Polistes dominulaEuropean paper wasp

Ge­o­graphic Range

Eu­ro­pean paper wasps are na­tive to the Palearc­tic Re­gion around the Mediter­ranean, north­ern Africa, south­ern Eu­rope, the Mid­dle East, and east­ern China. They were in­tro­duced to the Nearc­tic Re­gion of North Amer­ica in the 1970s and 1980s. Al­though ini­tially found pri­mar­ily on the east coast of the United States, they have since spread into the Mid­west and re­cently to­wards the west­ern and south­west­ern United States. Eu­ro­pean paper wasps were in­tro­duced to Canada, in­clud­ing On­tario, Nova Sco­tia, and British Co­lum­bia in the 1990’s. They have also been in­tro­duced to Chile and Ar­gentina in the Neotrop­i­cal Re­gion as well as west­ern Aus­tralia in the Aus­tralian Re­gion. (Cervo, et al., 2000; Ja­cobs, 2011)

Habi­tat

Eu­ro­pean paper wasps live in tem­per­ate and ter­res­trial habi­tats in­clud­ing chap­ar­ral, for­est and grass­land bio­mes. They re­side in urban, sub­ur­ban, and agri­cul­tural lo­ca­tions. They tend to re­side close to human civ­i­liza­tion be­cause they nest in human struc­tures. They also live in forests and on plants where they can feed and nest. When nest­ing, they choose spaces cre­ated by farm ma­chin­ery and recre­ational struc­tures. Dur­ing win­ter, im­preg­nated queens re­side in pro­tected lo­ca­tions such as within house walls or in hol­low trees. These fe­males then cre­ate nests in these lo­ca­tions or nearby at the be­gin­ning of spring. (Buck, et al., Feb­ru­ary 19, 2008; Ja­cobs, 2011; Rusina, et al., 2011)

Phys­i­cal De­scrip­tion

Eu­ro­pean paper wasps are black wasps with yel­low mark­ings in­clud­ing rings along the ab­domen and spots lo­cated in var­i­ous lo­ca­tions on the clu­peus (face) and ab­domen, as­so­ci­ated with gen­der and pos­si­bly dom­i­nance. Dif­fer­ences in the clypeal col­oration pat­terns may sug­gest ag­gres­sion and sta­tus lev­els. Some in­di­vid­u­als have one spot on their clypeus, and a greater quan­tity of black on the clypeus may sug­gest dom­i­nance and also in­di­cate phys­i­cal size. The fla­gella (an­ten­nae) are bright red­dish-or­ange. (Buck, et al., 2008a; Buck, et al., 2008b; Buck, et al., Feb­ru­ary 19, 2008; Green and Field, 2010)

Eu­ro­pean paper wasps are smaller than the na­tive North­ern paper wasps and mea­sure 2.0 cm in length on av­er­age. They are bi­lat­er­ally sym­met­ri­cal, and their bod­ies are made up of tag­mata in­clud­ing a head, tho­rax, and ab­domen, with a thin con­stric­tion be­tween the tho­rax and ab­domen. They have six legs, a pair of an­tenna, and a pair of wings. There is an in­den­ta­tion lo­cated in the meso­pleu­ron, as well as ridg­ing along the propodeum (the first ab­dom­i­nal seg­ment that is fused to the tho­rax). Com­pared to other species in their genus (Polistes), Eu­ro­pean paper wasps are known to fly with their hind legs po­si­tioned slightly below their body. (Buck, et al., 2008a; Buck, et al., Feb­ru­ary 19, 2008; Cran­shaw, 2008)

Col­oration and size ex­hibit sex­ual di­mor­phism. The ven­tral sur­face is black in fe­males and yel­low in males. Fe­males tend to be larger than males. Fe­male forewing length ranges from 9.5 to 13.0 mm, whereas males' forewings range from 8.5 to 12.0 mm. (Buck, et al., 2008a; Buck, et al., 2008b)

  • Sexual Dimorphism
  • female larger
  • sexes colored or patterned differently
  • Average length
    2.0 cm
    0.79 in
  • Range wingspan
    8.5 to 13.0 mm
    0.33 to 0.51 in

De­vel­op­ment

Meta­mor­pho­sis takes place as eggs de­velop into lar­vae, which are fed by both the queen and worker wasps that col­lect food. The lar­vae pu­pate and then trans­form into adults. If the tem­per­a­ture and food con­di­tions are op­ti­mal, the eggs de­velop into adults in 40 days. Vi­bra­tions caused by adult Eu­ro­pean paper wasps in­flu­ences whether the de­vel­op­ing lar­vae be­come work­ers or founders. This vi­bra­tion is caused by an­ten­nae drum­ming which bio­chem­i­cally af­fects the lar­vae and al­ters gene ex­pres­sion in the lar­val stage. (Surya­narayanan, et al., 2011)

Ju­ve­nile Hor­mone (JH) is found in many Polistes species and func­tions in reg­u­lat­ing de­vel­op­ment, di­a­pause, and re­pro­duc­tion. Whereas in other species JH re­spon­sive­ness may dif­fer be­tween foundresses and work­ers, in Eu­ro­pean paper wasps, JH re­spon­sive­ness is con­di­tion-de­pen­dent and there are no clear dif­fer­ences in re­spon­sive­ness be­tween foundresses and work­ers. (Tib­betts, et al., 2011)

Re­pro­duc­tion

Within colonies of Eu­ro­pean paper wasps, one queen gen­er­ally mates with mul­ti­ple males. Res­i­dent males often cop­u­late with the fe­male more often than tran­sient males. Res­i­dent males de­fend one land­mark for a few con­sec­u­tive days, while tran­sient males may fly be­tween mul­ti­ple land­marks. Their pre­ferred land­marks are large trees. They may mark their leaf and stem perches by rub­bing their legs and last ster­n­ite (com­po­nent of the ster­num on the ven­tral por­tion of the ab­domen) on them. In choos­ing males, a queen flies over the dif­fer­ent males' ter­ri­to­ries and tends to choose a res­i­dent male be­cause res­i­dence often sug­gests that a male is large, ag­gres­sive, and sex­u­ally ac­tive. Once the lar­vae de­velop, the work­ers as­sist the foundress in pro­vid­ing food for the lar­vae through co­op­er­a­tive breed­ing. (Beani and Turil­lazzi, 1987; Beani, et al., 1992)

Queen Eu­ro­pean paper wasps are more likely to cop­u­late with non-nest­mate males than nest­mate males. On the con­trary, males do not dis­crim­i­nate be­tween nest­mate and non-nest­mate fe­males. Fe­males re­ject a sig­nif­i­cant num­ber of cop­u­la­tion ef­forts by males. Fe­males are known to mate with sev­eral males, thus sug­gest­ing a form of post-cop­u­la­tory choice in the fe­male. (Liebert, et al., 2010)

Eu­ro­pean paper wasps are dioe­cious and cop­u­late through in­ter­nal sex­ual re­pro­duc­tion. Fe­males are iteroparous and mate through­out the spring, cre­at­ing nu­mer­ous broods. In the fall, re­pro­duc­tive fe­males mate one final time and find a place to take shel­ter and hi­ber­nate for the win­ter. They store sperm post-cop­u­la­tion at the end of the sea­son which helps to delay fer­til­iza­tion until the spring. They re­side in spaces within houses, in walls, and out­doors. In April and May, they es­tab­lish nests typ­i­cally in dark, pro­tected lo­ca­tions. They often choose man-made struc­tures to build nests, and also pro­tec­tive rocks. Eu­ro­pean paper wasps nest in man-made struc­tures more often than north­ern paper wasps (Polistes fus­ca­tus). Fe­male wasps some­times nest alone with­out other fe­male foundresses. Al­ter­na­tively, fe­males choose to nest with other fer­til­ized fe­males and cre­ate mul­ti­ple-foundress nest­ing sites. Oc­ca­sion­ally, fe­males choose to wait in their win­ter hi­ber­na­tion lo­ca­tion until later in the sea­son, when they can take over a nest that has been aban­doned by an­other foundress. Foundresses at­tach their paper nests, con­structed from wood fibers and paper, to wood, metal, or rock. At the be­gin­ning of the sea­son, the fer­til­ized fe­male lays eggs within the few paper cells of the nest. Over the course of the sea­son, worker wasps help to re­con­struct and in­crease the size of the nest. (Cran­shaw, 2008; Zanette and Field, 2011)

Fe­males lay eggs within cells of the nest that hatch, pu­pate, and de­velop into adults in 3 to 4 weeks after they are laid. The eggs hatch into lar­vae in 3 to 5 days, and the queen wasp feeds them mas­ti­cated cater­pil­lars. The lar­vae cre­ate a silk cap over the top of a nest cell and pu­pate within this closed nest sec­tion. Within 3 to 4 weeks, the pupal wasps de­velop into full-sized adults. As adult wasps de­velop, they be­come work­ers within the colony and are sub­or­di­nate to the fe­male queen. In the sum­mer and later part of the sea­son, sev­eral dozen wasps com­prise the colony. These worker wasps help the foundress gather re­sources to feed fu­ture lar­vae and con­struct suf­fi­cient nest space for them. Some of the work­ers be­come re­pro­duc­tive males and fe­males to­wards the end of the sea­son. These wasps re­pro­duce, but fer­til­ized fe­males are the only wasps to sur­vive the win­ter into the next sea­son. (Cran­shaw, 2008)

  • Breeding interval
    European paper wasps breed numerous times each year, and have numerous broods.
  • Breeding season
    European paper wasps typically mate in spring, summer and fall.
  • Average gestation period
    3-5 days

Fe­male Eu­ro­pean paper wasps pro­tect and pro­vi­sion their off­spring from be­fore they are fer­til­ized until the wean­ing process. Eggs de­velop in their bod­ies until there is a nest to lay them in. Once the eggs hatch, the foundress feeds the lar­vae chewed up in­sects like cater­pil­lars. The first brood of the sea­son be­come worker wasps that help to con­struct the nest and col­lect food for the fol­low­ing broods. The queen fo­cuses her en­ergy on lay­ing more eggs while the work­ers pro­vide the nec­es­sary de­fense and re­sources for the colony. (Bartelt, 2011)

  • Parental Investment
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • post-independence association with parents

Lifes­pan/Longevity

The lifes­pan of the queen wasp is longer than worker be­cause work­ers pro­tect the queen and she hi­ber­nates dur­ing the win­ter. Colonies with one or more queens are founded in late March or April. These ini­tially start with the sin­gle or mul­ti­ple foundresses that were fer­til­ized in the fall. They lay their eggs, which even­tu­ally de­velop into the first set of work­ers of the sea­son. Nei­ther males nor non-re­pro­duc­tive fe­males sur­vive the win­ter; their lifes­pan is less than a year. They only sur­vive the spring, sum­mer, and fall sea­sons. Re­pro­duc­tive fe­males may sur­vive mul­ti­ple sea­sons. (Cran­shaw, 2008; Strass­mann, et al., 2004)

Be­hav­ior

Eu­ro­pean paper wasps are mo­bile and move by fly­ing and walk­ing. They are so­cial and live in colonies that have dom­i­nance hi­er­ar­chies be­tween the queen and the work­ers, in­clud­ing hi­er­ar­chies within the work­ing wasps. Among the work­ers, four typ­i­cal be­hav­iors occur. These in­clude: gen­eral move­ment and be­hav­ior on the nest, search­ing for prey, fur­ther con­struc­tion of the nest, and sit­ting on the nest. Eu­ro­pean paper wasps are di­ur­nal and re­turn to the nest for the night. (Kar­sai, et al., 1996; Ther­aulaz, et al., 1990)

Since ag­gres­sive be­hav­iors are costly, hav­ing a dis­tinc­tive fa­cial phe­no­type for clypeal col­oration may de­crease the level of com­pe­ti­tion from other wasps. Eu­ro­pean paper wasps may be more likely to chal­lenge a weaker rival based on its clypeal pat­tern. (Cervo, et al., 2008; Green and Field, 2010; Shee­han and Tib­betts, 2009; Ther­aulaz, et al., 1992; Tib­betts and Dale, 2004)

Fe­male queens are fer­til­ized in the fall, go into hi­ber­na­tion, and pro­duce eggs in the early spring. Once these eggs be­come adults after 3 to 4 weeks, they be­come work­ers within the colony and are sub­or­di­nate to the fe­male queen. In the sum­mer and later part of the sea­son, sev­eral dozen wasps com­pose the colony. Within wasp colonies, the first born wasps of a brood may be the ones that be­come an in­te­gral part of the colony, whereas wasps born later in the brood may have lower so­cial sta­tus or may be ex­cluded from the colony. In small colonies, the work­ers be­sides the ini­tial six from a brood often have lit­tle in­ter­ac­tion with higher ranked wasps. (Cran­shaw, 2008; Kar­sai, et al., 1996)

A fer­til­ized fe­male Eu­ro­pean paper wasp that starts a colony dur­ing the spring builds a nest along with the help of her first brood of off­spring. The wasps con­tinue to use this nest through­out the sea­son al­though they add to it as the colony gets larger. The fol­low­ing sea­son the fer­til­ized fe­male, whether the same or dif­fer­ent in­di­vid­ual from the pre­vi­ous year, es­tab­lishes a new nest and rarely re­turns to the nest of the pre­vi­ous year. Foundresses visit sig­nif­i­cantly less area on the comb than work­ers. Each wasp vis­its roughly 12% of the comb sur­face within a day re­gard­less of the total time that they spend on the nest that day. While work­ers vis­ited on av­er­age 52% of the comb sur­face over many days, foundresses only vis­ited 50%. (Barac­chi, et al., 2010; Cran­shaw, 2008)

Home Range

There is lit­tle in­for­ma­tion about the home range of Eu­ro­pean paper wasps.

Com­mu­ni­ca­tion and Per­cep­tion

Recog­ni­tion be­tween Eu­ro­pean paper wasp in­di­vid­u­als in­creases the level of co­op­er­a­tion and si­mul­ta­ne­ously de­creases the level of ag­gres­sion be­tween wasps. How­ever, there may be no in­di­vid­ual recog­ni­tion bbe­cause quan­ti­ties of ag­gres­sive be­hav­iors are the same re­gard­less of whether the wasps have met be­fore or not. (Shee­han and Tib­betts, 2010)

In order to dif­fer­en­ti­ate be­tween nest­mates and non-colony mem­bers, Eu­ro­pean paper wasps may rely on iso­lated cu­tic­u­lar hy­dro­car­bons lo­cated on the epi­cu­ti­cle, the out­side waxy layer of the cu­ti­cle. These hy­dro­car­bons serve as a chem­i­cal iden­ti­fi­ca­tion sig­nal for in­di­vid­u­als to dis­crim­i­nate be­tween mem­bers of their colony and other colonies. How­ever, polar com­pounds lo­cated on the epi­cu­ti­cle do not play a role in this iden­ti­fi­ca­tion. (Br­us­chini, et al., 2011)

Eu­ro­pean paper wasps use tac­tile com­mu­ni­ca­tion in two forms of dom­i­nance be­hav­ior: mount­ing and box­ing. The more ex­treme form, mount­ing, in­volves a more dom­i­nant wasp drum­ming his or her an­ten­nae on an­other wasp's head, re­sult­ing in the re­cip­i­ent wasp low­er­ing its head and an­ten­nae in re­sponse. Box­ing con­sists of a wasp using its front legs to bat at an­other wasp and grap­pling with it by curl­ing and spi­ral­ing its body around its op­po­nent to try to gain dom­i­nance. Eu­ro­pean paper wasps also lightly bat other wasps’ heads with their an­ten­nae or drum on their ab­domens as a more sub­tle form of an­a­lyz­ing them. (Izzo, 2011)

Food Habits

Eu­ro­pean paper wasps are om­niv­o­rous, but some­times preda­tory, feed­ing on in­sect lar­vae and cater­pil­lars. They also con­sume aphids, hon­ey­dew, and nec­tar from flow­ers. Eu­ro­pean paper wasps con­sume a wider va­ri­ety of in­sects than north­ern paper wasps, which spe­cial­ize in cater­pil­lars. (Cervo, et al., 2008)

  • Animal Foods
  • insects
  • Plant Foods
  • fruit
  • nectar

Pre­da­tion

Eu­ro­pean paper wasps look sim­i­lar to other species in the genus, which may make it dif­fi­cult for preda­tors to dis­cern be­tween two species. (Bartelt, 2011)

Ecosys­tem Roles

Both na­tive and in­tro­duced colonies of Eu­ro­pean paper wasps play a role in pol­li­nat­ing plants. They also con­sume in­sects, par­tic­u­larly cater­pil­lars, as both adults and lar­vae. In cer­tain areas where Eu­ro­pean paper wasps are an in­tro­duced species, there are sig­nif­i­cant con­cerns re­gard­ing its ef­fects on the na­tive North­ern paper wasp. Since Eu­ro­pean paper wasps es­tab­lish their nests be­fore na­tive species in the spring, they can ex­pand their colonies with­out sig­nif­i­cant com­pe­ti­tion. By nest­ing early in the sea­son, they avoid bird preda­tors, thereby in­creas­ing their sur­vival rate and the like­li­hood that early broods sur­vive and de­velop into work­ers that can pro­tect lar­vae. Un­like na­tive mem­bers of their genus (Polistes), Eu­ro­pean paper wasps con­sume cater­pil­lars as well as other foods, and their var­ied diet ben­e­fits lar­val de­vel­op­ment due to in­creased nu­tri­tional vari­a­tion. Their re­pro­duc­tive suc­cess may have neg­a­tive ef­fects on na­tive (Polistes) species pop­u­la­tions, as well as other in­sects. (Cervo, et al., 2000; Cran­shaw, 2008; Ja­cobs, 2011; Stahlhut, et al., 2006)

Na­tive Eu­ro­pean paper wasps in the Mediter­ranean are often in­fected by par­a­sitoids in­clud­ing En­durus ar­gi­o­lus and Xenos ves­parum. Adult par­a­sitoids lay eggs on or in­side of a host or­gan­ism, which even­tu­ally hatch into lar­vae that in­fect, ster­il­ize, and some­times kill their hosts. These North Amer­i­can par­a­sitoids do not gen­er­ally in­fect Eu­ro­pean paper wasps in the United States be­cause they do not rec­og­nize them as hosts based on their re­cent in­tro­duc­tion. Wol­bachia is a par­a­sitic mi­crobe bac­te­ria that in­fects male and fe­male Eu­ro­pean paper wasps in both Italy and north­east­ern United States. (Cervo, et al., 2000; Ja­cobs, 2011; Stahlhut, et al., 2006)

  • Ecosystem Impact
  • pollinates
Com­men­sal/Par­a­sitic Species
  • bac­te­ria (Wol­ba­cia)
  • par­a­sitoid in­sects (Xenos ves­parum)
  • par­a­sitoid wasps (En­durus ar­gi­o­lus)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Eu­ro­pean paper wasps pol­li­nate plants, thereby ben­e­fit­ing hu­mans. They help con­trol in­sects such as cater­pil­lars, horn­worm lar­vae, cab­bage­worms, tent cater­pil­lars, and lar­vae of sawfly fam­i­lies Cim­bi­ci­dae, Dipri­on­idae, and Ten­thre­dinidae. (Bartelt, 2011; Cran­shaw, 2008)

  • Positive Impacts
  • pollinates crops
  • controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Eu­ro­pean paper wasps sting hu­mans and other an­i­mals when they get too close to their nests. Ad­di­tion­ally, na­tive wasp species may be threat­ened by them, which could even­tu­ally af­fect the fauna in the area. Eu­ro­pean paper wasps choose to nest in man-made struc­tures more often than other species in their genus (Polistes) and thus peo­ple often re­sort to using pes­ti­cides or other tech­niques to kill the wasps and re­move the nests. (Bartelt, 2011; Buck, et al., 2008a)

  • Negative Impacts
  • injures humans
    • bites or stings

Con­ser­va­tion Sta­tus

Eu­ro­pean paper wasps are not threat­ened or en­dan­gered.

Con­trib­u­tors

Eliza Stout (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor, Cather­ine Kent (ed­i­tor), Spe­cial Pro­jects.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

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

Neotropical

living in the southern part of the New World. In other words, Central and South America.

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.

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.

carnivore

an animal that mainly eats meat

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

cooperative breeder

helpers provide assistance in raising young that are not their own

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ectothermic

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

eusocial

the condition in which individuals in a group display each of the following three traits: cooperative care of young; some individuals in the group give up reproduction and specialize in care of young; overlap of at least two generations of life stages capable of contributing to colony labor

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

forest

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

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

hibernation

the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.

insectivore

An animal that eats mainly insects or spiders.

internal fertilization

fertilization takes place within the female's body

introduced

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

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

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.

motile

having the capacity to move from one place to another.

native range

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

nectarivore

an animal that mainly eats nectar from flowers

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

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

polyandrous

Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

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.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

vibrations

movements of a hard surface that are produced by animals as signals to others

visual

uses sight to communicate

Ref­er­ences

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