Nomia Melanderi
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The alkali bee, ''Nomia melanderi'', is a ground-nesting bee native to
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
s and semi-arid desert basins of the
western United States The Western United States (also called the American West, the Western States, the Far West, the Western territories, and the West) is List of regions of the United States, census regions United States Census Bureau. As American settlement i ...
. It was described by
Theodore Dru Alison Cockerell Theodore Dru Alison Cockerell (22 August 1866 – 26 January 1948) was an American entomologist and Systematics, systematic biologist who published nearly 4,000 papers, some of them only a few lines long. Cockerell's speciality was the insect or ...
in 1906. While solitary, these bees nest near each other and can form extremely dense aggregations in areas with favorable conditions. This bee nests in salt-saturated, or
alkali In chemistry, an alkali (; from the Arabic word , ) is a basic salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as a base that dissolves in water. A solution of a soluble base has a pH greater than 7.0. The a ...
ne, soil. Like some other bees such as ''
Megachile rotundata ''Megachile rotundata'', the alfalfa leafcutting bee, is a European bee that has been introduced to various regions around the world. As a solitary bee species, it does not build colonies or store honey, but is a very efficient pollinator of alfa ...
'', alkali bees are an effective pollinator of
alfalfa Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial plant, perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, ...
. The bee uses a specialized technique of opening alfalfa flowers for pollination by applying pressure to snap open the keel of the flower. Because of this and the fact that they prefer pollen to nectar, fly in a wide range of conditions, and perform well regardless of how well the field is watered, alkali bees are preferred to
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 ...
for alfalfa pollination but have been increasingly supplanted by '' M. rotundata'' in recent years. Due to the unusual nesting habits of this bee, farmers have developed methods to accommodate them with salty mud-fields where they can burrow and lay their eggs. Farmers started doing this after realizing that plowing up natural flats like these decreased the yield of alfalfa dramatically. Juvenile hormone (JH) analog methoprene can serve as a possible limiting factor for alkali bees since female bees are more likely to reach reproductive maturity with JH. Alkali bees are also considered solitary bees because females have their individual nests and live alongside other female alkali bees.


Biology


Appearance

The alkali bee is about two-thirds the size of a honeybee with black and metallic banding on the abdomen. Females have, but infrequently use, stingers and the males have white faces. Males and females in the northern part of the range are larger than their counterparts to the south. This is theorized to allow larval bees to survive for longer without food and may also be related to a slightly longer day in the summer, allowing for increased brood provisioning.


Predators and parasites

Larvae-specific threats include
bee flies The Bombyliidae are a family of flies, commonly known as bee flies. Some are colloquially known as bomber flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects. Over ...
(Bombyliidae) and specifically ''
Heterostylum robustum ''Heterostylum'' is a genus of flies belonging to the family Bombyliidae (bee-flies). There are 14 described species, distributed throughout the Americas. These are robust and very hairy flies with a body length of 10–15 mm. They can be di ...
'' as well as oil beetles ('' Meloe niger'') and both adults and larvae are susceptible to
pesticide Pesticides are substances that are used to control pests. They include herbicides, insecticides, nematicides, fungicides, and many others (see table). The most common of these are herbicides, which account for approximately 50% of all p ...
kills. Unusually, there are not any known kleptoparasites. It is also preyed upon by the conopid fly ''
Zodion obliquefasciatum ''Zodion obliquefasciatum'' is a species of thick-headed flies in the family Conopidae The Conopidae, also known as the thick-headed flies, are a family of fly, flies within the Brachycera suborder of Diptera, and the sole member of the supe ...
'' and is associated with the mite ''Imparipes apicola''. This mite has been found in association with alkali bees but not with other species within the genus '' Nomia''. The mites seem to oviposit in conjunction with emergence of fungi in cells (possibly ''
Ascosphaera ''Ascosphaera'' is a genus of fungi in the family Ascosphaeraceae. It was described in 1955 by mycologists Charles F. Spiltoir and Lindsay S. Olive. Members of the genus are insect pathogens. The type species, '' A. apis'', causes chalkbro ...
''). The fungi are either within the bodies of unhealthy larvae, but more commonly are found in the feces of healthy larvae. The fungi causes the fecal pellets to be strung together in a characteristic manner. After oviposition, the eggs develop, hatch, mate, feed on the fungus, then leave on the bee when it emerges. One generation of ''I. apicola'' is reared per generation of bee, so southern populations of bee which can rear multiple generations in a season are more likely to a have a high presence of mites. Adult females seem host far more mites than the males who seem to host relatively few, and they fall into cells during the provisioning stage where they carry out their life cycle, One study found that 60% of cells had mites, and the average mite count within these cells was 5.7. Although the mites can reach high densities, they don't appear to harm the bees which is possibly due to their fungivorous nature.


Distribution and habitat

The alkali bee lives in arid climates, primarily desert basins in the
western United States The Western United States (also called the American West, the Western States, the Far West, the Western territories, and the West) is List of regions of the United States, census regions United States Census Bureau. As American settlement i ...
from
central Washington Central Washington is a region of the U.S. state of Washington between the western and eastern parts of the state extending from the border with the Canadian province of British Columbia in the north to the border with the U.S. state of Oregon i ...
to
southern California Southern California (commonly shortened to SoCal) is a geographic and Cultural area, cultural List of regions of California, region that generally comprises the southern portion of the U.S. state of California. Its densely populated coastal reg ...
and east to western Colorado. They prefer to nest in moist,
silt Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed with sand or clay) or as sediment mixed in suspension (chemistry), suspension with water. Silt usually ...
y soils that have good drainage, a salty surface, and don't have vegetative groundcover. The
hydrology Hydrology () is the scientific study of the movement, distribution, and management of water on Earth and other planets, including the water cycle, water resources, and drainage basin sustainability. A practitioner of hydrology is called a hydro ...
of the soils appears to be the controlling factor for where nests are made. As soil makeup affects water retention and
capillary action Capillary action (sometimes called capillarity, capillary motion, capillary rise, capillary effect, or wicking) is the process of a liquid flowing in a narrow space without the assistance of external forces like Gravitation, gravity. The effe ...
, soils with some
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolinite, ). Most pure clay minerals are white or light-coloured, but natural clays show a variety of colours from impuriti ...
and
sand Sand is a granular material composed of finely divided mineral particles. Sand has various compositions but is usually defined by its grain size. Sand grains are smaller than gravel and coarser than silt. Sand can also refer to a textural ...
but mostly silt are preferred.


Behavior


Nesting behavior

While solitary, alkali bees frequently nest in high densities. Females lay 1 to 2 eggs a day, and spend most of the day building the cell for the egg. Immediately after emergence a female begins to excavate the main burrow which usually entails digging for about 12 hours. The dirt is removed and using the
mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
and foretarsi, formed into pellets which are then deposited around the entrance to the burrow. The burrows are usually about 15–20 cm in depth, but in laboratory conditions with looser soil, much deeper burrows have been created. In addition to the main vertical burrow, smaller passageways are dug horizontally. From these shorter tunnels, small vertical holes called cells are created for oviposition. Cell creation usually begins the night after the main burrow is excavated. First, the "mine burrow" is dug and then a soil lining applied to the sides which reduces the size of the cell. Lastly a clear,
lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...
-rich secretion from the
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 ...
(in the abdomen) is applied to the interior of the cell to waterproof it. After the cell is constructed, the female bee begins the process of provisioning. She brings back
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 mixes it with
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 ...
to form a ball which is then placed in the bottom of the cell. Any debris that makes its way into the cell during this period is meticulously removed. Once the pollen ball is completed, oviposition occurs. This is usually in the late afternoon. The egg is laid on top of the pollen ball, then the female leaves the cell without inspecting it and begins to cap the cell. The egg garners no further attention after this unless it becomes diseased. In this case, the cell will be opened and packed with dirt by the female. Three or four days after oviposition the egg hatches and the larva begins to consume the pollen ball. Once this has been consumed, the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
(now a prepupa) defecates and begins
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
. In the wild this process takes 11 to 12 days. In the southern region, eggs laid early in the season may emerge after this but for eggs later in the season or for those in the northern part of the range, the pupa will emerge as a bee in the following summer, cued by the warming soil temperatures.


Feeding behavior

Females must consume pollen as soon as they emerge in order for their
ovaries The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are endocr ...
to develop. They continue to consume pollen for at least two weeks. It is believed that because solitary bees have very large, lipid-rich eggs the pollen provides the lipids and
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
s required for making these eggs. Pollen consumption is larger in younger females, and primarily occurs in the afternoon and evening as pollen collected early in the day goes toward larval provisioning.


Mating behavior

In the first few weeks of the emergence period, the majority of emerging bees are male. Once they emerge, they begin patrolling for females who likely mate soon after emerging. Their attractiveness (driven by
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) decreases in the days after emerging. No evidence of courtship has been observed and the copulations last less than 30 seconds. While other males sometimes observe copulation, there isn't evidence for aggression or interference by these males


Conservation status

The alkali bee is not ranked for
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and manage ...
. Due to its commercially important role as a pollinator of the alfalfa crop in the western and northwestern United States, it is not a species of concern, unlike many native bees.


References

{{Authority control Insects of the United States Insects described in 1906 Hymenoptera of New Zealand Taxa named by Theodore Dru Alison Cockerell Nomia (bee)