Diapheromera femorataCommon American Walkingstick(Also: Northern Walkingstick)

Ge­o­graphic Range

Com­mon walk­ing­sticks have a broad ge­o­graphic range in North Amer­ica. They can be found all along the At­lantic coast and into north­ern Florida. This species can be found as far west as New Mex­ico in the United States (US) and as far north as Al­berta, Canada. They are found in 38 US states the north­ern most being Maine in the east, and North Dakota in the west. They are the only stick in­sects to occur in Canada, where they are found in Man­i­toba, On­tario, and Que­bec. (Giese and Knauer, 1977; Milne, 1980)

Habi­tat

Com­mon walk­ing­sticks are found in de­cid­u­ous woods and forests where their pre­ferred food sources (oak and hazel­nut) are abun­dant. They may also be found in agri­cul­tural fields, urban gar­dens and res­i­den­tial yards. (Milne, 1980)

Phys­i­cal De­scrip­tion

Com­mon walk­ing­sticks have very elon­gated bod­ies that are al­most cylin­dri­cal. The ab­domen and tho­rax are long and the ab­domen bears a pair of sin­gle seg­mented cerci that re­sem­ble palps and serve as a clasper. The head is small but bears an­ten­nae that are about 2/3 the length of the body. Legs are slen­der and the tarsi are five seg­mented.

Adult males av­er­age 75 mm in length, while fe­males are slightly larger at about 95 mm. Nymphs of the first five in­stars reach av­er­age lengths of about 11, 18, 25, 34, and 46 mm, re­spec­tively.

A dis­tin­guish­ing fea­ture is the supra-anal plate, which is a small and mem­bra­nous lobe above the anus. Their max­il­lae each con­tain a lacina with a tri­den­tate struc­ture. The species is apter­ous. Mem­bers of Di­a­pheromera femorata ex­hibit square-shaped heads. Males are brown, whereas fe­males have a hint of green to their brown color. There are other dis­tin­guish­ing char­ac­ter­is­tics that sep­a­rate the two sexes; the fe­murs of males tend to be banded, their sev­enth ab­dom­i­nal seg­ment is longer than their ninth, and they fea­ture cerci that lack spines. (Ar­ment, 2006; Giese and Knauer, 1977; Milne, 1980; Tilgner, et al., 1999; Walker, 1922)

  • Sexual Dimorphism
  • female larger
  • sexes colored or patterned differently
  • Average length
    Males 75mm, Females 95mm mm
    in

De­vel­op­ment

Im­ma­ture forms of the species are called nymphs. When nymphs hatch they are green, only to gain brown­ish color when they reach ma­tu­rity. The species typ­i­cally un­der­goes five molts, how­ever some male mem­bers of the species have been viewed to un­dergo four molts, whereas some fe­males have un­der­gone six molts. They are a hemimetabolous species, mean­ing that they have three life stages (egg, nymph, and adult) and do not have a pupal stage. Their life cycle is heav­ily de­pen­dent upon the syn­chro­niz­ing with host plant sea­sonal cy­cles.

Mem­bers of the species found in the north­ern United States and in South­ern Canada tend to be bian­nual with a new gen­er­a­tion every two years, whereas south­ern mem­bers tend to have a new gen­er­a­tion every year. Some pop­u­la­tions lay eggs in the fall, which then over­win­ter in the leaf lit­ter and hatch the fol­low­ing spring. Sea­sonal con­di­tions af­fect the lifes­pan of the nymph, cooler sea­sonal tem­per­a­tures lead the nymphs to reach the adult stage faster than in warmer sea­son tem­per­a­tures. Ac­cord­ing to one study, it took 74.7 days for nymphs to reach the adult stage, whereas the next sea­son it took 84.9 days for this to occur. In one study, hatch­ing oc­curred 11 to 17 days after the eggs were ov­posited. To reach the sec­ond in­star stage, 9 to 12 days were needed from the first molt, and to reach the third in­star, it re­quired 13 to 16 days. To reach the last nypmh stage 14 to 15 ad­di­tional days were re­quired, and from that to molt into an adult it re­quired 14 days. The vari­a­tions be­tween time lengths were due to cli­matic vari­a­tion through­out the sea­son. The first sea­son stud­ied was cooler than the sec­ond and due to the colder weather, the nymphs de­vel­op­ing in the first sea­son had one fewer in­star stage. The sec­ond sea­son re­quired roughly 17 days to reach the fifth in­star and an­other 19 days to be­come adults. The de­vel­op­ment and lifes­pan of nymphs is heav­ily de­pen­dent upon de­gree-days, they tend to re­quire an av­er­age of 1835 de­gree-days, to be­come adults. (Giese and Knauer, 1977; Milne, 1980)

Re­pro­duc­tion

Courtship rit­u­als seem to be ab­sent for Di­a­pheromera femorata. Exact mat­ing sys­tems are un­known for this species. In other species of Phas­mida, males will at­tach them­selves to a fe­male and ride on her back for sev­eral weeks until she is ready to mate. Mat­ing tends to be rather con­sis­tent amongst mem­bers of the order Phas­mida, in which the male climbs on the back of the fe­male, passes his ab­domen to hers, from ei­ther side to en­gage gen­i­talia. Males of most species of Phas­mida ex­tend their ab­domen down and around on the right side of the fe­male, how­ever, Di­a­pheromera femorata males pass theirs down from the left side of the fe­male. The male gen­i­talia form an asym­met­ri­cal struc­ture, which is slightly di­vided on the left side by an oblique groove, which is where the ejac­u­la­tory duct opens. The duct is po­si­tioned slightly to the right and opens at the ven­tral side of the in­sect. A chiti­nous plate pro­tects the dor­sal sur­face of the gen­i­talia, and gives rise to a blunt horn-like process on the left side of the gen­i­talia. Next to the me­dian groove, on the pos­te­rior-ven­tral side, arises two plates, which form a clasper. The two plates form an in­ward fac­ing apoph­ysis for mus­cle at­tach­ment, al­low­ing for the clasper to grab onto the fe­male dur­ing mat­ing. (Huber, et al., 2007; Walker, 1922; Huber, et al., 2007; Walker, 1922)

Ovipo­si­tion be­gins in late Au­gust, peaks dur­ing mid-Sep­tem­ber, and tails off at the end of Sep­tem­ber, where it con­tin­ues until late Oc­to­ber until most of the green fo­liage is no longer pre­sent. The process of ovipo­si­tion tends to ini­ti­ate be­tween noon and 3:00 P.M., with peak ac­tiv­ity rang­ing from the times of 3:00 and 9:00 P.M., thus mak­ing the process de­pen­dent upon sun­light. The fe­males of the species drop eggs one at a time. The eggs drop­ping from the trees sound like of droplets of rain. The eggs over­win­ter on the ground amongst the leaf lit­ter until spring when the nymphs hatch. They push through ends of the egg and then crawl up the near­est tree dur­ing the night to find food. Hatch­ing in oak forests oc­curs in mid June, typ­i­cally after full ex­pan­sion of the black oaks (Quer­cus ve­lutina) oc­curs and typ­i­cally lasts through­out July, how­ever, Di­a­pheromera femorata hatch­ing has oc­curred in Sep­tem­ber. Hatch­ing fol­lows a daily pe­ri­odic pat­tern, in which it oc­curs from the range of 4:00 P.M. to 7:00 A.M. with peak hatch­ing ac­tiv­ity oc­cur­ring within the time frame of 10:00 P.M. to 6:00 A.M. It has also been noted that hu­mid­ity plays a strong role in Di­a­pheromera femorata egg hatch­ing. Hatch­ing tends to occur when the hu­mid­ity is 80 per­cent or higher be­cause the mois­ture in the air serves as a lu­bri­cant, al­low­ing for the nymphs to get out of their eggs eas­ily. (Giese and Knauer, 1977; Milne, 1980)

  • Breeding interval
    Common walkingsticks breed once yearly.
  • Breeding season
    Breeding occurs between late August and late October.
  • Range eggs per season
    145.85 in a meters squared area to 787.06 in a meters squared area
  • Average eggs per season
    407.12 in a meters squared area
  • Average gestation period
    11-17 days
  • Average age at sexual or reproductive maturity (female)
    80 days
  • Average age at sexual or reproductive maturity (male)
    80 days

This species does not ex­hibit parental care. Eggs are dropped, usu­ally from great heights, down to the leaf lit­ter where they are left to over­win­ter. When the nymphs hatch they fend for them­selves. (Giese and Knauer, 1977)

  • Parental Investment
  • no parental involvement

Lifes­pan/Longevity

Cur­rently, lit­tle in­for­ma­tion is known re­gard­ing the lifes­pan of com­mon walk­ing­sticks. (Coul­son and Wit­ter, 1984; Giese and Knauer, 1977)

Be­hav­ior

Com­mon walk­ing­sticks use cam­ou­flage as their main means of de­fend­ing them­selves from birds and other preda­tors. This species' phys­i­cal ap­pear­ance has a strong re­sem­blance to a twig, which al­lows them to blend in with their sur­round­ings. Newly hatched nymphs tend to walk up the first ver­ti­cal ob­ject they en­counter, see if it is a host tree for feed­ing, and if not they leave the ob­ject and move in a dif­fer­ent di­rec­tion. Com­mon walk­ing­sticks are ac­tive at dif­fer­ent times of the day for dif­fer­ent ac­tiv­i­ties. Feed­ing and nymph hatch­ing take place typ­i­cally at night, where mat­ing usu­ally takes place dur­ing the day. Aside from mat­ing, they are soli­tary crea­tures. (Giese and Knauer, 1977; Milne, 1980)

Home Range

Ter­ri­tory size is cur­rently un­known for com­mon walk­ing­sticks.

Com­mu­ni­ca­tion and Per­cep­tion

Com­mu­ni­ca­tion and per­cep­tion meth­ods are cur­rently poorly un­der­stood for this species. In­di­vid­u­als likely use pheromones dur­ing mat­ing.

Food Habits

Di­a­pheromera femorata feeds on the fo­liage of trees and shrub­bery. They are par­tic­u­larly fond of oak and hazel­nut trees. They are her­biv­o­rous and have mandibles to cut pieces of the leaves, stems or flow­ers. Newly hatched nymphs tend to feed mainly on hazel (Cory­lus amer­i­cana) and black cherry (Prunus serotina), but in en­vi­ron­ments where these plants are not in high abun­dance they tend to eat white oak (Quer­cus alba). They have also been noted to con­sume sweet­fern (Comp­to­nia pere­g­rina), var­i­ous straw­berry and blue­berry plants, and beaked hazel (Cory­lus cor­nuta), from the time frame of May to mid-June. The nymphs eat by con­sum­ing all but the major veins of a leaf. Adults tend to feed pri­mar­ily on black oak (Quer­cus ve­lutina) at all times of the day, but peak feed­ing ac­tiv­ity was found to occur at 9:00 P.M. to 3:00 A.M., and their feed­ing habits tend to in­clude feed­ing on a sin­gle leaf for quite some time, stop, move about, and then begin feed­ing on an­other leaf. They are leaf skele­toniz­ers. (Clark, 1974; Coul­son and Wit­ter, 1984; Giese and Knauer, 1977; Milne, 1980)

  • Plant Foods
  • leaves
  • flowers

Pre­da­tion

Sev­eral in­sec­tiv­o­rous bird species are preda­tors of Di­a­pheromera femorata in­clud­ing crows and Amer­i­can robins. Com­mon walk­ing­sticks show a re­mark­able abil­ity for re­gen­er­at­ing legs that are lost by at­tacks from preda­tors. When preda­tors are pre­sent, they re­main mo­tion­less with their legs close to their bod­ies, thus re­sem­bling a twig.

Di­a­pheromera femorata eggs are sim­i­lar to that of other Phas­mid eggs in which they greatly re­sem­ble seeds of var­i­ous plants, and in the case of D. femorata they re­sem­ble seeds of legumes. This may be a de­fense mech­a­nism, the eggs con­tain a ca­pit­u­lum, a struc­ture that re­sem­bles a plant elia­some, which at­tracts var­i­ous species of ants. In other species of stick in­sects, the ants take the egg think­ing it is a seed, re­move the ca­pit­u­lum as food, and dis­card the rest of the egg at the bot­tom of their nest, thus pro­tect­ing the egg from out­side par­a­sites or preda­tors. ("For­est In­sect & Dis­ease Leaflet 82: Walk­ing­stick", 1971; Hughes and West­oby, 1992; Mill­ron, 1950; Milne, 1980; Sev­erin, 1910)

  • Anti-predator Adaptations
  • mimic
  • cryptic

Ecosys­tem Roles

Di­a­pheromera femorata is a vo­ra­cious her­bi­vore that can cause sig­nif­i­cant de­fo­li­a­tion when pop­u­la­tions num­bers are high. Dur­ing out­break years, this species may cause en­tire tree branches to die. So far, in­sec­tiv­o­rous birds con­sume enough dur­ing out­breaks to suf­fi­ciently con­trol the dam­age. Abun­dance of prey items dur­ing out­breaks may re­sult in pop­u­la­tion booms for the in­sec­tiv­o­rous bird species that con­sume them. Di­a­pheromera femorata fe­males pre­fer cer­tain tree species (black cherry, black and white oaks) as hosts to lay their eggs. Sev­eral species of par­a­sitic in­sects uti­lize Di­a­pheromera femorata as a host. The wasp species Me­si­tiopterus kahlii is a par­a­site to Di­a­pheromera femorata eggs. Two species of flies, Bio­mya genalis and Phas­mophaga an­ten­nalis de­stroy walk­ing­stick lar­vae. ("For­est In­sect & Dis­ease Leaflet 82: Walk­ing­stick", 1971; Giese and Knauer, 1977; Hughes and West­oby, 1992; Mill­ron, 1950; Milne, 1980; Sev­erin, 1910)

Species Used as Host
  • black oaks Quer­cus ve­lutina
  • white oaks Quer­cus alba
  • black cher­ries Prunus serotina
Com­men­sal/Par­a­sitic Species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Cur­rently there are no known pos­i­tive af­fects of Di­a­pheromera femorata on hu­mans.

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Com­mon walk­ing­sticks are her­bi­vores that com­monly "skele­tonize" leaves, or eat every part of the leaf but the vein. This causes sig­nif­i­cant dam­age to the trees and dur­ing out­breaks, this species may cause the death of en­tire tree branches. This in­sect species is par­tially re­spon­si­ble for the de­fo­li­a­tion and re­duc­tion of oak forests in the Ozark Moun­tains of Arkansas and Mis­souri in the United States. (Stephen, et al., 2001)

Con­ser­va­tion Sta­tus

The species is abun­dant and wide­spread, so no con­ser­va­tion meth­ods are being im­ple­mented. If any­thing, fu­ture ef­forts may be made to re­duce the pop­u­la­tion in re­gions suf­fer­ing from sig­nif­i­cant for­est de­fo­li­a­tion.

Con­trib­u­tors

Lind­say Har­ring­ton (au­thor), Rut­gers Uni­ver­sity, Dave San­nino (au­thor), Rut­gers Uni­ver­sity, David V. Howe (ed­i­tor), Rut­gers Uni­ver­sity, Rachelle Ster­ling (ed­i­tor), Spe­cial Pro­jects.

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

agricultural

living in landscapes dominated by human agriculture.

arboreal

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

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.

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

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

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

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.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

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

nocturnal

active during the night

oviparous

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

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

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

solitary

lives alone

suburban

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

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.

urban

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

Ref­er­ences

U.S. De­part­ment of Agri­cul­ture For­est Ser­vice. For­est In­sect & Dis­ease Leaflet 82: Walk­ing­stick. 82. Wash­ing­ton D.C.: U.S. Gov­ern­ment Print­ing Of­fice. 1971. Ac­cessed March 30, 2011 at http://​www.​na.​fs.​fed.​us/​spfo/​pubs/​fidls/​walkingstick/​walkingstick.​htm.

Ar­ment, C. 2006. Stick In­sects of the Con­ti­nen­tal United States And Canada: Species And Early Stud­ies. Lan­disville, Penn­syl­va­nia: Coach­whip Pub­li­ca­tions.

Clark, J. 1974. Stick and Leaf In­sects. Great Britain: Barry Shur­lock.

Coul­son, R., J. Wit­ter. 1984. For­est En­to­mol­ogy: Ecol­ogy and Man­age­ment. USA: Wi­ley-IEEE.

Giese, R., K. Knauer. 1977. Ecol­ogy of the Walk­ing­stick. For­est Sci­ence, 23(1): 45-63.

Gre­gory, T. 2002. Genome size of the north­ern walk­ing­stick, Di­a­pheromera femorata (Phas­mida: Het­erone­mi­idae). Cana­dian Jour­nal of Zo­ol­ogy, 80(7): 1303-1305.

Huber, B., B. Sin­clair, M. Schmitt. 2007. The Evo­lu­tion of Asym­met­ric Gen­i­talia in Spi­ders and In­sects. Bi­o­log­i­cal Re­views of the Cam­bridge Philo­soph­i­cal So­ci­ety, 82(4): 647-698.

Hughes, L., M. West­oby. 1992. Ca­pit­ula on Stick In­sect Eggs and Elaio­somes on Seeds: Con­ver­gent Adap­ta­tions for Bur­ial by Ants. Func­tional Ecol­ogy, 6(6): 642-648.

Mill­ron, H. 1950. The Iden­tity of a Clep­tid Egg Par­a­site of the Com­mon Walk­ing Stick, Di­a­pheromera femorata Say (Hy­menoptera: Clep­ti­dae). The Pro­ceed­ings of the En­to­mo­log­i­cal So­ci­ety of Wash­ing­ton, 52(1): 47.

Milne, L. 1980. Na­tional Audubon So­ci­ety Field Guide to North Amer­i­can In­sects & Spi­ders. New York: Al­fred A, Knopf, Inc..

Sev­erin, H. 1910. A Study on the Struc­ture of the Egg of the Walk­ing Stick Di­a­pheromera femorata Say; and the Bi­o­log­i­cal Sig­nif­i­cance of the Re­sem­blance of Phas­mid Eggs to Seeds. An­nals of the En­to­mo­log­i­cal So­ci­ety of Amer­ica, 3(1): 83-93.

Stephen, F., V. Sal­is­bury, F. Oliv­e­ria. 2001. Red Oak Borer, Enaphalodes ru­fu­lus (Coleoptera: Ce­r­am­by­ci­dae), in the Ozark Moun­tains of Arkansas, U.S.A.: An un­ex­pected and Re­mark­able For­est Dis­tur­bance. In­te­grated Pest Man­age­ment Re­views, 6(3-4): 247-252.

Tilgner, E., T. Kise­ly­ova, J. McHugh. 1999. A mor­pho­log­i­cal study of Timema cristi­nae vick­ery with im­pli­ca­tions for the phy­lo­ge­net­ics of phas­mida. Deutsche En­to­mol­o­gis­che Zeitschrift, 46(2): 149-162.

Walker, E. 1922. The Ter­mi­nal Struc­tures of Or­thopteroid In­sects: A Phy­lo­ge­netic Study. An­nals of the En­to­mo­log­i­cal So­ci­ety of Amer­ica, 15(1): 1-71.