Lasioglossum malachurum

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Lasioglossum malachurum
Lasioglossum malachurum female 1.jpg
Female
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Halictidae
Tribe: Halictini
Genus: Lasioglossum
Species:
L. malachurum
Binomial name
Lasioglossum malachurum
(Kirby, 1802)
Synonyms
  • Melitta malachurumKirby, 1802
  • Lasioglossum longulumCurtis, 1833

Lasioglossum malachurum, the sharp-collared furrow bee, is a small European halictid bee. [1] This species is obligately eusocial, with queens and workers, though the differences between the castes are not nearly as extreme as in honey bees. [2] Early taxonomists mistakenly assigned the worker females to a different species from the queens. They are small (about 1 cm), shiny, mostly black bees with off-white hair bands at the bases of the abdominal segments. [1] 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. [3]

Contents

Taxonomy and phylogeny

L. malachurum was described by the entomologist William Kirby in 1802. This species of bees fall within the genus Lasioglossum, which is the largest bee genus. [1] Lasioglossum falls within the family Halictidae, which includes small to midsized bees and is commonly referred to as the sweat bee family because the Halictidae are frequently attracted to human perspiration. [4] L. malachurum falls within the order Hymenoptera, a large order of generally winged insects containing wasps, bees, and ants. One species of lasioglossum that is closely related to the L. malachurum is L. hemichalceum. [5]

Description

Lasioglossum.malachurum Lasioglossum.malachurum.-.lindsey.jpg
Lasioglossum.malachurum

Caste dimorphism

Physical size is a major distinguishing feature between queens and gynes versus female worker bees. Queens consistently are larger in body size than workers. Queens also tend to have worn wings and worn mandibles as a result of higher activity, in addition to greater ovarian development correlating with the reproductive capacity of queens. Queens also generally have abundant fat stores. [3] This physical dimorphism persists throughout all stages of the lifecycle, from pupae to adult. Rarely, queens can be abnormally small compared to the size of workers, but abnormally large workers have never been observed in the wild. [1]

Nest identification

The sweat bee has a simple nesting architecture consisting of a vertical tunnel into the ground ending in a compartment reserved for brood-rearing. [3] They nest underground, like Lasioglossum zephyrus , but they do not form the same complex structures. [6] Nest entrances are frequently concealed using scattered foliage. [7] For L. malachurum, thousands of nests can be found located within a small region in nesting aggregations, which suggests that the bees likely have some sort of identification mechanism for conspecific nests. In fact, nest entrances and brood cell linings are both impregnated with secretions from the Dufour's gland of the bees, which plays a role in nest orientation. [8] Nests are dug into hard, compact soil and can be sealed from within by the queen using her abdomen. Because nests are dug into hard soil, construction of a nest represents a significant energy expense to a queen, which explains why gynes frequently usurp other nests rather than founding one of their own. [9] L. malachurum is a Western Palaearctic species and nests can be found across Southern England and the Channel Islands, most of continental Europe, and North Africa. [10] [11]

Distribution and habitat

L. malachurum is found across England, continental Europe, and northern Africa, so is capable of inhabiting a broad range of habitats and climates. Differences in the climes of the various habitats can frequently impact the social behavior of L. malachurum within that habitat. The length of summer, for example, can affect the length of the breeding season and thereby impact the number of broods a queen may have. The longer breeding seasons in Southern Europe can also lead to reproductive capacity in worker bees, making L. malachurum in those regions not purely eusocial. [3] Female workers in southern European colonies of L. malachurum consequently have significantly more developed ovaries than their counterparts in northern European colonies. [1]

Due to the relatively broad range of nesting habitats of the species, L. malachurum is subject to a myriad of climate-based selective pressures that cause a differential in behavior dependent upon location. Researchers have identified a tendency for L. malachurum in southern European climes to be characterized by more activity and the production of more worker broods prior to the production of a gyne brood, whereas L. malachurum in northern European climes exhibit less activity and only a single worker brood prior to the gyne brood. [1] [12]

Colony cycle

The queens of L. malachurum, following fertilization the previous year, begin to appear in the spring, when food sources are plentiful to sustain them after the long overwintering period. Although several females usually outwinter in the same burrow with little conflict, they start to act aggressively until a single female is left in possession of the burrow, leaving the evicted females to obtain or excavate burrows of their own. [1]

Each female with a nest tunnel then begins to build brood cells in short side passages which she excavates to the side of the main passage. Immediately following construction, each brood cell is mass-provisioned with a mixture of pollen and nectar in the form of a firm, doughy mass. An egg is laid on each pollen mass and the individual cell sealed by the female. Each egg takes 22 days to develop from an egg to a full adult. [13] She then goes on to construct more, similar cells containing eggs and pollen masses. [1] During this time, before the first worker brood has emerged, containing only about five workers, the foundress queen frequently leaves the nest to find provisions to build the brood cells of the nest. These nest absences are accompanied by high risk of intraspecific usurpation. [2] Once the brood is provisioned, the queen seals the nest synchronously with all other queens in the aggregation, and all above-ground activity ceases for three weeks. [9]

Larvae from the earliest eggs are full grown and start pupation by the end of May in Central Europe (or much earlier in warmer climates), emerging from their cells by mid-June. For bees in cooler climates with shorter breeding seasons, these adults are all non-reproductive female workers, somewhat smaller than their mother. The original maternal female bee remains within the nest and guards the entrance to the burrow, now acting as a queen while her non-reproductive daughters act as workers; they go out foraging for food and help in the construction of new brood cells, in which the queen lays new eggs. [1] For bees in warmer climates with longer breeding seasons, some of the female workers have reproductive capacity and can help breed up to two subsequent worker broods before the eventual gyne brood. [3]

Males begin to emerge from the final brood of the season by the beginning of August in Central Europe, or May–June in the Mediterranean region. Several days later, reproductive females begin to emerge, as well, which are morphologically similar to their queen. During sunny weather, the nest aggregation becomes a lek and the males vie for territory on the ground. [7] The males mate with the new reproductive females (from both their own and separate nests), although they do not attempt mating with the non-reproductive females. Impregnated females may continue to live in their mothers' nests, although it is thought that they only forage for their own food and do not contribute to the rest of the nest. [1]

With the arrival of the colder autumn weather, the males and non-reproductive females die off, and the impregnated reproductive females go on to spend the winter in diapause and repeat the lifecycle the following year. The lifecycle is longer, with two successive broods of workers, in southern Europe. [1] [12] Soil temperature specifically has been linked to decreasing the overall length of the nesting cycle. [13] Workers in warmer soil tend to require less time to provision a new brood, possibly because they have more active time each day. [13]

Behavior

Dominance hierarchies

L. malachurum colonies are monogynous. Although the queen coordinates the workers within the nest, the workers can exert pressures on the queen, as well. Because queens frequently are replaced through usurpation and queen turnover, the workers within a nest are less related to the queen than expected. As such, the workers are less likely to cooperate with the queen based on maximizing their own fitness. To get the workers to remain with the nest and continue to aid her own offspring, the queen often cedes reproduction to alien workers. [2] These workers can produce males or gynes and have high rates of reproduction. They do not, however, overwinter, nor are they morphologically similar to a queen or gyne. [3] The length of lifespan for the queens may be a factor that leads to more reproductive workers in southern climes that have longer breeding seasons; in northern climes, where the queen is generally alive for production of gynes, workers are less likely to be reproductive. In southern climes, where the queen frequently does not survive to gyne production, workers are more likely to be reproductive. [1]

Communication and recognition

Male-female communication

Communication between males and females of L. malachurum is mediated using pheromones. [2] Virginal queens attract males through a sex hormone that mated queens lack; thus, males are much more attracted to younger queens. The chemical distinctions between the pheromones released by virginal and mated queens are significant in terms of proportions of hydrocarbons and lactones. Whereas isopentenyl esters, unsaturated fatty acids, and hydrocarbons are far more abundant in the chemical makeup of a virginal queen's hormones, the old queen's hormones are more likely to be made up of largely macrocyclic lactones. This chemical distinction between virginal and old females allows males to distinguish between the two, thus enabling male choice in sexual partner. They are also capable of discriminating between familiar and unknown queens, and l generally are more attracted to queens of a foreign colony, which likely arose as an outbreeding mating strategy, which increases genetic diversity of the population and decreases the probability that the next generation would be subject to genetic disease. [8]

Nest recognition

Because L. malachurum generally nests in aggregations of thousands of colonies within a small region, the ability of bees to recognize their own nests is crucial. Outside the entrances to the nest, guards are posted. These guards mark the nest entrances with Dufour's gland secretions to create olfactory cues to guide colony specific bees back to the nest. These secretions are largely made up of alkanes and alkenes. [8] Bees of the species L. malachurum also rely on visual cues and landmarks to determine where their nests are, which they establish through orientation flights and continuously update through reorientation flights. [14]

Nestmate discrimination

Nestmates are able to identify other nestmates using olfactory cues, as well. Chemical similarities, specifically in nonvolatile alkenes and alkanes, provide this cue. Notably, the component of discrimination is not based on kinship, but on shared nesting site, and individual bees of L. malachurum do not behave differently toward related nestmates and unrelated nestmates. In a given nest, the many different compounds mix together to form a gestalt odor that marks all of the members of the nest. This form of scent blending is found commonly among other members of Hymenoptera including certain bees, wasps, and ants. The gestalt bouquet of odors differs significantly in terms of chemical composition between different nests. [15]

Usurpation

Queens frequently can be usurped by intraspecific gynes, because the creation of a new nest is a significant energy investment for gynes. [9] The probability of a colony usurpation increases steadily as the season progresses, most likely because the probability of a gyne having the time and resources to found a new colony and produce new workers decreases. [2] The increasing risk of usurpation may play a role in the small worker broods, because the queen ceases foraging and seals her nest in mid-May to protect from usurping gynes. [2] Early nest closure has been favored by pressure from usurpation and has developed into an evolutionarily stable strategy. [9] It might also explain why queens are highly aggressive toward other conspecific queens. [2] In confrontations between foundresses and usurping gynes, female size is a significant determinant of outcome because it gives the queen or usurper a significant advantage. Although they have a physical advantage in usurpation, larger queens are more likely to found their own nests rather than usurp another. [9]

Mating behavior

The nest aggregation of thousands of colonies that persists throughout the year becomes a lek mating system on warm summer days in the latter half of the breeding season (July through September). The daily operational sex ratio has far more males than females, which is common in a lek mating system and among other bees. [7] The lek mating is a system in which a large group of males clusters at sites visited by females for mating. [16] Males are attracted during mating by a mix of olfactory cues produced in multiple glandular sources from the female. Males seek out unrelated, unmated females; because females are only receptive for a short period following their emergence, males must seek out and find an unmated gyne before the large number of other males. [7] Mating can occur either in the female's nest or on flowers. Queens can be polyandrous, but only rarely do they actually mate with multiple males. Most colonies arise from only one queen and one male. [2]

Aggression

Aggression is commonly seen between conspecifics of all halictid bees; especially between usurper females, drawn out fights can occur that last for nearly a half-hour and result in damage to or loss of limbs and body parts. Guard bees of the L. malachurum species, which are workers that defend the nest, also demonstrate antagonistic behavior toward non-nestmate conspecifics, by orienting their stingers toward the intruders or blocking the entrance to the nest with their abdomens. They are able to discriminate between non-nestmates and nestmates, which they allow to pass with ease. Foragers, though, are either unable to discriminate between nestmate and non-nestmate conspecifics or are uninterested in pursuing aggression against non-nestmate conspecifics, so exhibit high levels of tolerance toward all conspecifics. [17]

Pollination

L. malachurum bees are polylectic, meaning that they collect pollen from a broad range of unrelated plant species. Although L. malachurum displays opportunism when selecting flowers from which to extract pollen, they will generally narrow their selection of pollen during a given collection period. The species demonstrates floral consistency, and generally only collects from one pollen source during a given pollen flight. The pollination behavior of L. malachurum may be useful for humans to cultivate and develop, because the bee has been implicated in pollinating species of plants that are commonly used by humans for food and for medication. [18]

Parasites

L. malachurum colonies are parasitized by the kleptoparasitic species of the genus Sphecodes. The bees of this genus have developed a strategy for invading the nest of host species that involves avoiding interaction with any host individuals. The parasites generally parasitize L. malachurum nests during the early spring when the foundress queens need to leave their nests unattended to forage. Other species enter guarded nests during the second brood cycle and kill the guard, as well all other present host individuals, before laying eggs in the brood cells. After oviposition, the parasite closes the brood cells to prevent the host female from returning and replacing the parasite eggs. Because the parasites in the genus Sphecodes parasitize a broad range of other bee species, few adaptations to the specific L. malachurum bee colonies exist. [19]

Related Research Articles

<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. They are the group for which the term 'eusocial' was first coined by entomologist, Suzanne Batra.

<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 entered 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</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>Polistes metricus</i> Species of wasp

Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes metricus can be found attached to the sides of buildings, trees, and shrubbery.

<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>Lasioglossum zephyrus</i> Species of bee

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". It is considered a primitively eusocial bee, although it may be facultatively solitary. The species nests in burrows in the soil.

<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>Polistes biglumis</i> Species of wasp

Polistes biglumis is a species of social wasp within Polistes, the most common genus of paper wasp. It is distinguished mainly by its tendency to reside in montane climates in meadows or alpine areas. Selection pressure from the wasp's environment has led to several idiosyncrasies of its behavior and lifecycle with respect to its relative species in the genus Polistes. It alone among paper wasps is often polyandrous. In addition, it has a truncated nesting season that gives rise to unique competitive dynamics among females of the species. P. biglumis wasps use an odor-based recognition system that is the basis for all wasp-to-wasp interaction of the species. The wasp's lifecycle is highly intertwined with that of Polistes atrimandibularis, an obligate social parasite wasp that frequently invades the combs of P. biglumis wasps.

<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 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.

<i>Nannotrigona testaceicornis</i> Species of bee

Nannotrigona testaceicornis is a eusocial stingless bee species of the order Hymenoptera and the genus Nannotrigona. Its local common name is abelhas iraí. This species has a large geographic distribution and occupies different biomes, including urban areas, around Neotropical America. The bees of this species nest in trees or artificial cavities because of this broad distribution. N. testaceicornis is important for agriculture because it will pollinate a vast number of plant species year round.

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>Augochlorella aurata</i> Species of insect

Augochlorella aurata is a primitively eusocial species of sweat bee in the family Halictidae. It is one of three species of Augochlorella found east of the Rocky Mountains in North America. The body is a brilliant green metallic color, diffused to varying extents with a copper, red, or yellow color. A. aurata is a generalist pollen feeder and likely an important pollinator for some horticultural crops. A common name is golden green sweat bee.

References

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  4. ↑ "Sweat Bees". Missouri Department of Conservation. Archived from the original on 2015-09-26.
  5. ↑ Danforth, Bryan N.; Eardley, Connal; Packer, Laurence; Walker, Kenneth; Pauly, Alain; Randrianambinintsoa, Fano José (2008-01-01). "Phylogeny of Halictidae with an emphasis on endemic African Halictinae". Apidologie. 39 (1): 86–101. doi: 10.1051/apido:2008002 . ISSN   0044-8435.
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  10. ↑ "Lasioglossum malachurum (Kirby, 1802) | BWARS". www.bwars.com. Retrieved 2015-09-26.
  11. ↑ "Lasioglossum malachurum -- Discover Life". www.discoverlife.org. Retrieved 2015-09-26.
  12. 1 2 Soro, A.; Ayasse, M.; Zobel, M. U.; Paxton, R. J. (2008-10-25). "Complex sociogenetic organization and the origin of unrelated workers in a eusocial sweat bee, Lasioglossum malachurum". Insectes Sociaux. 56 (1): 55–63. doi:10.1007/s00040-008-1037-y. ISSN   0020-1812. S2CID   9799543.
  13. 1 2 3 Weissel, N.; Mitesser, O.; Liebig, J.; Poethke, H.-J.; Strohm, E. (2006-11-09). "The influence of soil temperature on the nesting cycle of the halictid bee Lasioglossum malachurum". Insectes Sociaux. 53 (4): 390–398. doi:10.1007/s00040-005-0884-7. ISSN   0020-1812. S2CID   6326295.
  14. ↑ Brünnert, U.; Kelber, A.; Zeil, J. (1994-09-01). "Ground-nesting bees determine the location of their nest relative to a landmark by other than angular size cues". Journal of Comparative Physiology A. 175 (3): 363–369. doi:10.1007/BF00192995. ISSN   0340-7594. S2CID   45287998.
  15. ↑ Soro, Antonella; Ayasse, Manfred; Zobel, Marion U.; Paxton, Robert J. (2010-10-22). "Kin discriminators in the eusocial sweat bee Lasioglossum malachurum: the reliability of cuticular and Dufour's gland odours". Behavioral Ecology and Sociobiology. 65 (4): 641–653. doi:10.1007/s00265-010-1066-1. ISSN   0340-5443. S2CID   19379999.
  16. ↑ Beehler, Bruce; Foster, Mercedes (February 1988). "Hotshots, Hotspots, and Female Preference in the Organization of Lek Mating System". The American Naturalist. 131 (2): 203–219. doi:10.1086/284786. S2CID   85295271.
  17. ↑ Polidori, Carlo; Borruso, Luigimaria (2011-06-19). "Socially peaceful: foragers of the eusocial bee Lasioglossum malachurum are not aggressive against non-nestmates in circle-tube arenas". Acta Ethologica. 15 (1): 15–23. doi:10.1007/s10211-011-0103-z. ISSN   0873-9749. S2CID   254165941.
  18. ↑ Polidori, Carlo; Rubichi, Alice; Barbieri, Valeria; Trombino, Luca; Donegana, Marta (11 December 2009). "Floral Resources and Nesting Requirements of the Ground-Nesting Social Bee, Lasioglossum malachurum (Hymenoptera: Halictidae), in a Mediterranean Semiagricultural Landscape". Psyche: A Journal of Entomology. 2010: 1–11. doi: 10.1155/2010/851947 .
  19. ↑ Sick, M.; Ayasse, M.; Tengö, J.; Engels, W.; Lübke, G.; Francke, W. (1994-01-01). "Host-parasite relationships in six species of Sphecodes bees and their halictid hosts: Nest intrusion, intranidal behavior, and Dufour's gland volatiles (Hymenoptera: Halictidae)". Journal of Insect Behavior. 7 (1): 101–117. doi:10.1007/BF01989830. ISSN   0892-7553. S2CID   21839345.
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