Lasioglossum zephyrus

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Lasioglossum zephyrus
Sweat Bee (Lasioglossum zephyrum) and a Cuckoo Wasp (8084530876).jpg
L. zephyrus (bottom) with a cuckoo wasp
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Hymenoptera
Family:
Halictidae
Genus:
Lasioglossum
Species:
L. zephyrus
Binomial name
Lasioglossum zephyrus
(Smith, 1853) [1]
Synonyms
  • Halictus zephyrusSmith, 1853
  • Halictus (Chloralictus) academicusSandhouse, 1924
  • Halictus (Chloralictus) vintonensisSandhouse, 1924
  • Lasioglossum zephyrumAuctt. (Missp.)
  • Lassioglossum [sic] zypherumFalin, 2003 (Missp.)

Lasioglossum zephyrus is a sweat bee of the family Halictidae, found in the U.S. and Canada. It appears in the literature primarily under the misspelling "zephyrum". [1] It is considered a primitively eusocial bee (meaning that they do not have a permanent division of labor within colonies), [2] although it may be facultatively solitary (i.e., displaying both solitary and eusocial behaviors). [1] [3] The species nests in burrows in the soil.

Contents

Taxonomy and phylogeny

The order Hymenoptera contains more eusocial species than any other order. [4] Eusociality has many origins and is present in many degrees within Hymenoptera, with L. zephyrus being one of the more primitive examples. [2] [4] The genus Lasioglossum is predominantly eusocial, but has a high level of variation in sociality. [4] For instance, Lasioglossum zephyrus, as well as another species of this family of bees, Halictidae, Halictus ligatus , are known for their characteristic primitively eusocial behavior. This unique type of eusociality gives insight about the evolutionary background of eusociality of these species. [5] Six species within this genus have reverted to solitary life, social polymorphism, or parasitism. [4]

Description and identification

Lasioglossum zephyrus is characterized by its dark green metallic color, reddish abdomen, and a hairier face than most other species. Males are 6 to 7 millimeters long, slightly larger than females. Males are distinguished from females by their brighter green color and redder abdomens. [6] Since the castes vary along a spectrum, there is no definite way to distinguish a worker from a queen based on appearance alone. [7] However, queens can be identified by their behavior, pushing subordinates down in the nest and nudging them to inhibit their reproduction. [8]

Distribution and habitat

L. zephyrus has been found throughout the United States, in the months of March through October. [6] [9] Nests are usually constructed in April along the south facing edges of streams, and are clustered in aggregations of up to 1,000 nests. [7] [10] It nests in burrows that are typically constructed by young females. However, older females may also contribute to burrowing if their nests have been destroyed. These females excavate primarily at night, but activity has been reported throughout the day. Workers use their mandibles to loosen the soil, then carry it a short distance to be picked up by another bee. This soil is ultimately smoothed over the walls or used to fill evacuated old burrows. Females excavate cells and line them with a liquid produced in their enlarged Dufour's glands and secreted from the apex of the abdomen. [9]

Colony cycle

Lasioglossum zephyrus nests consist of less than 20 individuals. [9] During the spring, around April, new nests are established by one or more females. [7] [9] Some of these nest may never gain more members, and the female will remain solitary. If they do expand, the colony will reach its maximum size of 10–20 around August. [3] [9] Growth of the colony is gradual throughout the summer. [7] Inseminated young queens overwinter in their nests so that the cycle may be repeated. [3] [7] However, overwintered queens are likely to die early on in the summer. After the death of an old queen, a new queen takes over within hours. [9] L. zephyrus colonies have a high mortality rate, especially when inhabited by a solitary bee lacking guards. [7] [8]

Behavior

Dominance hierarchy

Roles within the colony are indirectly determined by age or physical differences, but direct determination is made by dominance interactions. [9] This type of hierarchy is very different from other species of bees, such as Lasioglossum hemichalceum , which is characterized by its egalitarian behavior and lack of aggression. All female L. hemichalceum are able to reproduce and cooperate to raise their young. [11] Queens have been shown to back workers into the depths of the burrows, preventing them from foraging and growing larger queen sized ovaries. [8] The tendency to display these behaviors is not influenced by the genetic relatedness of the subordinate bee, but it does decrease with time. In general, within colonies more closely related, there tends to be less aggression between bees. [12] When the queen dies, another worker takes her place and begins reproduction. [8] Among females, queens are at the top of the dominance hierarchy with their developed ovaries. Second are the guards, with some ovarian development. Foragers are the most subordinate group. [12]

Division of labor

Young females are responsible for constructing the nest. As adults, Lasioglossum zephyrus females become either mated egg-layers or workers who forage, guard, and make cells. [9] In a typical Lasioglossum zephyrus colony, the oldest females tend to be mated queens. However, among similarly aged bees, the largest is generally the queen. [3] The queen does not collect pollen or construct cells after her first batch of workers has hatched. [13] Queens and workers are not very physically distinct, since bee size varies on a continuum. [3]

Male behavior

Males are not considered productive members of the nest since they do not remain in their home nests. Instead, they form swarms around clusters of nests as early as 2 days after hatching. [7] [14] [15] Early in the spring, the males are solitary or clustered in small groups, but by late summer they gather in swarms of thousands of bees. Within these swarms there are head-on collisions between bees as they search for females and flowers. [15] While foraging, L. zephyrus searches for flowers, sometimes even forcing them open in order to get first access to the pollen. [16] Males have been seen landing on small pebbles and small objects as if they were females and have been observed mating as young as 4 days old up until death. In the laboratory, males were observed to have specific patrolling routes, flying between specific objects in sequence. They sleep in holes, plants, or abandoned burrows, solitarily or in small groups. [15]

Reproductive suppression

Most queens will mate during their lifetimes, but some workers may mate as well since they are physiologically capable. [3] In fact, about 18% of workers mate successfully between the months of June and August. When a queen is present, workers are not typically receptive to mating. [14] Females have been shown to partake in egg cannibalism, eating the eggs of nest mates, thus inhibiting their reproduction, [9] as well as the aggressive behaviors described in the dominance hierarchy subsection.

Mating behavior

Mating tends to take place near the nest's entrance, where the males typically fly. [14] Males will pursue females on land, but not in the air, and have been shown to aggressively attack females at the nest's entrance. Males will approach and occasionally grasp bees of other species, but no heterospecific matings have been documented. Females secrete an odor that functions as an aphrodisiac and a sex attractant. This odor is produced by female virgins between the ages of 2 and 8 days. The courtship period is short, and copulation occurs with the male above the female. In the field, L. zephyrus has been seen to copulate for 10–42 seconds. Most of the males' attempts at mating were met with resistance, but the females tend to remain relatively motionless during copulation. Many females are successful at preventing unwanted matings, suggesting that they have at least some control over their choice of mate. As many as four males have been observed stacked on top of a copulating male in attempt to mate with that female. [17]

Communication and recognition

Individual bees can recognize their nest and nest mates by their odors, which helps to prevent parasitism by unrelated L. zephyrus as well as other species. [18] [19] This recognition has been documented in both males and females. Males can identify individual females by their unique odors and learn to avoid females who are unreceptive to mating. [19] L. zephyrus has also been shown to leave foreign nests even without being attacked, which supports the theory that they use odors to identify their home nests. [18]

Kin selection

Genetic relatedness

Haplodiploid sex determination Haplodiploid-sex-determination-system3.png
Haplodiploid sex determination

In Hymenoptera species, like Lasioglossum zephyrus, females are diploid and males are typically haploid, in a system called haplodiploidy. However, in inbred L. zephyrus populations, and also in other hymenopteran species, diploid males have been discovered. [20] Hymenopteran workers are expected to be related by a coefficient of 3/4, assuming the queen only mates only once. [10] [14] However, in those nests studied, the relatedness coefficient was found to be lower, around 0.7. This could be explained by the insemination of multiple queens. With a relationship this high, significant altruism is likely. [10] The workers share only half of their genes with the queen, so they have more genes in common with their sisters. Since workers are more closely related to their sisters than their own offspring, whose relationship coefficient would be 0.5, kin selection has been proposed as an additional reason why workers mate so infrequently. They can pass down more genes through sisters than the same number of direct offspring. [14] Nests have been shown to be more genetically similar to nests in close proximity. [10]

Kin recognition and discrimination

Once an individual has learned the odor of its kin, it can successfully identify kin it has never met, since odors are genetically determined. Additionally, it can determine whether this relative is a close or distant relative. [21] While learning does play a role in kin recognition, the genetic differences in odor are thought to be more critical in kin recognition. Kin recognition is beneficial for defense of the entire nest, as well as at the individual level. Individuals benefit by contributing only to the fitness of their own related nest mates. [18]

Diet

Adult Lasioglossum zephyrus feed their larva pollen from surrounding plants. Since they forage at a variety of different flowers, they collect multiple types of pollen. Some types are more protein dense than others, but the adults do not compensate for the difference in pollen quality. All offspring receive the same amount of pollen, which results in offspring of different sizes. Size is an advantage for females because it gives them a better chance of becoming a queen. They also tend to produce more eggs of better quality, and greater size increases the chance that a female survives the winter. Larger males are better able to defend territories and compete for mates. Size is also a factor for both species in regards to flight. Larger body size leads to an increased body temperature and the ability to fly sooner in the season and with greater frequency. [22] This species has been observed forcing open unbloomed flowers of species Xyris tennesseensis to extract the pollen, ensuring first and exclusive access. [16]

Interactions

Parasites

L. zephyrus has been parasitized by diverse organisms including bacteria, fungi, protozoa, and other species of hymenoptera. Nematodes have been found in the abdominal cavities of L. zephyrus females, and have led to a reduction in ovarian development. Parasites that attack in the spring during nest founding tend to be most successful. If threatened, the bee may try to attack it or decapitate the intruder. Guard bees are often responsible for the protection of the nest and will send one member to attack while the others block the entrance with their abdomens.

Nematode removed from the abdominal cavity of a hornet Asian Hornet (Vespa velutina) & Mermithid nematode (black background).JPG
Nematode removed from the abdominal cavity of a hornet

Nests containing only infected females are in danger of dying out. While infected bees are able to excavate burrows, they are less active than healthy bees and do not collect pollen to make cells.

Gregarine protozoa have been found generally in older specimens of L. zephyrus. It is thought that once a bee is infected, it takes time for the spores to fully develop. These parasites have only been found in females, although it is unclear why males remain unaffected. [7]

The bee species Lasioglossum cephalotes is a kleptoparasite of L. zephyrus. [7] The mutillid Pseudomethoca frigida is also a parasitoid, and known to engage in aggressive fights with female L. zephyrus. Instead of stinging its opponent, L. zephyrus attempts to decapitate it. The mutillid retreats, unable to defend itself. [7]

Related Research Articles

<span class="mw-page-title-main">Bee</span> Clade of insects

Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamily Apoidea. They are presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families. Some species – including honey bees, bumblebees, and stingless bees – live socially in colonies while most species (>90%) – including mason bees, carpenter bees, leafcutter bees, and sweat bees – are solitary.

<span class="mw-page-title-main">Halictidae</span> Family of bees

Halictidae is the second-largest family of bees with nearly 4,500 species. They are commonly called sweat bees, as they are often attracted to perspiration. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every continent except Antarctica. Usually dark-colored and often metallic, halictids are found in various sizes, colors and patterns. Several species are all or partly green and a few are red, purple, or blue. A number of them have yellow markings, especially the males, which commonly have yellow faces, a pattern widespread among the various families of bees. The family is one of many with short tongues and is best distinguished by the arcuate basal vein found on the wing. Females in this family tend to be larger than the males.

<i>Halictus rubicundus</i> Species of bee

Halictus rubicundus, the orange-legged furrow bee, is a species of sweat bee found throughout the Northern Hemisphere. 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 at times of low sea level during the Pleistocene epoch.

<i>Lasioglossum malachurum</i> Species of bee

Lasioglossum malachurum, the sharp-collared furrow bee, is a small European halictid bee. This species is obligately eusocial, with queens and workers, though the differences between the castes are not nearly as extreme as in honey bees. Early taxonomists mistakenly assigned the worker females to a different species from the queens. They are small, shiny, mostly black bees with off-white hair bands at the bases of the abdominal segments. L. malachurum is one of the more extensively studied species in the genus Lasioglossum, also known as sweat bees. Researchers have discovered that the eusocial behavior in colonies of L. malachurum varies significantly dependent upon the region of Europe in which each colony is located.

<i>Lasioglossum</i> Genus of insects

The sweat bee genus Lasioglossum is the largest of all bee genera, containing over 1800 species in numerous subgenera worldwide. They are highly variable in size, coloration, and sculpture; among the more unusual variants, some are cleptoparasites, some are nocturnal, and some are oligolectic. Most Lasioglossum species nest in the ground, but some nest in rotten logs.

<span class="mw-page-title-main">Eusociality</span> Highest level of animal sociality a species can attain

Eusociality, the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform at least one behavior characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.

<i>Ropalidia marginata</i> Species of insect

Ropalidia marginata is an Old World species of paper wasp. It is primitively eusocial, not showing the same bias in brood care seen in other social insects with greater asymmetry in relatedness. The species employs a variety of colony founding strategies, sometimes with single founders and sometimes in groups of variable number. The queen does not use physical dominance to control workers; there is evidence of pheromones being used to suppress other female workers from overtaking queenship.

<span class="mw-page-title-main">Halictinae</span> Subfamily of bees

Within the insect order Hymenoptera, the Halictinae are the largest, most diverse, and most recently diverged of the four halictid subfamilies. They comprise over 2400 bee species belonging to the five taxonomic tribes Augochlorini, Thrinchostomini, Caenohalictini, Sphecodini, and Halictini, which some entomologists alternatively organize into the two tribes Augochlorini and Halictini.

<i>Halictus ligatus</i> Species of bee

Halictus ligatus is a species of sweat bee from the family Halictidae, among the species that mine or burrow into the ground to create their nests. H. ligatus, like Lasioglossum zephyrus, is a primitively eusocial bee species, in which aggression is one of the most influential behaviors for establishing hierarchy within the colony, and H. ligatus exhibits both reproductive division of labor and overlapping generations.

<i>Megalopta genalis</i> Species of bee

Megalopta genalis is a species of the family Halictidae, otherwise known as the sweat bees. The bee is native to Central and South America. Its eyes have anatomical adaptations that make them 27 times more sensitive to light than diurnal bees, giving it the ability to be nocturnal. However, its eyes are not completely different from other diurnal bees, but are still apposition compound eyes. The difference therefore lies purely in adaptations to become nocturnal, increasing the success of foraging and minimizing the danger of doing so from predation. This species has served as a model organism in studies of social behavior and night vision in bees.

<i>Lasioglossum cressonii</i> Species of insect

Lasioglossum cressonii is a species in the sweat bee genus Lasioglossum, family Halictidae. Halictidae exhibit eusocial hierarchy behavior which is interesting given that eusociality in this group is hard to evolve and easy to lose. L. cressonii is found throughout North America. L. cressonii have been shown to be important pollinators for apple trees and many other North American native plants.

<i>Lasioglossum vierecki</i> Species of bee

Lasioglossum vierecki, also known as Dialictus vierecki and Halictus vierecki, is a sand sweat bee and is part of the family Halictidae of the order Hymenoptera. It is found in the eastern half of North America from Minnesota to the New England States down to Georgia and Louisiana and up in Manitoba and Ontario. Commonly found in sandy areas, it pollinates various flowers such as grass-leaved goldenrod and rattlesnake master.

<i>Lasioglossum hemichalceum</i> Species of bee

Lasioglossum hemichalceum, which has sometimes been confused with L. erythrurum, is a sweat bee endemic to Australia. Large numbers of unrelated females will typically share a single nest, a behavior referred to as "communal". Nests are constructed underground by the independent efforts of the females. L. hemichalceum will typically begin creating new colonies during the summer, with brood production from late November through the first few months of spring. Members of this species do not demonstrate aggressive behavior towards one another. As the size of the colony increases, the reproductive potential of each female does not change, unlike many species of bees.

Lasioglossum figueresi, formerly known as Dialictus figueresi, is a solitary sweat bee that is part of the family Halictidae of the order Hymenoptera. Found in Central America, it nests in vertical earthen banks which are normally inhabited by one, though sometimes two or even three, females. Females die before their larvae hatch. It was named after José Figueres Ferrer, a famous Costa Rican patriot, and studies of its behavior are now general models for social behavior studies.

Lasioglossum aeneiventre, also known as Dialictus aeneiventre, is a social sweat bee and is part of the family Halictidae of the order Hymenoptera. Found in Central America, it nests mostly on flat ground though sometimes in vertical banks. It is often compared to L. figueresi.

<i>Augochlora pura</i> Species of insect

Augochlora pura is a solitary sweat bee found primarily in the Eastern United States. It is known for its bright green color and its tendency to forage on a variety of plants. Inhabiting rotting logs, this bee can produce up to three generations per year. Both males and females have been observed licking sweat from human skin, most likely seeking salt

<i>Halictus sexcinctus</i> Species of bee

Halictus sexcinctus, commonly referred to as the six-banded furrow bee, is a species of sweat bee found throughout Europe and as far east as Asian Turkey and Iraq.The H. sexcinctus can be easily confused with the closely related species, Halictus scabiosae, due to very similar morphological features. H. sexcinctus show a social polymorphism in which different colonies can exhibit solitary, communal, or eusocial structure. Due to this large variance in social organization, it was suspected that it was not one species at all, but rather multiple, cryptic species. However, genetic analysis was able to confirm these varying populations as one species. H. sexcinctus will forage from multiple flower species, but prefers plant species with wide-open flowers. Their nests can be found dug into the ground in loamy or sandy soil.

<span class="mw-page-title-main">Dialictus</span> Subgenus of insects

Dialictus is a subgenus of sweat bees belonging to the genus Lasioglossum. Most of the members of this subgenus have a metallic appearance, while some are non-metallic. There are over 630 species worldwide. They are commonly found in the Northern Hemisphere and are found in abundance in North America. Members of this subgenus also have very diverse forms of social structure making them model organisms for studying the social behavior of bees.

<i>Augochlorella</i> Genus of bees

Augochlorella is a genus in the bee family Halictidae, commonly called sweat bees. They display metallic coloration, ranging from reddish to gold to bluish green, as is typical for other genera in the tribe Augochlorini.

<i>Bombus vancouverensis</i> Species of bee

Bombus vancouverensis is a common species of eusocial bumblebee of the subgenus Pyrobombus. B. vancouverensis inhabits mountainous regions of western North America, where it has long been considered as a synonym of Bombus bifarius, and essentially all of the literature on bifarius refers instead to vancouverensis. B. vancouverensis has been identified as one of the two species of bumblebee observed to use pheromones in kin recognition. The other is the frigid bumblebee, Bombus frigidus.

References

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  3. 1 2 3 4 5 6 Interactions in colonies of primitively social bees: Artificial colonies of Lasioglossum zephyrum . PNAS. Retrieved 08-27-2011.
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  5. ↑ Pabalan, N.; Davey, K.G.; Packer, L. (2000). "Escalation of Aggressive Interactions During Staged Encounters in Halictus ligatus Say (Hymenoptera: Halictidae), with a Comparison of Circle Tube Behaviors with Other Halictine Species'". Journal of Insect Behavior. 13 (5): 627–650. doi:10.1023/A:1007868725551. S2CID   20773517.
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  11. ↑ Kukuk, Penelope F. (1992-01-12). "Social Interactions and Familiarity in a Communal Halictine Bee Lasioglossum (Chilalictus) hemichalceum". Ethology. 91 (4): 291–300. doi:10.1111/j.1439-0310.1992.tb00870.x. ISSN   1439-0310.
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  13. ↑ Michener, Charles D.; Brothers, Denis J.; Kamm, Dwight R. (1971-04-01). "Interactions in Colonies of Primitively Social Bees: II, Some Queen-Worker Relations in Lasioglossum zephyrum". Journal of the Kansas Entomological Society. 44 (2): 276–279. JSTOR   25082413.
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  16. 1 2 Wall, M. A.; Teem, A. P.; Boyd, R. S. (Mar 2002). "Floral Manipulation by Lassioglosssum zephyrum (Hymenoptera: Halictidae) Ensures First Access to Floral Rewards by Initiating Premature Anthesis of Xyris tennesseenis (Xyridaceae) Flowers". Florida Entomologist. 85 (1): 290–291. doi: 10.1653/0015-4040(2002)085[0290:fmblzh]2.0.co;2 .
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  20. ↑ Kukuk, Penelope F.; May, Bernie (1990-09-01). "Diploid Males in a Primitively Eusocial Bee, Lasioglossum (Dialictus) zephyrum (Hymenoptera: Halictidae)". Evolution. 44 (6): 1522–1528. doi:10.2307/2409334. JSTOR   2409334. PMID   28564310.
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  22. ↑ Roulston, T'ai H.; Cane, James H. (2002-01-01). "The effect of pollen protein concentration on body size in the sweat bee Lasioglossum zephyrum (Hymenoptera: Apiformes)". Evolutionary Ecology. 16 (1): 49–65. doi:10.1023/A:1016048526475. ISSN   0269-7653. S2CID   23594452.
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