Mutillidae

Mutillidae sp/

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Hymenopterida
Ordo: Hymenoptera
Subordo: Apocrita
Superfamilia: Pompiloidea
Familia: Mutillidae
Subfamilia (8): Dasylabrinae – Mutillinae – Myrmillinae – Myrmosinae – Pseudophotopsinae – Rhopalomutillinae – Sphaeropthalminae – Ticoplinae

Genera: Aglaotilla – Allotropidia – Ancistrotilla – Bidecoloratilla – Bimaculatilla – Bischoffiella – Carinomyrmosa – Chilemutilla – Darditilla – Dasymutilla – Ephuta – Ephutomorpha – Frigitilla – Gogoltilla – Indratilla – Invreiella – Lynchiatilla – Macromyrme – Mutilla – Nuristanilla – Odontomutilla – Odontophotopsis – Orientidia – Pherotilla – Promecidia – Pseudomethoca – Rasnitsynitilla – Rimulotilla – Sinotilla – Sphaeropthalma – Spinulomutilla – Timulla – Tobantilla – Tricholabiodes – Xystromutilla – Zavatilla – Zeugomutilla
Name

Mutillidae Latreille, 1802
References

Latreille, P.A. 1802. Histoire naturelle, générale et particulière des crustacés et des insectes. Ouvrage faisant suite à l’histoire naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par C.S. Sonnini, membre de plusieurs sociétés savantes. Familles naturelles des genres. Tome troisième. F. Dufart, Paris, xii + pp. 13–467 + [1 (errata)]. BHL Reference page.
Barthélémy, C., Lelej, A.S. & Loktionov, V.M. 2018. A contribution to the Mutillidae (Hymenoptera) of Hong Kong. Zootaxa 4514(2): 230–242. DOI: 10.11646/zootaxa.4514.2.6 Paywall Reference page.
Bartholomay, P.R., Waldren, G.C. & de Oliveira, M.L. 2017. Observation of a mixed-sex, mixed-species aggregation of velvet ants, genus Timulla Ashmead, 1899 (Hymenoptera: Mutillidae) in the Brazilian Amazon, Roraima, with a new synonymy. Zootaxa 4362(1): 135–140. DOI: 10.11646/zootaxa.4362.1.8 Reference page.
Bartholomay, P.R., Williams, K.A., Luz, D.R. & Morato, E.F. 2015: Frigitilla gen. nov., a new genus of Amazonian Mutillidae (Hymenoptera). Zootaxa 3957(1): 49–58. DOI: 10.11646/zootaxa.3957.1.3. Preview (PDF) Reference page.
Brothers, D.J. 2015. Revision of the Rhopalomutillinae (Hymenoptera, Mutillidae): 1, generic review with descriptions of three new genera. Journal of Hymenoptera Research 46: 1–24. DOI: 10.3897/JHR.46.5733. Reference page.
Brothers, D.J. 2018. Aglaotilla, a new genus of Australian Mutillidae (Hymenoptera) with metallic coloration. Zootaxa 4415(2): 357–368. DOI: 10.11646/zootaxa.4415.2.6 Paywall Reference page.
Brothers, D.J. & Lelej, A.S. 2017. Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research 60: 1–97. DOI: 10.3897/jhr.60.20091 Reference page.
Cambra, R.A., Brothers, D.J. & Quintero, D. 2016. Review of Frigitilla Williams in Bartholomay et al., 2015, with description of a new species from Panama (Hymenoptera: Mutillidae). Zootaxa 4189(2): 395–400. DOI: 10.11646/zootaxa.4189.2.13. Reference page.
Cambra, R.A., Buschini, M.L.T., Arias, D.Q., Brozoski, F. & Lustosa, P.R. 2017. Ephuta icema Casal, 1969 and its host Auplopus subaurarius Dreisbach, 1963 (Hymenoptera: Mutillidae, Pompilidae) from Brazil. Zootaxa 4272(2): 285–290. DOI: 10.11646/zootaxa.4272.2.9. Reference page.
Cambra, R.A., Queiros, C.N., de Deus, J.P.A., Wiliams, K.A., Bartholomay, P.R., Iantas, J., Nether, M.C., Nishida, K., Añino, Y.J., Saavedra, D. & Buschini, M.L.T. 2021. Description of the male of Xystromutilla bucki Suárez, 1960 (Hymenoptera, Mutillidae), including new information on the biology of the genus. ZooKeys, 1011: 73–84. DOI: 10.3897/zookeys.1011.60944 Open access Reference page.
Das, D., Kumar, G.P. & Bhadra, K. 2017. Description of a new species of the genus Odontomutilla Ashmead (Hymenoptera: Mutillidae) from India. Zootaxa 4290(2): 373–376. DOI: 10.11646/zootaxa.4290.2.7. Reference page.
Lelej, A.S.; Brothers, D.J. 2008: The genus-group names of Mutillidae (Hymenoptera) and their type species, with a new genus, new name, new synonymies, new combinations and lectotypifications. Zootaxa 1889: 1–79. Abstract & excerpt Reference page.
Lelej, A.S., Williams, K.A., Loktionov, V.M., Pang, H. & Xu, Z-F. 2017. Review of the genus Zeugomutilla Chen, 1957 (Hymenoptera, Mutillidae, Mutillini), with description of two new species. Zootaxa 4247(1): 1–15. DOI: 10.11646/zootaxa.4247.1.1. Reference page.
Soliman, A.M. 2016. A new species of Tricholabiodes Radoszkowski, 1885 (Hymenoptera: Mutillidae) from Saudi Arabia. Zootaxa 4067(1): 95–100. DOI: 10.11646/zootaxa.4067.1.8. Reference page.
Zhou, H-T., Lelej, A.S., Williams, K.A. & Liu, J-X. 2018. Revision of the Oriental genus Zavatilla Tsuneki (Hymenoptera, Mutillidae, Trogaspidiini), with descriptions of two new species. Zootaxa 4418(2): 101–120. DOI: 10.11646/zootaxa.4418.2.1 Paywall Reference page.

Additional references

Bartholomay, P.R., Williams, K.A., Cambra, R.A. & de Oliveira, M.L. 2019: Does the genus Dasymutilla Ashmead occur in South America? The new genus Quwitilla, new combinations, and new distribution records from Neotropical velvet ants (Hymenoptera: Mutillidae) Zootaxa 4623(2): 261–282. DOI: 10.11646/zootaxa.4623.2.3. Reference page.
Boehme, N.F., Tanner, D.A., Williams, K.A. & Pitts, J.P. 2012: Faunal study of velvet ants (Hymenoptera: Mutillidae) and their activity patterns and habitat preference at Ash Meadows National Wildlife Refuge, Nye County, Nevada, USA. Zootaxa 3587: 1–45. Preview reference page
Brabant, C.M.; Williams, K.A.; Pitts, J.P. 2010: True females of the subgenus Photomorphina Schuster (Hymenoptera: Mutillidae). Zootaxa, 2559: 58–68. Preview
El-Torkey, A.M. et al. 2011: Review of the Palaearctic species of Macromyrme Lelej 1984 (Hymenoptera: Mutillidae) with description of a new species from Egypt. Zootaxa, 2863: 63–68. Preview
El-Torkey, A. & Soliman, A.M. 2015. Discovery of the genus Vanhartenidia Lelej, 2006 (Hymenoptera: Mutillidae) in Egypt, with description of a new species. Zootaxa 3999(2): 298–300. DOI: 10.11646/zootaxa.3999.2.10. Preview (PDF) Reference page.
Lelej, A.S. 1995: A review of Palaearctic and Oriental species of the genus Nemka Lelej with description of Oriental genus Mickelomyrme gen. n. (Hymenoptera, Mutillidae). Far Eastern entomologist, 6: 1–20. PDF
Lelej, A.S. 1995: To the knowledge of East Asian species of the tribe Smicromyrmini Bischoff, 1920 (Hymenoptera, Mutilidae) with description of four new genera and eight new species. Far Eastern entomologist 13: 1–28. PDF
Lelej, A.S. 1995: To the knowledge of the velvet ants (Hymenoptera, Mutillidae) of Rajasthan, Western India. Far Eastern entomologist, 20: 1–11. PDF
Lelej, A.S. 1996: To the knowledge of the East Asian species of the tribe Trogaspidiini Bischoff, 1920 (Hymenoptera, Mutillidae) with description of eight new genera and two new species. Far Eastern entomologist, 30: 1–24. PDF
Lelej, A.S. & K.V. Krombein, 1999: Two remarkable new genera of mutillid wasps (Hymenoptera: Mutillidae, Sphaeroptalminae, Pseudomethocini) from Thailand. Far Eastern entomologist, 79: 1–8. PDF
Lelej, A.S. & P.G. Nemkow, 1997: Phylogeny, evolution and classification of Mutillidae (Hymenoptera). Far Eastern entomologist, 46: 1–24. PDF
Lelej, A.S. & Harten, van A., 2014: A new species of the genus Physetopoda Schuster 1949 (Hymenoptera, Mutillidae) from the United Arab Emirates. Zoologicheskii zhurnal, 93 (1): 205-?
Lelej, A.S.; Yildirim, E. 2009: A review of the Mutillidae (Hymenoptera) of Turkey. Zootaxa, 2160: 1–28. Abstract & excerpt
Lelej, A.S., Zhou, H-T., Loktionov, V.M. & Xu, Z-F. 2016. Review of the genus Promecidia Lelej, 1996, with description of two new species from China (Hymenoptera, Mutillidae, Trogaspidiini). ZooKeys 641: 103–120. DOI: 10.3897/zookeys.641.10765. Reference page.
Ljubomirov, T.; Ghahari, H. 2012: An annotated checklist of Mutillidae (Insecta: Hymenoptera) from Iran. Zootaxa 3449: 1–25. Preview Reference page.
Pitts, J.P.; Wilson, J.S.; Williams, K.A.; Boehme, N.F. 2009: Velvet ants (Hymenoptera: Mutillidae) of the Algodones sand dunes of California, USA. Zootaxa, 2131: 1–53. Abstract & excerpt
Pitts, J.P.; Wilson, J.S.; Williams, K.A.; Boehme, N.F. 2010: Nocturnal velvet ant males (Hymenoptera: Mutillidae) of Deep Canyon, California including four new species and a fifth new species from Owens Lake Valley, California. Zootaxa, 2553: 1–34. Preview PDF
Tu, B-B., Lelej, A.S. & Chen, X-X. 2015. Review of the genus Taiwanomyrme Tsuneki, 1993 (Hymenoptera, Mutillidae, Mutillinae), with description of two new species from China. Zootaxa 4020(3): 588–600. DOI: 10.11646/zootaxa.4020.3.10.Preview (PDF) Reference page.
Turrisi, G.F., Palmerini, M.M. & Brothers, D.J. 2015. Systematic revision and phylogeny of the genera Blakeius Ashmead, 1903 and Liomutilla André, 1907, with description of two new genera (Hymenoptera: Mutillidae, Myrmillinae). Zootaxa 4010(1): 1–78. DOI: 10.11646/zootaxa.4010.1.1. Preview (PDF); ISBN 978-1-77557-781-2 (paperback); ISBN 978-1-77557-782-9 (Online edition) Reference page.
Zhou, H-T., Lelej, A.S. & Xu, Z-F. 2017. Discovery of the subfamily Rhopalomutillinae (Hymenoptera: Mutillidae) in China and description of Pherotilla brothersi sp. nov. Zootaxa 4273(4): 597–594. DOI: 10.11646/zootaxa.4273.4.8. Reference page.

Links

Mutillidae – Taxon details on National Center for Biotechnology Information (NCBI).

Vernacular names
Deutsch: Ameisenwespen
English: Velvet ants
日本語: アリバチ
norsk: Maurvepser
Diné bizaad: Naʼazǫ́ǫzii
polski: Żronkowate
русский: Осы-немки
svenska: Spindelsteklar
Türkçe: Karınca arıları

The Mutillidae are a family of more than 7,000 species of wasps whose wingless females resemble large, hairy ants. Their common name velvet ant refers to their dense pile of hair, which most often is bright scarlet or orange, but may also be black, white, silver, or gold. Their bright colors serve as aposematic signals. They are known for their extremely painful stings, (the sting of the species Dasymutilla klugii rated a 3 on the Schmidt pain index and lasts up to 30 minutes[1]), and has resulted in the common name "cow killer" or "cow ant" being applied to the species Dasymutilla occidentalis.[2] However, mutillids are not aggressive and sting only in defense. In addition, the actual toxicity of their venom is much lower than that of honey bees or harvester ants.[3] Unlike true ants, they are solitary, and lack complex social systems.[4]

Distribution

Mutillidae can be found worldwide with approximately 230 genera or subgenera and around 8,000 species worldwide. Over 400 species occur in the North American southwest.[5]

North American Mutillidae have eight phenotypically distinct and geographically limited Müllerian mimicry rings (Desert, Eastern, Madrean, Texan, Red-headed Timulla, Black-headed Timulla, Tropical, and Western) making up one of the largest Müllerian mimicry complexes on the planet.[6] These mimicry rings are the result of repeated convergent evolution of aposematic traits between co-occurring velvet ant species, rather than shared, phylogenetic history.[7] Through the evolution of aposematic traits in velvet ant species in the same ring, local predators have learned to avoid this well-defended species.
Description

The exoskeleton of all velvet ants is unusually tough (to the point that some entomologists have reported difficulty piercing them with steel pins when attempting to mount them for display in cabinets). This characteristic allows them to successfully invade the nests of their prey and also helps them retain moisture. Mutillids exhibit extreme sexual dimorphism. Like some related families in the Vespoidea, males have wings, but females are wingless. The males and females are so distinct in their morphology that entomologists often find it very hard to determine whether a given male and female belong to the same species, unless they are captured while mating.[8] In some species, the male carries the smaller female aloft while mating, which is also seen in the related family Thynnidae.
A female eastern velvet ant

As is the case for all aculeates, only female mutillids are capable of inflicting a sting. The stinger is a modified female organ called an ovipositor, which is unusually long and maneuverable in mutillids. In both sexes, a structure called a stridulitrum on the metasoma is used to produce a squeaking or chirping sound when alarmed. Both sexes of mutillids also bear hair-lined grooves on the side of the metasoma called felt lines. Only two other vespoid families, (Bradynobaenidae and Chyphotidae) have felt lines, but the females of these families have a distinct pronotum, with a transverse suture separating it from the mesonotum; in female mutillids, these two thoracic segments are completely fused. Members of the family Myrmosidae, formerly classified as a subfamily of mutillids, also have a distinct pronotum in females, but lack felt lines in both sexes.
Behavior

Adult mutillids feed on nectar. Although some species are strictly nocturnal, females are often active during the day. Females of Tricholabiodes thisbe are sometimes active up to two hours before sunset. Guido Nonveiller (1963) hypothesized the Mutillidae are generally stenothermic and thermophilic; they may not avoid light, but rather are active during temperatures that usually occur only after sunset.
Defense mechanisms

Predation is one of the strongest forces natural selection uses to drive the evolution of an organism’s morphology, physiology, and behavior.[9][10][11] During this coevolution, the prey either being consumed by the predator or escaping has resulted in a plethora of impressive defensive strategies in prey species to improve the likelihood of escape. Velvet ants avoid predation using the following defense mechanisms; a venomous sting (if female), aposematic coloration, a stridulatory organ in their abdomen, an alarm secretion from their mandibular gland, and a durable exoskeleton. This array of defenses has contributed to the velvet ants being attributed the title of "the indestructible insect." This title was given to the velvet ants after experimental experiments of interactions between velvet ants and their potential predators that resulted in the survival of the ant and the ultimate avoidance by the predator. [12]

The venom that velvet ants inject through their stinger has an unknown composition. According to one researcher, the painfulness of the sting of Dasymutilla klugii outscored 58 other species of stinging insects tested; the only species this researcher rated as having a more painful sting were the Paraponera clavata (bullet ant), Synoeca septentrionalis (warrior wasp), Pepsis spp. and Hemipepsis spp. (tarantula hawks).[13] In an experimental setting, only two lizard species (one whiptail and one side-blotched lizard) attacked a velvet ant it was exposed to.[12] In both cases the velvet ants were exhibiting rapid lateral and vertical movements to ward off an attack. Once the attack occurred the velvet ants would immediately sting the lizards. This sting resulted in the dropping of the ants in both cases and avoidance for the remainder of the trial.[14] The side-blotched lizard was found dead in its tank 24 hours later.[12] The side-blotched lizard is a natural predator of velvet ants, while the whiptail is not.[12]

The aposematic coloration of velvet ants often corresponds to a specific Müllerian mimicry ring consisting of dozens of species. This offers protection because many local predators have learned to avoid prey with this same coloration.[6] To test the aposematic coloration on birds, mealworms were painted to resemble a velvet ant. During these trials, none of the painted mealworms were consumed, while all the control mealworms were consumed immediately.[12] However, the painted mealworms were attacked by the birds, but the birds immediately ceased the attack.[12] These experiments provide evidence that the aposematic coloration of velvet ants causes their predators to hesitate, acting as a visual defense mechanism.

The stridulatory organ that velvet ants possess produces an audible squeaking when the abdomen is contracted.[15] This mechanism is an auditory cue warning predators that are about to attack to stay away. In shrews, every time they would get within 1 meter of a velvet ant, the velvet ant would begin stridulating.[14] Stridulations became more frequent as the predator moved closer to the velvet ant, and the shrew never attempted to attack the velvet ant. However, different scenarios with shrews have shown that the velvet ant would also stridulate after the shrew attacked it. Every time this occurred the shrew dropped the wasp.[14]

The exoskeleton of the velvet ant is remarkably strong. When compared to the exoskeleton of a honeybee, the velvet ant’s required 11 times more force to crush using a force transducer.[15] As well as being durable, the exoskeleton is also round, making it more difficult for predators to pierce it with attempted stings or bites. During all the trials that led to the fracture of a velvet ant’s exoskeleton, a total of 4 times, resulted in the death of that velvet ant within 24 hours. Aside from protection from predators, the exoskeleton also serves a function in maintaining moisture control.[15]

Due to these strong defense mechanisms, local predators generally avoid the velvet ants, so it has been difficult to determine their predators. [12] One study found tropical/ subtropical iguanian lizards (dactyloidae) to be a local predator of velvet ants in the Black-headed Timulla and Tropical mimicry rings. [16]
Life cycle
A female of Nemka viduata viduata (Pallas, 1773) looks for a nest of Bembix oculata to deposit her eggs.

Male mutillids fly in search of females; after mating, the female enters a host insect nest, typically a ground-nesting bee or wasp burrow, and deposits one egg near each larva or pupa. Only a few species are known to parasitize other types of hosts;[17] exceptions include the European velvet ant, Mutilla europaea, one of the only species that attacks social bees (e.g., Bombus), and the genus Pappognatha, whose hosts are tree-dwelling orchid bees. The mutillid larvae then develop as idiobiont ectoparasitoids, eventually killing their immobile larval/pupal hosts within a week or two. Velvet ants exhibit haplodiploid sex determination, as do other members of the superfamily Vespoidea.
See also

List of Mutillidae genera

References

Evans, David L.; Schmidt, Justin O. (1990). "Hymenopteran venoms: striving toward the ultimate defense against vertebrates". Insect Defenses: Adaptive Mechanisms and Strategies of Prey and Predators. SUNY Press. pp. 387–419. ISBN 978-0-88706-896-6.
"Red velvet ant; cow killer". Arthropod Museum. University of Arkansas: Division of Agriculture. Archived from the original on May 26, 2012. Retrieved September 12, 2011.
Meyer, W.L. (1996). "Most Toxic Insect Venom". Book of Insect Records. University of Florida.
Hunt, J.H. (1999). "Trait mapping and salience in the evolution of eusocial vespid wasps" (PDF). Evolution. 53 (1): 225–237. doi:10.1111/j.1558-5646.1999.tb05348.x. PMID 28565172. S2CID 205781311.
"Mutillidae — velvet ants". Featured Creatures. University of Florida/IFAS.
Wilson, J.S.; Jahner, J.P.; Forister, M.L.; Sheehan, E.S.; Williams, K.A.; Pitts, J.P. (2015). "North American velvet ants form one of the world's largest known Mullerian mimicry complexes". Current Biology. 25 (16): R704-6. doi:10.1016/j.cub.2015.06.053. PMID 26294178.
Wilson, Joseph S.; Williams, Kevin A.; Forister, Matthew L.; von Dohlen, Carol D.; Pitts, James P. (2012-12-11). "Repeated evolution in overlapping mimicry rings among North American velvet ants". Nature Communications. 3 (1): 1272. doi:10.1038/ncomms2275. ISSN 2041-1723. PMID 23232402.
Goulet, Henri; Huber, John T. (1993). Hymenoptera of the world : an identification guide to families. Agriculture Canada. ISBN 978-0660149332. OCLC 28024976.
Lima, S.L.; Dill, L.M. (1990). "Behavioral decisions made under the risk of predation: A review and prospect". Canadian Journal of Zoology. 68 (4): 619–640. doi:10.1139/z90-092.
Lelej, A.S. (2002). Catalogue of the Mutillidae (Hymenoptera) of the Palaearctic region (PDF). Vladivostok: Dalnauka. ISBN 978-5-8044-0280-9.
Nonveiller, G.; Suarez, F.J. (1990). Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta: Hymenoptera. Hymenopterorum catalogus. Vol. 18. SPB Academic. ISBN 978-9051030488. OCLC 889162931.
Gall, Brian G.; Spivey, Kari L.; Chapman, Trevor L.; Delph, Robert J.; Brodie, Edmund D.; Wilson, Joseph S. (2018). "The indestructible insect: Velvet ants from across the United States avoid predation by representatives from all major tetrapod clades". Ecology and Evolution. 8 (11): 5852–5862. doi:10.1002/ece3.4123. ISSN 2045-7758. PMC 6010712. PMID 29938098.
Starr, C.K. (1985). "A simple pain scale for field comparison of hymenopteran stings". Journal of Entomological Science. 20 (2): 225–232. doi:10.18474/0749-8004-20.2.225.
Gall, B.G.; Spivey, K.L; Chapman, T.L.; Delph, R.J.; Brodie, E.D., Jr.; Wilson, J.S. (2018). "The indestructible insect: Velvet ants from across the United States avoid predation by representatives from all major tetrapod clades". Ecology and Evolution. 8 (11): 5852–5862. doi:10.1002/ece3.4123. PMC 6010712. PMID 29938098.
Schmidt, J.O.; Blum, M.S. (1977). "Adaptations and responses of Dasymutilla occidentalis (Hymenoptera: Mutillidae) to predators". Entomologia Experimentalis et Applicata. 21 (2): 99–111. doi:10.1111/j.1570-7458.1977.tb02663.x. S2CID 83847876.
Pan, Aaron D.; Williams, Kevin A.; Wilson, Joseph S. (2017-02-01). "Are diurnal iguanian lizards the evolutionary drivers of New World female velvet ant (Hymenoptera: Mutillidae) Müllerian mimicry rings?". Biological Journal of the Linnean Society. 120 (2): 436–447. doi:10.1111/bij.12894. ISSN 0024-4066.

Brothers, Denis J., "Phylogeny and classification of the aculeate Hymenoptera, with special reference to Mutillidae" (1975) University of Kansas Science Bulletin 50(11):483-648.

Sources

Brodie, E.D. Jr. (1968). "Investigations on the skin toxin of the adult rough-skinned newt, Taricha granulosa". Copeia. 1968 (2): 307–313. doi:10.2307/1441757. JSTOR 1441757.
Brodie, E.D. Jr.; Formanowicz, D.R. Jr.; Brodie, E.D. III (1991). "Predator avoidance and antipredator mechanisms: Distinct pathways to survival". Ethology Ecology & Evolution. 3 (1): 73–77. doi:10.1080/08927014.1991.9525390.
Endler, J.A. (1986). Natural selection in the wild. Princeton University Press.
Mickel, C.E. (1928). "Biological and taxonomic investigations on the mutillid wasps". United States National Museum Bulletin. 143 (143): 1–351. doi:10.5479/si.03629236.143.1.
Milne, Lorus J. (1980). National Audubon Society Field Guide to North American Insects and Spiders. Audubon Society Field Guide (Turtleback). Knopf. ISBN 978-0-394-50763-7.