Cicada Killer Bibliography


Introduction

What follows is a partially annotated bibliography of scientific literature referring to cicada killers (Sphecius). It is intended as a source for those interested in cicada killer biology. Those using military loans and student loans while studying entomology will find these references and resources to be quite helpful. Some students go into the military in order to afford their education, justmilitaryloans.com offers military loans for education in all fields, including entomology & biology. The research spans a time period of over one hundred years and covers topics ranging from basic natural history to molecular biology. The list may not be comprehensive, and some sources are not annotated because I have not yet obtained them. Annotations are mostly my own. Many thanks to Chuck Holliday for digging up some really obscure papers and providing commentary for them. If you know of a relevant reference that is not on this list, or find a significant error here, please The current number of references listed is 98.
If you want a bibliography of the genus Sphecius, there is a good one at the California Academy of Sciences site (pdf).

References

  1. Alcock, J. 1975. The behaviour of western cicada killer males, Sphecius grandis (Sphecidae, Hymenoptera).  Journal of Natural History 9:561-566.
    A comparison of male behavior between the western and eastern species.

  2. Allard, H.A. 1937. The periodical cicada. American Naturalist 71: 588-604.
    A good description of the biology of the periodical cicada. However, on p. 601 Allard states that cicada killers take "… their toll of the active [periodical cicada] adults." This is very unlikely because periodical cicadas emerge in May and June and mate, lay their eggs and die by early July, while female cicada killers do not usually emerge until mid July. Only two instances of cicada killers taking a total of three Magicicada have been reported.

  3. Arnett, R.H. and R.L. Jacques, Jr. 1981. Simon & Schuster's guide to insects. Simon & Schuster, New York, 511 pp.
    Normally, a field guide would not be included here, but this one is notable for its errors. The entry claims that cicada killers are known for their painful sting, and the wasp in the photo provided looks more like Sphecius grandis than S. speciosus, which is the name provided.

  4. Balduf, W.V. 1941. Take offs by prey-laden wasps (Hymen.: Pompilidae, Sphecidae). Entomological News 52:91-92.
    Contains an incredible account of a loaded female crawling up the author's body and taking off from the top of his head. Balduf also reports that a female cicada killer can probably carry a paralyzed cicada weighing four to six times her own weight. This statement needs to be taken with a grain of salt because the female wasp he weighed tipped the scale at only 356 mg and was, thus, very small (an average female cicada killer weighs about 950 mg). Further, she was not carrying a cicada when she was caught.

  5. Berenbaum, M.R. 1989. Ninety-nine gnats, nits and nibblers. University of Illinois Press, Urbana. 254 pp.
    A brief, entertaining summary of the life of female cicada killers is found on pages 143-144.

  6. Bohart, R. M. 2000. A review of the Gorytini in the neotropical region (Hymenoptera: Sphecidae: Bembicinae). Contributions on Entomology International 4(2):111-259.
    Contains a taxonomic key for the New World species of Sphecius on p. 172. Note that S. convallis and S. grandis are mistakenly reversed in this key. Also mentions "sham" stinging by males in several genera, including Sphecius (p. 114), and the fact that Sphecius spectabilis and Ligorytes joergenseni (both in South America) are known to provision their nests with cicadas. He also notes that the Australian wasps, Exeirus lateralis and Austrogorytes bellicosus (p. 116), as well as the South American wasps, Liogorytes rufulus and L. brasilicus (p. 208), may also provision their nests with cicadas.

  7. Boothby, M.T., S. Griffith, and J.R. Coelho. 2002. Wasp Roundup. Natural History 111(7):12.
    Two letters to the editor regarding Coelho's article in Natural History, and his reply.

  8. Bringer, S.L. 1996. Effects of envenomation by cicada killers (Sphecius speciosus) on longevity and physiology of annual cicadas (Tibicen spp.). M.S. Thesis, Western Illinois University.
    Stung cicadas live longer than unstung controls, apparently because of lower rates of water loss, not metabolism.

  9. Burroughs, J. 1881. Pepacton. Houghton, Mifflin and Company, Boston.
    On pages 135-139 a section entitled "The Sand Hornet" describes a great deal of cicada killer biology, including the fact that prey exceeds predator in body mass, and the presence of spurs on the female. Errors are present (and expected) in this, the earliest reference to Sphecius we have encountered.

  10. Cane, J. 1979. The hind tibiotarsal and tibial spur articulations in bees (Hymenoptera: Apoidea). Journal of the Kansas Entomological Society 52:123-137.
    The hind-tibial spurs in female eastern cicada killers arise from a small corner of the distal end of the hind tibia.

  11. Casteels, P., J. Romagnolo, M. Castle, K. Casteels-Josson, H. Erdjument-Bromage, and P. Tempst. 1994. Biodiversity of apidaecin-type peptide antibiotics. Journal of Biological Chemistry 269:26107-26115.
    Cicada killers injected with bacteria "produced a massive quantity of apidaecin (two different isoforms) but no significant levels of any other peptides."

  12. Champlain, A.B. 1929. Hunters of the treetops wherein the Cicada-killers do their stuff. Nature Magazine 14:176 (September).
    A florid popular account of the eastern cicada killer with several mistakes, chief among them that female cicada killers hunt male cicadas by homing in on their mating calls.

  13. Clark, A.H. 1937. Potent personalities--wasps and hornets. National Geographic Magazine 72:47-72.
    A nice, mostly accurate, popular treatment of a variety of wasps, including cicada killers. Has good descriptions of Sphecius drinking sap, and some color illustrations.

  14. Coelho, J.R. 1997. Sexual size dimorphism and flight behavior in cicada killers (Sphecius speciosus). Oikos 79:371-375.
    Females are twice the size of males because they carry cicadas in flight. Males have the same maneuverability (flight muscle ratio) as females, and can be produced at half the cost. About 10% of females carry cicadas that are larger than they should be able to carry; these females compensate by carrying their prey up vertical objects on foot and flying toward their burrows in descent.

  15. Coelho, J. R. 1998. An acoustical and physiological analysis of buzzing in cicada killer wasps (Sphecius speciosus). Journal of Comparative Physiology A 183 (6): 745-751.
    A loud (72 dB at 3 cm) defensive buzz is produced with many harmonics and occasional buzz pulses. Producing the buzz causes a rapid increase in thorax temperature.

  16. Coelho, J.R. 2001. Behavioral and physiological thermoregulation in male cicada killers (Sphecius speciosus) during territorial behavior. Journal of Thermal Biology 26(2):109-116. | Full text available for purchase at Science Direct
    Males use a large suite of behavioral and physiological strategies to maintain a relatively constant thorax temperature while perching and patrolling their territories.

  17. Coelho, J.R. 2002. Spurred on to greater depths. Natural History 111(6):20-22.
    A popularized treatment of the Coelho and Wiedman (1999) paper below.

  18. Coelho, J.R., J.M. Hastings and C.W. Holliday. 2012. Near-optimal foraging in the Pacific cicada killer Sphecius convallis Patton (Hymenoptera: Crabronidae). Insects 3(1):133-140.
    This population appears to be evolved to such a body size that the single prey species is the ideal size for females to carry.

  19. Coelho, J.R., J.M. Hastings, C.W. Holliday, G. Flure and K. Barnes. 2008. Sexual dimorphism of the femora, tibiae and hind tibial spurs in the eastern cicada killer, Sphecius speciosus Drury (Hymenoptera: Crabronidae) in the United States. Entomological News 119(1):11-18..
    Females not only have disproportionately larger hind tibial spurs than males, they have more robust hind tibiae.

  20. Coelho, J. R., J.M. Hastings, C.W. Holliday and A. Mendell. 2008. Load carriage during foraging in two species of solitary wasps. Journal of Hymenoptera Research 17:57-63.
    A Bembix species takes flies that are much smaller than it could potentially carry, but avoidance of prey theft may be an adaptive benefit. A Tachytes species takes almost precisely the ideal size of prey on average, but variation in prey size is great.

  21. Coelho, J.R. and C.W. Holliday. 2000. Effects of size and flight performance on intermale mate competition in the cicada killer, Sphecius speciosus Drury (Hymenoptera: Sphecidae). Journal of Insect Behavior 14(3):345-351. Abstract
    Males that are successful in mating are larger and more maneuverable (higher flight muscle ratio) than their unsuccessful competitors.

  22. Coelho, J.R. and C.W. Holliday. 2008. The effect of hind-tibial spurs on digging rate in female eastern cicada killers. Ecological Entomology 33:1-5.
    Females with intact spurs dig twice as fast as those without them because their soil loads are twice the size. Spurless females would lose significant foraging time to digging, which may have provided directional selection for enlarged spurs.

  23. Coelho, J.R., C.W. Holliday, J.M. Hastings, E. Maty, M. Swigart and A. Mendell. 2007. Thermoregulation in male western cicada killers (Sphecius grandis Say) in the Chihuahuan desert. Journal of Thermal Biology 32(5):270-275. Abstract
    S. grandis males use some of the same behaviors as S. speciosus to maintain their body temperature, but do not perch on the ground, and leave their territories much earlier in the day because of the much higher ambient temperature.

  24. Coelho, J. R. and K. Wiedman.  1999.  Functional morphology of the hind tibial spurs of the cicada killer (Sphecius speciosus Drury).  Journal of Hymenoptera Research 8(1):6-12.
    The spurs act as one-way flaps and are used as shovels in digging, but cannot be used in prey carriage.

  25. Curran, C.H. 1962. Insects in your life, Sheridan House Publ., NY.
    An account of cicada killer biology begins on p. 123. The account has many inaccuracies (females hunt cicadas by homing on the males' mating calls, females provision nests with more than a dozen cicadas, etc.), but Curran does make an impassioned plea for not killing the wasps because they control cicadas, which damage the delicate new growth on trees.

  26. Dambach, C.A. and E. Good. 1943. Life history and habits of the cicada killer in Ohio. The Ohio Journal of Science 43:32-41.
    Covers virtually all aspects of cicada killer natural history. An essential primer. However, the account of how female wasps remove dirt from the burrows they are digging (top of p. 34) is incorrect (see Coelho & Wiedman, 1999).

  27. Davis, W.T. 1891. Notes of habits of the larger digger wasp. Canadian Entomologist pp. 10-11 (January).
    Contains three accounts of Sphecius (its genus was Stizus in 1891) carrying paralyzed cicadas up trees to gain altitude for flights back to their burrows.

  28. Davis, W.T. 1920. Mating habits of Sphecius speciosus, the cicada-killing wasp. Bulletin of the Brooklyn Entomological Society 15:128-129.
    Davis incorrectly states that mating in Sphecius "takes place in the air and is very brief." He also reports that cicada killers drag paralyzed cicadas up trees to gain altitude for the flight back to their burrows and that, on Long Island, Sphecius takes four species of cicadas: Tibicen sayi, T. linnei, T. lyricen and T. canicularis, often placing more than one species in the same burrow.

  29. Davis, W.T. 1924. Cicada killing wasps and flies. Journal of the New York Entomological Society 32:113.
    This reference does not belong here because it makes no mention of Sphecius - the wasps are Polistes pallipes and Vespa crabro and they were seen eating live cicadas on the ground. The flies were Asilids and one was seen to attack a cicada in mid-air. The title is misleading.

  30. Denton, S.B. 1931. Habits of the cicada-killer. Bulletin of the Brooklyn Entomological Society 26:35.
    A one-paragraph account of males chasing females and then returning to their perches on blackberry leaves and a female which was found in a burrow Denton dug up.

  31. Dow, R. 1942. The relation of the prey of Sphecius speciosus to the size and sex of the adult wasp (Hym.: Sphecidae). Annals of the Entomological Society of America 35:310-317.
    Two cicadas are used to provision a female larva, one for a male. This pattern is apparently the proximate cause of sexual size dimorphism, females being about twice the size of males.

  32. Dow, R. 1942. The relation between prey and sex in the cicada killer. Proceedings of the Entomological Society of Washington 44(6 - June):127-128.
    Cocoons found in excavated burrows are of two sizes and correspond to the two sexes, with female cocoons being larger than those of males. All cocoons which hatched into females were taken from nest cells containing the remains of two cicadas and all but one or two hatched males came from cells with the remains of only one cicada.

  33. Eason, P.K., G.A. Cobbs, & K.G.Trinca. 1999. The use of landmarks to define territorial boundaries. Animal Behaviour 58(1):85-91. Abstract
    The presence of structures in the environment facilitates the formation of territorial boundaries, minimizes agonistic encounters between males, and lowers the cost of territorial defense.

  34. Ebeling, W. 1975. Urban entomology. Univ. of California Press, Berkeley (also available online at http://www.entomology.ucr.edu/ebeling/ebel9-2.html).
    In Chapter nine, p. 21 of the online edition, Ebeling states that Sphecius speciosus will not sting unless handled and gives the U.S. distributions of cicada killer species as follows:

    • Sphecius speciosus - east of the Rocky Mountains and south into Mexico;

    • S. convallis - Kansas to California and into Baja California;

    • S. grandis - Tennessee, Missouri, Arkansas and throughout the West;

    • S. hogardii - southern tip of Florida and West Indes.

  35. Evans, H. E. 1966. The evolution of prey-carrying mechanisms in wasps. Evolution 16:468-483.
    Contains a brief discussion of pedal prey carriage and function of hind tibial spurs in females (pp. 475-476).

  36. Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard University Press, Cambridge, Mass.
    A thorough review of cicada killer biology, including information on three U.S. Sphecius species and several other cicada-killing sphecids of the world, including notes on their cicada prey. Excellent labeled diagrams of head, thorax, wings, and male genitalia in Chapter I, female tarsi, burrows and cicada in Chapter V.

  37. Evans, H.E. 1966. The behavior patterns of solitary wasps. Annual Review of Entomology 11: 123-154.
    Places many aspects of the behavior of Sphecius speciosus in context with that of other solitary wasps. There is much food for thought in this excellent review.

  38. Evans, H.E. and C.S. Lin. 1956. Studies on the larvae of digger wasps (Hymenoptera : Sphecidae) Part II: Nyssininae. Transactions of the American Entomological Society 82:35-66 and plates.
    A formal description of the larva of Sphecius speciosus (pp. 46-47) with figures (plate V).

  39. Evans, H.E. and K.M. O’Neill. The sand wasps: natural history and behavior. Harvard, Cambridge, Mass., 340 pp.
    A thorough review of cicada killer biology is included.

  40. Evans, H.E. and M.J. West Eberhard. 1970. The wasps. University of Michigan Press, Ann Arbor.
    Brief mention of Sphecius in comparison with other wasps, such as in digging behavior.

  41. Felt, E.P. 1906. Notes on the cicada-killer. New York State Museum Memoir 8:603.
    Felt mentions that cicada killers had been "so abundant at Poughkipsie as to occasion complaints because its numerous burrows rendered lawns unsightly."

  42. Fox, W.J. 1895. Synopsis of the Stizini of Boreal America. Proceedings of the Academy of Natural Sciences of Philadephia July 2:264-268.
    Includes a really old key to the species of Sphecius and gives the U.S. distributions of cicada killer species as follows:

    • Sphecius speciosus - common in the eastern U.S., Texas, Mexico;

    • S. convallis - Kansas, Texas, California and into Baja California;

    • S. grandis - Tennessee, Missouri, Arkansas, Texas, New Mexico, Utah, Nevada, Washington;

    • S. hogardii - Key West in Florida and the Bahamas.

    Fox also doubts the existence of S. fervidus (Handlirsch) as a species.
  43. Genaro, J.A. and C.J. de Varona, 1998. Comportamiento de Sphecius hogardii durante la nidification (Hymenoptera : Sphecidae). Caribbean Journal of Science 34:323-324.
    An account (in Spanish) of the nesting behavior of S. hogardii in Cuba; 63 burrows were observed. Their behavior closely resembles that of S. speciosus, except that they dig burrows in the floors of caves opening into a semi-deciduous forest in Caguanes National Park in Sancti Spiritus province. They use only Uhleroides walkeri cicadas to provision their nests and three nest cells that were excavated had 4, 4 and 6 cicadas in them. The authors took 66 cicadas from females returning to their burrows and female cicadas (44) outnumbered males (22). One cocoon was found and it resembled that of S. speciosus, with the exception that the pores in the middle of the cocoon were on only one side and did not form a ring around the cocoon, as in S. speciosus.

  44. Grant, P.R. 2006. Opportunistic predation and offspring sex ratios of cicada-killer wasps (Sphecius speciosus Drury). Ecological Entomology 31(5):539–547.
    Predation on cicadas appears to be opportunistic and non-selective; larvae excavated from nests were male-biased, close to expectation from Fisher’s equal-investment model. Sex of larvae was unrelated to the sex of the prey.

  45. Hastings, J. 1986. Provisioning by female western cicada killer wasps, Sphecius grandis (Hymenoptera:Sphecidae): influence of body size and emergence time on individual provisioning success. Journal of the Kansas Entomological Society 59:262-268.
    Larger females are more successful foragers and medium-sized females have great difficulty carrying cicadas back to their burrows. Small females have virtually no provisioning success. Females captured slightly more male cicadas than females, perhaps due to the males’ mobility and tendency to perch on exposed branches.

  46. Hastings, J. 1989. The influence of size, age, and residency status on territory defense in male western cicada killer wasps (Sphecius grandis, Hymenoptera: Sphecidae). Journal of the Kansas Entomological Society 62:363-373.
    Larger males have greater ability to hold territories and presumably greater mating success. Most males observed mating had been active on their territories when their mates emerged. Younger males had an advantage in taking over territories from older males. Small males' territories were usually on the periphery of the breeding aggregation and small males predominated among "satellite" males using non-territorial mating tactics.

  47. Hastings, J. 1989. Protandry in western cicada killer wasps, (Sphecius grandis, Hymenoptera:Sphecidae): an empirical study of emergence time and mating opportunity. Behavioral Ecology and Sociobiology 25:255-260.
    Males emerge before females, and male emergence date appears to be a variable which can be subject to natural selection because it is shown to affect the males' mating opportunities. Males have remarkably short lifespans (10-14 days).

  48. Hastings, J.M., J.R. Coelho, & C.W. Holliday. 2008. Mating at high population density in a colonial territorial wasp, Sphecius speciosus Drury (Hymenoptera: Crabronidae). Journal of the Kansas Entomological Society 81(3):301-308.
    The size advantage of large males in aquiring mates partly breaks down at high male density at the largest cicada killer aggregation yet documented.

  49. Hastings, J. M. , C.W. Holliday and J. R. Coelho. 2008. Body size relationship between cicada killers, Sphecius speciosus Drury (Hymenoptera: Crabronidae), and their prey: Does prey size determine wasp size? Florida Entomologist 91(4): 657-663.
    The size of available cicada prey is a good predictor of the sizes of male and female cicada killers.

  50. Hastings, J.M., C.W. Holliday, A. Long, K. Jones and G. Rodriguez. 2010. Size-specific provisioning by cicada killers, Sphecius speciosus, (Hymenoptera: Crabronidae) in North Florida. Florida Entomologist 93(3):412-421.
    Females appear to choose cicadas appropriate to their own size.

  51. Hastings, J.M., P. J. Schultheis, M.K. Whitson, C.W. Holliday, J.R. Coelho, and A.M. Mendell. 2008. DNA barcoding of New World cicada killers (Hymenoptera: Crabronidae). ZooTaxa 1713:27-28.
    COX-1 sequences show that haplotypes of Sphecius grandis Say fall into two highly divergent clades, suggesting they are cryptic species. Two clades of S. convallis Patton were found, and the sequences of one clade are virtually identical to those of S. speciosus Drury, suggesting that this clade of S. convallis is conspecific with S. speciosus.

  52. Holliday, C.W. and J.R. Coelho. 2006. Improved key to New World species of Sphecius (Hymenoptera: Crabronidae). Annals of the Entomological Society of America 99(5): 793-798. PDF Reprint
    A key that reverses Bohart's error and provides more definitive characters for distinguishing among the species. It also gives geographic ranges for the five New World species of Sphecius.

  53. Holliday, C.W., J.R. Coelho and J. M. Hastings. 2010. Conspecific kleptoparasitism in Pacific cicada killers, Sphecius convallis (Hymenoptera: Crabronidae): Partially provisioned nest cells are appropriated in situ by other females.
    Poster presented at the 2010 ESA meeting. PDF of poster here.

  54. Holliday, C.W., J. M. Hastings and J.R. Coelho. 2009. Cicada prey of New World cicada killers, Sphecius spp. (Hymenoptera: Crabronidae). Entomological News 120(1):1-17.
    Cicada killers were recorded taking a huge variety of cicadas, suggesting that they are primarily opportunistic foragers.

  55. Howes, P.G. 1919. Insect behavior. Gorham Press, Boston. 176 pp.
    Contains a brief discussion of the function of hind tibial spurs for prey carriage in females (pp. 136-137) and erroneously states that, "it does not seem probable that an insect possesses the power of distinguishing her eggs to the extent of telling which are to bring forth the males and which the females." (p. 93)

  56. Hungerford, H.B. and F.X. Williams. 1912. Entomological News and Proceedings of the Entomological Section, Academy of Natural Sciences, Philadelphia 23:241-260.
    Notes that Sphecous speciosus (then named Stizus speciosus) feeds on the exudations of willow trees and another wasp, Chlorion (a mud dauber), violently competes with it for the exudations.

  57. Joos, B., and T.M. Casey. 1992. Flight energetics of the cicada killer wasp. American Zoologist 32:54A [abstract].
    Provides data on oxygen consumption and wingbeat frequency in hovering flight.

  58. Krombein, K.V. 1952. Biological and taxonomic observations on the wasps in a coastal area of North Carolina (Hymenoptera : Aculeata). Wassman Journal of Biology 10: 257-341.
    Sphecius speciosus were common on barrens and were small; he captured 5 males and two females in late June and early July at Kill Devil Hills and all were smaller than average (p. 334).

  59. Krombein, K.V. 1953. Kill-Devil Hills wasps. Proceedings of the Entomological Society of Washington 55: 113-135.
    Sphecius speciosus were seen visiting two species of oak (Querqus marilandica and Q. virginiana).

  60. Krombein, K.A. 1979. Catalog of Hymenoptera in America north of Mexico, Volume 2. Smithsonian Institution Press, Washington.
    Lists Sphecius genera, their taxonomic histories, and geographic distributions, as follows:

    • Sphecius speciosus - U. and L. Austr. Zones east of Rockies, south into Mexico;

    • S. convallis - U.S. west of 100th meridian, Baja California;

    • S. grandis - Tennessee, Missouri, Arkansas, Texas, Kansas, New Mexico, Utah, Nevada, Washington, California south to Central America;

    • S. hogardii - South Florida, West Indes; another subspecies occurs in the Bahamas.

    Also lists the species' parasites on their cicada nest provisions and their cicada prey species and has a good bibliography for each species.
  61. Levetter, K.J., A.K. Majumdar and C.W. Holliday. 1994. The ecology of the cicada-killer wasp, Sphecius speciosus. In: R.D. Yearout (ed) Proceedings of the National Conference on Undergraduate Research VIII. The University of North Carolina, Asheville, pp.933-937.
    Describes attempts to rear cicada killers in the laboratory. Females spend the night in their burrows and do not leave until the sun shines upon them in the morning.

  62. Lin, N. 1963. Observations of suspected density dependent fighting between females of the cicada killer wasp Sphecius speciosus. Bulletin of the Brooklyn Entomological Society 58:121-123.
    Reports infrequent fights between female Sphecius speciosus and notes that they may be fighting over paralyzed cicadas.

  63. Lin, N. 1964. Weather and the natural regulation of three populations of the cicada killer wasp. Ph.D. Dissertation, University of Kansas, Lawrence.
    This thesis can be obtained by interlibrary loan, but it's really expensive. In this and most of his subsequent studies of Sphecius speciosus nesting aggregations on the Parade Grounds in Brooklyn, NY, Lin censuses male and female wasp emergence by counting their emergence hole frequency and diameter. He reports that larval survival (realized natality) varies directly with rainfall totals during the nesting season (months of July and August). Population density appeared to have no effect on larval survival.

  64. Lin, N. 1966. Weather and the natural regulation of three populations of the cicada killer wasp, Sphecius speciosus. American Zoologist 6(3):208 Abstract.
    In this abstract Lin reports the results of his Ph.D. thesis (see above).

  65. Lin, N. 1966. The establishment of a territory about the human observer by the cicada killer wasp, Sphecius speciosus (Hymenoptera : Sphecidae). American Zoologist 6(3):582. [Abstract].
    Lin reports that male wasps frequently used his person as a perch in defending their mating territories as he observed them. Males would move with Lin up to 45 feet away and, thus, changed the boundaries of their territories as Lin moved; many returned to Lin at the same location two or more days in a row and even alighted on the same part of his anatomy that they had used the previous day!

  66. Lin, N. 1966. Copulatory behavior of the cicada killer wasp, Sphecius speciosus. Animal Behavior 14:130-131.
    Detailed description of mating based on 61 observed copulations. Findings are similar to those in the paper below.

  67. Lin, N. 1967. Role differentiation in copulating cicada killer wasps. Science 1334-1335.
    Describes the functional roles of males and female during 79 copulations. In both this and the preceding paper, Lin appears to have missed the interesting precopulatory behavior of the male (he strokes and shakes the female's head with his forelegs while mounted on her back and taps her head and antennae with his antennae) that was reported by Reinhard (1929). Alcock (1975) reports similar male precopulatory behavior in S. grandis

  68. Lin, N. 1967. Territorial behavior in the cicada killer wasp Sphecius speciosus (Drury)(Hymenopteraa: Sphecidae) I. Behaviour 20:115-133.
    Describes male territoriality in a Sphecius speciosus population on Brooklyn, NY, playing fields, and notes that "Localizing of territories in places of high emergence hole density fosters male-female interaction because the owners are more likely to contact emerging females." Yup, ya gotta be there when the phone rings!

  69. Lin, N. 1978. Contributions to the ecology of the cicada killer, Sphecius speciosus (Hymenopteraa: Sphecidae). Journal of the Washington Academy of Science 68(2):75-82.
    Wasps do not emerge via the burrow hole dug the previous year by their mothers, but tunnel directly from the nest chamber to the surface; hard soil delays emergence. Lin notes that there is significant male-female overlap in emergence hole diameter. Life span is about 15 days for males and approximately 30 days for females.

  70. Lin, N. 1979. Differential prey selection for the sex of offspring in the cicada killer Sphecius speciosus (Hymenoptera: Sphecidae). Proceedings of the Entomological Society of Washington 81:269-275.
    A reexamination of data from Dow (1942) and Dambach and Good (1943) which shows that females know in advance the sex of the egg they will lay.

  71. Lin, N. 1979. The weight of cicada killer wasps Sphecius speciosus and the weight of their prey. Journal of the Washington Academy of Science 68:159-163.
    Notes a slight overlap in body masses of male and female wasps. Two new species of Tibicen used as prey are reported; T. figurae and T. resh. Previous claims of cicadas weighing 4-6 times more that the wasps carrying them appear to be greatly exaggerated.

  72. Lin, N. and C.D. Michener. 1972. Evolution of sociality in insects. The Quarterly Review of Biology 47:131-159
    Suggests that the early beginnings of sociality may be present in nest aggregations and that as many as four female Sphecius speciosus share the same burrow. On p. 142 they postulate three classes of female: 1) strong fliers which can lift large cicadas and which tend to lay female eggs in nest chambers provisioned with two cicadas, 2) "An intermediate type, some of them being able to carry only light cicadas Such wasps lay mostly male-producing eggs" and 3) "Wasps that neither provision nor start new nests." These wasps may parasitize their larger sisters by laying eggs in and closing nest open chambers they find containing one cicada. Field studies show that females remain in one class. They also report observations of "Type 2" wasps abandoning attempts to return to their own burrows with a paralyzed cicada and, instead, entering a nearby burrow.

  73. Manee, A.H. 1915. Notes on cicada-killer. Entomological News 26:266.
    Reports seeing Sphecius speciosus feeding on ash and oak sap oozing from gashes in the bark made by beetles. cicada killers also were seen to eat gum on oak acorn buds and to feed on persimmon fruit on the ground.

  74. Marlatt, C.L. 1907. The periodical cicada. U.S. Bureau of Entomology, Bulletin 71.
    Under the title "Hymenopterous Enemies" (pp. 130-135), Marlatt summarizes Riley's account (1892) of the biology of Sphecius speciosus. On p. 128 he states that cicada killers may prey on late-emerging periodical cicadas.

  75. Matheson, R. 1951. Entomology for introductory courses. Comstock Publ., Ithaca, NY.
    Contains a brief, accurate account of Sphecius speciosus biology on pp. 517-518.

  76. O'Neill, K.M. 2001. Solitary wasps: behavior and natural history. Cornell University Press, Ithaca. 406 pp.
    Describes cicada killer biology many times in context and comparison to other species.

  77. Packard, A.S. 1867. Revision of the fossorial hymenoptera of North America. Proceedings of the Entomological Society of Philadelphia 6:353-444.
    On pp. 442-443 Packard describes and gives the geographic distribution of Sphecius speciosus (then known as Stizus speciosus), Sphecius grandis (then known as Stizus grandis) and Sphecius hogardii (then known as Stizus Hogardii).

  78. Pasquale, L. 2002. Spare that cicada killer. Natural History 111 (8):8.
    Letter to the editor regarding Coelho's article in Natural History

  79. Pate, V.S.L. 1936. Studies in the nyssonine wasps. II. The subgenera of Sphecius (Hymenoptera : Sphecidae : Gorytini). Bulletin of the Brooklyn Entomological Society 31:198-200.
    Pate established a new subgenus of Sphecius: Nothosphecius. A taxonomic key to distinguish Sphecius from Nothosphecius and Spheciennus is provided. Nothosphecius has since been changed back to Sphecius but Spheciennus still stands.

  80. Pfennig, D.W. and H.K. Reeve. 1989. Neighbor recognition and context-dependent aggression in a solitary wasp, Sphecius speciosus (Hymenoptera:Sphecidae). Ethology 80:1-18.
    Larger females tend to win burrow disputes, but there is a resident advantage as well. Females are less tolerant of intrusion when there is an exposed cicada in the burrow and when the intruder is smaller. Females are more tolerant of their near neighbors than they are of females from more distant burrows.

  81. Pfennig, D.W. and H.K. Reeve. 1993. Nepotism in a solitary wasp as revealed by DNA fingerprinting. Evolution 47:700-704.
    Neighboring females are related, thus providing a kin selection argument for the greater tolerance to burrow intrusion.

  82. Rau, P. and N. Rau. 1918. Wasp studies afield. Princeton University Press, Princeton. 372 pp.
    Contains a brief, anecdotal account of an encounter with cicada killers foraging on sap of sunflowers.

  83. Reinhard, E.G. 1929. The witchery of wasps. Century, New York.
    A florid but accurate account of cicada killer biology. He notes the unusual precopulatory behavior of males, the digging behavior of females and the fact that larvae hatching from nest cells with two cicadas are females, while those hatching from cells with only one cicada are males.

  84. Riley, C.V. 1892. The larger digger wasp. Insect Life 4:248-252.
    This is a classic paper by the grandfather of American entomology documenting numerous aspects of cicada killer biology, including drawings of burrow structure and of the pupae. Most of the information is still considered to be correct.

  85. Riley, C.V. 1893. On the larva and some peculiarities of the cocoon of Sphecius speciosus. Procedings of the Entomological Society of Washington. 2:171-172.
    Riley notes the unusual "tubercules" woven into the silk cocoon by the larva before the cocoon hardens and opines that they may function in the larva’s breathing.

  86. Ross, K.G. and R.W. Mathews. 1991. The social biology of wasps. Cornell University Press, Ithaca, New York.

  87. Salt, G.L. and C.W. Holliday. 1992. The ecology of the cicada-killer wasp, Sphecius speciosus. In: R.D. Yearout (ed) Proceedings of the National Conference on Undergraduate Research VI, Volume III. The University of North Carolina, Asheville, pp.1399-1404.
    Describes male territorial behavior, mating. Suggests the presence of a mating pheromone and for outcrossing--females rarely return to their natal nesting aggregation.

  88. Savin, W.M. 1923. A wasp that hunts cicadas. Natural History 23:569-575.
    An accurate popular account of Sphecius speciosus biology. Contains the earliest report of the orientation flights made by females as they leave their burrows to hunt (p. 571).

  89. Schmidt, J.O. 1990. Hymenoptera venoms: striving toward the ultimate defense against vertebrates. In: D.L. Evans and J.O. Schmidt (eds) Insect defenses: Adaptive mechanisms and strategies of prey and predators. SUNY Press, Albany, pp. 387-419.
    Contains LD50 of cicada killer venom to mice (not very lethal) as well as other Hymenopteran venoms.

  90. Smith, F.F. 1925. Note on damage to formal garden by cicada-killer. Journal of Economic Entomology 18:836.
    Smith reports a large Sphecius nesting aggregation on a Philadelphia estate and opines, wrongly, that such large aggregations are uncommon. The wasp’s burrows are alleged to have been killing the boxwoods under which they were dug and a teaspoon of potassium cyanide was placed in each burrow to kill the wasps; it worked, but don’t try this at home! It’s possible that the boxwoods “... had died out in many places above these [cicada killer] tunnels...” because the gardener had tried cyanide the year before.

  91. Smith, J.B. 1901. Notes on some digger wasps. Journal of the New York Entomological Society 9:29-40, 52-72.

  92. Storer, J.H. 1953. The web of life. Signet Key Books, New York. 128 pp.
    The burrow of a cicada killer is shown in a plate in the center of the book to illustrate the benefits of invertebrates to the soil.

  93. Strange, C. K. 2000. The Cicada Killers of Florida (Hymenoptera: Sphecidae). Entomology Circular No. 402, September/October 2000, Florida Department of Agriculture and Conservation Services, Division of Plant Industry.
    A brief and accurate account of the biology of cicada killers in Florida, with a new report of Sphecius hogardii from Lover's Key in the Ft. Meyers area.

  94. Tashiro, H. 1987. Turfgrass insects of the United States and Canada. Cornell University Press, Ithaca, NY, 391 pp.

  95. Teale, E.W.  1962.  The strange lives of familiar insects. Dodd, Mead & Company, New York.  208 pp.
    Chapter 12 is "The life of the cicada-killer wasp," a popular, well written account of the life cycle of the animal, without too many misconceptions.

  96. Trinca, K.G. and P.K. Eason. 1994. The use of visual landmarks as territorial boundaries by the cicada killer wasp. American Zoologist 34:63A [Abstract].
    The presence of structures in the environment facilitates the formation of territorial boundaries, minimizing agonistic encounters between males. Full paper is Eason et al. (1999) above.

  97. Vittum, P.J., M.G. Villani and H. Tashiro. 1999. Turfgrass insects of the United States. Cornell Univ. Press., Ithaca.
    Pages 266-268 contain a brief and accurate account of Sphecius speciosus biology.

  98. Walton, R. 2011. Solitary wasps: behavior and identification. An introduction for naturalists. rkwalton.com
    An outstanding page with many video articles on digging wasps, including the Eastern cicada killer. A must-see site for all.

  99. Weaver, J.E. 1995. Life history, habits, and control of the cicada killer wasp in West Virginia. Circular 161, March 1995, Agricultural and Forestry Experiment Station, West Virginia University, 14 pp.
    Tests various insecticides and application methods (best is spraying into burrow) for control of cicada killers on golf courses. PDF Reprint.

  100. Werner, F. and C. Olson. 1994. Insects of the Southwest. Fisher Books, Tuscon, AZ.
    Contains a brief account of Sphecius grandis and S. convalis biology on p. 116.

  101. Wiedman, K.A. 1997. The function of spurs on the hind legs of cicada killers (Sphecius speciosus). Honors Thesis, Department of Biological Sciences, Western Illinois University, 11 pp.
    Hind tibial spurs act as one-way flaps that apparently function to aid in movement of soil during burrow excavation.

  102. Zim, H. S. and Cottam, C. 1951. Insects: a guide to familiar American insects. Golden Press, New York.
    Although a children's field guide, this thin book's (and subsequent editions) entry on cicada killers contains the only published range map of the eastern species. The map in the 2001 edition shows Sphecius as occurring everywhere in the U.S. and not in Mexico; for correct ranges of the five New World Sphecius species, see Holliday and Coelho, 2006.


The Life Cycle of the Cicada Killer | Help Cicada Killer Research | The Cicada Killer Thriller Page

[ 1998-2009 Joe Coelho]