insect origami: into the fold

210 AMERICAN ENTOMOLOGIST • Winter 2003

Origami is a Japanese word—“ori”meaning folding and “gami” mean-ing paper. It is a challenging art and

craft in which square sheets of paper areformed into specific objects by folding andcreasing. In the strict form of origami, asquare piece of paper, without any cutting,pasting, or decorating, is used. The first pa-pers that could be folded were made in Chinaabout 2,000 years ago (Jackson 1991). In538 A.D., a Buddhist priest brought paper-making methods to Japan from Korea(Honda 1983). By the eighth century,origami was established for the annual DollFestival in Japan, where children made pa-per dolls to throw into the river in the beliefthat evil spirits threatening children wouldbe swept away (Chiyo 1982, Harbin 1997).Origami soon became a practical art form,used to make boxes and small envelopes tohold offerings, spices, medicines, combs, fans,and hairpins (Boursin 2000). Since the early800s, origami has spread as a form of enter-tainment handed down from generation togeneration. Although an independent tradi-tion of paper folding has existed in Europefor hundreds of years (Robert J. Lang, per-sonal communication), the modern form ofthe art as an extension of Japanese origamibegan in the early 1950s in England andspread throughout Great Britain, the UnitedStates, and Europe (Mulatinho 1995).

Insect OrigamiAmong the most difficult models to fold

in origami are insects (Lang 1995). Thesemodels are challenging because their com-plex morphological features are difficult toreproduce from single, square sheets of pa-per; insects usually have three pairs of legs,two pairs of wings, one pair of antennae,and other specialized appendages. Detailedinsect models with all their appendages mayrequire more than 100 carefully plannedfolds.

A general design for origami models re-quires assembling many geometric folds intobasic shapes or “bases” (e.g., the bird, blintz,fish, frog, kite, pig, waterbomb, and wind-mill bases). A single base may be used formany origami models. For example, a fewinsect models can be folded using thewaterbomb base (Fig. 1). These bases, how-ever, may not have the correct geometry todifferentiate all the parts of a delicate insectmodel. In the past, only the blintzed birdbase (i.e., a combination of the bird base andthe blintz base) had enough folds to makeinsects with the correct number of append-ages (Fig. 2); therefore, insect models werefew (Lang 1995). In the 1980s, modern in-sect origami was started by enthusiastic pa-per folders who developed new techniques

to fold detailed insect models. For example,many modern paper folders have developedtheir own insect bases rather than using tra-ditional bases. The need for new insect basesfurther indicates the difficulty of insectorigami based on traditional bases and tech-niques.

Challenging Insect Morphologyin Origami

In addition to the proper number of ap-pendages for intricate insect models, specialtechniques are needed to make the segmen-tation of the abdomen and legs. The mostcommon procedures in origami, such as thevalley, mountain, and reverse folds (Engel1989), can be used to represent the segmen-tation of insects. The body segmentation and

Fig. 1. Evolution ofthree origamiinsects. Note thatthree origamiinsects wereoriginated from thesame base fold,the waterbombbase (seeasterisk). Thebutterfly (left) wasmodeled byYoshizawa(Jackson 1990),the fly (center)was modeled bythe Korean Jong IeJupgi Association(1988), and thedragonfly (right)was modeled byBarbour (2002).

Insect OrigamiInsect OrigamiInsect OrigamiInsect OrigamiInsect OrigamiInto the FoldInto the FoldInto the FoldInto the FoldInto the Fold

Yong-Lak Park and Jeff Bradshaw

Using the art of paper folding to stimulate an interest ininsect diversity and morphology.

Downloaded from https://academic.oup.com/ae/article-abstract/49/4/210/2474732by gueston 13 February 2018

AMERICAN ENTOMOLOGIST • Volume 49, Number 4 211

jointed legs and antennae described bySnodgrass (1935) and Chapman (1982) canbe modeled with mountain and valley folds(Fig. 3A) and reverse folds (Fig. 3B). Themodalities of antennae and legs also can bemodeled with other origami techniques, suchas the petal, squash, and rabbit-ear folds.With careful planning, more evolutionarilyderived structures that represent morpho-logical characteristics (i.e., “key” characters)for families or species of insects can be folded(Fig. 4). Knowledge of characters based ontaxonomic keys can be used when choosingor developing the base fold for an insectmodel from a square sheet of paper. For ex-ample, an origami model with a furca andcollophore represents a springtail(Collembola, Fig. 4A), a model with threecerci and a silver body represents a silverfish(Thysanura, Fig. 4B), and a model with rap-torial forelegs and a long pronotum repre-sents a praying mantis (Mantodea, Fig. 4C).

What Insects Have Been Folded?In the past, most insect origami models

were common insects, such as butterflies,dragonflies, grasshoppers, and beetles. How-ever, many insect models have been foldedthat can be identified to family (Table 1). Byorigami, 46 families in 20 orders ofHexapoda have been represented. Beetles arethe most abundant (at least 11 familiesfolded) and popular in the origami insectworld. Perhaps, in part, these origami taxarepresent the insects in the natural world thatare most obvious and abundant to the art-ist. Some examples of these delicate origamiinsects are shown in Fig. 5.

Origami for Entomological EducationThe impact of origami has expanded well

beyond art and craft. Engineers use origamito help them develop new structures (Boursin2000) or model complex molecules (Cucciaet al. 1995); mathematicians use origami tostudy geometry (Hull 1995); occupationaltherapists use origami for the rehabilitationof hands and fingers (Smith 1992, Cornelius1995, Boursin 2000); and teachers use

origami as an educational tool (Smith 1992,Cornelius 1995). Adding an entomologicaltwist to education with origami could im-prove a student’s understanding and appre-ciation of the natural world.

Origami is an interesting way to presentinsect morphology, embryology, and system-atics in a geometric manner. For example,origami may be used to help visualize theimportance of cuticular infolding or segmen-tation of insect bodies that cannot be ex-plained easily with diagrams in the classroom.Embryonic development of insects also couldbe better conceptualized in the classroom byillustrating cellular determination and dif-ferentiation with origami. Regions on a planeof paper are selected (Fig. 2) to form bodyparts (like developmental determination),and then creases and folds are made thatallow each region of paper to form morespecialized structures (like developmental dif-ferentiation). With origami, students couldunderstand the structure and function of

Fig. 2. Planninginsect origami withthe blintzed birdbase. It has thepotential for foldingpaper insects withthe correct numberof appendages.

Fig. 3. A reverse fold (A) creates jointed legsand antennae, and multiple mountain andvalley folds (B) generate the segmentation ofthe insect body of a termite (Isoptera:Termitidae) designed by Park (2003).

Fig. 4. Origami insects representing keycharacters of orders and families: (A)Poduridae (Collembola); (B) Lepismatidae(Thysanura); (C) Mantidae (Mantodea).Models were designed by Park (2003).



segments and appendages more easily (Fig.3) and learn about evolution and speciationby creating new insect models from the samebase fold (Fig. 1).

Insect Origami for the MassesAlthough origami has been very popu-

lar in the large cities where origami organi-zations are active, it is much less common insmall U.S. cities and towns. The first authorintroduced origami to the public in Ames,Iowa, through origami demonstrations,classroom activities, and art exhibits. Myinterest (the first author) in origami goesback to elementary school, when teachersin South Korea folded paper to demonstrategeometry and mathematics. Since arrivingin Iowa in 1999 for graduate school, I havefolded paper insects not only because I aman entomologist, but also because insectmodels are known to be the most difficultorigami. Each of these insects (Figs. 3 and4) took 2 to 3 months to create from de-signing to folding. My favorite insects tofold are microscopic ones, such asproturans, collembolans, and diplurans,because most of these have not been foldedyet and are not familiar to the public.I am now trying to fold the gladiator,Mantophasmatodea, which is scientificallyimportant, being the first new insect orderto be described since 1915 (Trivedi 2002),but it is not familiar to the public.

My interest (the second author) wasgained through Park’s efforts at Iowa StateUniversity. Besides a general awe for insectsmodeled from papers, I see great potentialto educate the public about entomologythrough origami. What better way to teachpeople who declare real insects to be “icky”than to use paper insects? We organized twosuccessful insect origami art shows in 2003with the ambitious help of the EntomologyGraduate Student Organization at IowaState University. The purpose of the exhib-its was cultural exchange by using insectorigami.

One exhibit was for Iowa StateUniversity’s annual VEISHEA (an acronymfor the colleges of Veterinary Medicine, En-gineering, Industrial Science, Home Eco-nomics, and Agriculture) festival, “thelargest student-run, alcohol-free celebra-tion of entertainment and education in thenation” (VEISHEA Inc. 2003). At this ex-hibit, we displayed insect behavior, system-atics, taxonomy, and evolutionaryprinciples, and the first author gave a how-to presentation in a multimedia classroomfor many visitors.

Because of the popularity of the exhibitat VEISHEA, we were invited to display ourexhibit at Reiman Gardens, Ames, Iowa(Reiman Gardens 2003). The exhibit, “A

Table 1. Phylogeny (Borror et al. 1989) of origami insects

Subclass Order Suborder Family References

Apterygota Protura Acerentomidae Park (2003)Collembola Poduridae Park (2003)Diplura Campodeidae Park (2003)Thysanura Lepismatidae Kakitsuka (2003), Park (2003)

Pterygota Odonata Anisoptera Aeshnidae Park (2003)Libellulidae Lang (1995), Kakitsuka

(2003), Park (2003)Orthoptera Caelifera Acrididae Lang (1995), Kakitsuka

(2003), Park (2003)Ensifera Gryllidae Kakitsuka (2003),

Park (2003)Mantodea Mantidae Lang (1995), Kakitsuka (2003),

Park (2003)Blattaria Blattellidae Lang (2002),

Kakitsuka (2003)Isoptera Termitidae Kakitsuka (2003),

Park (2003)Dermaptera Labiidae Kakitsuka (2003),

Park (2003)Phthiraptera Anoplura Pediculidae Park (2003)

Pthiridae Park (2003)Hemiptera Gerromorpha Gerridae Kakitsuka (2003)

Nepomorpha Nepidae Kakitsuka (2003)Belostomatidae Kakitsuka (2003)Notonectidae Kakitsuka (2003)

Pentatomorpha Pentatomidae Kakitsuka (2003),Park (2003)

Homoptera Auchenorrhyncha Cicadidae Kawahata (1994), Lang(1995), Kakitsuka (2003)

Membracidae Lang (1995)Cicadellidae Kakitsuka (2003)

Neuroptera Planipennia Chrysopidae Park (2003)Myrmeleontidae Kakitsuka (2003)

Coleoptera Polyphaga Staphylinidae Kakitsuka (2003)Lucanidae Lang (1995), Kakitsuka

(2003), Park (2003)Scarabaeidae Lang (1995), Kakitsuka

(2003), Park (2003)Buprestidae Kakitsuka (2003)Elateridae Park (2003)Erotylidae Payet (2002)Lampyridae Kakitsuka (2003), Park (2003)Coccinellidae Lang (1995), Kakitsuka

(2003)Cerambycidae Lang (1995), Nishikawa

(2002)Curculionidae

Kakitsuka (2003)Adephaga Dytiscidae Kakitsuka (2003)

Siphonaptera Pulicidae Kakitsuka (2003)Diptera Brachycera Tabanidae Park (2003)

Muscidae Kakitsuka (2003),Park (2003)

Calliphoridae Payet (2002)Trichoptera Integrepalpia Brachycentridae Park (2003)Lepidoptera Ditrysia Papilionidae Lang (1995),

Kakitsuka (2003), Park (2003)Nymphalidae Park (2003)Saturniidae Park (2003)Sphingidae Kakitsuka (2003)

Hymenoptera Apocrita Vespidae Lang (1995),Kakitsuka (2003)

Formicidae Lang (1995), Kakitsuka (2003)

Note: references listed are those that contain the most recently and frequently modeled representatives for theassociated family.



Paper Bug’s Life” (Park 2003), included 33insects created by the first author, along with1,800 paper butterflies and 550 paperdragonflies created by members of the En-tomology Graduate Student Organizationand their families (Figs. 6 and 7). The suc-cess and potential impact of these exhibitswas amazing. The exhibits led to six inter-views by various media (newspapers, T.V.,and university communications). The ex-hibit at Reiman Gardens, originally sched-uled for 2 weeks, was extended for anotherweek halfway through the show, duringwhich approximately 4,600 people visitedthe gardens.

Tips for Insect OrigamiIf you want to try to fold some insect

models, we suggest several tips for success-ful folding of detailed insects:(1) Start with simple models.(2) Exact folding is necessary because paper fold-

ing is based on geometry, and errors in theearly folds of complex models are amplifiedas the folding proceeds.

(3) Use big, thin, crisp, and strong papers; insectorigami requires many overlapping folds, soyou need paper that can be folded throughseveral overlapping layers. You can cut wrap-ping paper into squares to fold, or you canbuy origami paper from hobby stores or mailorder.

(4) Be patient, because learning insect origamimay take a long time.

Fig. 5. Origamiinsects: (A)

Pentatomidae(Hemiptera), dorsal;

(B) Pentatomidae(Hemiptera),

ventral; (C)Acrididae

(Orthoptera); (D)larvae of

Brachycentridae(Trichoptera); (E)

larva ofScarabaeidae

(Coleoptera); (F)pupae of

Papilionidae(Lepidoptera); (G)

Aeshnidae(Odonata), dorsal;

(H) Aeshnidae(Odonata), ventral;

(I) Danaidae, female(Lepidoptera).Models were

designed by Park(2003).

Fig. 6. Origamiart by studentsat Iowa StateUniversity:Butterfly (top),Convergence(bottom left), andCircling (bottomright). Thebutterfly models,as diagrammedby Yoshizawa(Jackson 1990),were folded byentomologystudents andtheir families andartisticallyarranged by PattiAnderson, JeffBradshaw, EmilyGartrell, andKate Kronback,(Department ofEntomology,Iowa StateUniversity,Ames).



Fig. 7. Origami(microlepidopterans)by entomologystudents at IowaState University,as diagrammedby Yoshizawa(Jackson 1990).The flower(upper leftcorner) wasfolded bySeon-Hye Park.

(5) An advantage of origami is that paper canbe unfolded, so if you make a mistake, youstart again.

(6) The best way to understand an origamimodel is to fold it and then completely un-fold it to investigate important creases onthe paper.

With practice, you may develop new pa-per insects by modifying the original models.

Suggested ReadingInsect origami books with diagrams.Lang, R. J. 1995. Origami Insects and

Their Kin. General Publishing Co., Toronto,Ontario.

Lang, R. J. 2003. Origami Insects II. Gal-lery Origami House, Tokyo (in English andJapanese).

Photo clips for folding Yoshizawa’s but-terfly (Jackson 1990).

Park, Y.-L. 2003. A Paper Bug’s Life. http://paper-insect.uu.st.

AcknowledgementWe thank Marlin Rice, Robert Lang, and

Julie Todd for reviewing this manuscript.



References CitedBarbour, A. F. 2002. Anita’s origami. http://

www.ulster.net/~spider/dragfly.htm.Borror, D. J., C. A. Triplehorn, and N. F. Norman.

1989. An introduction to the study of in-sects, 6th ed. Sanders College Publishing, FortWorth, TX.

Boursin, D. 2000. Advanced origami. FireflyBooks, Willowdale, Canada.

Chapman, R. F. 1982. The insects: structure andfunction. Harvard University Press, Cam-bridge, MA.

Chiyo, A. 1982. Origami for Christmas.Kodansha International, Tokyo.

Cornelius, V. 1995. Using origami to support abasic education, pp 125–209. In V. Cornelius[Ed.], Proceedings of the Second InternationalConference on Origami in Education andTherapy. Conference on Origami in Educa-tion and Therapy, 1995, New York, NY.Origami USA, New York.

Cuccia, L. A., R. B. Lennox, and F. M. Y.Ow. 1995. Molecular modeling offullerenes with modular origami, pp. 21–30. In Cornelius, V. [Ed.], Proceedings ofthe Second International Conference onOrigami in Education and Therapy. Con-ference on Origami in Education andTherapy, 1995, New York, NY. OrigamiUSA, New York.

Engel, P. 1989. Origami from angelfish to Zen.Dover Publications, New York.

Southland Products, Inc.

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Harbin, R. 1997. Secrets of origami. Dover Pub-lications, New York.

Honda, I. 1983. The world of origami. JapanPublications Trading Co., Tokyo.

Hull, T. 1995. Geometric constructions viaorigami, pp. 31–38. In V. Cornelius [Ed.],Proceedings of the Second International Con-ference on Origami in Education andTherapy. Conference on Origami in Educa-tion and Therapy, 1995, New York, NY.Origami USA, New York.

Jackson, P. 1990. Classic origami. Apple Press,London, UK.

Jackson, P. 1991. Festive Folding. Quintet Pub-lishing Ltd., London, UK.

Kakitsuka, T. 2003. Origami insect models. http://member.nifty.ne. jp/kakitsuka/gal lery/insects_e.html.

Kawahata, F. 1994. The insect war. http://www.origami.gr.jp/Model/Insect/index-e.html.

Korean Jong Ie Jupgi Association. 1988. Mar-velous origami. Jong Ie Nara Publishing,Seoul.

Lang, R. J. 1995. Origami insects and their kin.General Publishing Co., Toronto Ontario.

Lang, R. J. 2002. Origami alive; insect gallery.h t t p : / / t e c h n o s t e r o n e . c o m / o r i g a m i /model.jsp?key=024.

Mulatinho, P. 1995. Origami. Chartwell Books,Secaucus, NJ.

Nishkawa, S. 2002. Origami alive; insect gal-

lery. http://technosterone.com/origami/.Park, Y.-L. 2003. A paper bug’s life. http://

paper-insect.uu.st.Payet, G. 2002. Origami alive; insect gallery.

http://technosterone.com/origami/.Reiman Gardens. 2003. Reiman gardens, Iowa

State University, Ames. http://www.reimangardens.iastate.edu/

Smith, J. 1992. Proceedings of the first interna-tional conference on origami in educationand therapy, 1991, Birmingham, England.British Origami Society, Birmingham.

Snodgrass, R. E. 1935. Principles of insect mor-phology, McGraw-Hill, New York.

Trivedi, B. P. 2002. New insect order found insouthern Africa. National GeographicToday, 28 March 2002. http://news.nationalgeographic.com/news/2002/03/0328_0328_TVstickinsect.html.

VEISHEA Inc. 2003. Veishea 2003. Ames, IA.http://www.veishea.org/faq/faq.htm.

Note: All web sites were correct as of 26 Octo-ber 2003.

Yong-Lak Park earned his Ph.D. degree at Iowa

State University and currently he is a

postdoctoral scholar in the department of Ento-

mology at University of California, Riverside.

Jeff Bradshaw, is a graduate student co-ma-joring in entomology and plant pathologyat Iowa State University. 7


insect origami: into the fold

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