Trigona fulviventris
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''Trigona fulviventris'', known by the common names ''culo-de-vaca'', ''culo-de-señora'', ''mu'ul-kab'', ''culo-de-buey'', and ''culo-de-vieja'', is a species of stingless bee found in Mexico and neotropical regions of Central and South America. It is one of the largest and most widespread bees of its genus. They exhibit complex foraging behaviors by integrating spatio-temporal learning and flower scents. ''T. fulviventris'' has traditionally been observed to abstain from aggressive behavior with other species; however, more recent analyses have shown that ''T. fulviventris'' emit pheromones that act as attack signals particularly when related individuals are captured by predators.


Taxonomy and phylogeny

''Trigona fulviventris'' was first described by Félix Édouard Guérin-Méneville in 1845. It is a member of 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 includes ants, bees, wasps, and sawflies and part of the family Apidae, which includes other bees such as bumble bees, honey bees, and
orchid bees The tribe Euglossini, in the subfamily Apinae, commonly known as orchid bees or euglossine bees, are the only group of corbiculate bees whose non-parasitic members do not all possess eusocial behavior. Description Most of the tribe's species ...
. It is further categorized in the genus '' Trigona'', a genus of stingless bees. Two subspecies of ''T. fulviventris'' have been identified: ''T. f. fulviventris'' and ''T. f. guianae''. These subspecies are distinguished by coloration; morphological studies have shown that the subspecies cannot be easily separated by strictly morphological analyses.


Description and identification

''Trigona fulviventris'' individuals are morphologically characterized by the expression of two projections, called tubercles on their labra as well as the presence of four mandibular teeth. The two identified subspecies of ''T. fulviventris'', ''T. f. fulviventris'' and ''T. f. guianae'', can be distinguished by differences in metasomal coloration. ''T. f. fulviventris''
metasoma The metasoma is the posterior part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory system, and circula ...
are rust-colored with some differences in tone between individuals, while ''T. f. guianae'' metasoma and the rest of the body are black, sometimes expressing some reddish tones. ''T. fulviventris'' individuals are among the largest of their genus, ranging in size from 5 to 6.5 millimeters in length.


Diet

''Trigona fulviventris'' feeds mostly on pollen and nectar. Known species of plant from which ''T. fulviventris'' feeds are ''
Passiflora vitifolia ''Passiflora vitifolia'', the perfumed passionflower, is a species of ''Passiflora'', native to southern Central America (Costa Rica, Nicaragua, Panama) and northwestern South America (Venezuela, Colombia, Ecuador, Peru). Description It is a vi ...
, Pavonia dasypetala, Heliconia imbricata, Quassia amara, Dioclea,
Lantana camara ''Lantana camara'' (common lantana) is a species of flowering plant within the verbena family (Verbenaceae), native to the American tropics. It is a very adaptable species, which can inhabit a wide variety of ecosystems; once it has been introduc ...
, Tabebuia,
Asystasia The genus ''Asystasia'' belongs to the family Acanthaceae and comprises approximately 70 species found in the tropics, including the weedy species ''Asystasia gangetica''. Selected species * '' Asystasia africana'' (S. Moore) C.B. Clarke * ''As ...
, Insertia,
Psychotria ''Psychotria'' is a genus of flowering plants in the family Rubiaceae. It contains 1,582 species and is therefore one of the largest genera of flowering plants. The genus has a pantropical distribution and members of the genus are small understor ...
,
Stromanthe ''Stromanthe'' is a genus of flowering plants in the family Marantaceae, native to the tropical portions of the Americas from Mexico to Trinidad to northern Argentina.Idárraga-Piedrahita, A., R. D. C. Ortiz, R. Callejas Posada & M. Merello. (eds ...
,
Justicia aurea ''Justicia aurea'', the Brazilian plume or yellow jacobinia, is an ornamental shrub native to the Cerrado vegetation of Brazil. This plant may be propagated by herbaceous stem cutting, and it can usually get to 1,50 - 2,50 m tall. They flou ...
,
Heliconia tortuosa ''Heliconia tortuosa'' is an herbaceous tropical perennial commonly found in secondary succession in montane forests in Central America and southern Mexico ( Chiapas and Tabasco Tabasco (), officially the Free and Sovereign State of Tabasco ( ...
,
Hibiscus rosa-sinensis ''Hibiscus rosa-sinensis'', known colloquially as Chinese hibiscus, China rose, Hawaiian hibiscus, rose mallow and shoeblack plant, is a species of tropical hibiscus, a flowering plant in the Hibisceae Tribe (biology), tribe of the family (biol ...
,
Impatiens walleriana ''Impatiens walleriana'' ( syn. ''Impatiens sultanii''), also known as busy Lizzie (British Isles), balsam, sultana, or simply impatiens, is a species of the genus '' Impatiens'', native to eastern Africa from Kenya to Mozambique. The Latin ...
'', and ''
Fuchsia ''Fuchsia'' () is a genus of flowering plants that consists mostly of shrubs or small trees. The first to be scientifically described, '' Fuchsia triphylla'', was discovered on the Caribbean island of Hispaniola (Haiti and the Dominican Republi ...
''. However, they have also been observed to forage opportunistically on fungi, dead animals, flesh, and fecal matter as well.


Nests


Exterior

''Trigona fulviventris'' nests are made in the ground, often near buttresses and roots of large trees, and have also been found in urban areas (particularly in crevices in walls of buildings). There is one entryway to each nest, which is made large enough for many individuals to pass through at once. Their nest cavities are enclosed by a thick
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
bitumen. Nests are cylindrical in shape and very rigid, with no documented ornamentation.


Interior

Within the nest, storage pots for pollen and nectar are between 0.7 and 1.0 centimeters in diameter and are partially separated from each other. These storage pots are located to the sides of and underneath the brood. Brood cells are oval in shape and are organized in regular, horizontal combs; there are between twenty and thirty combs of brood cells in a nest on average. Brood cell construction in ''T. fulviventris'' is
asynchronous Asynchrony is the state of not being in synchronization. Asynchrony or asynchronous may refer to: Electronics and computing * Asynchrony (computer programming), the occurrence of events independent of the main program flow, and ways to deal with ...
.


Distribution and habitat

''Trigona fulviventris'' is one of the most widely distributed bees of the genus ''Trigona'' and is found in
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
,
Belize Belize (; bzj, Bileez) is a Caribbean and Central American country on the northeastern coast of Central America. It is bordered by Mexico to the north, the Caribbean Sea to the east, and Guatemala to the west and south. It also shares a wate ...
,
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
,
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
,
El Salvador El Salvador (; , meaning " The Saviour"), officially the Republic of El Salvador ( es, República de El Salvador), is a country in Central America. It is bordered on the northeast by Honduras, on the northwest by Guatemala, and on the south b ...
,
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by H ...
,
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. The republic of Honduras is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Oce ...
,
Nicaragua Nicaragua (; ), officially the Republic of Nicaragua (), is the largest country in Central America, bordered by Honduras to the north, the Caribbean to the east, Costa Rica to the south, and the Pacific Ocean to the west. Managua is the cou ...
,
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
, and the Panama Canal Zone. It is more commonly found at low- and mid-altitudes and has been observed to withstand a wide range of humidities. Habitats in which ''T. fulviventris'' colonies make their homes include both tropical dry and tropical wet forests.


Colony cycle

New ''T. fulviventris'' colonies are established in the spring every year when one or more workers from a previous colony leave their nest and begin scouting divots in tree trunks for a new nest location. These bees have been shown to mark these sites with pheromones, often leaving odor trails to lead to desirable nest locations. However, these pheromones can also attract rival ''T. fulviventri''s colonies, which can lead to aggressive encounters between the attracted workers and the new nest-initiation workers (further described in Nest Initiation Aggression below). Once it is determined which workers will inhabit the new area, gynes (reproductive females) will mate with a swarm of males mid-flight and enter the new nest to initiate a new colony.


Behavior


Foraging


Spatio-temporal learning

Spatio-temporal (also called time-place) learning behavior, which is the ability of an individual to associate the time and place of an event especially in foraging, has been documented in ''T. fulviventris''. This behavior has been observed in species only when it is beneficial for the species ecologically, and spatio-temporal learning is not observed in species where resources that individuals forage for are made continuously. The presence of spatio-temporal learning in foraging ''T. fulviventris'' bees suggests that it is an ecologically favorable behavior. ''T. fulviventris'' bees were observed to learn the locations and times of feeding events, and even arrive up to thirty minutes before the feeding event in anticipation of the food reward and stay in the location up to thirty minutes after the event. However, not all ''T. fulviventris'' workers have been observed to express this spatio-temporal learning behavior. This variation in the behavior can be attributed to differences in foraging strategies among workers in the same colony, which may suggest that differing foraging behaviors within a colony may be a more evolutionarily stable and efficient strategy.


Flower constancy

Individual ''T. fulviventris'' foragers were evaluated for their flower constancy with respect to floral scent and were shown to have definite and specific preferences for a single floral scent. In one study, between 78 and 87 percent of foragers were observed to visit flowers that had the same distinct floral scent during successive foraging events, suggesting that chemical cues are important to flower constancy. Furthermore, ''T. fulviventris'' foragers were more likely to favor the same floral scent as the first forager to return to the nest, indicating that ''T. fulviventris'' foragers carry and relay food odors to other foragers inside the nest. When presented with flowers of different coloration, foraging ''T. fulviventris'' individuals seemed to distinctly prefer one color to the others, indicating that visual components are also important to flower constancy.


Scent marking

Scent marking, in which bees mark flowers that have already been foraged from with a pheromone, increases foraging efficiency and has been documented in various bees including honey bees, bumblebees, and sweat bees. ''T. fulviventris'' individuals have been observed to scent-mark flowers they have already visited and reject flowers that have been visited in the preceding forty-five minutes. However, this behavior was not observed with every foraging situation; one study showed that scent-marking occurred when ''T. fulviventris'' workers visited ''Priva mexicana'' flowers, but not when they visited ''Crotalaria cajanifolia'' flowers. This indicates that scent-marking is not always the most favorable foraging strategy and is context-dependent. For instance, ''P. mexicana'' flowers are deep and require bees to climb inside the flower to reach the nectar, which is costly in terms of both time and energy; therefore, it is not worth expending energy and time on foraging on a ''P. mexicana'' flower that has already been visited. However, C. cajanifolia flowers are relatively easier to forage from, so visiting a ''C. cajanifolia'' flower that has already been visited is not as costly for a ''T. fulviventris'' worker.


Defense and aggression

''Trigona fulviventris'' individuals have been observed to abstain from engaging in aggressive behaviors with individuals of other species, particularly larger species like humans. However, ''T. fulviventris'' engages in aggressive behavior with smaller
arthropod 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 ...
s as well as other ''T. fulviventris'' individuals, particularly those of other colonies, especially during nest initiation.


Alarm signals

The heads of ''T. fulviventris'' workers have been analyzed for the presence of chemicals that can act as pheromones and attack signals. These chemicals are stored in the individuals’ mandibular glands, and the most prominent chemicals expressed by ''T. fulviventris'' individuals are nerol (at a rate of about fifty percent) and octyl caproate (at a rate of about twenty percent). Nerol release has been shown to decrease the number of bees leaving the nest by nearly half, while increasing behaviors such as biting and wing vibration. These chemicals can also act as attraction chemicals, attracting members of the colony to an individual captured by a predator outside the nest.


Attacking

Many bees of the genus ''Trigona'' have been identified as expressing aggressive defense behaviors, such as biting and attacking, mediated by the release of attack pheromones from workers when intruders invade their nests. It has been widely reported that, even when large intruders invade ''T. fulviventris'' nests or nests are presented with a crushed ''T. fulviventris'' individual, there are no attacking or biting behaviors observed. However, later studies have shown that alarm responses do exist in ''T. fulviventris'' individuals, particularly in response to smaller arthropods capturing individuals outside the nest, although to a lesser extent than other, more aggressive ''Trigona'' species. These responses, which are mediated by pheromones, include leaving the nest to follow or swarm a predator that has captured an individual, as well as biting predators.


Nest initiation aggression

While ''T. fulviventris'' does not regularly engage in interspecific aggression, there have been many reports of aggression within the species, especially as it relates to nest initiation. As explained earlier, nest initiation occurs when one or more workers mark an attractive site with pheromones. These pheromones can attract workers from other colonies, which results in aggression between the two groups. If the two opposing colonies are represented by about the same number of workers, then the encounter results in a "stalemate" and neither group inhabits the marked site. If, however, one group is represented in much greater numbers, they are permitted to establish a new colony at the marked site. This pheromone-marking and aggression leads to the even distribution of ''T. fulviventris'' nests so as to minimize the amount of aggression between colonies (i.e. as a result of pheromone signals, no two nests are established in close proximity to lessen the likelihood of intraspecific aggression).


Nestmate recognition

''Trigona fulviventris'' individuals are capable of distinguishing nestmates from non-nestmates through recognition of a range of compounds. These compounds include
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ex ...
s and
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, fr ...
s that are present in ''T. fulviventris'' wax as well as locally available floral oils that are present in plant material used to construct nests. These odor cues are important to judging which individuals are nestmates and which are not, which is necessary in determining which individuals to engage in aggressive encounters with. The amount of time that passes between the first encounter of two bees and the start of aggression between them, called mean latency, is negatively correlated with the occurrence of aggression. In other words, the longer it takes two bees to initiate aggressive behaviors, the less likely it will be that there will be aggression between them. This relationship could be a result of variations in the magnitudes of the differences in
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
signals two bees put out. For instance, if one ''T. fulviventris'' bee has only a slightly different signal than another bee, it will take longer to analyze this difference and engage in an aggressive behavior. In contrast, a large difference in signal is more likely to be noticed immediately and aggressive behaviors can occur more quickly, decreasing the mean latency.


Stinging

''Trigona fulviventris'', like other ''Trigona'' bees, does not display stinging behavior. Some bees in the genus ''Trigona'' have been shown to harbor
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
stinging accessories; these vestigial structures are largely absent in ''T. fulviventris'' individuals.


Human importance

The sticky resin used by ''T. fulviventris'' workers in building their nests has been prized by Colombian fisherman as an effective means of caulking fishing canoes that have sprung leaks.


References

{{Taxonbar, from=Q2132968 fulviventris Hymenoptera of North America Insects of Mexico Hymenoptera of South America Insects described in 1845 Taxa named by Félix Édouard Guérin-Méneville