Halictus rubicundus

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Halictus rubicundus

Scientific classification

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Hymenoptera

Family:

Halictidae

Genus:

Halictus

Species:

H. rubicundus

Binomial name

Halictus rubicundus
(Christ, 1791)

Halictus rubicundus is a species of sweat bee found throughout the Northern Hemisphere. It is
small (~1 cm), dark brown, with fine white bands across the apices of the abdominal segments. The
males are more slender, with longer antennae and yellow markings on the face and legs; they are
distinguished from males of similar species by the absence of an apical hair band on the terminal
abdominal segment.
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H. rubicundus was introduced into North America from the Old World during one of two main
invasions of Halictus subgenera. These invasions likely occurred via the Bering land bridge during
the Pleistocene during times of low sea level.
[2]

The species exhibits different social behaviors depending on climate it is a solitary species in
cooler regions, but is eusocial in warmer areas, sometimes with solitary and eusocial colonies
appearing simultaneously in the same population. The sweat bees are known for its variability in
social behavior, which has become a model for social plasticity. This variability has contributed to an
understanding of social evolution in behavior.
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Contents
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1 Taxonomy and phylogeny

3 Distribution and habitat

4 Nesting

2 Description and identification

4.1 Nesting based on soil hardness

4.2 Nesting based on temperature

4.3 Nesting based on soil water content

5 Colony cycle

6 Behavior

6.1 Dominance hierarchy

6.2 Division of labor

6.3 Mating behavior

6.4 Nest-site fidelity

6.5 Gregarious nesting

7 Kin selection

7.1 Genetic telatedness within colonies

7.1.1 Genetic telatedness among colonies of different behaviors

7.2 Pheromone tecognition

7.3 Costs and benefits of sociality

7.4 Sex ratio in different colonies

7.5 Sex ratio effects on sociality

7.6 Hierarchy conflict

8 Body size as a result of temperature

9 Parasites

10 References

Taxonomy and phylogeny[edit]

Halictus rubicundus is a species of the order of Hymenoptera and family Halictidae, more commonly
known as sweat bees. They get this common name from their known attraction to perspiration.
This species is unique because of their polymorphic social behavior that varies by environmental
conditions, but other species of the Halictidae family are though to also have this variability in
sociality. The genus Halictus was described by Latreille in 1804.
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Description and identification[edit]

The family of Halictidae is unique and easy to identify by its metallic nature. They are known as a
type of sweat bee, as they are often attracted to perspiration. H. rubicundus specifically are less
metallic, but have white stripes on the bottom of their abdominal segment and yellow-orange legs.
The bee is about 10mm long in body length. In social populations, females of the first brood, the
workers, can be identified because they are slightly smaller than the second brood females, the
foundresses.
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Both solitary and eusocial types of the bee have nests in southward facing slopes built for the
entrance of their burrows. This slope maximizes the heat absorption from the sun, making the nest
warmer. The in-ground nests are built in isolated areas, consisting of sand or soil. The nests with a
favorable slope were thought to increase foraging efficiency of adults and development of larvae with
a stable thermal environment.
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Distribution and habitat[edit]

H. rubicundus live in multiple locations worldwide, including North America and the UK, mostly in
temperate climates. It is believed that the differences in latitude actually contribute to their social
behavior. Those living in more northern geographic locations are often more solitary in behavior than
those in southern areas. This difference is widely studied, as it provides insight into the evolutionary
social behavior from solitary to social. Nests are haplometrotic, meaning that they're founded by
single females. The species lives in burrowed nests in the ground, where there is a constructed
slope to provide thermal regulation of the nest. This regulation is important for the development of
the both the egg and the larvae. Stones or areas of vegetation are usually found near the nest
entrance, likely because of the heating properties of these objects. A higher temperature increases
the rate at which the larvae reaches the threshold to gain the ability to fly. Therefore, a warmer nest
leads to increased rate of development of the offspring.
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The nests can be up to 120 mm deep, and are constructed in a wide range of soil types. These
common bees are well studied for the evolution of their social behavior. Because social and social
nests produce more offspring than solitary nests, social nests will burrow further into the ground, as
the second brood of the social population will be nested beneath the first brood. Females typically
nest in dense packs, likely because the nesting females are relatives and demonstrate philopatric
behavior.
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Nesting[edit]
Halictus rubicundus is distributed throughout the Holarctic region. There are two nest types, social
and solitary. Social populations typically nest in warmer regions, such as Kansas and southern
Ontario, while solitary populations nest in cooler regions, such as Scotland and Alaska. In marginal
regions, both social and solitary behavior can be found in different nests of the same population. The

solitary phenotype is expressed as a response to colder environments because the warm season is
not long enough to produce sequential worker and production broods.
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Nesting based on soil hardness[edit]

The nests are burrowed into the ground in loam soil. H. rubicundus has a high tolerance for soil
hardness. Soil hardness affects the density of nesting. Females prefer to nest in softer ground as
they spend less energy and time excavating the nest. Foundresses will choose to build their nests in
patches of softer ground until they reach the critical nearest-neighbor distance of about 50 mm, at
which point the close spacing poses a high risk of the nests collapsing. At this point, further
foundresses would be forced to build their nests in harder soils where the nests could be built closer
together without compromising nest architecture. Foundresses may test the hardness of the soil by
biting into the surface or performing a short test dig.
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Nesting based on temperature[edit]

Nest temperature determines egg development and foraging active time of females. As long as
temperatures dont reach lethal thresholds, developmental rates of offspring will increase with
temperature. Higher temperatures will also increase the thoracic temperature and allow females to
fly more rapidly. With increased speed in flight, females are allowed more time for foraging, mating,
and excavating nests.
The desire for increased temperature in nesting sites in the reason most nests of H. rubicundus are
south facing and sloped. Based on the distribution of this species, facing the south maximizes the
period of time sunlight is shining directly on the nest. Sloping substrates increase the surface area of
the nest and allow for higher absorption of sunlight. In order to test the temperature of the substrate,
females often spend several seconds basking at various points on the ground while searching for
nesting sites.
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Nesting based on soil water content[edit]

The water content of the substrate in which foundresses build the nest is highly important.
Waterlogging must be avoided by using well-drained soils, which provides another advantage to
building in sloping ground. However, there must be an adequate moisture level to prevent
desiccation of the brood cells. For this reason, soil samples of the nests of H. rubicundus have a
relatively high humidity.
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Colony cycle[edit]
The colony cycle of H. rubicundus is an annual cycle based on the seasonal schedule of hibernation
and mating. After hibernation during the winter, female foundresses who mated the previous cycle
emerge in the spring. They each create their own nests in late spring, where they rear a single
brood. Females are more likely to build nests where there is a warmer surface temperature, as this
indicates a warmer interior of the nest for the offspring to develop more quickly. The worker bees
provide pollen as food to the larvae. The gyne will continue to forage for 35 weeks, after which she
will stop provisioning food to the brood cell. The brood cells are left inactive for 12 weeks before
the emergence of the first brood. The first brood emerges in mid to late June. Most females
emerging from the first brood will stay with their natal nest and act as foraging workers.
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After the first brood emerges, the daughters will collect pollen on which the foundresses will resume
laying eggs. Immediately after mating, the females undergo hibernation for the winter again, and the
colony cycle begins again.
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The annual cycle differs slightly for eusocial and solitary bees, in terms of the number of worker bees
and foundresses that are born in the broods. For example, in solitary populations, the females first
brood rears 40% females who are all able to mate before the next hibernation season. However, in
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eusocial populations, the emergence from hibernation occurs about a month earlier, and results in a
brood with mostly worker females. Nesting for solitary populations begins between May and June.
The absence of a brood of female workers defines this nest type as solitary, so solitary populations
produce only one reproductive brood that is provisioned by a gyne. The emergence of this brood is
at approximately the same time as the emergence of the second brood in the social colony cycle.
Upon emergency, the offspring mate and then females enter hibernation away from the nesting site.
As in social colonies, the males and nest foundresses die at the end of the season.
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Behavior[edit]
H. rubicundus is widely studied for their variability in behavior depending on geographic location.
Those in the south are known to exhibit eusocial behavior, while those in the north are known to be
solitary.
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Dominance hierarchy[edit]
There is a caste-like system in H. rubicundus. The dominant motherly behavior often drives away
the offspring, but often times, female offspring stay despite this aggressive behavior. These
categories are not thought to be genetically specified, but rather decided by mating behaviors and
social factors in the first few days of adulthood. In this hierarchy, there is a foundress, which is a
gyne that is always mated and starts their own colony after hibernation each cycle. She is
considered the foundress queen if she is the dominant one in the colony who reproduces. A gyne is
any female with the potential to become a nest foundress. Below that is a non-gyne, which is a
female that stays in an existing colony and often does not mate or reproduce. Within these nongynes, there are some who are considered replacement queens, which can sometimes take the
place of a foundress queen in the colony. At the bottom of the caste system is the worker, which
helps maintain the functions of the colony, including foraging.
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Division of labor[edit]
In the eusocial and solitary dichotomy, the behaviors toward division of labor is different. In eusocial
colonies, they behave similarly to other eusocial bees. In these colonies, the first brood is primarily
worker-females, which in turn help the foundress rear her second brood. The second brood yields
gynes and males, which breed to repeat the cycle over. However, in solitary behaving colonies, the
first brood can yield gynes as well as non-gynes, but usually not workers. The cooperative breeding
behavior that the worker bees in eusocial colonies can exhibit benefits them because their actions
maintain the colony and the fitness of the queen foundress. By helping their mother raise the
second brood, they can benefit by kin selection hypothesis. This means that they help with foraging
and feeding the larvae, as well as maintaining the proper functioning of the nest and colony so that
the second brood can develop healthily and reproduce to pass down the genes that they share.
However, in solitary colonies, the offspring do not serve as workers and do not help the mother
establish a second brood, but rather go off to try to establish nests of their own.
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There is no evidence for predetermined morphological or physiological difference in caste for H.

rubicundus. The differentiation into different castes is based on behavior. Females that do not mate
immediately after emergence become workers or replacement queens while the others become
gynes. Another factor that may dictate the role of a female is the relative abundance of males to
newly emerged females. There will be a higher percentage of gynes relative to non-gynes when
there are an abundance of males.
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Mating behavior[edit]
Unlike other bee species that mate in the air, mating in H. rubicundus occurs on the ground surface
in and around the nest aggregation. Males hover around their natal nest and wait to encounter

females that are entering or leaving a surrounding nest. After mating, females enter diapause and
restart the cycle the following spring.
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In this species, there are foundresses, gynes, non-gynes (who can be replacement queens), and
worker females. The foundresses are those who are virtually guaranteed to mate before the winter
hibernation begins. The gynes and non-gynes are distinct groups, as gynes are likely to mate and
become foundresses, while non-gynes are not. However, it is theorized that male abundance could
possibly play into this distinction early in life. If there is an abundance of males, a virgin female
could mate early in life, before diapause, and make a caste switch into being a gyne. If she is left
unmated, however, she would likely stay a non-gyne. This caste-determination occurs in the first
few days of the female's life, and highly depends on male availability. The first brood that a
foundress has usually rears gynes, non-gynes, and some males. The second brood has gynes and
males only. H. rubicundus is one of the first species discovered to have first broods with both gynes
and non-gynes. It was previously thought that the non-gynes were all produced in the first brood
and the gynes in the second brood.
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Nest-site fidelity[edit]
Nest-site fidelity may be due to one of three reasons.
1. Philopatry is the tendency for adult bees to nest near parental nest. Returning to the natal
nest is beneficial because the nest must have been successful enough to produce adults for
one year, so it is assumed to be in good enough condition to raise another brood. This
prevents H. rubicundus from taking the risk of settling in a failing nest.
2. Habitat learning describes the process through which females recognize characteristics of
the nest from which she came and chooses to nest in similar conditions. Although this is
different from philopatry in that she wont purposely choose to be near her previous nest and
will make selections based on environmental factors, the nest the female chooses will often
still be near her original nest.
3. Social facilitation may influence the nesting location chosen by a female because the
benefits of nesting close to other bees may outweigh the costs of finding a new location with
a suitable substrate.
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Gregarious nesting[edit]
Dense nesting tendencies of H. rubicundus are most likely due to the following three factors:
1. There is a limited amount of suitable substrate in which the bees can build their nests, so
they must build many nests packed tightly together without compromising the structural
integrity of the nest.
2. As mentioned earlier, philopatry is an important factor in maintaining an aggregation. The
search for a new nesting site requires a lot of resources, so females will likely limit their
dispersal and stay near their natal nest sites.
3. Hymenopteran and dipteran species may attach the ground nest of H. rubicundus. Although
it would seem that aggregation of nesting would increase the mortality due to parasitism as
they would be more conspicuous, it is likely that there is a dilution effect that reduce
mortality by parasitism.
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Kin selection[edit]

Genetic telatedness within colonies[edit]

Depending on the geographic location of the colony and its behavior, genetic relatedness within the
colonies differs once again. In colonies in the north that exhibit solitary behavior, the genetic
relatedness is different because the first brood does not yield worker bees who help the mother raise
the following brood. Therefore, there are no workers to help the mother and each of the gynes goes
to establish its own nest, which means the colonies are not genetically related. In colonies that are
established further south, where they practice eusocial behavior, they are genetically related within
the colonies. For example, the first brood, which yields several workers, helps the foundress in her
colony. The first brood therefore stays in the colony, and is directly genetically related to the mother
by half, and helps to raise the second brood, which is also related to them by half.
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Genetic telatedness among colonies of different behaviors

[edit]

There are colonies both of eusocial and solitary behavior that have been studied for environmental
differences and genetics. It has been shown that there is a stronger link of genetic relatedness
between two colonies with similar behavioral patterns, than those of closer geographic distance and
different social behaviors. This does not necessarily mean that social behavior is governed by
certain genes, but it could be linked to certain genetic lineages that are more suited for certain
environments. Although there is much more studying that must be done on the correlation between
genetics and the environment and social behavior, it has been recorded that there is some sort of
link between the three parameters. Evolutionary characteristics are involved in the crosslink
between the northern populations of H. rubicundus having more solitary behavior, and the southern
populations being eusocial. It is a possibility, however, than this link was brought on by
environmental control of sociality, rather than a purely genetic standpoint.
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Pheromone tecognition[edit]
Halictid bees have a gland known as the Dufours gland that extends throughout the abdomen. It is
found primarily in female Hymenoptera. The Dufours gland, which is associated with the sting
structure, secretes fluids that are important for socioecological functioning. In H. rubicundus, the
Dufours gland produces pheromones that may aid females in recognizing brood cells as well as
other individuals in the nest.
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Costs and benefits of sociality[edit]

In each different environment, there have been adaptations to minimize the costs and maximize the
benefits of their social behavior. In bees with solitary behavior, the cold environments have likely
allowed for this behavior because of shorter breeding seasons. Having shorter growing seasons
would degrade the possibility of having a second brood in the same season, which changes the
behavior of the worker bees. There would be no benefit to the worker bees to stay with the mother
and help her maintain the colony, as she would not be able to produce another brood that season,
so there would be kinship selection benefits. Instead, the first brood in a colony of solitary behavior
consists of many gynes, which are potential foundresses, who must mate before the short season is
over in order to establish their own colony the following year. On the other end of the spectrum, the
populations that exhibit eusocial behavior had completely different costs and benefits to their
sociality. As discussed in the sections about genetic relatedness, there are benefits to worker bee
behaviors in eusocial colonies, as they are genetically related to the foundress and to next brood.
Thus, by kin selection, it is beneficial to the workers who cannot mate to help the mother
reproduce a healthy second brood that can then pass on half of the same genes they share.
However, it is costly to the workers to be in this social environment because they cannot reproduce
themselves and directly pass down the genes that they have. In both situations, however, it is more
beneficial to exhibit the certain social behavior that they do, than to find an alternative strategy of
sociality.
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Sex ratio in different colonies[edit]

Social colony:
In the social colony where there are two broods in one cycle, there are different sex ratios for each
brood. The first brood will contain 75100% females to provide plenty of workers for the nest to help
the mother produce a second brood. The second brood is slightly male-biased, resulting in a sex
ratio of about 60% males. The products of the second brood will act as reproductives (both the
males and females).
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Solitary colony:
In solitary populations, nesting begins later and only one brood is produced. The brood has a sex
ratio made up of 60% males, similar to that of the second brood in social colonies. This brood is also
produced at about the same time that the second brood is produced in social populations so that the
colony cycle ends at about the same time.
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Sex ratio effects on sociality[edit]

Male abundance and ability to mate have a major effect on deciding the social behavior of a nongyne versus gyne bee. Therefore, the sex ratio is important to consider. Warmer temperatures for
the first brood of a foundress in the spring leads to a higher ratio of male to female offspring. The
best predictor for a female's fate in being a gyne or a non-gyne depends on the male abundance in
proportion to virgin females. This sex ratio is affected by temperature and photoperiod of the male
egg production. It is proposed that this male bias largely decides the demographic of the colony or
population, which influences the sociality that is seen in that population, whether it be largely
populated by gynes who go off to create their own nests, or non-gynes who stick around to stay in
their nesting colony with their foundress.
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Hierarchy conflict[edit]
There is a social caste system in the H. rubicundus species that is affected not solely by genetics,
but behaviors in the first few days of adult life. In studying these bees however, it was found that
foundresses were never worker bees, and only about five percent of all foundresses ever carry
pollen. The future foundresses were observed leaving the nest within a few days of their adulthood.
Males do not overwinter, so they must find a mate before the female hibernates for the winter.
Because there is no predetermined physiological change in the females that decide what level in
the caste system they will be, it can be inferred that this is determined by behavioral conditions.
Therefore this hierarchy is established by the solitary or eusocial behavior that the population
exhibits and the foundress's control over the behavior of its broods. The females must actively seek
a mate within the first few days of its life if it wants to leave it's mother's nest and have a life as a
gyne and potential foundress.
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Body size as a result of temperature[edit]

Temperature affects the size of offspring in Halictus rubicundus, but there are a couple of
hypotheses that offer explanations as to why such is the case:
1. At optimum temperatures, the maximum offspring size will be produced. Smaller sizes will be
the result of both higher and lower temperatures based on differing stresses. So varying
conditions around one optimum temperature will lead to different offspring sizes.
2. Extreme temperatures may reduce flowering and therefore have an indirect effect on cell
provisioning. If temperatures are lower than optimum, lower flowers will be produced and
fewer resources will be available to increase growth of offspring.
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Parasites[edit]
Halictus rubicundus are parasitized by other hymenopteran species. Although it would seem as
though nesting in dense groups would draw attention to aggregations and increase mortality by
parasitism, such is not observed to be the case. Because the sweat bees nest densely, we can
assume that there is a large dilution effect that proportionally decrease mortality rates by parasites.

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