A three-celled nest of Liris argentatus was excavated and examined in upstate New York. The nest was constructed from the terminus of an abandoned cicada-killer burrow and the cells contained 1-4 incompletely paralyzed Gryllus pennsylvanicus as prey. Two of the three cells were cleptoparasitized by the satellite fly Senotainia trilineata. The components of the ecology and behavior of L. argentatus we observed are compared with those from previous studies on this species

Kurczewski, Frank E

Spofford, Margery G

English

ValpoScholar

Further Observations on the Nesting Behavior of \u3ci\u3eLiris Argentatus\u3c/i\u3e (Hymenoptera: Sphecidae)

Valparaiso University

The Great Lakes Entomologist Volume 20 Number 3 - Fall 1987 Number 3 - Fall 1987 Article 3 October 1987 Further Observations on the Nesting Behavior of Liris Argentatus (Hymenoptera: Sphecidae) Frank E. Kurczewski State University of New York College of Environmental Science and Forestry Margery G. Spofford State University of New York College of Environmental Science and Forestry Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Kurczewski, Frank E. and Spofford, Margery G. 1987. "Further Observations on the Nesting Behavior of Liris Argentatus (Hymenoptera: Sphecidae)," The Great Lakes Entomologist, vol 20 (3) Available at: https://scholar.valpo.edu/tgle/vol20/iss3/3 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 1987 THE GREAT LAKES ENTOMOLOGIST 121 FL-:RTHER OBSERVATIONS ON THE NESTING BEHAVIOR OF URIS ARGENTATUS (HYMENOPTERA: SPHECIDAE) Frank E. Kurczewski and Margery G. Spoffordl ABSTRACT A tlJre<!-celled nest of Uris argentatus was excavated and examined in upstate New Yori.:. The nest was constructed from the terminus of an abandoned cicada-killer burrow and the cells contained 1-4 incompletely paralyzed Gryllus pennsylvanicus as prey. Two of the tlJre<! cells were cleptoparasitized by the satellite fly Senotainia trilineata. The components of the ecology and behavior of L. argentatus we observed are compared with those from pre"ious studies on this species. Despirc the fact that Uris argentatus (Beauvois) has been studied for nearly a century, there is much to learn about its nesting behavior. Although many prey species of Gryllidae are listed foc L. argentatus (Ashmead 1894, Williams 1914, Rau and Rau 1918, Rau 1922. Krombein and Evans 1955, Krombein 1958, Kurczewski and Kurczewski 1971, Steiner 19:6, O'Brien and Kurczewski 1982a), the list has not been exhausted. The number of p ey per cell varies from only a single, rather large, late instar Gry lus veletisI Alexander and Bigelow) in each of 13 fully-provisioned cells (O'Brien and Kurczewski 198:a •. to as many as six, small, immature crickets per cell (Ashmead 1894). Nest locariOll5 of L argentatus are variable, with some wasps using pre-existing cavities and 0i.ber'S .::onstrocting burrows from the sand surface (Bohart and Menke 1976, O'Brien and Kurczcllioski 198241. Our observations on L. argentatus supplement the recent study by O'Brien and Kurczewski (1982a). Specifically, they were made later in the year than theirs. in"oIH~ a different generation of wasps, and add new i formation to the behavior and ecol~' of this species. On 9 August 1985. at the edge of a field bordering a man-made sand pit near Owasco Lake. just south of Auburn, Cayuga County, New York, we observed a female L. argelfU1!JLS hunting for crickets in holes and depressions on a south-facing, moss-covered slope. This habitat was nearly identical to that described by O'Brien and Kurczewski f 198241. Tbe next day, on a bare sandy road leading into the sand pit, we observed a slightly smaller L. rgentatus transporting prey. The cricket was grasped by the bases of its antennae Ilioith !be wasp's mandibles, ventral side up, and carried to the nest in low flights. 8-12 em in length. During each of three subsequent provisioning trips this wasp lost her Ilio'ay and made several circles, 30-50 cm in diameter, in the vicinity of her entt'alll.""e. After tra"'ersing 15-20 cm she would climb to the top of a low plant, pause 'l\bile retaining her grasp of the prey, and look around, perhaps a form of navigation. During ODe such circle the provisioning wasp attracted a female of the satellite fly S(!1l(}rainia rri.liru!ata (Wulp) (Sarcophagidae: Miltogramminae) which trailed in flight at a d.ist.ance of 15 em. When the provisioning wasp paused, the fly alit on the cricket's abdominal stemites and larviposited. As the female resumed transport to the nest, the fly 'Enyirollmenol and ~1 Biology. State Cniyersity of New York College f Environmental Science and ~'. S~TlIC1l5e. !'.'Y 13210. 1Kurczewski and Spofford: Further Observations on the Nesting Behavior of <i>Liris ArgentatPublished by ValpoScholar, 1987122 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No.3 1C 1 A/,)\~ly B l. 	Abandoned burrow of Sphecius speciosus. lateral view, and cells of Uris argentatus; m=maggots. rode on the cricket's sternites for 4 cm, flew off, landed, and remained motionless on the sand. The wasp proceeded unimpeded into the nest entrance, seemingly unaware of the maggot(s) on the prey. The nest entrance being used by the female L. argentatus was that of an abandoned Sphecius speciosus (Drury) (Sphecidae) burrow, 2.35 cm in diameter, which was also being used by the sphecid Philanthus politus Say and the pompilid Anoplius marginatus (Say). On IO August, amidst the repeated nesting activities of the three species of wasps, two females of Phrosinella aurifacies Downes (Sarcophagidae; Miltogramminae) were seen to enter the Sphecius entrance and one of the flies larviposited. The P. politus and A. marginatus nests were not excavated so we cannot ascertain the larvipositional frequency o  P. aurifacies on their cells (prey), but no L. argentatus cell contained a P. aurifacies maggot. The abandoned cicada-killer burrow was 25 cm long and terminated at a depth of 9 cm beneath the sand surface (Fig. 1). Three cells of L. argentatus were unearthed just beyond the terminus of the Sphecius burrow at depths of 13, 14, and 18 cm. Cell A was located 4 cm almost directly beneath the terminus of the cicada-killer burrow and separated laterally from cell B by 2 cm; cells B and C were separated by 4 cm. The cells varied in shape and size: A, 1.1 cm wide and high, and 2.6 cm long; B, 1.3 em wide, 1.5 em high, and 1.7 cm long; C, 1.5 cm wide and high, and 2.4 cm long. All prey were placed in the cells head inward and ventral side up or on the side. They were only lightly paralyzed and moved their antennae, mouthparts, and legs considerably; some actually walked or jumped out of the cells when unearthed. Cell A contained a single nymph of Gryllus pennsylvanicus Burmeister, weighing 152 mg, with a small wasp larva feeding at the left forecoxal corium and placed transversely across the cricket's thoracic sternum between the bases of the first and second pairs of legs. Cell B held two smaller nymphs of G. pennsylvanicus with one bearing an even smaller wasp larva in the same position as above and the other, three The wasp larva disappeared within 24 h. Cell C contained thre  small n one immature female of G. pennsylvanicus, and four maggots. No egg or wasp larva was found in this cell which appeared to be the oldest of the three. The smallest cricket in the cell weighed only 48 mg. All seven maggots formed puparia and three females and two males of Senotainia 2The Great Lakes Entomologist, Vol. 20, No. 3 [1987], Art. 3https://scholar.valpo.edu/tgle/vol20/iss3/31987 THE GREAT LAKES ENTOMOLOGIST 123 trilinea1l1 emerged on 27 and 28 August 1985. The two other flies were determined as a female and male of S. trilineata from their pupal remains. On 15 August this wasp was observed transporting a paralyzed cricket, as before, into another cicada-killer entrance, only the nest was active. (The female S. speciosus took in a par lyz d cicada shortly after the provisioning female L. argentatus entered). The nest was exca,·ated on 19 August. The S. speciosus burrow was traced for 135 cm to a depth of 35 cm. It led to six fully-provisioned cicada-killer cells, but no L. argentatus cells were unearthed during the excavation. DISCUSSION 1be location of L. argentatus nests varies between study sites. Williams (1914) noted females nesting in the cracks of a sidewalk. Rau (1922) observed a wasp flying into a hole in a day bank. A.ll of the nests in O'Brien and Kurczewski's (1982a) study were started from the sand surface. In our study of L. argentatus, females nested from abandoned and actiye cicada-killer burrows. 1be depdls of L. argentatus cells reflect the variation in nest locations. Rau and Rau ( 1918 1 found a single cell of this species at a depth of 6.5 cm and another in a clay bank only 3.2 em deep. Rau (1922) recorded a third cell of this species at a depth of 15 cm. O'Brien and Kurczewski (1982a) reported L. argentatus cells from four nests at rather uniform depdls of from 9.0 to 14.5 cm. The maximum and minimum cell depths recorded in their study were from a lO-celled nest. Three cells from the nest in the present study were unearthed at depths of 13-18 cm. The terminus of the associated cicada-killer burrow was only 9 cm deep, indicating L. argentatus had constructed side burrows Iearum! to its cells. 1be-spe.."ies of prey of L. argentatus reflect the time of year and the varied habitats in which t e wasps nest. For example, Krombein (1958) took a female with a nymph of Orocharis saJlaJor l.Thler in North Carolina in August, and Kurczewski and Kurczewski 119""'11. two wasps with a male Gryllus rubens (Scudder) in Florida in March and an oyervoimcring nymph of Miogryllus verticallis (Serville) in Kansas in May, respectively. In ~e'" 'lad:. L. argentatus preyed upon Gryllus veletis in May and G. pennsylvanicus in August IO'Brien and Kurczewski 1982a, present study). 1bere is often an inverse relationship between the number of prey per cell and the Size1SI of the cricket(s) in L. argentatus. O'Brien and Kurczewski (1982a) reported a single. large G. veleris in each of 13 cells. Rau and Rau (1918) noted four adult male G. penllS)l\'aniC1l5 in a single cell and two G. pennsylvanicus nymphs and one Allonemobius fasciaIus IDe Geer) in another cell. The present study reveals from one large to four small nymphs of G. penllS)'lmnicus per cell. We found no correlation between cell size and prey mass nor between cell depth and age of wasp progeny in L. argentatus. O'Brien and Kurczewski ,1982a), likewise, noted no correlation between cell length and prey weight (size I in this species. but found that shallower cells held older larvae in one 10-celled nest. Se,'eral author"s have commented on the incomplete paralysis of the prey of L. argen1i:lnLS. We found the prey crickets to be extremely lively, exhibiting considerable IIlOwment of the appendages, and, in some cases, walking or jumping from the cells ",nen unearthed. The reason for the light paralysis of prey in L. argentatus is unknown, but y~' iD.."OIDJllete paralysis is well-documented for other larrine wasps such as Tachytes 1TU'rf[ILJ Fox and T. intermedius (Viereck) (Krombein and Kurczewski 1963, Kurczewski and-Kurcze"'ski 1984). Steiner (1976), too, found only temporary paralysis with partial re..."'OYe1Y of motory reflexes in three species of Liris. He believed that, because such wasps are rather primitive sphecids (Bohart and Menke 1976), their venom has only a limited eff~""t both in time and space. He (1976, 1984) noted further that the mole cricketprey of anoc:bcr primitive larrine, genus Larra, recovers even more completely from the effects of the wasp' s '·enom. The mole crickets behave normally, as if they had not been stung. Steiner 1.1976. 1984) related this complete recovery to the "weak venom 3Kurczewski and Spofford: Further Observations on the Nesting Behavior of <i>Liris ArgentatPublished by ValpoScholar, 1987124 THE GREAT LAKES ENTOMOLOGIST Vol. 20, No.3 properties" of the wasp and (or) the considerable size difference between the wasp and its enormous prey. The 13 cells of L. argentatus excavated by O'Brien and Kurczewski (1982a) were devoid of cleptoparasites, yet two of the three cells we unearthed in the present study each contained several miltogramrnine maggots (Senotainia trilineata). The fact that O'Brien and Kurczewski's (1982a) study was done in May when such flies are virtually absent, and our study was made in August when satellite flies are abundant, suggests adult overwintering in sphecid and pompilid wasps is a means to avoid such cleptoparasitism (O'Brien and Kurczewski 1982b, Aim and Kurczewski 1984). In L. argentatus and an unrelated sphecid, Podalonia luctuosa (Cresson), which preys upon cutworms (O'Brien and Kurczewski 1982b), the spring-nesting (overwintering) generations appear to be larger in size than the mid to late summer-nesting generations in upstate New York. However, the spring-nesting generations f these species may be more obvious to the observer because there are practically no other wasps nesting at this time of year. M. F. O'Brien (pers. comm.) believes there may be two different (isolated) gene pools of "L. argentatus" comprising spring and summer sibling species that are synchronized with two sibling species of host crickets, Gryilus veletis and G. pennsylvanicus, whose adults are also allochronic (Alexander and Bigelow 1960, Alexander 1968). One larval L. argentatus that spun its cocoon in New York on 15 August 1985, died during late summer before pupating and did not provide us with the evidence to support O'Brien's hypothesis. Two sphecids that do not overwinter in the adult stage, Ammophila harti (Fernald) and Oxybelus bipunctatus Olivier, also have large spring and small late summer generations, based upon actual numbers of nesting females observed (Hager and Kurczewski 1986, pers. observ., respectively). Many individuals of each of these species have been reared to ascertain that a certain proportion f the spring population goes into the makeup of later generations of the same year. However, we do not know what proportion of the spring populations converts directly to spring generations of the subsequent year(s). In Senotainia trilineata and S. vigilans Allen, two sarcophagid flies cleptoparasitic on sphecid wasps, some larvae convert to adults in just a few weeks while others emerge as adults after undergoing diapause for 1-3 years (Spofford et al. 1986, pers. observ .). After reviewing and evaluating all of this information and examining the enormous amount of collecting data presented by Krombein and Gingras (1984), we believe that L. argentatus is at least bivoltine in New York State. ACKNOWLEDGMENTS We thank R. D. Alexander and M. F. O'Brien, Division of Insects, Museum of Zoology, The University of Michigan, for identifying Grylius pennsylvanicus and for r viewing the manuscript; and A. S. Menke, BBII, ARS, USDA, for confirming the identity of Uris argentatus. LITERATURE CITED Alexander, R. D. 1968. Life cycle origins, speciation, and related phenomena in crickets. Quart. Rev. BioI. 43:1-41. Alexander, R. D., and R. S. Bigelow. 1960. Allochronic speciation in field crickets, and a new species, Acheta veletis. Evolution 14:334-346. Aim, S. R., and F. E. Kurczewski. 1984. Ethology of Anop/ius tenebrosus (Cresson) (Hymenoptera: Pompilidae). Proc. Entomol. Soc . Washington 86: II 0-119. Ashmead, W. H. 1894. The habits of the aculeate Hymenoptera. Ill. Psyche 7:59-66. Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. Univ. California Press, Berkeley. 695 pp. Hager, B. J., and F. E. Kurczewski. 1986. Nesting behavior of Ammophila harti (Fernald) (Hymenoptera: Sphecidae). Amer. MidI. Natur. 116:7-24. 4The Great Lakes Entomologist, Vol. 20, No. 3 [1987], Art. 3https://scholar.valpo.edu/tgle/vol20/iss3/31987 THE GREAT LAKES ENTOMOLOGIST 125 Krombein. K. Y. 1958. Biological notes on some wasps from Kill Devil Hills, North Carolina, and addiriollS to the faunal Jist (Hymenoptera, Aculeata). Proc. Entomo!' Soc. Washington 60:97-110. Krombein. K. Y.. and H. E. Evans. 1955. An annotated list of wasps collected in Florida, March 20 to April 3. 1954 (Hymenoptera, Aculeata). Proc. Entomo!. Soc. Washington 57:223-235. Krombein. K. V .. and S. S. Gingras. 1984. Revision of North American Uris F bricius 'Hyme.DOp(era: Sphecoidea: Larridae). Smithson. Contrib. Zoo1. 404:1-96. Krombein. K. V and F. E. Kurczewski. 1963. Biological notes on three Floridian wasps .Hymenoptera. Sphecidae). Proc. BioL Soc. Washington 76:139-152. Kurcz.ewski. F. E.. and E. J. Kurczewski. 1971. Host records for some species of Tachytes and other Larri.nae. 1. Kansas Entomo!. Soc. 44:131-136. 1984. ~fating and ncsting behavior of Tachytes intermedius (Viereck) (Hymenoptera: Spba."i(\aeL Proc. Entomo!' Soc. Washington 86:176-184. O·Brien. M. F. and F. E. Kurczewski. 1982a. Nesting and overwintering behavior ofUris argentata 'Hyme.DOp(era: Larridae). 1. Georgia Entomo!. Soc. 17:60-68. ___.1982b. Ethology and overwintering of Podalonia luctuosa (Hymenoptera: Sphecidae). Great Lake;; Emomol. 15:261-275. Ran. P. 1922. Ecological and behavior notes on Missouri insects. Trans. Acad. Sci. St. Louis 2·U--1. Ran. P .. and~. Ran. 1918. Wasp studies afield. Princeton Vnlv. Press, Princeton. 372 pp. SpoffOfd. ~t G .. F. E. Kurczewski, and D. J. Peckham. 1986. Cleptoparasitism of Tachysphex lerminaru.s (Hymenoptera: Sphecidae) by three species of Miltogrammini (Diptera: Sarcophagidael. Ann. Entomo!. Soc. Amer. 79:350-358. Steiner ....... 1976. Digger wasp predatory behavior (Hymenoptera, Sphecidae). II. Comparative study of dosely related wasps (Larrinae: Uris nigra, Palearctic; L. argentata and L. aequalis, Nearctic) that all paralyze crickets (Orthoptera, Gryllidae). Z. Tierpsychol. 42:343-380. 1984. Wfiy can mole crickets stung by Larra wasps (Hymenoptera, Sphecidae: Larrinae) res:tIIIli! IlOrnlal activities? The evolution of temporary paralysis and permanent deactivation of the prey 1. Kansas Entomol. Soc. 57:152-154. Wi.l.liaJ:ns. F. X. 1914. Monograph of the Larridae of Kansas. Vniv. Kansas Sci. Bull. 8: 117-213. 5Kurczewski and Spofford: Further Observations on the Nesting Behavior of <i>Liris ArgentatPublished by ValpoScholar, 1987

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Further Observations on the Nesting Behavior of \u3ci\u3eLiris Argentatus\u3c/i\u3e (Hymenoptera: Sphecidae)

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