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''Tuberolachnus salignus'', the giant willow aphid, is a species of
aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...
, in the genus ''
Tuberolachnus ''Tuberolachnus'' is a genus of aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group include ...
''. They are reputed to be the largest aphids, with a body length of up to 5.8mm. First described by
Johann Friedrich Gmelin Johann Friedrich Gmelin (8 August 1748 – 1 November 1804) was a German natural history, naturalist, chemist, botanist, entomologist, herpetologist, and malacologist. Education Johann Friedrich Gmelin was born as the eldest son of Philipp F ...
in 1790, it feeds on many species of
willow Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...
(''Salix'' species), and has one known host-specific parasite, ''Pauesia salignae''. ''Tuberolachnus salignus'' does not require a male for reproduction. Only females have ever been recorded, which suggests that reproduction is
parthenogenetic Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
and all offspring produced are genetically identical
clones Clone or Clones or Cloning or Cloned or The Clone may refer to: Places * Clones, County Fermanagh, Ireland * Clones, County Monaghan, a town in Ireland * Clones railway station, Ireland Biology * Clone (B-cell), a lymphocyte clone, the massiv ...
.


Description

Adults exhibit a pear-shaped body with dark brown/grayish coloration. This aphid species has distinct black dots that form rows along its body. Along the dorsal side of their abdomen, about 2/3 of the way down, ''T. salignus'' displays a defining single large conical tubercle or bump that resembles a thorn which can grow to 0.2-0.3 mm. Closer to the posterior and along the sides of the body are two smaller black cone structures ( siphunculi). ''Tuberolachnus salignus'' reproduces solely through parthenogenesis so all aphid adults are female. Adults can be either winged or wingless. The two pairs of wings are clear with a dark brown edge along the forewing. When looking at ''T. salignus'' under a microscope the presence of numerous tiny hairs is apparent. This gives the abdomen a grayish-gold hue. Adult ''T. salignus'' has three pairs of legs that are primarily a dark brownish color, but smaller, red/orange sections are also present. They also have a pair of antennae that are less than half of the aphid's body length.


Range


Natural global range

''Tuberolachnus salignus'' is a
cosmopolitan Cosmopolitan may refer to: Internationalism * World citizen, one who eschews traditional geopolitical divisions derived from national citizenship * Cosmopolitanism, the idea that all of humanity belongs to a single moral community * Cosmopolitan ...
species. They exist anywhere that willow species grow naturally or are cultivated. The giant willow aphid originated in parts of Asia but has now spread to all parts of the globe (except Antarctica). There is no known exact place of origin for this aphid species, but it is believed to be somewhere within Japan and Korea. The only known
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
of this species is endemic to Japan, leading to the belief that this may be the place of their origin. ''Tuberolachnus salignus'' is now found in all parts of New Zealand where willow species are present. It only recently made it to New Zealand, with the first confirmed sighting in December 2013 in Auckland.


Habitat

This species is almost entirely dependent on willow trees. They live on the bark of these trees and feed on their sap. These giant aphids survive year-round with the most activity being in late summer and early fall. Their population numbers are known to be largely weather and temperature related. Following instances of large amounts of rain, ''T. salignus'' populations have been known to increase rapidly.


Ecology


Life cycle/phenology

''Tuberolachnus salignus'' has a generation period of 2–3 weeks, which is largely dependent on temperature. Once hatched, the aphid larvae go through four stages of development until they reach their adult forms. Ideal temperatures for ''T. salignus'' generation range from 17.5 to 22.5 degrees Celsius. It takes two weeks for nymphs to become adults, with adults living an additional five weeks after that. Mature aphids can produce anywhere from 35 to 71 offspring. Generally, ''T. salignus'' is most active in the warmer months, with populations peaking in late summer and early autumn. In colder months, the overall fecundity decreases but colonies are still active. In the spring, colonies are usually not visible. During this time, it is thought ''T. salignu''s retreats behind the willow tree bark or into the ground to escape the harsh climate. ''Tuberolachnus salignus'' reproduces via parthenogenesis. Resulting in a population consisting of all females. All offspring are identical genetic clones of their mothers. In turn, there are no mating behaviors that occupy these aphids as they do not participate in sexual reproduction.


Diet and foraging

''Tuberolachnus salignus'' is known to feed primarily on willow and ''Salix'' tree species such as the white willow (''
Salix alba ''Salix alba'', the white willow, is a species of willow native to Europe and western and Central Asia.Meikle, R. D. (1984). ''Willows and Poplars of Great Britain and Ireland''. BSBI Handbook No. 4. .Rushforth, K. (1999). ''Trees of Britain an ...
''), the common sallow (''
Salix cinerea ''Salix cinerea'' (common sallow, grey sallow, grey willow, grey-leaved sallow, large grey willow, pussy willow, rusty sallow) is a species of willow native to Europe and western Asia.Meikle, R. D. (1984). ''Willows and Poplars of Great Britain ...
')'' and the Indian willow (''
Salix tetrasperma ''Salix tetrasperma'', commonly called Indian willow, is a medium-sized tree of wet and swampy places, shedding its leaves at the end of monsoon season. It flowers after leafing. The bark is rough, with deep, vertical fissures and the young shoot ...
''). It has also been recorded on Quince (''
Cydonia oblonga The quince (; ''Cydonia oblonga'') is the sole member of the genus ''Cydonia'' in the Malinae subtribe (which contains apples, pears, and other fruits) of the Rosaceae family. It is a deciduous tree that bears hard, aromatic bright golden-yell ...
''), Apple (''
Malus ''Malus'' ( or ) is a genus of about 32–57 species of small deciduous trees or shrubs in the family Rosaceae, including the domesticated orchard apple, crab apples (sometimes known in North America as crabapples) and wild apples. The genus i ...
spp.'') and Poplar (''
Populus ''Populus'' is a genus of 25–30 species of deciduous flowering plants in the family Salicaceae, native to most of the Northern Hemisphere. English names variously applied to different species include poplar (), aspen, and cottonwood. The we ...
spp.'') Like many herbivorous Hemiptera, ''T. salignus'' uses modified mouthparts to pierce the
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
of plants and feed on its sap. As the name suggests, giant willow aphids prefer willows of the ''Salix'' species but have been known to feed on poplar, apple, and pear species in New Zealand. They prefer to feed during the day and during the warmer hours. When feeding, the aphid will climb to the tip of the willow where the shoot begins. The flesh of the plant is softer here, allowing the aphid to inject its stylets into the plant's phloem. ''Tuberolachnus salignus'' uses two stylets to extract the sap flowing within the phloem. One is used to inject saliva into the phloem, while the other is used to ingest the sap and saliva mixture. Willow sap is an extremely nutritious food source, as it provides critical sugars and water to various parts of the willow tree. As the aphid feeds, it is ingesting an incredible amount of these sugars and water, so much so that it needs to excrete them to avoid overfeeding. These are excreted as a sugary liquid known as honeydew, which attracts various other insect species such as honeybees, wasps, and ants. These other species will feed on the honeydew that is excreted from ''T. salignus''.


Predators, prey, and parasites

Due to the almost exclusive feeding of willow sap by ''T. salignus'', the honeydew that is produced as a result attracts numerous types of insects. Wasps, bees, ants, flies and coccinellids have all been observed to feed on honeydew of this species. These insects feed only on the honeydew, not the actual aphid. In New Zealand, ''Pauesia nigrovaria'' was introduced as a biocontrol parasitoid of ''T. salignus'' in 2020. Elsewhere, like North America and parts of Europe, parasitic wasps, lady beetles, hover fly maggots, lace wings are frequent predators.


Interactions with humans

''Tuberolachnus salignus'' has only recently come into the limelight in New Zealand. Upon arriving here in 2013, cause for alarm was immediately raised. As previously noted, these giant aphids feed almost exclusively on willow species. As ''T. salignus'' populations begin to rapidly grow on the willow tree, many negative impacts start to become apparent. Willow trees are an important group of plants in New Zealand. While they are not native, they are used in many facets of society in order to increase land stabilization and reduce soil erosion. Historically, willows have been planted along riverbanks and on particularly sloping land to help keep the soil beneath from eroding away. Furthermore, willows play an important roll in honey production throughout New Zealand. The flowers of willows are rich with pollen and nectar and attract honeybees. In other parts of the world, such as the U.K., large amounts of willows are cultivated for biomass production. With the presence of large ''T. salignus'' colonies on willow trees, many of the contributions listed above are severely hindered. Additionally, honeybees that forage on ''T. salignus'' honeydew have poorer health and produce a lower quantity and quality of honey.


References


Further reading

* * * {{Taxonbar, from=Q14086166 Lachninae Hemiptera of Europe Insects described in 1790 Taxa named by Johann Friedrich Gmelin