Eucera
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''Eucera'' is a genus of bees in the family
Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long antennae, especially in males. Eucera species can be found in diverse habitats, including meadows, fields, and urban gardens, primarily in the
Palearctic The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. Th ...
and
Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...
regions, covering parts of Europe, Asia, North Africa, and North America. Eucera bees play a crucial role in pollinating a wide range of flowering plants. They are solitary bees, meaning that each female builds and provisions her own nest without forming social colonies like
honeybees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the cur ...
. Female Eucera bees construct their nests in the ground, typically in well-drained, sandy soils. They lay their eggs within the nest and provide a food supply for their offspring by collecting
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
and
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
from flowers. Eucera bees are active from spring to fall, and their flight period often coincides with the blooming period of their preferred flowering plants. They are generalist
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
s, meaning they visit a wide variety of flowers, but some species show preferences for specific plants or families, such as
legume Legumes are plants in the pea family Fabaceae (or Leguminosae), or the fruit or seeds of such plants. When used as a dry grain for human consumption, the seeds are also called pulses. Legumes are grown agriculturally, primarily for human consum ...
s or
sunflowers ''Helianthus'' () is a genus comprising around 70 species of annual and perennial flowering plants in the daisy family Asteraceae commonly known as sunflowers. Except for three South American species, the species of ''Helianthus'' are native to ...
.


Description

As in most members of the tribe
Eucerini The Eucerini (often called longhorn bees or long-horned bees) are the most diverse tribe in the family Apidae, with over 32 genera worldwide that were previously classified as members of the family Anthophoridae. All species are solitary, though ...
, the antennae of males are very long. Old World ''Eucera'' can be identified through having five or six maxillary palpomeres, with the first
flagellomere An antenna (plural: antennae) is one of a pair of appendages used for sensing in arthropods. Antennae are sometimes referred to as ''feelers''. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in ...
shorter than the scape and the
clypeus The clypeus is one of the sclerites that make up the face of an arthropod. In insects, the clypeus delimits the lower margin of the face, with the labrum articulated along the ventral margin of the clypeus. The mandibles bracket the labrum, but ...
protruding in front of the
compound eye A compound eye is a Eye, visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidium, ommatidia, which are tiny independent photoreception units that consist of a cornea, lens (anatomy), lens, and p ...
by at least the width of the eye in side view. These characteristics are found in both sexes. Additionally, the males have convergent carinae on their sixth ventral abdominal segment (
sternite The sternum (: sterna) is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the su ...
). ''Eucera'' species nest in the ground.Michener, C. D. (2007) The bees of the world, second edition. Johns Hopkins University Press, Baltimore and London, 953 pp


Morphology

''Eucera'' have a thick layer of fur and are dark in color. They have a protuberant
clypeus The clypeus is one of the sclerites that make up the face of an arthropod. In insects, the clypeus delimits the lower margin of the face, with the labrum articulated along the ventral margin of the clypeus. The mandibles bracket the labrum, but ...
and are typically around 0.4 to 0.7 inches long. The size of bees in the genus ''Eucera'' range from 11 to 18 mm. When emerging from nests, an immature ''
Eucera nigrilabris ''Eucera'' is a genus of bees in the family Apidae, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long Antenna (biology), antennae, especially in males. Eucera species can ...
'' male will be somewhat red in color and a bit sluggish. A mature ''Eucera nigrilabris'' male will appear grey in color and more active.


Sexual dimorphism

In ''
Eucera berlandi ''Eucera'' is a genus of bees in the family Apidae, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long antennae, especially in males. Eucera species can be found in diver ...
'', males have long antennae containing three times the amount of
neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s for olfaction and ten times more pore plates than females.


Distribution

''Eucera'' is
holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
. Research shows that the ''Eucera'' complex originated in the Nearctic region in the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
and dispersed twice. The first dispersal having occurred as far as  24.2-16.6 million years ago during the warmer summer season and later again 13.9-12.3 million years ago during the springtime, allowing ''Eucera'' to thrive in cooler regions.


Phylogeny

''Eucera'' belongs to the tribe Eucerini and makes up roughly 50% of the tribe. Historically, the genus had 219 species classified in five subgenera, of which 78 species were known from Europe. However, six genera have recently been added to the genus ''Eucera'' as new subgenera: ''
Tetralonia ''Tetralonia'' is a genus of insects belonging to the family Apidae. The genus has almost cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the ...
,
Peponapis ''Peponapis'' is a genus of bees belonging to the family Apidae. They are Squash bees, specialized pollinators of squashes and related plants. The species of this genus are found in North America. Species: *'' Peponapis apiculata'' *'' Pepon ...
,
Syntrichalonia ''Syntrichalonia'' is a genus of exquisite long-horned bees in the family Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, i ...
,
Cemolobus ''Eucera (Cemolobus) ipomoeae'' was historically treated as a monotypic genus of long-horned bees in the family Apidae, but was recently included in the genus Eucera. There is one described species in the subgenus, ''C. ipomoeae,'' an oligol ...
, Xenoglossodes'' and ''
Xenoglossa ''Xenoglossa'' is a genus of large squash bees in the family Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bum ...
.''


Behavior

In the
Middle East The Middle East (term originally coined in English language) is a geopolitical region encompassing the Arabian Peninsula, the Levant, Turkey, Egypt, Iran, and Iraq. The term came into widespread usage by the United Kingdom and western Eur ...
, ''Eucera'' are active in the months of February to May.


Flight behavior

''Eucera'' have a spring flight season.


Nesting behavior

''Eucera'' are solitary bees that nest within the ground. They tend to nest in areas that are composed of clay or sand. It is a characteristic of all bees of the genus ''Eucera'' to have vertical and elongated cells within nests. There are around two to three cells per nest, which are found branching off of the main tunnel. In ''
Eucera nigrilabris ''Eucera'' is a genus of bees in the family Apidae, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long Antenna (biology), antennae, especially in males. Eucera species can ...
'', each tunnel harbors four to six cells. The first few cells are false cells and are not used for reproductive purposes. The cells below the false cells are used to lay eggs. Also, the entirety of the tunnel is lined with wax. For nesting, ''Eucera nigrilabris'' prefers soil of lower sodicity and salinity that has low calcium carbonate concentrations. They make lined nests that are about 85 cm into the ground. There has not been any observed occurrences of
kleptoparasitism Kleptoparasitism (originally spelt clepto-parasitism, meaning "parasitism by theft") is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct f ...
around ''Eucera nigrilabris'' nesting sites. '' Eucera palaestinae'' use their
Dufour's gland Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed a ...
to secrete a mixture of
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and Hydrophobe, hydrophobic; their odor is usually fain ...
s, methyl esters, and unsaturated fatty acids which provide the nest with an odor. This odor helps the bees locate their home when living in a dense aggregation of nests.


Mating behavior

'' Eucera kullenbergi'' is known to fall victim to sexual deception by '' Ophrys leochroma'' flowers that mimic the sex
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s of female bees. ''Eucera palaestinae'' live in dense nest aggregations with males emerging from their nests in the ground about a week before the females to take a look around the nesting site. Once the females emerge, males of the species will engage in aggressive competition to mate with them. Shortly after mating, the female becomes unreceptive. The initial attraction towards a female is by sight, but there is additional research suggesting that virgin females are distinguished by their specific scent and therefore are sought after more aggressively. It is thought that the cause of this sex attraction is due to small glands located on
tergites A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'margin'. ...
on the abdomen. In ''
Eucera nigrilabris ''Eucera'' is a genus of bees in the family Apidae, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long Antenna (biology), antennae, especially in males. Eucera species can ...
'', the males also emerge a few days earlier than females. Once the female emerges from the nest, males will fight with each other to mate. Mating time in this species occurs for 3–6 minutes, and once the female has mated, she becomes unresponsive.


Foraging behaviors

''Eucera'' can be generalists or specialists in foraging preference. They are able to pollinate both agricultural and naturally occurring plants. They pollinate plants in the deserts of Israel and the Mediterranean. ''Eucera,'' like other efficient foragers, avoid going back to the same food source after it has been previously depleted. Research suggests that ''Eucera'' use a combination of reward-based-patch-leaving rule and scent marking strategy to avoid returning to previously visited sites. ''Eucera'' resemble bumblebees in this matter but it is believed that they use different strategies (bumblebees use a numerical strategy) suggesting that the difference in strategies may reflect the lower learning capabilities of solitary bees like ''Eucera'' when compared to the social bumblebee. ''Eucera cinerea'' has specialized thick bristles that curve and are used for foraging for pollen within flowers that have hidden anthers. ''Eucera'' from southwest France are known to mainly forage from
Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...
and
Brassicaceae Brassicaceae () or (the older but equally valid) Cruciferae () is a medium-sized and economically important Family (biology), family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous pla ...
plants. ''Peponapis'' and ''Xenoglossa'' are known to be specialists for squash plants, such as ''
Cucurbita pepo ''Cucurbita pepo'' is a cultivated plant of the genus ''Cucurbita''. It yields varieties of winter squash and pumpkin, but the most widespread varieties belong to the subspecies ''Cucurbita pepo'' subsp. ''pepo'', called summer squash. It has b ...
''. ''
Peponapis pruinosa ''Peponapis pruinosa'' is a species of solitary bee in the tribe Eucerini, the long-horned bees. Its common name is the eastern cucurbit bee. It may be called the squash bee, but this name can also apply to other species in its genus, as well as ...
'' specializes on squash plants, and so the current distribution of this species in North America is due to the expansion of plant cultivation throughout North America. The species moved from
Mesoamerica Mesoamerica is a historical region and cultural area that begins in the southern part of North America and extends to the Pacific coast of Central America, thus comprising the lands of central and southern Mexico, all of Belize, Guatemala, El S ...
into the more temperate regions of North America.


Sociality

''Eucera'', similar to the rest of the tribe Eucerini, are solitary by nature. In some highly eusocial bee species, such as honey bees, males are raised and fed in their colonies. Males of ''Eucera'' live their lives independently.


Alarm signaling

''Eucera'' do not produce or respond to alarm pheromones as social bees do. Social bees are able to identify and avoid inflorescences that have the smell of dead bees of their species. This is possibly due to injured social bees releasing signals known as alarm signals to warn others of danger. ''Eucera'' do not showcase this behavior of avoiding inflorescences marked by dead bees and instead respond similarly to flowers that have predation alarm signals and flowers that do not have such alarm signals.


Kleptoparasitism

The bee genus ''
Nomada With over 850 species, the genus ''Nomada'' is one of the largest genera in the family Apidae, and the largest genus of cuckoo bees. Cuckoo bees are so named because they enter the nests of a host and lay eggs there, stealing resources that the h ...
'', a genus that typically lays their eggs in the nests of other bees, is most likely to kleptoparasitize ''Eucera'' bees.


Species

List of ''Eucera'' species


References

*


External links

* * {{Taxonbar, from=Q3734126 Apinae Bee genera Fauna of the Palearctic realm Fauna of the Nearctic realm