], and it appears that these wasps prefer more temperate locations characterized by open, less humid, grassland environments [15][13][16].

File:Brachygastra.lecheguana3.jpg.png
B. lecheguana nest. Image from Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)

Colony Cycle

As a member of the polistine wasp tribe Epiponini, B. lecheguana is a swarm-founding species [5]. Their nests are polygynous [5], with reproductively active females numbering anywhere from 1% [9] to possibly even 17% of a colony [17]. Reports of the total number of individuals in a given colony have been recorded by a couple different sources as ranging from 13800 [9] to 15000 [1], although it appears that a limited number of colony counts are documented in the scientific literature. The male to female ratio was reported as 0 in a colony collected in May [9], 1:15 in a colony collected in January, and 1:1 in a colony observed in the fall [1].

Nests are recorded to be perennial, often lasting several years [1]. New colonies are formed in the spring when several fertile queens, accompanied by a few worker wasps in a swarm, establish new nest sites [1].

Caste Differences

In a study that characterized the morphological difference between different castes in B. lecheguana, researchers used ovary development to classify individuals as workers, queens, or “intermediates” [9]. Workers had either undeveloped or slightly developed ovaries, and constituted around 39% of the study's pool of individuals, while queens, with well developed ovaries containing 6-12 oocytes, accounted for around 0.7% of the individuals [9]. A third group, with moderately developed ovaries containing 1-5 oocytes, was identified as an intermediate caste, and accounted for around 60.3% of the colony [9]. While the ovaries belonging to queen individuals had evidence of insemination, there was no detectable sperm in the reproductive organs of the intermediate individuals [9].

Anecdotal accounts of B. lecheguana nests suggest that queens are larger than workers [1]. However it appears that a more careful statistical analysis indicates that there is no overall size difference between queens, workers, or intermediates [9]. There is no difference in overall size, but the queens have a set of anatomical proportions distinct from the intermediates and workers, smaller in some structures, and larger in others [9].

While it appears that caste might be discernible through ovary development [9], the absence of morphological differences between individuals in B. lecheguana suggests that there is no meaningful morphological caste [5]. However it is clear that individuals can be designated reproductives and nonreproductives [5].

Mimicry

Bequaert describes the extensive color and shape similarities between B. lecheguana and a number of other species, using the word “homeochromic” to name these similarities instead of “mimic” [1]. He lists a number of vespid wasps, potter wasps belonging to the genera Pachodynerus, Odynerus, and Ancistrocerus, and several crabronid wasps of the genera Gorytes and Cerceris [1]. Along with wasps, there are several bee species that are homeochromic, including mason bees of the genera Megachile, Anthidiellum, Hypanthidium, and Dianthidium, Cuckoo bees belonging to the genus Epeolus, stingless Trigona bees, and sweat bees from the genus Halictus [1]. He also lists a soldier fly of the genus Stratiomys as a homeochromic species [1]. While it is unclear what species he is referring to, he cites Stelis costaricensis as a homeochromic species, and while Stelis is a genus of orchid, he did not elaborate enough to rule out the possibility that he was simply using an outdated name for another insect [1].

While B. lecheguana, Bequaert suggests, is the most abundant in this group of homeochromic species, he does not make the claim that these similarities arose from mimicry in the classic sense, but he suggests that further research would be helpful [1].

Diet

Nectar

B. lecheguana is known to visit a number of plants to obtain nectar resources [2]. This wasp has been documented visiting the flowers of Baccharis spp. [18], Erythrina crista-galli [19], Ziziphus cotinifolia, Solanum paniculatum, Sidastrum paniculatum, and Erythroxylon catingae [13] and more [1]. In addition to obtaining nectar from flowers, B. lecheguana has been reported as a visitor of extrafloral nectaries on such species as Banisteriopsis malifolia [2].

Predation

This wasp does not rely solely on nectar for food; it is a well-documented predator of a number of different arthropod species [20]. Among its prey are beetles of the genus Anthonomus [2][20] and the coffee leaf miner Leucoptera coffella [11]. It is known to forage on a number of plant species including Eugenia uniflora fruits [21], Banisteriopsis malifolia [2], and several agriculturally relevant plants [22][23][12].

Species Interactions

Competition

This species has been documented in competition for food resources with several other animal species [2][12]. B. lecheguana has been hypothesized to be in a mutualistic relationship with the plant Banisteriopsis maliflora, since the plant supplies nectar reserves to the wasp, and the wasp preys on herbivorous insect larvae living on the plant [2]. However, B. lecheguana is competing with the ant species Camponotus blandus for the plant’s resources, and consequently the wasp is typically found on ant-free plants, and has been known to interrupt in its foraging when an ant approaches [2].

Another study suggests that B. lecheguana may compete with parasitoids for coffee leaf miner larvae in an agricultural setting [12].

Pollination

Although this wasp visits a diverse collection of flowering plants [1], it is only reported as a pollinator in a limited number of instances. B. lecheguana is documented as a pollinator of Baccharis spp. [18]. Additionally, as an alien species to the Galapagos Islands, there have been reports that catalog B. lecheguana as an important pollinator on the island of Santa Cruz [14].

Parasites

The limited information on this species documents very few instances of parasitism, although it is unlikely that this wasp is free from parasites [1]. One report suggests that B. lecheguana is subject to parasitism by Strepsiptera [1].

Predators

B. lecheguana is a prey target for a number of different animals [24]. The lizard Tupinambis teguixin is known to prey on this wasp, having easy access to its nests since they are low to the ground and easily accessible [1]. A number of birds are suspected to prey on B. lecheguana, although there are relatively few documentations of this in the literature [1][4]. Also the army ant Eciton dulcius is reported to prey on this wasp [24]. Another source of predation is by asilids, or robber flies [1].

Human Interactions

Agriculture

B. lecheguana is a known natural enemy of the coffee leaf miner Leucoptera coffeella, along with several other species of wasp including Polistes versicolor, Polybia paulista, [25] Polybia occidentalis [11], Polybia scutellaris, and Protonectarina sylveirae [12]. They are often used as biological controls in coffee plantations in order to prevent the proliferation of the devastating leaf miner [10]. In order to test the effect pesticides have on the environment, and to test the efficacy of using pesticides in conjunction with natural predators, the effect that several pesticides have on the health of B. lecheguana and other wasps was assayed [10]. This wasp species is found to be highly susceptible to chlorpyrifos and other organophosphate pesticides [10]. Even at half the recommended dosage of a number of these pesticides, the wasp was found to have nearly 100% mortality [11].

In addition to coffee plantations wasp has also been documented foraging in cashew farms [22] and inhabiting eucalyptus plantations [23] where it may play a role in pest control.

Stings and Aggressiveness

The aggressiveness of this wasp is disputed [1][3][4]. While some report that the wasp is rather unaggressive even when disturbed [1], others warn of its aggressive behavior towards human victims [3][4]. Having a moderately painful sting [26], B. lecheguana uses an autotomous stinging strategy, employing a barbed stinger specialized for venom delivery and release of the stinger in the sting site [27].

Honey

Some sources indicate that the honey from the B. lecheguana hives is harvested regularly and consumed [1]. In some places in Mexico it appears that B. lecheguana is maintained in a state of semi-domestication [4]. It is warned, however, that the honey can be poisonous at times [4]. The toxicity of the honey is a result of the wasp collecting nectar during the Datura blooming season [1].

Medicine

Ethnoentomologists and anthropologists have documented use of B. lecheguana as a medical treatment in certain communities [28][28]. Some eat both the honey and the larvae of the wasp [4], and others use the adult wasps themselves [28]. It appears that the honey has been used to treat coughing or asthma in the Pankararé and Matinha communities in Brazil [28]. The wasp itself has been used in the Serrinha area as a pain relief for stings [28].

References

  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al Bequaert, J.Q. (1932). “The Nearctic social wasps of the subfamily polybiinae (Hymenoptera; Vespidae). Entomologica Americana. 13(3):87-150.
  2. ^ a b c d e f g h i Alves-Silva, E., Baronio, G.J., Torezan-Silingardi, H.M., Del-Claro, K. (2013). “Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system.” Entomological Science. 16:162-169.
  3. ^ a b c d e f Henrique-Simões, M., Cuozzo, M.D., Frieiro-Costa, F.A. (2011). “Social wasps of Unilavras/Boqueirão Biological Reserve, Ingaí, state of Minas Gerais, Brazil.” Check List: Journal of Species Lists and Distribution. 7(5)
  4. ^ a b c d e f g Costa Neto, E.M. (2004). “La etnoentomología de las avispas (Hymenoptera, Vespoidea) en el poblado de Pedra Branca, estado de Bahia, nordeste de Brasil.” Boletín de la Sociedad Entomológica Aragonesa. 34:247-262.
  5. ^ a b c d e f g Noll, F.B., Wenzel, J.W., Zucchi, R. (2004). “Evolution of caste in Neotropical swarm-founding wasps (Hymenoptera: Vespidae; Epiponini).” American Museum Novitates. 3467:1-24.
  6. ^ a b c d e f g Andena, S.R. (2012). “A phylogenetic analysis of the social wasp genus Brachygastra Perty, 1833, and Description of a new species (Hymenoptera: Vespidae: Epiponini).” American Museum Novitates. 3753:1-38
  7. ^ details of the citation
  8. ^ Zhu, Y., Queller, D.C., Strassman, J.E. (1999). “A phylogenetic perspective on sequence evolution in microsatellite loci.” Journal of Molecular Evolution. 50:324-338.
  9. ^ a b c d e f g h i j k Shima, S.N., Noll, F.B., Zucchi, R. (2000). “Morphological caste differences in the Neotropical swarm-founding Polistine wasp, Brachygastra lecheguana (Hymenoptera: Vespidae, Polistinae, Epiponini).” Sociobiology. 36(1):41-52.
  10. ^ a b c d Fernandes, F.L., Da Silva, P.R., Gorri, J., Pucci, L.F., Da Silva, I.W. (2013). “Selectivity of old and new organophosphate insecticides and behavior of Vespidae predators in coffee crop.” Sociobiology. 60(4):471-476.
  11. ^ a b c d Gusmão, M.F., Picanço, M., Alfredo, H.R.G., Moura, M.F. (2000). “Selectividade fisiológica de insecticidas a vespidae predadores do bicho-mineiro-do-cafeeiro.” Pesquisa Agropecuária Brasileira. 35(4):681-686.
  12. ^ a b c d e Reis Jr., R., DeSouza, O., Vilela, E.F. (2000). “Predators impairing the natural biological control of parasitoids.” Anais da Sociedade Entomológica do Brasil. 29(3):507-514.
  13. ^ a b c Santos, G.M.D.M., Aguiar, C.M.L., Gobbi, N. (2006). “Characterization of the social wasp guild (Hymenoptera: Vespidae) visiting flowers in the Caatinga (Itatim, Bahia, Brazil).” Sociobiology. 47(2):1-12.
  14. ^ a b Traveset, A., Heleno, R., Chamorro, S., Vargas, P., McMullen, C.K., Castro-Urgal, R., Nogales, M., Herrera, H.W., Olesen, J.M. (2013). “Invaders of pollination networks in the Galápagos Islands: emergence of novel communities.” Proceedings of the Royal Society B. 280:1-9.
  15. ^ Hermes, M.G. & Köhler, A. (2006). “The flower-visiting social wasps (Hymenoptera, Vespidae, Polistinae) in two areas of Rio Grande do Sul State, southern Brazil.” Revista Brasileira de Entomologia. 50(2):268-274.
  16. ^ De Souza, M.M., Pires, E.P., Prezoto, F. (2014). “Seasonal richness and composition of social wasps (Hymenoptera, Vespidae) in areas of cerrado biome in Barroso, Minas Gerais, Brazil.” Bioscience Journal. 30(2):539-545.
  17. ^ Sugden, E.A. & McAllen, R.L. (1994). “Observations on foraging and nest biology of the Mexican honey wasp, Brachygastra mellifica (Say) in Texas (Vespidae: Polybiinae).” Journal of the Kansas Entomological Society. 67(2):141-155.
  18. ^ a b Freitas, L., & Sazima, M. (2006). “Pollination biology in a tropical high-altitude grassland in Brazil: interactions at the community level.” Annals of the Missouri Botanical Garden. 93(3):465-516.
  19. ^ Galetto, L., Bernardello, G., Isele, I.C., Vesprini, J., Speroni G., Berduc, A. (2000). “Reproductive biology of Erythrina crista-galli (Fabaceae).” Annals of the Missouri Botanical Garden. 87(2):127-145.
  20. ^ a b Reyes-Rosas, M.A., López-Arroyo, J.I., Buck, M., Loera-Gallardo, J. (2011). “First report of a predaceous wasp attacking nymphs of Diaphorinacitri (Hemiptera: Psyllidae), Vector of Hlb.” Florida Entomologist. 94(4):1075-1077.
  21. ^ Souza, G.K., Pikart, T.G., Jacques, G.C., Castro, A.A., DeSouza, M.M., Serrão, J.E., Zanuncio, J.C. (2013). “Social wasps on Eugenia uniflora Linnaeus (Myrtaceae) plants in an urban area.” Sociobiology. 60(2):204-209.
  22. ^ a b Santos, G.M.D.M. & Presley, S.J. (2010). “Niche overlap and temporal activity patterns of social wasps (Hymenoptera: Vespidae) in a Brazilian cashew orchard.” Sociobiology. 56(1):121-131.
  23. ^ a b De Souza, A.R., Venancio, D.F.A., Zanuncio, J.C., Prezoto, F. (2011). “Sampling methods for assessing social wasps species diversity in a Eucalyptus plantation.” Journal of Economic Entomology. 104(3):1120-1123.
  24. ^ a b Jeanne, R.L. (1975). “The adaptiveness of social wasp nest architecture.” The Quarterly Review of Biology. 50(3):267-287.
  25. ^ Scalon, J.D., Avelar, M.B.L., Alves, G.F., Zacarias, M.S. (2011). “Spatial and temporal dynamics of coffee-leaf-miner and predatory wasps in organic coffee field in formation.” Ciencia Rural, Santa Maria. 41(4):646-652.
  26. ^ Starr, C.K. (1985). “A simple pain scale for field comparison of Hymenopteran Stings.” Journal of Entomological Science. 20(2):225-232.
  27. ^ Shorter, J.R. & Rueppell, O. (2012). “A review on self-destructive defense behaviors in social insects.” Insectes Sociaux. 59:1-10.
  28. ^ a b c d e Costa Neto, E.M. (2002). “The use of insects in folk medicine in the state of Bahia, Northeastern Brazil, with notes on insects reported elsewhere in Brazilian folk medicine.” Human Ecology. 30(2):245-263
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