Detection and Identification of Two New Native Hymenopteran Associated with the Exotic Sirex noctilio in North America Author(s): Christopher R. Standley, E. Richard Hoebeke, Dylan Parry, Douglas C. Allen and Melissa K. Fierke Source: Proceedings of the Entomological Society of Washington, 114(2):238-249. 2012. Published By: Entomological Society of Washington DOI: http://dx.doi.org/10.4289/0013-8797.114.2.238 URL: http://www.bioone.org/doi/full/10.4289/0013-8797.114.2.238

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DETECTION AND IDENTIFICATION OF TWO NEW NATIVE HYMENOPTERAN PARASITOIDS ASSOCIATED WITH THE EXOTIC SIREX NOCTILIO IN NORTH AMERICA

CHRISTOPHER R. STANDLEY,E.RICHARD HOEBEKE,DYLAN PARRY,DOUGLAS C. ALLEN, AND MELISSA K. FIERKE

(CRS, DP, DCA, MKF) Department of Environmental and Forest Biology, State University of New York, Environmental Science and Forestry (SUNY-ESF), Syracuse, New York 13210, U.S.A. (e-mail: [email protected]); (ERH) Department of Entomology, Cornell University, Ithaca, New York 14853, U.S.A.; current address: Georgia Museum of Natural History, University of Georgia, Athens, Georgia 30602, U.S.A. (e-mail: [email protected]).

Abstract.—Rhyssa crevieri (Provancher), a primary of Siricidae , and the Holarctic poemeniine ichneumonid, Pseudorhyssa nigricornis (Ratzeburg), a clep- toparasitoid of Siricidae via its parasitoids (: ), were reared from two stands of Scots pine (Pinus sylvestris L.) and red pine (P. resinosa Ait.) in- fested with the exotic siricid Sirex noctilio F. near Tully (Onondaga Co.), New York, in 2010. Previously, P. nigricornis has been recorded from the primary siricid parasitoids persuasoria (in and North America) and R. howdenorum Townes and Townes and R. lineolata (Kirby) (in North America). Peak emergence of P. nigricornis occurred in early May concurrently with R. persuasoria and R. crevieri.Asecondpeak occurred in late May, which overlapped peak emergence of R. lineolata and nortoni (Cresson). Although 14 individuals of two native siricids, Sirex nigricornis F. and S. edwardsii Brulle´, emerged from sampled trees, the number of S. noctilio recovered was far higher (372), suggesting cleptoparasitism of the exotic woodwasp rather than native siricids. Approximately 26% of rhyssine parasitoids in pine stands were clepto- parasitized by P. nigricornis. This study marks the first association of R. crevieri and P. nigricornis with S. noctilio in North America. A diagnosis, color images of characters, and a key are provided to aid in the identification of all parasitoids in this study. Key Words: cleptoparasitism, , Rhyssa lineolata, Rhyssa crevieri, Megarhyssa nortoni, morphological characters, identification key, rearing data DOI: 10.4289/0013-8797.114.2.238

Sirex noctilio F. (: Siricidae) et al. 2007). Its primary hosts are stressed is a relatively large wood-boring trees within the Pinus (Spradbery native to Eurasia and North Africa (Borchert and Kirk 1978). However, under outbreak population levels, S. noctilio possesses the ability to attack relatively healthy * Accepted by Michael W. Gates trees (Spradbery 1973), disperse rapidly VOLUME 114, NUMBER 2 239

(Neumann et al. 1987, Tribe and Cillie´ The genus Pseudorhyssa contains 2004, Corley and Villacide 2009), and four species: the Eurasian P. alpestris cause great economic damage (Haugen (Holmgren) (northwest and central Europe 1990, Hurley et al. 2007). Previous in- and Japan), P. acutidentata Kusigemati vasions have been documented through- (Japan), and P. maculiventris Sheng and out the southern hemisphere (Hurley et al. Sun (China), as well as the Holarctic P. 2007). In 2004, a single adult female was nigricornis (Fitton et al. 1988, http:// captured in Fulton, NY, eliciting concerns www.taxapad.com/global.php). Species for native North American pine forests of Pseudorhyssa cleptoparasitize rhyssine (Hoebeke et al. 2005). Ichneumonidae, consuming the primary In areas where S. noctilio has become parasitoid larvae and feeding on the siricid an economically important invasive, Rhyssa, larvae paralyzed by the primary parasitoid Megarhyssa (Ichneumonidae: Rhyssinae), (Kerrich 1966, Spradbery 1969). Although and Ibalia (Cynipoidea: Ibaliidae) para- mostoftheliteraturedealingwithP. sitoids have been introduced for biological nigricornis uses the name P. maculicoxis control (Miller and Clark 1935, 1937; (Kreichbaumer), Horstmann (1999) con- Rawlings 1951; Zondag 1959; Zondag cluded P. maculicoxis should be synony- and Nuttall 1961; Cameron 1965; Nuttall mized with P. nigricornis. 1972; Taylor 1976, 1978; Murphy 1998). Several studies indicate P. nigricornis Species of Rhyssa and Megarhyssa attack is associated exclusively with conifer- late-instar larvae (Murphy 1998) by dril- feeding siricid larvae via their rhyssine ling into wood after hosts (Chrystal and parasitoids (Spradbery 1969, Schimitschek Myers 1928a, b). Ibaliid wasps attack eggs 1974, Kusigemati 1984). In Europe, P. and early-instar siricid larvae (Murphy nigricornis (cited as P. maculicoxis)iswell 1998) using drill shafts left by ovipositing documented as a cleptoparasitoid of the siricids. siricids Sirex juvencus L., S. cyaneus F., The parasitoid guild of native siricid and S. noctilio F. via the Holarctic R. per- woodborers in eastern North American suasoria (L.) (Spradbery 1969, 1970). The conifers (Taylor 1976, Murphy 1998) North American range of P. nigricornis includes primary parasitoids of the includes Quebec south to western North ichneumonid genera Rhyssa [R. crevieri Carolina, west to Alberta and California (Provancher), R. lineolata (Kirby), R. and southwest to northern Arizona howdenorum Townes and Townes, and (Townes and Townes 1960; Kirk 1974, R. persuasoria (L.)] and Megarhyssa [M. 1975; Carlson 1979). Kirk (1974, 1975) nortoni (Cresson)], and the ibaliid genus documented the association of P. nig- Ibalia [I. leucospoides ensiger (Norton)]. ricornis with R. persuasoria in Arizona This parasitoid community also includes and with R. persuasoria, R. lineolata, a secondary parasitoid or cleptoparasitoid, and R. howdenorum in western North the Holarctic poemeniine ichneumonid Carolina. Porter (2001) noted an abun- Pseudorhyssa nigricornis (Ratzeburg). dance of R. howdenorum and P. nig- Two recently published studies (Long ricornis (cited as P. maculicoxis)on et al. 2009, Eager et al. 2011) have docu- loblolly pine (Pinus taeda L.) on the mented the hymenopteran parasitoids as- Eastern Shore of Maryland; the sir- sociated with S. noctilio-infested pine trees icids Urocerus cressoni Norton, S. nig- in North America as Ibalia l. ensiger, ricornis F., and S. edwardsii Brulle´ on Rhyssa persuasoria, R. lineolata, and the same trees were the likely hosts of R. Megarhyssa nortoni. howdenorum. 240 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Female P. nigricornis are receptive to parasitoids and the cleptoparasitoid P. males two or three days post emergence nigricornis associated with S. noctilio. (Spradbery 1969). Gravid females ob- We also provide emergence phenologies serve R. persuasoria drilling for siricid for S. noctilio and the complex of primary larvae and, following completion of dril- rhyssine parasitoids, including the pre- ling, use the drill shafts to access and viously unreported R. crevieri and the cleptoparasitize hosts (Spradbery 1969, cleptoparasitoid P. nigricornis. Townes 1969). Females of P. nigricornis, however, do not necessarily have to ob- METHODS AND MATERIALS serve drilling by the primary parasitoid, In April 2010, pine trees in two pure as they can locate drill shafts several days stands of P. res ino sa Ait. and P. s ylv es tr is after their completion (Spradbery 1969). L. near Tully, NY, were examined thor- The ovipositor of P. nigricornis is smaller oughly for the diagnostic resin beads as- in cross-section and less sclerotized than sociated with S. noctilio oviposition drills primary rhyssine attackers, which facili- along the bole. Within each stand, five tates successful access to drill shafts and P.resinosa and five P.sylvestris trees with siricid hosts (Spradbery 1969). clear signs of infestation were felled. Oviposition by P. nigricornis occurs Trees ranged from 14.5 to 23.0 cm di- close to primary parasitoid eggs and, fol- ameter at breast height (dbh), with crown lowing eclosion, a battle ensues between conditions ranging from recently dead larvae of P. nigricornis and those of the with a few brown needles to dying with primary parasitoid (Spradbery 1969, sparse green needles. 1970). First-instar larvae of P. nigricornis Each tree was subsampled using are heavily armored and larger than methods developed by Eager et al. (2011). rhyssine larvae, with exaggerated, sickle- Nine 0.5 m bolts were removed systemat- shaped mandibles, and paired caudal ap- pendages that aid in defeating primary ically from each of the ten felled trees. If < species (Spradbery 1969, 1970). During trees were 10 m tall, the whole tree was the initial interaction, P. nigricornis lar- taken. All sample bolts were transported to vae rear on their hind, paired appendages a rearing facility at the State University of to kill rhyssine larvae with their large New York, College of Environmental Sci- mandibles (Spradbery 1969). No rhyssine ence and Forestry (SUNY ESF), Syracuse, larvae, irrespective of instar, prevailed NY. Ends of bolts were sealed with Wax- TM against attacking first instars of P. nig- lor end sealant (Willamette Valley Com- ricornis. After consuming the rhyssine pany) to prevent desiccation and placed in larva, P. nigricornis consumes the siricid cardboard emergence tubes. Tubes were host paralyzed by the adult rhyssine held at ambient environmental conditions before emerging the following spring in an outdoor insectary. All emerging in- (Spradbery 1969). sects were collected and recorded multiple Although two previous studies (Long times per day. To reduce noise when re- et al. 2009, Eager et al. 2011) documented porting emergence phenologies, daily data several ichneumonid parasitoids associated were summed over weekly time periods. with S. noctilio in North America, neither Identity of all rhyssine Ichneumonidae recorded P. nigricornis or R. crevieri from and Pseudorhyssa reared in this study infested trees. Here, we summarize and was confirmed morphologically using illustrate diagnostic characters and provide keys in Townes and Townes (1960) and a key to aid in identification of rhyssine in Kerrich (1966) and Wahl (1993), VOLUME 114, NUMBER 2 241 respectively, and/or by comparison with 10). In contrast, species of Rhyssa are identified research specimens housed in recognized by the fore wing with vein the Cornell University Collection cu-ajoiningCudistaltothebifurcation (Ithaca, NY) and the United States of M and Cu (Fig. 5), the occipital carina National Museum of Natural History is incomplete mediodorsally (Fig. 9), the (Washington, DC). Reared specimens of last visible tergite of the female gaster is S. noctilio, its ichneumonid parasitoids, extended into a polished, truncate horn and P. nigricornis, are deposited in the (Fig. 7), and tergite 2 of the gaster lacks SUNY ESF Entomology Museum Col- defined anterolateral grooves. Also, gastral lection (Syracuse, NY), Cornell Univer- segment 1 in Pseudorhyssa has a sharp sity Insect Collection (Ithaca, NY), and lateral longitudinal carina extending from the University of Georgia Museum of the spiracle to the posterior margin, while Natural History Collection of in species of Rhyssa gastral segment 1 (Athens, GA). lacks this distinct lateral longitudinal Percent cleptoparasitism by P. nigricor- carina (Wahl 1993). nis was calculated as the number of P. The Palearctic Pseudorhyssa alpestris, nigricornis divided by the total number of also reared from S. noctilio in Europe rhyssines plus P. nigricornis. (Schimitschek 1974), has not been de- tected in North America. It differs from P. RESULTS nigricornis by dorsal segments 1-3 of the gaster usually with a distinct reddish- Diagnosis brown mark near the posterior margin Species of the Rhyssinae and the ge- and by the face of the female often with nus Pseudorhyssa are easily recognized obscure brownish marks (Fitton et al. by the presence of strong transverse 1988). In contrast, the gaster and face of ridges on the mesoscutum (Fig. 11). No the female of P. nigricornis are black other Nearctic ichneumonid has this without any brownish markings. unique and distinctive feature. The fol- Among the species of Rhyssa reared lowing morphological characters differ- in this study, R. persuasoria is readily entiate P. nigricornis from any species of recognized by the flagellum of the an- Rhyssa. Conspicuous yellow or cream tenna being entirely black (Fig. 2). In markings on the head, thorax, propodeum, both R. crevieri and R. lineolata,thefla- and hind margins of the gastral segments gellum of the antenna has a broad white (Figs. 1, 2) characterize species of Rhyssa, band (Fig. 1), except in occasional small whereas the head, thorax, propodeum, and males (Townes and Townes 1960). Col- gaster in P. nigricornis lack any yellow or oration distinguishes these two latter cream markings (Fig. 3). Pseudorhyssa species. In R. crevieri, the white mark on nigricornis is further distinguished by the the metapleuron is subtriangular (Fig. 12), fore wing with vein cu-a joining at the prolonged forward along the upper mar- bifurcation of M and Cu (Fig. 4), the oc- gin. This subtriangular mark varies sub- cipital carina of the head capsule is com- stantially in females and is often absent or plete mediodorsally (Fig. 8), the last much reduced in some males. The hind visible tergite of the female gaster is coxa has no white mark dorsobasally broadly rounded apically (Fig. 6), and ter- (Fig. 12) or sometimes a very small gite 2 of the gaster (both sexes) possesses white mark limited to its constricted a moderately deep, smooth anterolateral base. In contrast, the white mark on the groove on each side of the midline (Fig. metapleuron in R. lineolata is rounded 242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1–3. Habitus, lateral view. 1, Rhyssa lineolata.2,Rhyssa persuasoria.3,Pseudorhyssa nig- ricornis. Scale bar = 5 mm. VOLUME 114, NUMBER 2 243

Figs. 4–10. Parasitoid characters. 4–5, Fore wing. 4, Pseudorhyssa nigricornis; note vein cu-a at bifurcationofMandCu(arrow).5,Rhyssa lineolata; note vein cu-a joining Cu distal to bifurcation of M and Cu (arrow). 6–7, Apex of gastral segments, dorsal view. 6, Pseudorhyssa nigricornis.7,Rhyssa lineolata. 8–9. Upper part of head, posterior view. 8, Pseudorhyssa nigricornis; note occipital carina complete mediodorsally (arrow). 9, Rhyssa lineolata; note occipital carina incomplete mediodorsally (arrow). 10. Base of gastral segments, dorsal view, of Pseudorhyssa nigricornis; note deep anterolateral grooves on tergite 2 (arrows).

(Fig. 13), constricted next to the upper usually in the form of a small dash but in margin. This mark is absent or reduced in larger females often enlarged and irregular some males. The hind coxa of R. lineolata in shape or covering the entire upper sur- has a white mark dorsobasally (Fig. 13), face of the coxa. Although both species 244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 11–13. Thoracic characters. 11, Rhyssa lineolata, mesoscutum; note strong transverse ridges. 12–13, Color markings of metapleuron and hind coxa. 12, R. crevieri. 13, R. lineolata. are recognized currently as valid, a foot- deepanterolateralgrooves(Fig.10)...... note in the “Catalog of Hymenoptera . . . Pseudorhyssa nigricornis (Ratzeburg) North of Mexico” (Carlson 1979) states: [synonyms: Pseudorhyssa maculicoxis (Kreichbaumer 1889), Pseudorhyssa sternata “It remains to be proven that crevieri is Merrill 1915, and Pseudorhyssa praealpina distinct from lineolata.” (Gyorfi 1946)] The following key, based on characters – Occipital carina incomplete mediodorsally given in the diagnosis above, allows for (Fig. 9). Female: last visible tergite of gaster the separation of all rhyssine ichneu- extended into a polished, truncate horn monids and Pseudorhyssa reared from (Fig. 7). Gastral tergite 2 without anterolateral grooves...... 2. Sirex noctilio-infested trees in the north- 2. Clypeus with median tooth or projection. eastern U.S. Female: sternite 2 of gaster with 2 median tubercles near middle ...... Key to Species of Ichneumonidae With ...... Rhyssa spp...... 3. a Strongly Transversely Rugose – Clypeus truncate. Female: sternite 2 of gas- Mesoscutum ter with 2 median tubercles near base ...... Megarhyssanortoni(Cresson) [wings (Fig. 11) unicolorous, subhyaline; stigma yellowish; (Adapted from Fitton et al. 1988 and female: tergites 4–6 with conspicuous, yel- Townes and Townes 1960) low, round, lateral spots on each side; thorax and gaster light brown to blackish, with 1. Occipital carina complete mediodorsally (Fig. conspicuous yellow markings] 8). Female: last visible tergite of gaster not 3. First 4 tergites of gaster with narrow whitish extended as horn (Fig. 6). Gastral tergite 2 with apical margin, of even width, and continuous VOLUME 114, NUMBER 2 245

across middle, or slightly narrower or inter- S. nigricornis and S. edwardsii, < 4% of ruptedmedially...... the total siricid emergence from sampled ..Rhyssa howdenorum Townes and Townes trees(Table2,Fig.15). (not found in this study) – First 4 tergites of gaster with white apical spots, not forming a continuous band across DISCUSSION middle...... 4. The rearing and identification of R. 4. Flagellum of antenna with broad white band crevieri as well as P. nigricornis from (Fig. 1) ...... 5. – Flagellum of antenna entirely black (Fig. 2) Sirex noctilio-infested trees at our study ...... Rhyssa persuasoria (L.) site in central New York marks the first 5. Metapleuron with subtriangular white spot, association of both this primary parasitoid prolonged forward along upper margin (Fig. and the cleptoparasitoid with S. noctilio in 12); hind coxa without white spot dorsoba- sally (Fig. 12), but sometimes with small North America. Previous studies in this white spot at base ...... region (Long et al. 2009, Eager et al. 2011) ...... Rhyssa crevieri (Provancher) recorded only the rhyssines R. persuasoria – Metapleuron with rounded white spot, con- and R. lineolata. However, morphological stricted next to upper margin (Fig. 13); hind similarity between R. crevieri and R. coxa with white spot dorsobasally (Fig. 13), often enlarged and irregular in shape..... lineolata may explain why R. crevieri ...... Rhyssa lineolata (Kirby) went unnoticed in both studies. Both species contain broad, white bands on the Emergence phenology.—Twenty adult antennae, except for some small males R. crevieri (9 males, 11 females) and (Townes and Townes 1960), and similar fifty-one adult P. nigricornis (30 males, white highlights on the rest of the body. 21 females) emerged from sample bolts Despite these similarities, differences in taken from five of the ten felled pine emergence phenologies of these parasit- trees (Table 1). Peak emergence of P. oids (Fig. 14) provide further support for nigricornis overlapped with its pre- their separate species status. viously recorded host, R. persuasoria, It is less likely that P. nigricornis would and with R. crevieri, a previously un- be misidentified as a species of Rhyssa recorded host (Fig. 14). A second peak based on distinguishing morphological in P. nigricornis emergence (Fig. 14) characteristics as noted in the key. Timing overlappedwithtwoothernativerhys- of sampling may account for the absence sines, R. lineolata and Megarhyssa of this species in one of the prior studies. nortoni. Cleptoparasitism of rhyssines by Long et al. (2009) collected samples from P. nigricornis was ; 26%. The introduced Sirex-infested trees in late June, which, S. noctilio wasbyfarthedominant according to our emergence phenology siricid with 372 individuals emerging (see Fig. 14), is late for recovery of Rhyssa from the infested bolts and only 14 native and the associated cleptoparasitoid. In

Table 1. Tree data associated with Sirex noctilio-infested pines harboring Rhyssa crevieri and Pseudorhyssa nigricornis near Tully, NY, in 2010.

Tree species Height (m) DBH (cm) Crown condition No. R. crevieri No. P. nigricornis

P. resinosa 11.3 17.5 sparse green needles 0 1 P. resinosa 16.2 18.5 needles all brown 1 7 P. resinosa 12.7 23 few brown needles 13 19 P. sylvestris 9.0 13 sparse green needles 1 0 P. sylvestris 8.9 17 sparse green needles 5 24 246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 14. Emergence phenology of Pseudo- Fig. 15. Emergence phenology of pooled rhyssa nigricornis, Rhyssa persuasoria, R. line- rhyssine primary parasitoids (Rhyssa persuasoria + olata, R. crevieri, and Megarhyssa nortoni reared R. lineolata + R. crevieri + M. nortoni), Pseudo- from pine trees felled near Tully, NY, in 2010. rhyssa nigricornis, Sirex noctilio, and native Siricidae populations near Tully, NY, in 2010. another study (Eager et al. 2011), col- In North America, P. nigricornis has lections from the same field site used in been associated with primary parasitoids our study (Heiberg Forest) did not yield of the native siricids Sirex abbotii Kirby P. nigricornis, perhaps because it focused (= cyaneus F.), S. nigricornis F., and S. exclusively on larval sampling. Charac- cyaneus in the southeastern U.S. (Kirk teristics distinguishing P. nigricornis lar- 1974) and of S. cyaneus, S. californicus vae from rhyssine larvae are not obvious (Ashmead), and S. longicauda Middlekauff (Spradbery 1970) and individuals of this in the southwestern U.S. (Kirk 1975). The species could have been easily over- strong overlap in emergence phenology looked. Alternatively, P. nigricornis may of P. nigricornis with the four rhyssine exhibit a density dependent relationship species suggests that all (R. persuasoria, with its rhyssine hosts as Eager (2010) R. lineolata, R. crevieri, and possibly M. recorded relatively low numbers of rhys- nortoni) may be suitable hosts. Initial sines in 2009. In 2011, sampled trees at peak emergence of P. nigricornis (early Heiberg Forest also had low numbers of May in 2010) was concurrent with both rhyssines and failed to yield any P. nig- R. persuasoria and R. crevieri, while the ricornis (C. Standley, unpublished). later peak coincided with R. lineolata and M. nortoni (Fig. 14). Emergence phe- Table 2. Number of emerged Pseudorhyssa nigricor- nologies of P. nigricornis supports uti- nis, rhyssine Ichneumonidae, and species of Siricidae lization of R. persuasoria and R. lineolata during the 2010 flight season from 0.5 meter logs as reported by Spradbery (1969) and Kirk removed from four trees near Tully, NY. (1974). Phenologies also indicate R. Species No. emerged crevieri is likely being exploited and it is possible P. nigricornis is utilizing M. Pseudorhyssa nigricornis 51 nortoni. When M. nortoni is excluded Rhyssa lineolata 70 Rhyssa crevieri 20 from calculations, percent cleptoparasi- Rhyssa persuasoria 31 tism of the remaining rhyssines increases Megarhyssa nortoni 27 from 26 to 30%. Sirex noctilio 372 Our data suggest P. nigricornis may Sirex nigricornis 8 be a significant source of mortality for Sirex edwardsii 6 R. persuasoria, R. lineolata,andR. 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