Schwarziana Quadripunctata

Schwarziana quadripunctata - Wikipedia, the free encyclopedia
https://en.wikipedia.org/wiki/Schwarziana_quadripunctata
From Wikipedia, the free encyclopedia
Schwarziana quadripunctata is a small, stingless bee found in a stretch

of the South American Amazon from Gois, Brazil, through Paraguay, to
Misiones, Argentina.[1] This highly eusocial insect constructs earthen
nests in the subterranean level of the subtropical environment, an
unusual feature among other stingless bees. The species ranges in sizes
from 6.0 to 7.5 millimeters and feeds on a diverse diet of flowering
plants found abundantly on the forest floor, including guacatonga
(Casearia sylvestris) and the mistletoe species Struthanthus
concinnus.[2]
Schwarziana quadripunctata
Schwarziana quadripunctata at the

nest
1 Taxonomy and phylogeny

2 Description
2.1 Bees
2.2 Nests
3 Distribution and habitat
4 Colony cycle
5 Behavior
5.1 Dominance hierarchy
5.2 Division of labor
5.3 Reproduction and reproductive suppression
5.4 Communication
5.5 Navigation
5.6 Mating behavior
6 Kin selection
6.1 Genetic relatedness within colonies
6.2 Kin selection and discrimination
6.3 Worker-queen conflict
6.4 Queen-queen conflict
7 Diet
8 Defense
9 References
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Hymenoptera
Family:
Apidae
Genus:
Schwarziana
Species:
S. quadripunctata
Binomial name
Schwarziana quadripunctata
(Lepeletier, 1836)
Distribution of S. quadripunctata
S. quadripunctata was first described by the French entomologist and former president of the French
Entomologist Society Amd Louis Michel le Peletier in 1836. Although originally placed in the genus
Trigona, more recent taxonomic evaluations have since placed it under its current genus, following the 1943
work of Padre J.S. Moure,[1] occasionally referred to as the Father of Brazilian Bee Taxonomy." However,
some still consider the term Schwarziana as a subgenus rather than genus and instead incorrectly classify it
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under the closely related genus Plebeia.

The closest relative of S. quadripunctata is the species Schwarziana mourei, determined through morphological
and genetic evidence collected in the early twenty-first century. Until recently (2015) it was believed that these
were the only two existing species under the genus Schwarziana in the world. However, two more species were
discovered in the high altitude environment of South America S. bocainensis in southeastern Brazil and S.
chapadensis in central Brazi.[3]
Bees
The stingless bee S. quadripunctata varies in size from 6.0 to 7.5 millimeters.[4] Worker bees and dwarf queens
tend to be on the lower end of this spectrum while queens tend to lie on the higher end. Dwarf queens and
workers tend to have an average weight of about 30 mg, but have been known to weigh as much as 40 mg or as
little as 22 mg. Mated queens (those in charge of the nest) are, in comparison, much larger. The average queen
bee weighs in at about 130 mg. However, some have been measured at over 160 mg,[5] about a quarter of the
weight of an average paperclip. Colonies contain a larger queen with greater fecundity than dwarf queens,
causing the size discrepancy between the two.[5] Unusual for most other eusocial insects, worker bees and dwarf
queen bees tend to be similar in weight and size.[4] The bees have a pale coloration of brown or reddish-brown
with occasional yellow markings on the head.[3] They maintain a punctate thorax and abdomen and a dorsal
thoracic area sporting a few hairs.[1] Glands are present on the head and thorax. The glands are larger in the
heads of general worker bees and larger in the thorax for nursing worker bees.[6]
Nests
Nests are entirely underground on the subtropical floor and consist of vast and expansive cavities. Each nest is
occupied by a single colony consisting of an individual queen and several thousand workers. Brood cells are
arranged in spiral combs, each housing only one individual. Cells housing queens are typically much larger than
the surrounding cells housing males and workers.[4]
The distribution of S. quadripunctata ranges from the central highlands of Brazil down towards the northern
edge of Argentina on the eastern side of South America. Here, the environment consists of a wide variety of
biodiversity living within lush tropical and subtropical rainforests. Altitudes can reach up to fifteen hundred
meters (1500 m) in some areas. Lying so near the equator, temperatures are fairly warm year-round, averaging
twenty-five degrees Celsius (25 C).[1]
Nests are built into the fertile topsoil of the subtropical rainforests.[4] The niche breadth of S. quadripunctata
includes a wide variety of flowering plants that overlaps with many other eusocial bees native to the area,[2]
including members of the tribes Meliponini and Trigoni, as well as Africanized honeybees. Each nest features a
single, simple rounded entrance, as opposed to a triangular entrance or one with a tube network, several layers,
or multiple openings. S. quadripunctata nest entrances in So Paulo, Brazil consisted of an average entrance
area of 14.5 square millimeters (mm2), relatively small compared to the 2.1 square millimeter (2.1 mm2) body
of the bee. This may account for the lower traffic levels of the stingless bee through the entryway, an average of
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17.4 bees per minute per nest.[7]

Members of this species are fairly common in their large South American region, with nests containing over
several thousand members each.[2] However, their population is believed to have declined by nearly sixty-five
percent (65%) over the past few decades due to competition with the introduced Africanized honeybees[2] along
with human deforestation. This decline is projected to continue into the future.
Colonies are founded by the swarming of individuals consisting of a single queen followed by several thousand
worker bees.[4] This can only be achieved if both the queen and her workers align themselves towards the
same goal in production of fertile females. Once a colony has been established, housing combs are produced by
the workers, with larger ones allocated to potential queen larvae and the more numerous, normal-sized cells
produced for the worker bee eggs.[8] Dwarf queens are also reared in worker cells. Queen cells are typically
located near the edge or periphery of the comb while worker and dwarf queen cells are randomly aggregated
towards the center. Although less than one percent (1%) of dwarf queens were seen to emerge from worker
cells, they account for nearly eighty-six percent (86%) of the total queen population.[4]
A colony will continue to grow in size until the queen bee dies or swarms to a new location to begin a new
colony.[8] In rare cases, two queens may swarm to the same location, due to balanced attractiveness, to begin a
new colony. In these special cases, the younger queen typically produces more eggs than the older queen and
begins to take over the colony, phasing out the older member.[9]
Dominance hierarchy
In the stingless bee S. quadripunctata, as in many social insects, queens reign at the top of the dominance
hierarchy, followed by dwarf queens and finally workers. However, dwarf queens have been observed to lead
nearly one in five colonies. Although, this accounts for only seventeen percent (17%) of dwarf queens and is
significantly lower than the rate at which they were reared (constituting 86% of all females). This does show,
however, that becoming a dwarf queen can be beneficial.[4]
Dwarf queens have a lower fecundity and reproduction rate than normal queens. This appears to be the
consequence of a reduced egg-laying rate[10] and a lower average ovary weight. A queens ovary is nearly four
times larger than a dwarf queens.[4]
In some aspects, however, workers occasionally have a slight advantage over queens and dwarf queens. Worker
bees are in charge of delivering food supplies to developing larvae. Thus, workers can force underdeveloped
dwarf queen larvae and queen larvae to be converted into a worker bee, or to be eradicated completely, through
extreme limitations of their food rations.[4]
Division of labor
The division of labor often, in some way or another, reflects the evolutionary sex ratio of a species. This is
especially true in S. quadripunctata, where worker bees can outnumber queens by about one thousand to one
(1000:1).[4]
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The main purpose of the colonial queen, or gyne, is to lay eggs within the brood cells housed in the combs.
They also play a vital role in the establishment of new colonies. Using their attractive senses, queen bees are
able to lead a swarm of workers and the occasional dwarf queen to new locations.[8] Until fairly recently, it was
hypothesized that these dwarf, or miniature, queens were the result of developmental mistakes. However,
through Tom Wenseleers' "Caste Conflict Hypothesis," it is believed that dwarf queens are evolutionarily
advantageous in that they develop in order to avoid becoming a worker bee.[10] Their role was found to become
the head of a colony in the absence of viable, full-sized queens.[4]
The roles of the worker bee in S. quadripunctata include the maintenance and protection of larvae (queen,
dwarf, or worker) placed within the brood cells. They are responsible for delivering nutrients to the developing
larvae until they hatch.[10] Worker bees are also responsible for accompanying the establishments of new
colonies by a queen.[8]
Reproduction and reproductive suppression

Unlike typical honeybees, which practice multiple mating, stingless bees have been suspected of single mating
between drones and queen bees. This, along with the consequential pairing of the high relatedness rates within
colonies of S. quadripunctata, serves to explain the high degree of kin selection among stingless bees.[11]
After a female queen mates and lays her brood, the worker bees package them into specifically designed cells
queens in larger royal cells and workers and dwarf queens in smaller ones. Each cell is then capped with a
layer of wax-like substance. With the queens role in production complete, the combs are left to the tending of
worker bees, which distribute nutrients and vital resources to the growing larvae.[4] With so much control,
worker bees occasionally discriminate or display reproductive suppression tendencies against dwarf queens by
either uncapping the cell before the dwarf hatches, halting the supply of nutrients, or killing the newly
developed individual upon emergence of the cell.[10]
Communication
A major strength of any eusocial insect is the ability to communicate amongst one another. Often, this method
of communication comes in the form of chemical signals between members of the same colony. In particular,
cuticular hydrocarbon levels are thought to be of great importance to S. quadripunctata communication and can
vary greatly according to age, sex and caste.[12]
Hydrocarbons in the stingless bee can vary in length from nonadecane with nineteen carbons to tritriacontane
with thirty-three carbons, as discovered in a study conducted by Nunes et al. (2009). It was found that cuticles
of older worker bees contained higher concentrations of heptacosene (C27) and hentriacontene (C31) than
younger workers and virgin queens, indicating that these hydrocarbons play an important role in nest mate
recognition. It was also found that younger stingless bees lacked some of the major hydrocarbons found in older
individuals, most likely due a result of less exposure to the wax present in the nest. The relative concentrations
of hydrocarbons in members of a colony differed between unrelated sites, alluding to the hypothesis that the
wax of each nest contains a unique chemical blend from which the developing larvae absorb.[12]
Navigation
Along with, and often closely related to, communication, effective navigation within an environment is vital to
the successful reproduction of eusocial insects. Although some organisms methods of navigation seem quite
apparent (i.e. through modes of sight and scent) the navigational system of Schwarziana quadripunctata had
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remained elusive until quite recently. It has been determined that S. quadripunctata navigates, like many other
bees, through the use of magnetoreception utilizing magnetic fields in the atmosphere to recognize differences
in location, altitude and directionality.[13]
This notion of magnetic navigational skills was first observed in a 2005 study conducted by Esquivel et al. on a
S. quadripunctata nest near Rio de Janeiro, Brazil. The study applied various levels of applied magnetic fields
across the entrance of the nest. The exiting angles (both vertical and horizontal) of the individual bees were then
measured as they emerged from the nest. The primary control experiment confirmed that the majority of the
nest had a preferential exiting angle in accordance with the local geomagnetic field. Over the course of four
months, the study group measured the two angles in comparison to various magnitudes of applied magnetic
fields and compared them to observed angles in natural geomagnetic fields. A significant correlation was found
between the two, definitively showing that the stingless bee relies on magnetoreception for navigational
purposes.[13]
A similar 2005 study by Lucano et al. observed the same nest of Schwarziana quadripunctata to determine the
location of the bodily magnetoreceptors. After separating the head, thorax, abdomen and antennae of a dozen
stingless bees, magnetic fields were applied to each body part. It was found that the head, thorax, and abdomen
present a diamagnetic contribution (inducing a magnetic field opposite to the applied field), while the antennae
displayed a paramagnetic one (inducing a magnetic field in the same direction as the applied field). The
antennae of the stingless bee, therefore, exist as a complex sensory organ used for orientation and navigation
through the combined use of thousands of hair-shaped sensilla.[14]
Mating behavior
Like most eusocial insects, S. quadripunctata reproduction relies heavily on the established hierarchy and level
of environmental quality. Usually, the physogastric queen lays the majority of the eggs. However, the miniature
(or dwarf) queens may also lay fertile eggs, although they tend to be fewer in number and smaller in size.[15]
As multiple virgin queens approach the age of reproductive success, they begin to accumulate the
reddish-brown pigmentation associated with adult members. At this time, male workers will begin individually
courting them on the layers of wax on the comb or on the comb itself. Courting occurs in the form of buccal
contact between the males and females through a lateral hole in the thorax of the female.[15] Some virgin queens
will begin manipulating the cerumen (wax) layers[15] in preparation for the construction of brood cells by the
workers.[8] If the physogastric queen deems a virgin queen is too much of a sexual threat, it may be imprisoned
in a specialized cell in groups of one to five along with one or two worker bees. This is done in order to limit
interference to the cell provisioning and oviposition processes of the physogastric queen.[15]
Future queen eggs are positioned near the periphery of the comb in larger, specialized cells while workers and
dwarf queens (in a ratio of about 1000 to 6, respectively) are randomly placed about the interior in smaller
brood cells. The eggs are cared for by the workers.[4]
Genetic relatedness within colonies

Since S. quadripunctata reproduction typically relies on a single queen, genetic relatedness within individual
colonies is expected to be relatively high. In an analyzation of the genetic relatedness within four separate
colonies with an average of eight different worker genotypes per colony, an average relatedness of .792 was
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found among worker bees within each colony.[11] This value has been replicated in several studies, including
one conducted by Toth et al. (2001) among eighteen colonies in Brazil (relatedness value = 0.75 0.04 among
worker bees). In the same study, males (being haploid) had a relatedness value of 1.0 to queens, while workers
only shared approximately fifty percent (50%) of the queens genes (relatedness value = 0.48). Relatedness
between workers and males were also about half (relatedness value = 0.51). The data was collected and
calculated on the evaluation of seven polymorphic microsatellite loci, each with several attributable alleles.[16]
Kin selection and discrimination

Kin recognition often goes hand in hand with communication. For instance, researchers compared the relative
amounts of various hydrocarbon compounds present in the epicuticles, serving as a chemical odor to other
individuals, of S. quadripunctata. The study showed that a diverse array of these chemicals were present in each
member. They then charted these hydrocarbon differences and found a distinct separation according to colony
location, age, sex, and hierarchal position. In particular, older workers, which are in charge of colony protection,
had higher concentrations of the alkenes heptacosene and hentriacontene. It was thus deduced that these alkenes
were vital parts of kin recognition. Some of these essential hydrocarbons were also found to be lacking in
younger individuals, alluding to the idea that the chemicals are acquired as the exposure to the waxy comb,
specific to each colony, increases.[12]
One case sought to further test the importance of hydrocarbon concentration. In a group of related worker bees,
half were injected with an alien alkene. Aggressive behaviors towards these individuals by the non-injected
worker bees increased significantly. They were unable to be recognized by their unique odor and were
subsequently treated as intruders.[12]
The recognition of various odors associated with hydrocarbon concentrations is also responsible for the ability
of worker bees to determine the location of their home nest.[12]
Worker-queen conflict
Many stingless bee colonies, those of S. quadripunctata included, are repopulated by a single queen who mates.
This should, in theory, create a conflicting rift between queens and the worker bees due to variations in genetic
relatedness. Queens produce haploid males that are genetically identical to them. In contrast, workers only share
fifty percent (50%) of their genes with males, leading to an evolutionary conflict of interest. However, worker
bees were not observed to increase their aggressive behaviors towards newly reproduced males.[16]
A few possible explanations arose from the study in an attempt to explain this lack of aggression. One
hypothesis[16] couples the production of males in a stingless bee society with an increase in food reproduction.
In this case, the benefit of a stable diet would overcome the workers desire to produce females. Another
possibility[16] is that aggression is beneficial no matter the sex being produced. The aggression between workers
and queens may just be residual left over from a past evolutionary benefit.
Queen-queen conflict
More distinctive to the intra-communal dynamics of the Schwarziana quadripunctata nest is the existence of
multiple statuses of queen the physogastric queen (those that are larger in size and more involved in
reproduction) and multiple dwarf queens. The number of these dwarf queens varies in accordance to resource
availability, colony conditions, and the physical state of the current physogastric queen. However, the dwarf
queens are typically produced year round, with several hatching simultaneously.[15]
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Mate attractiveness to the newly born queens coincides with pigmentation. The more brightly colored queens
were more likely to lay more eggs than their paler counterparts. As the newborn queens begin to age,
pigmentation increased as well as the number of courtships from male stingless bees. In the case of an
encounter between the physogastric queen and the newborn virgin queens, the larger queen may initiate
buccal contact with the smaller queen. A rapidly growing virgin queen may incite her own imprisonment in
specialized prison chambers. In one case, an abnormally large and pigmented virgin queen became overly
agitated in her prison cell. The release of this queen prompted the interruption of brood cell construction,
though the physogastric queen maintained her normal routine. Within a couple days, the virgin queen was found
dead and was being carried off by workers.[15]
The defining aspect of queen-queen conflict appears to rely heavily on size differentiation. The dominant queen
can reach sizes over three times larger than that of the dwarf queens. However, multiple queen production
serves a vital role in the survival of the colony, ensuring that, should one queen unexpectedly succumb to injury
or illness, another one is readily available to take its place in maintaining the nest.[5]
The subtropical environment of the South American Amazon provides a vast array of dietary options for a
stingless bee such as S. quadripunctata. With such a densely populated niche of pollenating insects, it remains
of utmost importance to balance competition and resource acquisition. It is often adverse weather conditions,
rather than complete depletion of pollen and nectar from the resource, that terminate the length of harvesting by
the stingless bees.[2]
In total, S. quadripunctata utilizes over 35 different species of flowering plants. Many of these produce a large
quantity of flower material, limiting interspecies and intraspecies competition. Over ten percent (10%) of their
diet comes from the species Casearia sylvestris (called guaatonga) and Myrcia tomentosa (commonly referred
to as goiaba-brava) alone. Other popular flowering plants comprising their diet include Mikania catharinensis,
Piptocarpha oblonga (or brao-do-rei in Brazil), and Cordia trichoclada.[2]
The entrance to a stingless bee nest serves as a particularly vulnerable area. It serves as the threshold that
separates the viability of the species from the outside world. The size of the entryway is critical. It must remain
large enough to allow the easy to-and-fro passage of the bee, yet small enough to dissuade potential predators
(such as orb spiders) from causing havoc inside the nest. Evolutionarily, a size compromise must be reached.[7]
The nest entrance of Schwarziana quadripunctata consists of a simple, round opening. This is a distinct
comparison to the complex entryways consisting of layered entrances or closable doorways observed in some
other related species. Through the observation and measurement of several S. quadripunctata nests near So
Paulo, Brazil, entrances were discovered to have an average area of 14.5 square millimeters (mm2). In
comparison to the average stingless bee having an area of 2.1 mm2, this gives a ratio of 6.8 opening to bee. This
smaller entrance provides more than enough space for the relatively low enter and exit traffic of 17.4
individuals per minute.[7]
S. quadripunctata appears to rely heavily on the small entryway size as their main source of defense. Although
three to five guards are typically seen guarding the entryway, they act in a fairly non-aggressive manner
(dipping inside the inner entrance) when presented with an intruder. Perhaps they act as an alarm for the rest of
the nest rather than an attack force.[7]
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1. Michener, C.D. (2007). The Bees of the World. Baltimore: Johns Hopkins University Press.
2. Wilms, Wolfgang; Imperatriz-Fonseca, Vera L.; Engels, Wolf (1996). "Resource Partitioning between Highly
Eusocial Bees and Possible Impact of the Introduced Africanized Honey Bee on Native Stingless Bees in the
Brazilian Atlantic Rainforest" (PDF). Studies on Neotropical Fauna and Environment 31 (3-4): 137151.
doi:10.1076/snfe.31.3.137.13336.
3. Melo, Gabriel A.R. (2015). "New Species of the Stingless Bee Genus Schwarziana (Hymenoptera, Apidae)". Revista
Brasileira de Entomologia.
4. Wenseleers, Tom; Ratnieks, Francis L.W.; de F Robiero, Marcia; de A Alves, Denise; Imperatriz-Fonseca, Vera-Lucia
(2005). "Working-Class Royalty: Bees Beat the Caste System". Biology Letters 1 (2): 125128.
doi:10.1098/rsbl.2004.0281. PMC 1626201. PMID 17148145.
5. Ribiero, Mrcia De F.; Alves, Denise De A. (2001). "Size Variation in Schwarziana quadripunctata Queens
(Hymenoptera, Apidae, Meliponini)" (PDF). Revista de Etologia 3 (1): 5965.
6. Cruz-Landim, C.; Reginato, R.G. (2001). "Exocrine Glands of Schwarziana quadripunctata (Hymenoptera, Apinae,
Meliponini)". Brazilian Journal of Biology 61 (3): 497505. doi:10.1590/s1519-69842001000300020.
7. Couvillon, M.J.; Wenseleers, T.; Imperatriz-Fonseca, L.; Nogueira-Neto, P.; Ratnieks, F.L.W. (2007). "Comparative
Study in Stingless Bees (Meliponini) Demonstrates that Nest Entrance Size Predicts Traffic and Defensivity". Journal
of Evolutionary Biology 21 (1): 194201. doi:10.1111/j.1420-9101.2007.01457.x.
8. Santos-Filho, Prsio de Souza; Alves, Denise de Araujo; Eterovic, Andr; Imperatriz-Fonseca, Vera Lucia; Kleinert,
Astrid de Matos Peixoto (2006). "Numerical Investment in Sex and Caste by Stingless Bees (Apidae: Meliponini): A
Comparative Analysis". Studies on Neotropical Fauna and Environment 37 (2): 207221.
doi:10.1051/apido:2006015.
9. Engels, Wolf (ed.) (1990). Social Insects: An Evolutionary Approach to Castes and Reproduction. New York:
Springer-Verlag.
10. Ribeiro, Mrcai de F.; Wenseleers, Tom; Santos-Filho, Prsio de S.; Alves, Denise de A. (2006). "Miniature Queens
in Stingless Bees: Basic Facts and Evolutionary Hypotheses". Adipologie 37 (2): 191206.
doi:10.1051/apido:2006023.
11. Peters, John M.; Queller, David C.; Imperatriz-Fonseca, Vera L.; Roubik, David W.; Strassmann, Joan E. (1998).
"Mate Number, Kin Selection and Social Conflicts in Stingless Bees and Honeybees" (PDF). Proceedings of the Royal
Society of London 266 (2): 379384. PMC 1689682.
12. Nunes, T.M.; Turatti, I.C.C.; Mateus, S.; Nascimento, F.S.; Lopes, N.P.; Zucchi, R. (2009). "Cuticular Hydrocarbons
in the Stingless Bee Schwarziana quadripunctata (Hymenoptera, Apidae, Meliponini): Differences between Colonies,
Castes and Age" (PDF). Genetics and Molecular Research 8 (2): 589595. doi:10.4238/vol8-2kerr012.
13. Esquivel, Darci M.S.; Wajnberg, E.; do Nascimento, F.S.; Pinho, M.B.; Lins de Barros, H.G.P.; Eizemberg, R. (2005).
"Do Magnetic Storms Change Behavior of the Stingless Bee Guiriu (Schwarziana quadripunctata)?".
Naturwissenschaften 94 (2): 139142. doi:10.1007/s00114-006-0169-z.
14. Lucano, M.J.; Cernicchiaro, G.; Wajnberg, E.; Esquivel, D.M.S. (2005). "Stingless Bee Antennae: A Magnetic
Sensory Organ?" (PDF). Biometals 19 (3): 295300. doi:10.1007/s10534-005-0520-4.
15. Nogueira-Ferreira, F.H.; Silva-Matos, E.V.; Zucchi, R. (2009). "Interaction and Behavior of Virgin and Physogastric
Queens in Three Meliponini Species (Hymenoptera, Apidae)". Genetics and Molecular Research 8 (2): 703708.
doi:10.4238/vol8-2kerr008.
16. Tth, Eva; Strassmann, Joan E.; Imperatriz-Fonseca, Vera L.; Queller, David C. (2001). "Queens, not Workers,
Produce the Males in Stingless Bee Schwarziana quadripunctata quadripunctata" (PDF). Animal Behavior 66 (2):
359368.
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