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Euglossa hyacinthina
A female Euglossa hyacinthina shaping resin along the rim of the growing nest envelope
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Apidae
Subfamily:	Apinae
Tribe:	Euglossini
Genus:	Euglossa
Species:	E. hyacinthina
Binomial name
Euglossa hyacinthina Dressler, 1982

The Euglossa Hyacinthina (Long-Tongued Bee) is a species of an orchid bee subfamily (Apinae) in the family Apidae . While it may not be the most familiar specie, the Hyacinthina is one that needs analyzing. With a tongue that can get up to as long as 4cm, the orchid bee specie is found in Central America countries . In a neotropical climate, the Hyacinthina has adapted to hot and humid weather. With wings that have dark shades and are translucent, the bee also consists of a metallic, glossy blue skeleton.

"Medium sized, large body stature, long-tongued, and fast," the Euglossa Hyacinthina is characterized by its eusociality and solitary life-style that goes against most other bee species . Additionally, this specie has no worker or queen bees and runs by an atypical social hierarchy dominated by females. As a result, the interactions between male and female bees are very intriguing and one can trace the origins of sociality by analyzing the Euglossa Hyacinthina.

Description

Euglossa is a genus of a larger species known as euglossine bees. Euglossine are a subfamily of Apinae (orchid bees) and exhibit very little social behavior; mostly characterized as solitary . Eulaema Hyaccinthina is a common species that lives in the ranges of Central America (Costa Rica) all the way to Northern Argentina . It is commonly found in the Amazon forest and plays an important role in the pollination of various flowers including a sweet passion fruit that is dependent on the Euglossa Hyacinthina for cross pollination.

Taxonomy and Phylogeny

Named after their 4 cm long tongues, Euglossines are sometimes commonly known as the long-tongued bees. Euglossini comprise of a diverse and very widespread Neotropical taxon that comprises of "5 genera and nearly 200 species," one being the Euglossa . Additionally, Euglossa is the most specious genus with over 129 identified species including the Hyacinthina. From the kingdom of Animalia, this orchid bee is a subset of the phylum arthropoda. More specifically, it is in the insecta class and is considered to be in the order of hymenoptera. Among the distinct species of Euglossine bees, the Euglosaa genus is known for its bright, metallic skeleton and can in some Neotropical forests make up 25% of the total bee population. The Neotropical Euglossa bees are close relatives with Apini (honey bees) and Bombini (bumble bees) .

Researchers have identified from four genes (16S, 28S, cytochrome b, LW Rh) that the Bombini and Meliponini (Stingless Bee) form a clade, and there is uncertainty on whether Euglossini is a sister group to either Apini or Bombini/Meliponini . Recently, research has strongly proved that Euglossini is indeed a sister group to Apini.

Specifically, there are five genera of Euglossini five hypotheses have been formulated on the relationships of these genera (Aglae, Exaerete, Euglossa, Eufriesea, and Eulaema) . Interestingly, not one definite conclusion has been made because of the incongruency at the root. For all five hypotheses, the Algae is the base group, Eufriesea/Eulaema are sister groups to either Exaerate/Euglossa. The one group that is the most variable is the Euglossa group and this group is where the Hyacinthina specie resides.

Description and Identification

The figure above clearly displays the glossy metallic blue that the Hyacinthina is known for. The wings are of a darker shade and translucent with females having pollen baskets in their hind legs. A study done that was published in 2003 by Elizabeth Capaldi showed that females and males are generally display monomorphism, with the exception of the thorax. The female bees had larger thoraxes compared the male bees and had a small patch of elongated, plumous hair that the male bees didn’t display. Because this specie is of the hymoneptera order, there was a significant deviation from the 1:1 ratio; in fact, 82% of the population consisted of females . One can generalize that females will be found in larger ratios and larger sizes because female orchid bees can control the sex of their offspring.

As opposed to other species, the Euglossa Hyacinthine nest’s are usually dispersed across a wide range. Usually composed of mud or resin, aerial nests are either placed on the underside of leaves or on plant stems. Initially, the nests are of a light orange color that signifies recent construction of the nest; as time passes, the light orange turns to a dark brown and the vertical ridges signify differences in age. The nests are around 6.0 cm long (.7 cm variability) and 4.2 cm wide (.8cm variability). In the nest are cells; the function of cells includes food storage and housing of the larvae, which is commonly known as “brooding”. As the number of cells increase, we can predict that the number of bees living in that particular hive will increase at a linear rate.

Distribution/Habitat

As mentioned before, the Euglossa Hyacinthina species is found in a Neotropical habitat. The Euglossini are most abundant in wet forests, some are found in the Savannas and forests near rivers/lakes. Euglossa hyacinthine habitats solely in Central America in countries such as Panama and Costa Rica. The nests are usually around 1500-1600 m above sea level and they are able to live in such high pressure environments.

Life cycle

Colony Initiation

Interestingly, the only tribe in the apide class/group that do not form large colonies are the Orchid Bees (Euglossine) because they tend to move independently. Because of this distinction, there are no worker bees or queen bees because there wouldn’t be need for such roles in independent movement. This was proven through the analysis of the number of females in comparison to the number of cells in the hive. Because the number of females outweigh the number of cells, once can assume that there are some female bees who are reproductively inactive. The number of female bees greatly outnumbers the number of male bees and presumably, the females are the ones also starting/colonizing existing nests. When creating a new nest, a female bee will make foraging trips for resin in order to construct the hives and this process will last anywhere from 8 minutes to 15 minutes. The construction of the hive itself usually takes anywhere from 10 minutes to 22 minutes. As a result, the process of foraging and contructing a nest takes a total of 18 minutes to 37 minutes for a female.

Colony Growth

Nesting is a seasonal venture for the Euglossa Hyacinthina. In a study done be Eberhard in 1988, the data clearly showed a skew in nest formation for certain seasons. In October of 1985, there 13 out of the 26 nests observed were formed and in October of 1987, five more were formed. Interestingly, there were none formed in the months of April, May or June. As a result, Euglossa Hyacinthina grow in numbers during the fall season, while declining in the summer season. This is most likely due to the number of predators during each season and the amount of available resources. In the fall season, the rate of predator appearance was lower and an increase in resin availability, while the summer season brought an increase in predators and decrease resin availability.

Behavior

A female Euglossa hyacinthina working on the construction of her nest envelope.

Graphical representation of work effort showing how the bee moves from one side of the nest to another while constructing the envelope. Each arc represents the area the bee worked in a given work session, each lasting approximately from 1 to 2 min. The concentric arcs represent 13 sequential work bouts during a 26 min session of construction; the outer-most arc is the starting position at the beginning of the session and the inner-most one is the ending position.

Euglossa hyacinthina constructs aerial out of resin. These nests attached to the side of secondary-growth stems of tall plants. Before 2012, the structure of these nests had been described, but the actual nesting behavior had never been studied. In 2012, a team of scientists documented this behavior for the first time. Video recordings of the bees constructing their nests were made using a GoPro HD camera mounted about 25 cm from the nest. The video is to the left of this paragraph.

Dominance Hierarchy

Because Euglossini do not have worker bees or queen bees and the females have power over male species, the dominance hierarchy has variations that from traditional bee hierarchies. Successful nests are usually founded by single, solitary females and these females stay until the emergence of brood. Once the original founder is done with the nest, there is nest reactivation that occurs by associated females of the same generation and high relatedness. The origin of association of females along with various behavior patterns result in social patterns within nests. The term eusociality perfectly describes the hierarchy within Euglossa Hyacinthina because in a colony consisting of a mother and a group of daughters, the evolutionarily stable strategy is often for the mother to cheat and the guard the new offspring while children forage for food. In this sense, the mother would be the “queen” and the daughters would be foraging for siblings rather than offspring and would be considered the “worker bees”. This concept also gives rise to the concept age-based dominance hierarchy as the older female bees seem to be in a position of power.

Communication

Male bees interact with female bees through their fragrances. After collecting various fragrances throughout its lifetime, the male is ready unleash this resource to females when wanting to reproduce. Reproductive communication is accomplished through sharing of fragrances.

Reproduction Suppression

Often times, when a female leaves its nest, subsequent female bees eat the existing eggs for nutrients and to produce its own eggs. This allows for consumption of high quality nutrients, which improves fecundity/longevity of life and exploit the advantages of parasitism.

Mating Behavior

This species follows a single mating policy, which is expected because of its taxonomy. Monandry within the Euglossini is consistent with the idea that females select a single mate with the best genes and best fitness based on male fragrance phenotypes. Males of many species of orchid bees establish display sites for fragrance signaling. These display sites are usually centered around perches – usually tree trunks - where males repeatedly hover over this perch to attract females. As a result, males are investing much of their lives trying to stock up on fragrances so that they can one day obtain a mate. This process is very cumbersome because females mate only once; there is a lot of energy and risk taking that males must take to be successful. Males are constantly fighting over fragrances and the best perches to attract females. The ability for males to store fragrance is a direct reference to their genetic quality. Males, in combination with fragrance quality, hover around a selected perch to share its fragrance. Through female choice, the female bee then chooses which male bee she would like to mate with.

Kin Selection

Since there are no worker bees and queen bees, the most important relationships are between mother/children. This specie is of the Hyneptomera order, and kin selection theory states that offspring of female bees will be .75 related to mothers and .25 related to father because the mothers only mate once. This leads to a higher relatedness among mother and daughters of each individual species. This theory also supports the eusociality displayed by Euglossa Hyacinthina because the bees are trying to increase genetic relatedness in offspring and consequently increase efforts in caring for the young.

Nesting

Interior view of a nest of Euglossa hyacinthina under construction - JHR-029-015-g001B

Euglossa Hyacinthina are the only tribe of the Apine clade that do not aggregate in large groups or have queen and worker bees. As a result, we can trace back social behavior of bees through the examination of this specific species. Often times, nests are inhabited by multiple females and there is always a trade-off between group-nesting and solitary nesting. Euglossa Hyacinthina must assess whether living in a group will lower chances of predation while having lower reproductive success due to multiple female habitation. An experiment done by Eberhard (1988), shows that Euglossa Hyacinthina often live in groups with an average of 2.1 females (1.2 variability) and often consist of same generation (sister-sister) or different generation (mother-daughter) nesting. There is kin selection associated with this behavior and the methods of resuse of nests; usually the daughter will inherit a preexisting nest. Although the sex ratio of 1:1 suggests there is solely individually acting bees in nests, there is also evidence that suggests there is some form of communication between nest mates. When females seal the nest during the night or inclement weather, females will always until all inhabitants of the nest are safely back. Additionally, there has been very little noticeable aggression among nest mates.

Advantages to nesting include increased vigilance against predation due to the increased inhabitants. If nests are solely inhabited, there is a great chance that predators will seek to take advantage of nests when the female bee is out foraging for food. In contrast, when nests are co-inhabited, other female bees are able to take care of the brood during foraging times and unfortunate death.

Mimicry/Parasitism

Mimicry

Known as the Mullerian mimicry, there have been many taxonomic problems arising from very similar resemblance of various species. Müllerian mimicry is when two or more poisonous species that are not closely related adapt to each other and start mimicking each other's warning signals. Similar color patterns are found in both the Eufriesea and Eulaema, and there was a lot of confusion regarding Eulaema tropica, Eufriesea surinamensis. While Eularma and Eufriesa mimic each other through color scheme and patterns, mimicry within Euglossa is much less obvious because there is less variation. Still, it is possible that many of the green species in any given area are, to some extent, Mullerian mimics. With very similar structure, the only way to distinguish Euglossa species are by skeleton color. While green Euglossas are found most frequently, Bronze is the major color in Costa Rica and in Panama, blue becomes much more frequent as altitude increases. As a result, these patterns found on these bees are suggestive of mimetic complexes.

Parasitism

While Euglossa Hyacinthina usually exhibit cooperative behavior, its relatives display definite forms of parasitism, which may signal the rare cases of parasitism even within the cooperative Hyacinthinas. “Aglae has been reared from nests of Eulaema nigrita. Exaerete frontalis and Ex. smaragdina parasitize Eulaema species, while Ex. dentata has been reared from nests of Eufriesea” (38)." “A number of parasites were also noted from euglossine nests including megachilid bees, meloid beetles, and mutillid wasps.” Interestingly, the Euglossine nests did no harm to human intruders, while avoiding wasps and beetles.

Interaction with other species

Diet

Eats nectar from orchid flowers.

Pollen Sources

• 1. Bixa (Bixaceae)
• 2. Cochlospermum (Cochlospermaceae)
• 3. Oncoba (Flacourtiaceae)
• 4. Clusia (Guttiferae)
• 5. Psidium (Myrtaceae)
• 6. Swartzia (Leguminosae)
• 7. Cassia (Leguminosae)
• 8. Sauvagesia (Ochnaceae)
• 9. Solanum (Solanaceae)
• 10. Xiphidium (Haemodoraceae)

Defense

Female orchid bees have a stinging apparatus. Although getting stung is very painful, people are usually lucky to get stung by a bee because of its rarity. Since orchid bees exhibit solitary behavior and do not defend their nests. Males, on the other hand, do not have any stinging mechanism or apparatus, which gives the female an evolutionary advantage over the males. As a defense mechanism, the stinging apparatus explains the sex ratio in favor of the females.

Parasites

• 1. Megachild Bees
• 2. Meloid Beetles
• 3. Mutillid Wasps

Human Importance

For neotropical habitats, Orchid Bees are important in maintaining homeostasis through the pollination of various orchid flowers. With one Euglossa species pollinating near 74 different plant species belonging to 41 families, Euglossa Hyacinthina also pollinate various flowers within Costa Rica. As Euglossa species can fly up to 23 km, various species are responsible for cross-pollinating which brings about genetic diversity. Historically, Euglossins bees have accounted for the pollination of all the neotropical region.

References

1. Capaldi, E.; Flynn, C. (2007). "Sex Ratio and Nest Observations of Euglossa hyacinthina (Hymenoptera: Apidae: Euglossini)" (PDF). Journal of the Kansas Entomological Society. 80 (4). Kansas Entomological Society: 395–399. Retrieved September 22, 2015.

2. Wcislo, D.; Vargas, G. (2012). "Nest construction behavior by the orchid bee Euglossa hyacinthina" (PDF). Journal of Hymenoptera Research. 29. the International Society of Hymenoptera: 15–20. doi:10.3897/JHR.29.4067. Retrieved September 22, 2015.{{cite journal}}: CS1 maint: unflagged free DOI (link)

3. Soucy, L.; Giray, T.; Roubik, D. (2003). "Solitary and group nesting in the orchid bee Euglossa hyacinthina (Hymenoptera, Apidae)" (PDF). Insectes Sociaux. 50. International Union for the Study of Social Insects: 248–255. doi:10.1007/s00040-003-0670-8. Retrieved September 22, 2015.

4. Augosto, S.C.; Garofalo, C.A. (2004). "Nesting biology and social structure of Euglossa (Euglossa) townsendi Cockerell (Hymenoptera, Apidae, Euglossini)" (PDF). Insectes Sociaux. 51. International Union for the Study of Social Insects: 400–409. doi:10.1007/s00040-004-0760-2. Retrieved September 22, 2015.

5. Eltz, T.; Schmid, M.; Roubik, D. (1997 April). "Haploid Karyotypes of Two Species of Orchid Bees (Hymenoptera: Apidae, Euglossini)" (PDF). 70 (2). Kansas Entomological Society: 142–144. Retrieved September 22, 2015. {{cite journal}}: Check date values in: |date= (help); Cite journal requires |journal= (help)

6. Eberhard, W. (1988). "Group Nesting in Two Species of Euglossa Bees (Hymenoptera: Apidae)" (PDF). Journal of the Kansas Entomological Society. 61 (4). Kansas Entomological Society: 406–411. Retrieved September 22, 2015.

7. Cameron, S. (2004). "PHYLOGENY AND BIOLOGY OF NEOTROPICAL ORCHID BEES (EUGLOSSINI)". Annual Revolution of Entomology. 49: 377–404. doi:10.1146/annurev.ento.49.072103.115855. Retrieved September 22, 2015.

8. Zimmermann, Y.; Roubik, D.W. (2009). "Single mating in orchid bees (Euglossa, Apinae): implications for mate choice and social evolution" (PDF). Insectes Sociaux. 56. International Union for the Study of Social Insects: 241–249. doi:10.1007/s00040-009-0017-1. Retrieved September 22, 2015.

9. Eltz, T.; Roubik, D.W.; Whitten, M. (2003). "Fragrances, male display and mating behaviour of Euglossa hemichlora: a flight cage experiment" (PDF). Physiological Entomology. 28 (251–260). The Royal Entomological Society. Retrieved September 22, 2015.

10. Andradee Silva, A.; Nascimento, Fabio (2012). "Multifemale nests and social behavior in Euglossa melanotricha (Hymenoptera, Apidae, Euglossini)". Journal of Hymenoptera Research. 26. International Society of Hymenoptera: 1–16. doi:10.3897/JHR.26.1957. {{cite journal}}: |access-date= requires |url= (help)CS1 maint: unflagged free DOI (link)

11. Dressler, R. (1982). "BIOLOGY OF THE ORCHID BEES (EUGLOSSINI)". Annual Review of Ecology. 13. Annual Reviews Inc.: 373–394. Retrieved September 22, 2015.

12. Ramirez, S. (15 December 2009). "Orchid bees" (PDF). Current Biology. Elsevier Inc. Retrieved September 22, 2015.

External links

• http://homepage.ruhr-uni-bochum.de/thomas.eltz/
• http://www.life.illinois.edu/scameron/
• http://www.bucknell.edu/majors-and-minors/biology/faculty-and-staff/elizabeth-capaldi-evans.html
• http://www.ots.ac.cr/rbt/attachments/volumes/vol30-2/04-Dressler-Euglossa3.pdf