Goldenrod gall fly

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Goldenrod gall fly
Eurosta solidaginis female.jpg
E. solidaginis female
Eurosta solidaginis puparium.jpg
E. solidaginis puparium
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Tephritidae
Subfamily: Tephritinae
Tribe: Dithrycini
Genus: Eurosta
Species:
E. solidaginis
Binomial name
Eurosta solidaginis
(Fitch, 1855) [1]
Synonyms
  • Ortalis nupheraHarris, 1835 [2]
  • Tephritis asterisHarris, 1841 [3]
  • Acinia solidaginis Fitch, 1855 [1]
  • Eurosta asteri Johnson, 1930 [4]
  • Eurosta subfascipennisStrickland, 1938 [5]

The goldenrod gall fly (Eurosta solidaginis), also known as the goldenrod ball gallmaker, is a species of fly native to North America. The species is best known for the characteristic galls it forms on several species in the Solidago , or goldenrod, genus. The fly's eggs are inserted near the developing buds of the plant. After hatching, the larvae migrate to an area below the plant's developing buds, where they then induce the plant's tissues to form into the hardened, bulbous chamber referred to as a gall. E. solidaginis’s interactions with its host plant(s) and insect, as well as avian, predators have made it the centerpiece of much ecological and evolutionary biology research, and its tolerance of freezing temperatures has inspired studies into the anti-freeze properties of its biochemistry.

Contents

Taxonomy

Eurosta solidaginis is in the order Diptera and the family Tephritidae. [6] The Tephritidae are commonly referred to as fruit flies, a common name they share with the Drosophilidae family. [6] The goldenrod gall fly was first put into the genus Eurosta by D. W. Cocquillet in 1910. [7] Two subspecies exist: E. solidaginis subsp. solidaginis and E. solidaginis subsp. fascipennis, distinguished morphologically by differences in hyaline regions of the wing margin. [8] The former subspecies can be further subdivided into two host races, one of which forms galls in Solidago altissima and the other in S. gigantea. [9] [10]

Distribution

E. solidaginis is widely distributed across the United States, ranging from Washington all the way to the eastern seaboard. [8] [9] The two subspecies occupy different ranges, with E. solidaginis subsp. solidaginis being found from the east coast to Minnesota and the Dakotas, up to the southeastern provinces of Canada, and down the southern border of the United States. [8] [9] E. solidaginis subsp. fascipennis, on the other hand, can be found as far west as Washington and as far east as Minnesota. [8]

Behavior and ecology

A Eurosta solidaginis larva in a freshly dissected gall. Dissected Eurosta solidaginis gall.jpg
A Eurosta solidaginis larva in a freshly dissected gall.

Adult E. solidaginis emerge from their galls in the spring, with the males emerging prior to the females. [11] [12] [13] The flies proceed to mate on goldenrod plants, and the females use their ovipositors to insert fertilized eggs into the buds of the goldenrod. [14] [9]

Though E. solidaginis has been reported to form galls on seven different species of goldenrod, only three appear to be common targets: Solidago canadensis, S. gigantea and S. altissima. [9] Once the larvae hatch, typically between 5 and 8 days after the egg is laid, the larva eats its way to the base of the goldenrod bud and induces a gall. [9] These galls serve as food sources and shelters from rain, wind and ice. [9] Despite being the larva's overwintering structure, the gall itself does not provide significant insulation. [14] [15] Instead, the larva itself has robust freezing tolerance. The larva feeds on the tissues of the gall and molts twice before excavating a narrow exit tunnel out of the gall in mid-September. [9] After digging its tunnel, without actually opening up the gall to the outside, the larva overwinters and, if it survives, molts into an adult and leaves the gall the following spring. [9]

A number of predators and parasites prey on the larvae of E. solidaginis. The black-capped chickadee (Poecile atricapillus) and the downy woodpecker (Picoides pubescens) target large galls, breaking them open and removing the larva living inside. [16] [17] The parasitic wasps Eurytoma obtusiventris and E. gigantea also target the gallmaker. The former injects its eggs directly into E. solidaginis larvae prior to gall formation, whereas the latter oviposits into the gall itself. [12] In both cases, the E. solidaginis larvae are consumed. There is also Mordellistena unicolor , a beetle whose larvae, after hatching on the surface of a gall, burrow their way in and feed off of its nutritive tissues. [18] M. unicolor typically kills the E. solidaginis larva inhabiting the gall, but this does not appear to be an essential part of its life cycle. [9] One of the upshots of these interactions is that larvae that produce galls of a moderate size – that is, not small and thin enough for wasps to easily penetrate, but not large enough to attract the attention of birds – have a fitness advantage. [9] In addition to these direct, predatory interactions, a negative correlation has been observed between goldenrod defoliation by Trirhabda beetles, suggesting that competition for plant resources may also have an important role to play in the Eurosta/Solidago system. [19]

Adult emerging from gall. Notice the ptilinum on the head Eurosta solidaginis ptilinum.jpg
Adult emerging from gall. Notice the ptilinum on the head

Physiology

The ability of E. solidaginis to survive the freezing temperatures of winter has been the subject of much research. In response to dropping temperatures and the senescence of surrounding plant tissues, the larva begins to synthesize and accumulate sorbitol and glycerol in its tissues. [20] [21] [22] These compounds help protect the larvae against freezing damage by lowering the melting point of their bodily fluids, thus reducing the amount of ice that can form. [23] Aquaporins, membrane proteins involved in the channeling of water, have also been shown to play a key role in E. solidaginis’ freezing tolerance. [24] [25] As ice forms in the larva's bodily fluids, solutes in the unfrozen liquid are concentrated, creating a strong osmotic gradient. In species like E. solidaginis that can channel water quickly enough in response to this freezing stress, water rapidly travels to the solute-rich extracellular environment, switching places with cryoprotectant molecules like glycerol, thus protecting the larva's tissues. [25] [26] Upregulation of these aquaporin proteins in the winter seasons corroborates the hypothesis that they play a key role in freezing tolerance. [24] It has also been found that mild winter temperatures are detrimental to the survival rate of E. solidaginis, with researchers speculating that cool or freezing temperatures may actually benefit the insect by allowing it to conserve energy [27]

Related Research Articles

<span class="mw-page-title-main">Tephritidae</span> Family of fruit flies

The Tephritidae are one of two fly families referred to as fruit flies, the other family being the Drosophilidae. The family Tephritidae does not include the biological model organisms of the genus Drosophila, which is often called the "common fruit fly". Nearly 5,000 described species of tephritid fruit fly are categorized in almost 500 genera of the Tephritidae. Description, recategorization, and genetic analyses are constantly changing the taxonomy of this family. To distinguish them from the Drosophilidae, the Tephritidae are sometimes called peacock flies, in reference to their elaborate and colorful markings. The name comes from the Greek τεφρος, tephros, meaning "ash grey". They are found in all the biogeographic realms.

<i>Solidago canadensis</i> Species of flowering plant

Solidago canadensis, known as Canada goldenrod or Canadian goldenrod, is an herbaceous perennial plant of the family Asteraceae. It is native to northeastern and north-central North America and often forms colonies of upright growing plants, with many small yellow flowers in a branching inflorescence held above the foliage. It is an invasive plant in other parts of the continent and several areas worldwide, including Europe and Asia. It is grown as an ornamental in flower gardens.

<i>Solidago juncea</i> Species of flowering plant

Solidago juncea, the early goldenrod, plume golden-rod, or yellow top, is a North American species of herbaceous perennial plants of the family Asteraceae native to eastern and central Canada and eastern and central United States. It grows from Nova Scotia west to Manitoba and Minnesota south as far as northern Georgia and northern Arkansas, with a few isolated populations in Louisiana and Oklahoma.

<span class="mw-page-title-main">Insect winter ecology</span> Survival strategies of insects during winter

Insect winter ecology describes the overwinter survival strategies of insects, which are in many respects more similar to those of plants than to many other animals, such as mammals and birds. Unlike those animals, which can generate their own heat internally (endothermic), insects must rely on external sources to provide their heat (ectothermic). Thus, insects persisting in winter weather must tolerate freezing or rely on other mechanisms to avoid freezing. Loss of enzymatic function and eventual freezing due to low temperatures daily threatens the livelihood of these organisms during winter. Not surprisingly, insects have evolved a number of strategies to deal with the rigors of winter temperatures in places where they would otherwise not survive.

<i>Solidago sempervirens</i> Species of aquatic plant

Solidago sempervirens, the seaside goldenrod or salt-marsh goldenrod, is a plant species in the genus Solidago of the family Asteraceae. It is native to eastern North America and parts of the Caribbean. It is an introduced species in the Great Lakes region. Similar plants found in the Azores are thought have evolved from a natural introduction of this species.

<i>Solidago rugosa</i> Species of flowering plant

Solidago rugosa, commonly called the wrinkleleaf goldenrod or rough-stemmed goldenrod, is a species of flowering plant in the family Asteraceae. It is native to North America, where it is widespread across eastern and central Canada and the eastern and central United States. It is usually found in wet to mesic habitats.

<i>Solidago altissima</i> Species of flowering plant in the family Asteraceae

Solidago altissima, the tall goldenrod or late goldenrod, is a North American species of goldenrod in the family Asteraceae which is widespread across much of Canada, the United States, and northern Mexico. It is common in much of its range and fairly tolerant of landscapes which have been disturbed by humans. It has become naturalized in many parts of the world.

<i>Solidago gigantea</i> Species of plant in the family Asteraceae native to North America

Solidago gigantea is a North American plant species in the family Asteraceae. Its common names include tall goldenrod and giant goldenrod, among others.

Mordellistena unicolor is a species of beetle in the genus Mordellistena of the family Mordellidae. It was described by John Lawrence LeConte in 1862. Larvae feed on gall tissue of the goldenrod gall fly and often consume the fly larvae within the gall as well.

o
<i>Eurosta</i> Genus of flies

Eurosta is a genus of gall maker flies in the family Tephritidae. There are seven species in the genus, all in North America.

<i>Solidago odora</i> Species of flowering plant

Solidago odora, the sweet goldenrod, anisescented goldenrod or fragrant goldenrod, is a North American species of goldenrod within the family Asteraceae. The plant is native to the United States and Mexico, found in every coastal state from Veracruz to New Hampshire and as far inland as Ohio, Missouri, and Oklahoma. It flowers from July through October.

<i>Solidago ulmifolia</i> Species of plant

Solidago ulmifolia, commonly known as elmleaf goldenrod, is a North American species of goldenrod in the family Asteraceae. It is found in Canada and the eastern and central United States.

<i>Gnorimoschema gallaesolidaginis</i> Species of moth

Gnorimoschema gallaesolidaginis, the solidago gall moth, goldenrod gall moth or goldenrod gallmaker, is a moth in the family Gelechiidae. It was described by Riley in 1869. It is found in North America, where it has been recorded from Alberta, British Columbia, California, Florida, Illinois, Indiana, Kentucky, Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, New Brunswick, New Hampshire, New York, North Carolina, Ohio, Oklahoma, Ontario, Pennsylvania, Quebec, Tennessee, Texas and Wisconsin.

<i>Rhopalomyia anthophila</i> Species of fly

Rhopalomyia anthophilathe Downy Flower Gall Midgeis a species of gall midges, insects in the family Cecidomyiidae. It induces galls on the host Solidago altissima across much of North America.

<i>Rhopalomyia solidaginis</i> Species of fly

Rhopalomyia solidaginis, the goldenrod bunch gall, is a species of gall midges, insects in the family Cecidomyiidae The galls of this species have the following host species of goldenrods:Solidago altissima, Solidago canadensis, Solidago rugosa They have been found across eastern North American.

<i>Asphondylia solidaginis</i> Species of fly

Asphondylia solidaginis is a species of gall midge (Cecidomyiidae) that induces galls on goldenrods in North America where it is widespread. It was first described by William Beutenmuller in 1907.

<i>Rhopalomyia clarkei</i> Species of fly

Rhopalomyia clarkei is a species of gall midges, insects in the family Cecidomyiidae. The larvae induce galls on two hosts: Solidago altissima and Solidago rugosa and are found in north-eastern and north central North America.

<i>Eurytoma obtusiventris</i> Species of insect

Eurytoma obtusiventris is a species of chalcid wasp in the family Eurytomidae. It is found in North America and is a parasitoid of Eurosta solidaginis. The wasp attacks after oviposition but prior to the formation of a gall. Eurytoma obtusiventris prompts the Eurosta solidaginis larvae to create a pupa, which the wasp will use after consuming its host. They typically consume the host in autumn and overwinter within the gall, emerging during the summer.

<i>Rhopalomyia capitata</i> Species of fly

Rhopalomyia capitata, the giant goldenrod bunch gall, is a species of gall midge in the family Cecidomyiidae. It is found across North America. They have been collected from Solidago canadensis, Solidago gigantea, and Solidago leavenworthiigallormers species page

References

  1. 1 2 Fitch, A. (1855). "Report on the noxious, beneficial and other insects of the State of New York. [I]". Transactions of the New-York State Agricultural Society. (1854) 14: 705–880.
  2. ↑ Harris, T.W. (1835). Insects. Pp. 553-602. In Hitchcock, E., Report on the geology, mineralogy, botany, and zoology of Massachusetts. 2nd Edition. Amherst: J. S. & C. Adams. pp. [4] + 702 pp., 19 pls, 3 maps.
  3. ↑ Harris, T.W. (1841). A report on insects of Massachusetts, injurious to vegetation. [1st edition]. Cambridge: Folsom, Wells & Thurston. pp. viii + 459.
  4. ↑ Johnson, C.W. (1930). "A list of the insect fauna of Nantucket, Massachusetts". Publ. Nantucket Maria Mitchell Assoc. 3 (2): 1–174.
  5. ↑ Strickland, E.H. (1938). "An annotated list of the Diptera (flies) of Alberta". Canadian Journal of Research. 16 (7): 175–219. doi:10.1139/cjr38d-012.
  6. 1 2 Borror, D. J.; White, R. E. (1970). Insects. Boston: Houghton Mifflin Company.
  7. ↑ Cocquillet, Daniel W. (1910). "The type species of the North American genera of Diptera" (PDF). Proceedings of the United States National Museum. 37. Government Printing Office: 534.
  8. 1 2 3 4 Ming, Y. (1989). A revision of the genus Eurosta Loew with a scanning microscopic study of taxonomic characters (Diptera: Tephritidae) (Master's thesis). Pullman, WA: Washington State University.
  9. 1 2 3 4 5 6 7 8 9 10 11 Abrahamson, Warren G.; Weis, Arthur E. (1997). Evolutionary ecology across three trophic levels: goldenrods, gallmakers, and natural enemies. New Jersey: Princeton University Press. ISBN   9780691012087.
  10. ↑ Waring, Gwendolyn L.; Abrahamson, Warren G.; Howard, Daniel J. (1990). "Genetic differentiation among host-associated populations of the gallmaker Eurosta solidaginis (Diptera: Tephritidae)". Evolution. 44 (6): 1648–1655. doi: 10.1111/j.1558-5646.1990.tb03853.x . PMID   28564317.
  11. ↑ Anderson, R. M.; McCrea, K. D.; Abrahamson, W. G.; L. M., Hartzel (1989). "Host genotype choice by the ball gallmaker Eurosta solidaginis (Diptera: Tephritidae)". Ecology. 70 (4): 1048–1054. Bibcode:1989Ecol...70.1048A. doi:10.2307/1941373. JSTOR   1941373.
  12. 1 2 Weis, A. E.; Abrahamson, W. G. (1985). "Potential selective pressures by parasitoids on the evolution of a plant-herbivore interaction". Ecology. 66: 1261–1269. doi:10.2307/1939179. JSTOR   1939179.
  13. ↑ Craig, T. P.; Itami, J. K.; Abrahamson, W. G.; Horner, J. D. (1993). "Behavioral evidence for host-race formation in Eurosta solidaginis". Evolution. 47 (6): 1696–1710. doi: 10.1111/j.1558-5646.1993.tb01262.x . PMID   28567992.
  14. 1 2 Uhler, Lowell D. (1951). Biology and ecology of the goldenrod gall fly: Eurosta solidaginis (Fitch). Vol. 300. Cornell University Agricultural Station Memoir. pp. 1–51.
  15. ↑ Layne, Jack R. (1993). "Winter microclimate of goldenrod spherical galls and its effects on the gall inhabitant Eurosta solidaginis (Diptera: Tephritidae". Journal of Thermal Biology. 18 (3): 125–130. Bibcode:1993JTBio..18..125L. doi:10.1016/0306-4565(93)90025-O.
  16. ↑ Schlichter, Lyanne (1978). "Winter predation by black capped chickadees and downy woodpeckers on inhabitants of the goldenrod ball gall". Canadian Field-Naturalist. 92: 71–74. doi: 10.5962/p.346636 .
  17. ↑ Abrahamson, Warren G.; Sattler, Joan F.; McCrea, Kenneth D.; Weis, Arthur E. (April 1989). "Variation in selection pressures on the goldenrod gall fly and the competitive interactions of its natural enemies". Oecologia. 79 (1): 15–22. Bibcode:1989Oecol..79...15A. doi:10.1007/BF00378234. ISSN   0029-8549. PMID   28312807. S2CID   206777734.
  18. ↑ Ping, Chi (1915). "Some inhabitants of the round gall of goldenrod". Journal of Entomology and Zoology: 161–179.
  19. ↑ Cunan, Ellery T.; Powell, Thomas H. Q.; Weis, Arthur E. (April 2015). "Evidence For Plant-mediated Competition Between Defoliating and Gall-forming Specialists Attacking Solidago altissima". The American Midland Naturalist. 173 (2): 208–217. doi:10.1674/amid-173-02-208-217.1. ISSN   0003-0031. S2CID   86758210.
  20. ↑ Baust, John G.; Lee, Richard E. (January 1982). "Environmental triggers to cryoprotectant modulation in separate populations of the gall fly, Eurosta solidaginis (Fitch)" (PDF). Journal of Insect Physiology. 28 (5): 431–436. Bibcode:1982JInsP..28..431B. doi:10.1016/0022-1910(82)90070-1.
  21. ↑ Rojas, Robert R.; Lee, Richard E.; Baust, John G. (December 1986). "Relationship of environmental water content to glycerol accumulation in the freezing tolerant larvae of Eurosta solidaginis (Fitch)". Cryobiology. 23 (6): 564. doi:10.1016/0011-2240(86)90117-3.
  22. ↑ Irwin, J.T.; Bennett, V.A.; Lee Jr, R.E. (26 March 2001). "Diapause development in frozen larvae of the goldenrod gall fly, Eurosta solidaginis fitch (diptera: tephritidae)" (PDF). Journal of Comparative Physiology B. 171 (3): 181–188. doi:10.1007/s003600000154. ISSN   0174-1578. PMID   11352100. S2CID   3056255.
  23. ↑ Muldrew, Ken; Acker, Jason P.; Elliott, Janet A. W.; McGann, Locksley E. (2004). "The water to ice transition: implications for living cells". In Fuller, Barry J.; Lane, Nick; Benson, Erica E. (eds.). Life in the Frozen State. pp. 67–108. doi:10.1201/9780203647073. ISBN   9780203647073.
  24. 1 2 Philip, Benjamin N.; Lee, Richard E. (July 2010). "Changes in abundance of aquaporin-like proteins occurs concomitantly with seasonal acquisition of freeze tolerance in the goldenrod gall fly, Eurosta solidaginis" (PDF). Journal of Insect Physiology. 56 (7): 679–685. Bibcode:2010JInsP..56..679P. doi:10.1016/j.jinsphys.2009.12.003. PMID   20005232.
  25. 1 2 Philip, B. N.; Yi, S.-X.; Elnitsky, M. A.; Lee, R. E. (2008). "Aquaporins play a role in desiccation and freeze tolerance in larvae of the goldenrod gall fly, Eurosta solidaginis". Journal of Experimental Biology. 211 (7): 1114–1119. doi: 10.1242/jeb.016758 . ISSN   0022-0949. PMID   18344486.
  26. ↑ Lee, Richard E. (1989). "Insect cold-hardiness: to freeze or not to freeze" (PDF). BioScience. 39 (5): 308–313. doi:10.2307/1311113. JSTOR   1311113.
  27. ↑ Irwin, Jason T; Lee, Richard E (May 2000). "Mild winter temperatures reduce survival and potential fecundity of the goldenrod gall fly, Eurosta solidaginis (Diptera: Tephritidae)" (PDF). Journal of Insect Physiology. 46 (5): 655–661. Bibcode:2000JInsP..46..655I. doi:10.1016/S0022-1910(99)00153-5. PMID   10742513.
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