Brachygastra | |
---|---|
Brachygastra mellifica | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Vespidae |
Subfamily: | Polistinae |
Tribe: | Epiponini |
Genus: | Brachygastra Perty, 1833 [1] |
Type species | |
Brachygastra lecheguana | |
Species | |
17 species |
Honey wasps are species in the genus Brachygastra of the family Vespidae. Brachygastra comprises 17 species of social paper wasps. The ancestral species are thought to have diverged about 32 million years ago within diverse Amazon rainforest. Subsequent speciation within the genus is thought to have mostly occurred between 23 Ma and 10 Ma, during the time of the Andean uplift when the landscape was significantly altered due to tectonic activity. The current cladistic organisation of the genus has been heavily reliant on morphological characteristics.
Brachygastra species live in colonies and construct arboreal paper nests in humid forest environments, although several species occur in open vegetation. They are widely distributed in Central and South America, and also occur in southwest areas of North America. They have a broad diet consisting of floral nectar and insect protein. Several species are known to collect and store nectar in large amounts as honey, a characteristic in the insect world only shared with various bee species, a few wasp species ( Polybia spp.), and a few ant species ( Myrmecocystus spp. and other genera). There are few historical reports of humans using honey produced by honey wasps, probably due to its small quantities and the wasp's fierce sting.
The species can provide important ecosystem services such as pollination, herbivore deterrence and predation of disease vectors.
Upon its introduction by Perty in 1833, the name Brachygastra was confused with the genus Brachygaster within the family Evaniidae. It was later changed to Nectarina, only to be confused with the genus of birds Nectarinia. Misspellings in publications further compounded the confusion. [2]
In 1991 Carpenter placed Brachygastra in a sister group of Chartergus based on morphological data. In 1993 Wenzel revised this and placed Brachygastra in a sister genus of Protonectarina + Polybia based on nest architecture. [2] Then, in 1994 Carpenter and Wenzel worked together, taking into account a combination of characteristics including adult morphology, nest architecture and larval morphology, and Brachygastra, again, was sister of Chartergus. [3] In 2004 whilst investigating the phylogeny of Epiponini, Noll et al. used discretised morphometric measurements of caste differentiation to reinforced the relationship between Brachygastra and Chartergus. [4] Since then both Arevalo et al. (2004) and Pickett and Carpenter (2010) have placed Barchygastra as a sister genus of Protonectarina after analysing molecular, morphological and behavioural data, however, both groups did not include Chartergus. [5] [6]
Brachygastra azteca is considered to be the most basal species, and a sister species to all others in the genus. [7] The rest of the genus Brachygastra has historically been divided into two main groups consisting of closely related species. The lecheguana group consists of B. mellifica, B. lecheguana, and B. borellii, and the smithii group of B. baccalaurea, B. bilineolata, B. smithii, B. propodealis, and B. buyssoni. [2] The scutellaris group, containting B. augusti, B. mouleae, B. fistulosa, B. cooperi, B. myseri and B. scutellaris, is composed of the remaining species that are apparently less closely related and not able to be placed in either of the previous groups. [7] These three groups are bound within a clade by shared characteristics of the scutellum. It is strongly angular, projects over the metanotum and is medially emarginated, usually in a V-shape. [2]
Initially the lechuguana group was thought to be one species named B. lecheguana, but in 1968, Naumann was able to discern the different species by looking at male features, giving rise to B. borellii and B. mellifica . [7] B. lecheguana and B. mellifica can only be separated by male genitalia features whilst B. borellii has distinct long hairs and deep punctation on the head and on the mesosoma. [2]
The three basal most species of the smithii group, B. bilineolata, B. moebiana, and B. smithii have a complex taxonomic history due to their morphological similarities and variations of colour patterns. B. smithii and B. moebiana were initially thought to be varieties of B. bilineolata based on colour pattern. [2] In 1968 Naumann elevated B. bilineolata and B. smithii to species level but still considered B. moebiana as a variety of B. bilineolata. Since then the occipital carina has been used to separate these species and establish B, bilineolata as the most basal species of the smithii group. [2]
The scutellaris group is composed of the remaining species that do not appear to be closely related and therefore do not fit into the lechuguana or smithii groups. [2]
The species of the genus Brachygastra are a common component of neotropical social wasp fauna. [8] [7] [9] The genus is broadly distributed throughout Central and South America, absent only from Chile, Uruguay and central and south Argentina. [9] One species, B. mellifica, occurs in southwest United States, in Texas and Arizona. [10] [7] [8]
Humid and forested environments are typical habitat for these wasps, although B. augusti, B. mouleae, B. moebiana and B. lecheguana can inhabit areas with open vegetation. [8] Two species, B. borellii and B. baccaraulea, live in upland areas. [8] The lecheguana group occurs in the Atlantic rainforest and Nearctic region, the scutellaris group lives mostly in the Atlantic forest and the smithii group is confined to the Amazon rainforest. [8] Finally, B. azteca , a sister species to all other species of the genus, occurs in Mexico. [8] [2]
The original diversification of Brachygastra probably began about 32 Ma. At that time diverse Amazonian rainforest extended all the way to the Caribbean coast of South America. [11] The Greater Antilles and Aves Ridge land bridge, GAARlandia preceded the formation of the Isthmus of Panama and is thought to have connected the Nearctic and Neotropics, allowing the ancestor of the lecheguana and azteca group to disperse between North, Central and South America. [8]
From 23 to 10 Ma, the Andean uplift accelerated and rapidly changed the Amazonian landscape, with the formation of the vast network of lakes, swamps and wetlands called the Pebas system. [11] It grew to approximately 1 million km2 in the northern portion of South America, [11] working to separate populations of ancestral Brachygastra. [8]
Fossils of social wasps are rare in the fossil record, probably due to behavioural characteristics and paper nest structures that do not lend themselves to fossilisation. [8]
Species are easily recognised by the very high scutellum that often projects over the metanotum. The metanotum and propodeum forms a flat, vertical posterior surface of the mesosoma. The abdomen is often described as truncate due to being wider than it is long (and the genus name means “short-bellied”). [6] [3] [7] Variation in the size and density of punctures on different parts of the body has been used to describe and identify species, subspecies, and forms of the genus. [2]
There are obvious morphological caste differences in many species with queens being larger than workers. Additionally, larger queens are generally more fecund and dominant that smaller counterparts. [12] Studies on morphological caste differences have been reported for B. augusti, B. bilineolata, B. lecheguana, B. moebiana, B. myersi and B. scutellaris. [10]
Nests are always arboreal and sessile. [13] They are made with various plant material consisting of minute chips and long fibres that are chewed and compacted into a paper like material. They are usually a brown or grey colour with a brittle texture. The entrances vary from circular holes or slits, sometimes with multiple entrances arranged chaotically. [13] [10] The primary comb is planar or spherical and is usually constructed in a spiral sequence. Nests can contain thousands of individuals and range in size depending on species and colony age or health. [13]
The diet of Honey wasps is made up of carbohydrates from nectar and honey, and protein attained from predation upon various insects such as beetles, weevils, lepidopterans, aphids and various larvae. [14] They have a wide feeding flexibility, probably due to the seasonal availability floral nectar and insect prey. [15] [16] [17] Floral nectar and insect protein are both stored within the comb of the nest. Brachygastra mellifica and B. lecheguana are well known for their large stores of nectar. B. azteca also stores nectar but in smaller quantities. In addition to nectar, species often collect honeydew from honey-dew producing cercopids and nymphal membracids. [9]
The nectar gathering activity of these wasp species allows them to provide pollination services to some plant types, such as myrtaceae in the Cerrado, [17] sunflower (Helianthus annuus L., Asteraceae) and mesquite in the Lower Rio Grande Valley of Texas and Avocado. [18] Additionally, honey wasps benefit some plant species by preying upon endophytic herbivorous insects. [15] [14] Damaged plants release volatile compounds that act as chemical and olfactory cues for predatory Brachygastra species, attracting them to the source of the grazing herbivorous insect. [19] They have been observed attacking many insect pest species such as Asian citrus psyllid ( Diaphorina citri), [15] [16] tomato leafminer (Tuta absoluta), [19] cotton boll weevil larvae ( Anthonomus grandis) , and the white coffee leaf miner larvae ( Perileucoptera coffeella) , [15] suggesting that they have potential to be used as a biological control agent for human crops.
Spiders commonly catch individual wasps in their webs. Opossums and woodpeckers have been observed attacking entire nests. [20]
Conservation of these wasps species is important to allow them to continue providing ecosystem services. Conservation practices include the protection of wasp nests, the maintenance of floral diversity, the preservation of forests and the use of pest control methods that do not harm these wasps. [8]
The Vespidae are a large, diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species, colonies usually last only one year, dying at the onset of winter. New queens and males (drones) are produced towards the end of the summer, and after mating, the queens hibernate over winter in cracks or other sheltered locations. The nests of most species are constructed out of mud, but polistines and vespines use plant fibers, chewed to form a sort of paper. Many species are pollen vectors contributing to the pollination of several plants, being potential or even effective pollinators, while others are notable predators of pest insect species, and a few species are invasive pests.
Paper wasps are a type of vespid wasps. The term is typically used to refer to members of the vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae and Stenogastrinae, which also make nests out of paper.
Polistes is a cosmopolitan genus of paper wasps and the only genus in the tribe Polistini. Vernacular names for the genus include umbrella wasps, coined by Walter Ebeling in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests, and umbrella paper wasps. Polistes is the single largest genus within the family Vespidae, with over 200 recognized species. Their innate preferences for nest-building sites leads them to commonly build nests on human habitation, where they can be very unwelcome; although generally not aggressive, they can be provoked into defending their nests. All species are predatory, and they may consume large numbers of caterpillars, in which respect they are generally considered beneficial.
The Polistinae is a subfamily of eusocial wasps belonging to the family Vespidae. They are closely related to the wasps and true hornets of the subfamily Vespinae, containing four tribes. With about 1,100 species total, it is the second-most diverse subfamily within the Vespidae, and while most species are tropical or subtropical, they include some of the most frequently encountered large wasps in temperate regions.
Brachygastra mellifica, commonly known as the Mexican honey wasp, is a neotropical social wasp. It can be found in North America. B. mellifica is one of few wasp species that produces honey. It is also considered a delicacy in some cultures in Mexico. This wasp species is of use to humans because it can be used to control pest species and to pollinate avocados.
The Epiponini are a large and diverse tribe of social wasps inhabiting the Neotropical region, with some species' ranges extending into the Nearctic region.
The Central American paper wasp is a nocturnal eusocial wasp. It is famous for its swarm based emigration behavior, and is native to the lowlands of Central and northern South America. This species has developed special night vision adaptations to facilitate their night-time swarming and foraging behavior and has important medicinal properties for the Pankararú people of Brazil.
Apoica flavissima is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens and workers. Unlike most Vespidae wasps, Apocia flavissima queens are smaller than their worker counterparts which results in unique intraspecies relationships.
Brachygastra lecheguana, formerly known as Nectarina lecheguana, is a species of dark paper wasp found across North and South America. It nests in underbrush in grassland-type environments, and produces honey, characteristic of the genus Brachygastra.
Agelaia vicina is a species of wasp in the genus Agelaia. They are neotropical social wasps known to have the largest colony sizes and nest sizes among social wasps, with some colonies exceeding over one million individuals. They are predators of land arthropods, consuming both insects and spiders alike. Recent sperm morphology studies have shown that although Vespidae belong to the superfamily Vespoidea, A. vicina may be more phylogenetically related to Apoidea.
Synoeca cyanea, commonly known as the marimbondo-tatu in Brazil, is a swarm-founding eusocial wasp. Native to Brazil and Argentina, S. cyanea is one of the largest and most aggressive species of social wasps and is feared in many rural areas. It begins its colony cycle in the early spring and continues until nest abandonment. Throughout its life, S. cyanea forage sugary substances and animal carcasses for food and wood pulp for its nest. S. cyanea is also known for its strong venom, which is enough to cause haemolytic activity.
Agelaia pallipes is a species of social paper wasp found from Costa Rica to Brazil, Argentina and Paraguay. A. pallipes is ground-nesting and is one of the most aggressive wasps in South America. This species is a predator of other insects, including flies, moths, and ground crickets, as well as baby birds.
Leipomeles dorsata is a neotropical paper wasp that is found across Central America and northern South America. It is a eusocial wasp with little differentiation between reproducing and non-reproducing females. In fact, workers can become temporary reproductives if the main reproductives are killed, allowing reproduction to continue until the main reproductive population recovers. The colony cycles through different ratios of main reproductive females and subordinate reproductive females, starting with few or no primary reproducing females, and increasing until there are only main reproductives.
Protopolybia exigua is a species of vespid wasp found in South America and Southern Brazil. These neotropical wasps, of the tribe Epiponini, form large colonies with multiple queens per colony. P. exigua are small wasps that find nourishment from nectar and prey on arthropods. Their nests are disc-shaped and hang from the undersides of leaves and tree branches. This particular species of wasp can be hard to study because they frequently abandon their nests. P. exigua continuously seek refuge from phorid fly attacks and thus often flee infested nests to build new ones. The wasps' most common predators are ants and the parasitoid phorid flies from the Phoridae family.
Polybia sericea is a social, tropical wasp of the family Vespidae that can be found in South America. It founds its colonies by swarming migrations, and feeds on nectar and arthropods.
Synoeca surinama is a Neotropical swarm-founding wasp of the tribe Epiponini. It is known for its metallic blue and black appearance and painful sting. S. surinama builds nests on tree trunks and can be found in tropical climates of South America. When preparing to swarm, there are a number of pre-swarming behaviors that members of S. surinama colonies partake in, such as buzzing runs and occasional brood cannibalism. In S. surinama, social environmental conditions determine the caste ranks of individuals in the developing brood. Unlike less primitive Hymenoptera species, S. surinama display little morphological variation between egg laying queens and workers. S. surinama wasps visit flowering plants and are considered pollinators. When these wasps sting, the stinger is left in the victim and the wasp ultimately dies.
Agelaia multipicta is a swarm-founding, highly eusocial wasp that lives in Mexico, Argentina, Trinidad and southern Brazil. It nests in natural cavities such as hollow trees and aggressively defends the nest from ants, who are brood predators. The workers and queens are morphologically distinguished by ovarian development as well as external features such as a larger petiole and gaster in the queen. Like other carrion-eating (necrophagous) wasp species, A. multipicta plays a scavenging role in the ecosystem. Agelaia multipicta was described by the Irish entomologist Alexander Henry Haliday in 1836.
Brachygastra scutellaris, a honey wasp, is a Neotropical, swarm-founding species that is found in South America and has a medium-sized population of 100–1000 individuals per colony. It stores large amounts of nectar in its nest for the production of honey, and it was even found that at certain times of the year, the nectar is toxic to humans, as they will extract nectar from hallucinogenic plants, depending on the season.
Protopolybia chartergoides, also known as Pseudochartergus chartergoides, is a species of wasp within the genus Protopolybia. It is a social wasp found in southern Central America and northern South America.
Parachartergus is a genus of epiponine social wasps belonging to the subfamily Polistinae. Species include:
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