Adelges piceae

Ge­o­graphic Range

Bal­sam woolly adel­gids (Adelges piceae) are na­tive to cen­tral Eu­rope and are thought to have spread to North Amer­ica in the early twen­ti­eth cen­tury. In North Amer­ica, these adel­gids are pri­mar­ily found in the Ap­palachian Moun­tains and north­east­ern United States. (Go­heen and Will­hite, 2006; Hrinke­vich, et al., 2016; Side­bot­tom, 2019)

Habi­tat

Bal­sam woolly adel­gids only live on fir and spruce trees in forested areas. Their pop­u­la­tions are largest in re­gions where the fir growth rate and qual­ity are the high­est. Higher adel­gid pop­u­la­tions are typ­i­cally found at lower el­e­va­tions, milder tem­per­a­tures, near for­est edges, and near moun­tain lakes and mead­ows. (Go­heen and Will­hite, 2006; Hrinke­vich, et al., 2016; Side­bot­tom, 2019)

Phys­i­cal De­scrip­tion

Bal­sam woolly adel­gid adults are very small and have a cov­er­ing of woolly wax that hides their body com­pletely. This cov­er­ing gives them the ap­pear­ance of a tiny piece of cot­ton on tree bark. Un­der­neath the cov­er­ing, adults are about 0.5 to 1 mm long, soft-bod­ied, and dark pur­ple or black in color. Nymphs are black and have an un­der­de­vel­oped wax cov­er­ing which grows as they ma­ture. Eggs are ob­long, am­ber-col­ored, and ap­prox­i­mately 0.3 mm long. Crawlers look very sim­i­lar to eggs but are slightly larger and have legs and eyes. (Go­heen and Will­hite, 2006; Ragen­ovich and Mitchell, 2006; Side­bot­tom, 2019; Varty, 1956)

  • Range length
    0.5 to 1 mm
    0.02 to 0.04 in

De­vel­op­ment

Al­most all bal­sam woolly adel­gids go through five stages of de­vel­op­ment: Egg, crawler, two stages of nymph, and adult. The first stage is the egg, fol­lowed by crawlers, which are some­times called neo­sis­tens. After set­tling, the crawler molts to a nymph. From there, nymphs molt two more times to reach adult­hood. It takes ap­prox­i­mately one month for a nymph to molt to an adult. In the wild, there are typ­i­cally two gen­er­a­tions per year, known col­lec­tively as sis­tens gen­er­a­tions. These two sis­tens gen­er­a­tions, heimo­sis­tens and aes­tivo­sis­tens, enter di­a­pause (a pe­riod of dor­mancy and de­layed de­vel­op­ment) dur­ing dif­fer­ent sea­sons. Hiemo­sis­tens enter di­a­pause in the win­ter. They hatch in the fall, set­tle, over­win­ter in the crawler stage, then con­tinue de­vel­op­ment in the spring. Aes­tivo­sis­tens enter di­a­pause in the sum­mer. They hatch in the sum­mer, set­tle, then aes­ti­vate as crawlers for three to six weeks be­fore ma­tur­ing. (Al­ston, et al., 2018; Mont­gomery and Havil, 2014; Side­bot­tom, 2019; Varty, 1956)

There is also a third gen­er­a­tion of bal­sam woolly adel­gids that is sel­dom seen in the wild called the pro­gre­di­ens gen­er­a­tion. This gen­er­a­tion hatches in the sum­mer from eggs laid by hiemo­sis­tens. They typ­i­cally hatch close to the be­gin­ning of the sea­son, be­fore or along­side the aes­tivo­sis­tens. Pro­gre­di­ens have six stages of de­vel­op­ment that are nearly iden­ti­cal those of the sis­tens gen­er­a­tions, but with one extra nymph stage. Un­like the sis­tens gen­er­a­tions, the pro­gre­di­ens gen­er­a­tions do not go through a pe­riod of di­a­pause. (Varty, 1956)

Re­pro­duc­tion

Bal­sam woolly adel­gids are all fe­male and re­pro­duce solely through partheno­gen­e­sis. (Side­bot­tom, 2019)

Bal­sam woolly adel­gids go through two to four gen­er­a­tions each year, de­pend­ing on tem­per­a­ture, cli­mate, and el­e­va­tion. Two gen­er­a­tions is most com­mon. The sum­mer gen­er­a­tion (aes­tivo­sis­tens) hatches in May through June. Aes­tivo­sis­tens reach ma­tu­rity by July and their ovipo­si­tion be­gins about a week after their final molt. The eggs laid by aes­tivo­sis­tens hatch in as lit­tle as a few days, de­pend­ing on tem­per­a­ture. The spring gen­er­a­tion (hiemo­sis­tens) hatches from these eggs. They then over­win­ter as crawlers and re­sume ma­tur­ing in March or April, reach­ing adult­hood in March through May. Bal­sam woolly adel­gids have been ob­served lay­ing up to 250 eggs in the lab, but only lay ap­prox­i­mately 100 eggs in the wild. The first few eggs that hiemo­sis­tens lay pro­duce pro­gre­di­ens, which al­most al­ways die be­fore they can re­pro­duce, but the vast ma­jor­ity of their eggs pro­duce aes­tivo­sis­tens. (Al­ston, et al., 2018; Mont­gomery and Havil, 2014; Ragen­ovich and Mitchell, 2006; Side­bot­tom, 2019; Varty, 1956)

  • Range eggs per season
    250 (high)
  • Average eggs per season
    100

Bal­sam woolly adel­gids do not take care of their off­spring, but their wax coat­ing may pro­tect their eggs until they hatch. (Side­bot­tom, 2019)

  • Parental Investment
  • no parental involvement

Lifes­pan/Longevity

The lifes­pan of bal­sam woolly adel­gids varies across gen­er­a­tions. Hiemo­sis­tens, the gen­er­a­tion that over­win­ters as nymphs, lives the longest on ac­count of their over­win­ter­ing. The pro­gre­di­ens have the short­est lifes­pan, sur­viv­ing only a few days in the wild. Neo­pro­gre­di­ens (pro­gre­di­entes crawlers) hatch in March through the be­gin­ning of May and can only feed on newly flushed leaves. These leaves do not ap­pear until late May, so all neo­pro­gre­di­ens die of star­va­tion. Dur­ing spring and sum­mer in Eu­rope, adults can live for up to two months, but rarely live that long in the wild due to heavy pre­da­tion. Ex­treme hot and ex­treme cold tem­per­a­tures in North Amer­ica may also shorten bal­sam woolly adel­gid lifes­pans. (Side­bot­tom, 2019; Varty, 1956)

Be­hav­ior

Bal­sam woolly adel­gids are com­pletely seden­tary for al­most their en­tire lives. The crawler life stage is the only stage that is ca­pa­ble of move­ment. Crawlers must lo­cate a suit­able place to feed within sev­eral days of hatch­ing or they will starve. Once a crawler has found a feed­ing lo­ca­tion, it set­tles, in­sert­ing its stylet into the tree. From this point it re­mains com­pletely seden­tary for the rest of its life. The sis­tens gen­er­a­tions typ­i­cally set­tle near the trunk of the tree. Crawlers can also be car­ried by the wind to an­other host tree. (Ragen­ovich and Mitchell, 2006; Side­bot­tom, 2019; Varty, 1956)

Home Range

Bal­sam woolly adel­gids spend al­most their en­tire life on a sin­gle spot on a fir tree. (Side­bot­tom, 2019)

Com­mu­ni­ca­tion and Per­cep­tion

Bal­sam woolly adel­gids have com­pound eyes for de­tect­ing light and can also de­tect tem­per­a­ture. They use these senses to help them find an op­ti­mal feed­ing lo­ca­tion dur­ing the crawler stage. After the crawler stage, the adel­gids keep their eyes but seem to have no fur­ther use for them. Bal­sam woolly adel­gids have no known forms of so­cial com­mu­ni­ca­tion. (Side­bot­tom, 2019; Varty, 1956)

Food Habits

Bal­sam woolly adel­gids feed ex­clu­sively on fir trees. To feed, these adel­gids in­sert their long, tube-like mouth­parts into the bark. They are most often found near the trunk of in­fested trees. In North Amer­ica, they pri­mar­ily feed on bal­sam firs (Abies bal­samea), Fraser firs (Abies fraseri), sub­apline firs (Abies la­sio­carpa), Pa­cific sil­ver firs (Abies am­a­bilis), and grand firs (Abies gran­dis). In Eu­rope, they are known to feed on Eu­ro­pean sil­ver firs (Abies alba). (Ragen­ovich and Mitchell, 2006)

  • Primary Diet
  • herbivore
    • eats sap or other plant foods
  • Plant Foods
  • sap or other plant fluids

Pre­da­tion

Bal­sam woolly adel­gids are prey to many an­i­mal species. Bee­tles in the genus Lari­cobus, the lady bee­tle genus Scym­nus, and the fly fam­ily Chamae­myi­idae are spe­cial­ist preda­tors on the adel­gid fam­ily Adel­gi­dae. Some of the most abun­dant gen­er­al­ist preda­tors of bal­sam woolly adel­gids are hover flies, mites, and lacewings. The ma­jor­ity of pre­da­tion is on the eggs. Bal­sam woolly adel­gids do not have any known anti-preda­tor be­hav­ioral adap­ta­tions, but the waxy, wool-like cov­er­ing on their back hides their real body from sight and helps them blend in with mosses and lichens on the tree bark. (Davis, et al., 2022; Hum­ble, 1994; Mitchell and Wright, 1967; Mont­gomery and Havil, 2014; Ragen­ovich and Mitchell, 2006)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

In North Amer­ica, bal­sam woolly adel­gids are an in­va­sive species and a deadly par­a­site to true fir trees, in­clud­ing bal­sam firs, Fraser firs, sub­alpine firs, Pa­cific sil­ver firs, and grand firs. Large adel­gid in­fes­ta­tions re­sult in the tree pro­duc­ing ab­nor­mal wood, which pre­vents the tree from get­ting proper water and nu­tri­ents. This de­fi­ciency can kill a host tree in two to nine years. By killing and dam­ag­ing these trees, bal­sam woolly adel­gids also con­tribute to de­for­esta­tion, which threat­ens other an­i­mal and plant species that are en­demic to those forests, in­clud­ing at least two species that are al­ready en­dan­gered and ten that are fed­eral species of con­cern. In Eu­rope, bal­sam woolly adel­gids have a com­men­sal re­la­tion­ship with fir and spruce trees, es­pe­cially Eu­ro­pean sil­ver firs. Be­cause bal­sam woolly adel­gids are na­tive to Eu­rope, these trees are re­sis­tant to adel­gid in­fes­ta­tions. (Boyn­ton and Kelly, 2007; Go­heen and Will­hite, 2006; Hrinke­vich, et al., 2016; Pot­ter, et al., 2005)

Species Used as Host
  • Bal­sam fir (Abies bal­samea)
  • Fraser fir (Abies fraseri)
  • Sub­alpine fir (Abies la­sio­carpa)
  • Pa­cific sil­ver fir (Abies am­a­bilis)
  • Grand fir (Abies gran­dis)
  • Eu­ro­pean sil­ver fir (Abies alba)
  • White fir (Abies con­color)
  • Noble fir (Abies pro­cera)
  • Shasta fir (Abies mag­nifica)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Bal­sam woolly adel­gids are prey to many other an­i­mal species. They pro­vide easy nu­tri­ents for their preda­tors, which ul­ti­mately helps sup­port bio­di­ver­sity in their en­vi­ron­ment. Gen­er­ally, greater bio­di­ver­sity in­di­cates a health­ier, more re­silient ecosys­tem, which is ben­e­fi­cial for both wildlife and hu­mans.

  • Positive Impacts
  • food

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Bal­sam woolly adel­gids cause se­vere eco­nomic costs to the Christ­mas tree in­dus­try. The ma­jor­ity of Christ­mas trees are fir trees, the pri­mary hosts of bal­sam woolly adel­gids. In­fes­ta­tions are dif­fi­cult to spot un­less the farmer knows where and what to look for. Fur­ther­more, in­fested trees are eco­nom­i­cally in­vi­able and must be culled to pre­vent fur­ther in­fes­ta­tions. Treat­ing an area for bal­sam woolly adel­gids can also be very ex­pen­sive, cost­ing famers up to $500 per acre. (Pot­ter, et al., 2005; Ragen­ovich and Mitchell, 2006)

  • Negative Impacts
  • crop pest

Con­ser­va­tion Sta­tus

There is in­suf­fi­cient data to de­ter­mine the abun­dance of bal­sam woolly adel­gids. Be­cause they are con­sid­ered in­va­sive in North Amer­ica, hu­mans have made nu­mer­ous at­tempts to con­trol their pop­u­la­tions through a va­ri­ety of meth­ods. The ma­jor­ity of these at­tempts have had seem­ingly lit­tle or no ef­fect on bal­sam woolly adel­gid pop­u­la­tions. The few meth­ods that are ef­fec­tive against bal­sam woolly adel­gids are costly and un­sus­tain­able. (Al­ston, et al., 2018; Ragen­ovich and Mitchell, 2006; Side­bot­tom, 2019)

Other Com­ments

Bal­sam wooly adel­gids are part of the Adelges piceae species com­plex. This com­plex con­sists of six adel­gid species: Adelges piceae, Adelges ne­bro­den­sis, Adelges prelli, Adelges merk­eri, Adelges schnei­deri, and Adelges nord­man­ni­anae. (Hav­ill, et al., 2020)

Most adel­gid species al­ter­nate be­tween a pri­mary host, which is a spruce tree, and an in­ter­me­di­ate host, which is a fir tree. Sex­ual gen­er­a­tions, which con­sist of both males and fe­males and re­pro­duce sex­u­ally, are only pro­duced on the pri­mary host. Bal­sam woolly adel­gids do not have a pri­mary host, so they do not have a sex­ual gen­er­a­tion and must re­pro­duce asex­u­ally. (Varty, 1956)

Bal­sam woolly adel­gids are some­times called sil­ver fir adel­gids in Eu­rope. (Varty, 1956)

Bal­sam woolly adel­gids from the pro­gre­di­ens gen­er­a­tions can be ei­ther apter­ous (with­out wings) or alate (with wings). Alate pro­gre­di­ens de­velop wing pads as nymphs and have fully formed wings as adults. These wings are trans­par­ent and ap­prox­i­mately 1 mm long. When alate pro­gre­di­ens sur­vive to adults, they be­come motile again and fly to a new host tree as soon as their wings harden. On this new host, they set­tle for a sec­ond time and once again be­come com­pletely seden­tary. Alate pro­gre­di­ens have not been ob­served in the wild. (Varty, 1956)

Con­trib­u­tors

Avery Gib­son (au­thor), Col­orado State Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

agricultural

living in landscapes dominated by human agriculture.

arboreal

Referring to an animal that lives in trees; tree-climbing.

asexual

reproduction that is not sexual; that is, reproduction that does not include recombining the genotypes of two parents

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

diapause

a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

infrared/heat

(as keyword in perception channel section) This animal has a special ability to detect heat from other organisms in its environment.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

native range

the area in which the animal is naturally found, the region in which it is endemic.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

parthenogenic

development takes place in an unfertilized egg

sedentary

remains in the same area

semelparous

offspring are all produced in a single group (litter, clutch, etc.), after which the parent usually dies. Semelparous organisms often only live through a single season/year (or other periodic change in conditions) but may live for many seasons. In both cases reproduction occurs as a single investment of energy in offspring, with no future chance for investment in reproduction.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

visual

uses sight to communicate

Ref­er­ences

Al­ston, D., R. Davis, D. McAvoy, L. Spears, D. Malesky, L. Hebert­son, C. Key­ees. 2018. "Bal­sam Woolly Adel­gid - Ad­vanced Fact Sheet" (On-line). Ac­cessed July 05, 2023 at https://​digitalcommons.​usu.​edu/​extension_​curall/​1838/​.

Boyn­ton, A., C. Kelly. 2007. "Car­olina North­ern Fly­ing Squir­rel" (On-line). NC­pe­dia. Ac­cessed July 07, 2023 at https://​www.​ncpedia.​org/​carolina-northern-flying-squirrel.

Davis, G., L. Lowrey, T. Eck­berg, J. Hicke, E. Smirnova. 2022. Char­ac­ter­iz­ing Bal­sam Woolly Adel­gid In­fes­ta­tions and As­so­ci­ated Tree Mor­tal­ity in Idaho. Jour­nal of Forestry, 120(4): 361-378. Ac­cessed July 10, 2023 at https://​doi.​org/​10.​1093/​jofore/​fvac007.

Go­heen, E., E. Will­hite. 2006. Field Guide to the Com­mon Dis­eases and In­sect Pests of Ore­gon and Wash­ing­ton Conifers. Port­land, Ore­gon: USDA For­est Ser­vice, Pa­cific North­west Re­gion. Ac­cessed July 02, 2023 at https://​www.​biodiversitylibrary.​org/​item/​152991#​page/​5/​mode/​1up.

Hav­ill, N., B. Grif­fin, J. An­der­sen, R. Foot­tit, M. Juste­sen, A. Cac­cone, V. D'Amico, J. Elk­in­ton. 2020. Species De­lim­i­ta­tions and In­va­sion His­tory of the Bal­sam Woolly Adel­gid, Adelges (Drey­fu­sia) piceae (Hemiptera: Aphi­doidea: Adel­gi­dae), Species Com­plex. Sys­tem­atic En­to­mol­ogy, 46(1): 186-204. Ac­cessed July 16, 2023 at https://​doi.​org/​10.​1111/​syen.​12456.

Hrinke­vich, K., R. Progar, D. Shaw. 2016. Cli­mate Risk Mod­el­ling of Bal­sam Woolly Adel­gid Dam­age Sever­ity in Sub­alpine Fir Stands of West­ern North Amer­ica. PLOS ONE, 11(10): 1-11. Ac­cessed July 02, 2023 at https://​journals.​plos.​org/​plosone/​article?​id=10.​1371/​journal.​pone.​0165094.

Hum­ble, L. 1994. Re­cov­ery of ad­di­tional ex­otic preda­tors of bal­sam woolly adel­gid, Adelges piceae (Ratze­burg) (Ho­moptera: Adel­gi­dae), in British Co­lum­bia. The Cana­dian En­to­mol­o­gist, 126(4): 1101-1103. Ac­cessed July 09, 2023 at http://​dx.​doi.​org/​10.​4039/​Ent1261101-4.

Mitchell, R., K. Wright. 1967. For­eign Preda­tor In­tro­duc­tion for Con­trol of the Bal­sam Woolly Adel­gid in the Pa­cific North­west. Jour­nal of Eco­nomic En­to­mol­ogy, 60(1): 140-147. Ac­cessed July 09, 2023 at https://​doi.​org/​10.​1093/​jee/​60.​1.​140.

Mont­gomery, M., N. Havil. 2014. "The Use of Clas­si­cal Bi­o­log­i­cal Con­trol to Pre­serve Forests in North Amer­ica" (On-line). Ac­cessed July 14, 2023 at https://​www.​fs.​usda.​gov/​nrs/​pubs/​jrnl/​2014/​nrs_​2014_​montgomery_​001.​pdf#:​~:text=It%20has%20two%20to%20­four%20­sis­tens%20%28­di­a­paus­ing%29%20­gen­er­a­tions,Balch%2C%201952%3B%20­Varty%2C%201956%3B%20Mitchell%20et%20al.%2C%201961..

Pot­ter, K., J. Framp­ton, J. Side­bot­tom. 2005. Im­pacts of Bal­sam Woolly Adel­gid on the South­ern Ap­palachian Spruce-Fir Ecosys­tem and the North Car­olina Christ­mas Tree In­dus­try. Third Sym­po­sium on Hem­lock Woolly Adel­gid in the East­ern United States, 1: 25-41.

Ragen­ovich, I., R. Mitchell. 2006. For­est In­sect & Dis­ease Leaflet 118: Bal­sam Woolly Adel­gid. Wash­ing­ton, D.C.: U.S. De­part­ment of Agri­cul­ture For­est Ser­vice.

Side­bot­tom, J. 2019. "Bal­sam Woolly Adel­gid" (On-line). NC State Ex­ten­sion. Ac­cessed July 02, 2023 at https://​content.​ces.​ncsu.​edu/​balsam-woolly-adelgid#​section_​heading_​4932.

Varty, I. 1956. Adelges In­sects of Sil­ver Firs. West Bromwich, Great Britain: Her Majesty's Sta­tionery Of­fice. Ac­cessed July 13, 2023 at https://​cdn.​forestresearch.​gov.​uk/​1956/​04/​fcbu026.​pdf.