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Brachygastra lecheguana

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Brachygastra lecheguana
File:Brachygastra.lecheguana1.jpg.png
Scientific classification
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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 is characterized, as noted in Andena & Carpenter (2012), by an anatomical projection over the posterior part of the insect. 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, and indeed some authors suggest that B. lecheguana and B. mellifica can be differentiated based solely on location [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 such entomologists as de Saussure, von Ihering, and du Buysson among numerous others, creating a source of descriptive information [1]. One study has focused on the morphological caste differentiation in this species [9] and another directly studied the foraging behavior of this wasp species [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 the males of B. mellifica have a comparatively long digitus [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, 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 [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 are 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 13800 [9] and 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 used both ovary development to classify workers, queens, and an “intermediate” group [9]. 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 [9]. 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 [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].

While anecdotal accounts of B. lecheguana suggest that queens are larger than workers [1], it appears through more careful statistical analysis 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, along with a number of 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 other insect [1].

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 [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, as it is a well-documented predator of a number of different arthropod species [20]. Among its prey is the beetle genus Anthonomus [2][20] and the coffee leaf miner Leucoptera coffella [11]. It is known to forage on a number of plant species seeking out insects including Eugenia uniflora fruits [21], Banisteriopsis malifolia [2], and a number of 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 of a number of pesticides was described for B. lecheguana and other wasps [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].

The wasp has also been documented foraging in cashew farms [22] and inhabiting eucalyptus plantations [23].

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], while 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 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 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