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''Protonectarina sylveirae'', commonly referred to as the Brazilian wasp, is a
neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
swarm-founding wasp species that ranges widely across South America. This species relies heavily on the consumption of animal protein rather than
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
. ''P. sylveirae'' preys heavily on agricultural pests to
coffee Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulant, stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world. S ...
crops, keeping pest populations low.


Taxonomy and phylogeny

''P. sylveirae'' belongs to the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
, which contains roughly 100,000 species of bees, ants, and wasps. The species belongs to the
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
tribe
Epiponini The Epiponini (formerly known as Polybiini) are a large and diverse tribe of social wasps inhabiting the Neotropical region, with some species' ranges extending into the Nearctic region. Selected species

* ''Apoica pallens'' * ''Leipomeles d ...
, which is characterized by complex nests, morphological differences between
castes Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
, and the occurrence of reproductive
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any o ...
. ''P. sylveirae'' is the only species within its genus, and is most closely related to genera ''
Polybia ''Polybia'' is a genus of eusocial wasps ranging from Central to South America (Mexico to Brazil, Argentina). Some produce enough honey to be collected and eaten by local people. Species *''Rutilotrixa lateralis'' (Walker, 1849) *'' Rutilotrix ...
'' and ''Protopolybia''.


Description

''Protonectarina'' is characterized by its relatively hairy eyes, and can be diagnosed by the width of the clypeus, or the front of the head. Queens have a lighter-colored body and a wider, brown spotted clypeus region compared to that of workers. The clypeus in workers features a black, horseshoe-shaped marking that varies widely among individuals. Queens are typically larger than workers, but this varies from colony to colony: a queen may even be identical in size to the workers in its colony. Age of an ''P. sylveirae'' individual can be determined from the pigmentation of the transverse
apodeme An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the l ...
across the hidden base of the fifth sternum. As a wasp ages, pigment gradually accumulates, and this region darkens.


Distribution and habitat

The distribution of ''P. sylveirae'' is wide and stretches from Brazil to Argentina. The species builds nests in trees, typically suspended from twigs. Nests are built from foraged
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
mashed with water, which forms a papery substance used to construct walls. Cylindrical nests are constructed as a series of enclosing envelopes. When building a nest, the first envelope is built from the substrate, and each subsequent envelope is added to that base. The seams between cells are characteristically irregular, and a simple entrance is built on the ventral surface of the structure. An average nest may have around 50 combs, but nests can be quite large, sometimes 30–40 centimeters long. This swarm-founding species sends several individuals to forage for pulp, and constructs the nest using the material that the pulp-foragers return with.


Colony cycle

''Protonectarina sylveirae'' colonies are typically large, which produces the need for behavioral specialization. They are swarm-founding wasps, meaning that new colonies are founded by a swarm of individuals, including multiple queens. It is common for a colony of 15,000 workers to have 250 or more queens.


Behavior


Dominance hierarchy

Queens of a colony are often tested by the workers, and their reactions to the worker's behavior can determine whether they are allowed to remain queen. Workers evaluate the number and quality of the offspring produced by the queen, and monitor the frequency of her aggressive behavior, as that suggests the capability of physical dominance. Clear morphological differences in the larger size of a potential queen compared to workers could suggest her capacity for physical dominance and reproduction, thereby suggesting her quality as a queen.


Division of labor

''Protonectarina sylveirae'' colonies are broken into two clear castes: queens and workers. These castes are morphologically different, but intermediate morphologies occur in some individuals, providing some social flexibility. The species shows diphasic
allometry Allometry is the study of the relationship of body size to shape, anatomy, physiology and finally behaviour, first outlined by Otto Snell in 1892, by D'Arcy Thompson in 1917 in ''On Growth and Form'' and by Julian Huxley in 1932. Overview Allom ...
, which means that two castes and roles within the colony are differentiated by size and shape. This suggests that caste is determined very early in development. Additional resources may be allocated to future queens so that they grow to larger size as adults.


Communication

Secretions and
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
are crucial for wasp-to-wasp
communication Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inquir ...
regarding all aspects of colony life. Dominance interactions, search for food and nest sites, reproductive behavior and defense are all mediated by these sensory compounds. Richardson's gland, a secretory gland found in the majority of wasps in the Epinonini tribe, produces organic signal molecules that travel from secretory cells to duct cells before being released into the air. ''P. sylveirae'' uses this gland to leave scent trails when traveling from the nest.


Foraging behavior

Most social wasps act as generalists in the
food web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...
s of their
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
. They collect and consume nectar and
exudate An exudate is a fluid emitted by an organism through pores or a wound, a process known as exuding or exudation. ''Exudate'' is derived from ''exude'' 'to ooze' from Latin ''exsūdāre'' 'to (ooze out) sweat' (''ex-'' 'out' and ''sūdāre'' 'to ...
s, but also prey upon other
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
. ''P. sylveirae'' is also known to scavenge for animal protein, going as far as to feed on animal carcasses. They are generalist foragers, but can learn and specialize to specific hunting and foraging locations.


Ovarian development

Three categories of ovarian development are determined in this species. Ovaries may vary from small or non-existent to mature and functional. An intermediate phase exists in which
oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
s are only developed in the upper portion of the ovary. This last type is associated with young queens, whose size is also smaller on average, making the queen closer in morphology to the working caste.


Interaction with other species

''P. sylveirae'' preys heavily on the coffee-eating moth, '' Leucoptera coffeella''. P. sylveirae attacks and consumes the moth larvae, which do a great deal of damage to coffee crops in South America. By keeping ''L. coffeella'' populations in check, the wasp can be very beneficial to both coffee plants and humans.


Human interaction


Stings

Stings of wasps, including that of ''Protonectarina sylveirae'', typically result in painful
inflammation Inflammation (from la, wikt:en:inflammatio#Latin, inflammatio) is part of the complex biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or Irritation, irritants, and is a protective response involving im ...
at the site of the wound. This response is due to
melittin Melittin is the main component (40–60% of the dry weight) and the major pain producing substance of honeybee (''Apis mellifera'') venom. Melittin is a basic peptide consisting of 26 amino acids. Function The principal function of melittin a ...
peptides Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. A p ...
Polybia-MP-I, N-2-Polybia-MP-I, Protonectarina-MP-NH2, and Protonectarina-MP-OH. These are specific to ''P. sylveirae'' and closely related wasps. 0.8-5% of humans will exhibit a generalized systemic reaction upon being stung by a wasp, including
anaphylaxis Anaphylaxis is a serious, potentially fatal allergic reaction and medical emergency that is rapid in onset and requires immediate medical attention regardless of use of emergency medication on site. It typically causes more than one of the follow ...
, which is life-threatening.


Effects of pesticides

Wasps like ''Protonectarina'' often prey upon insects that humans consider pests, such as the coffee-eating moth, ''L. coffeella''. This ecological interaction is vitally important to human populations, as an unchecked infestation of ''L. coffeella'' can cause significant damage to coffee crops, reducing crop yields by up to 50%.
Organophosphates In organic chemistry, organophosphates (also known as phosphate esters, or OPEs) are a class of organophosphorus compounds with the general structure , a central phosphate molecule with alkyl or aromatic substituents. They can be considered ...
, which are commonly used as insecticides for agricultural crops, are extremely toxic and have a large unintended effect on populations of ''P. sylveirae''. Insecticides reduce the occurrence and effectiveness of many aspects of ''P. sylveirae'' behavior, including orientation, motility, feeding, oviposition, and learning.


References

{{Taxonbar, from=Q10273223 Vespidae Monotypic Hymenoptera genera Hymenoptera of South America