THE SIRICID WOOD WASPS of CALIFORNIA (Hymenoptera: Symphyta)

Home , Rhyssa persuasoria, Sirex, Sirex cyaneus, Tremex, Urocerus, Xeris

Uroce r us californ ic us Nott on, f ema 1e. BULLETIN OF THE CALIFORNIA INSECT SURVEY

VOLUME 6, NO. 4

THE SIRICID WOOD WASPS OF CALIFORNIA (Hymenoptera: Symphyta)

BY WOODROW W. MIDDLEKAUFF (Department of Entomology and Parasitology, University of California, Berkeley)

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES 1960 BULLETIN OF THE CALIFORNIA INSECT SURVEY Editors: E. G: Linsley, S. B. Freeborn, P. D. Hurd, R. L. Usinget Volume 6, No. 4, pp. 59-78, plates 4-5, frontis. Submitted by editors October 14, 1958 Issued April 22, 1960 Price, 50 cents

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES CALIFORNIA

CAMBRIDGE UNIVERSITY PRESS LONDON, ENGLAND

PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA THE SIRICID WOOD WASPS OF CALIFORNIA

(Hymenoptera: Symphyta)

BY WOODROW W. MIDDLEKAUFF

INTRODUCTION carpeting. Their powerful mandibles can even cut through lead sheathing. The siricid wood wasps are fairly large, cylin- These insects are widely disseminated by drical insects; usually 20 mm. or more in shipments of infested lumber or timber, and length with the head, thorax, and abdomen of the adults may not emerge until several years equal width. The antennae are long and fili- have elapsed. Movement of this lumber and form, with 14 to 30 segments. The tegulae are timber tends to complicate an understanding minute. Jn the female the last segment of the of the normal distribution pattern of the spe- abdomen bears a hornlike projection called cies. the cornus (fig. 8), whose configuration is The Nearctic species in the family were useful for taxonomic purposes. This distinc- monographed by Bradley (1913). Since that date tive structure alsogives them the common name the description of one new North American of horntails. The ovipositor is exserted and in species in the genus Sirex has been published some species may be quite long, even exceed- (Middlekauff, 1948). The excellent papers by ing the body length. Males are normally smal- Benson (1943, 1951) in which he gives keys to ler than the females, with flattened tibiae and world genera and European species prompted tarsi and with the last abdominal sternite end- me to study more thoroughly the systematics ing in a sharp point. The size of adult siricids and distribution of the siricid fauna of Cali- is quite variable and may vaiy to twice the fornia. size in the same species. The wood-boring The family Siricidae in California is com- larvae are whitish cream or yellow, usually posed of two subfamilies: the Sirioinae con- cylindrical, weakly Sshaped with only ves- taining the genera Sirex, Urocerus, and Xeris, tigial thoracic legs and with a small horny and the Tremicinae containing the single genus spine at the posterior end. Abdominal prolegs Tremen. To date, 13 species and subspecies are absent. in these genera are definitely known from the In California siricids are not as a rule con- state. Sirez nigricornis Fab. is doubtfully re- sidered to be primary enemies of sound timber corded from California (Norton, 1869), and since usually they do not attack healthy stand- subsequent collections have failed to confirm ing trees. They will, however, readily attack it as being present in our fauna. fire-damaged or otherwise injured or dying trees and ones recently felled. Female Sirex areolatus have even been reported as attacking recently sawed redwood lumber. The larval BIOLOGY and adult borings reduce the quality of the Although a considerable amount of study has lumber causing it to be downgraded. Adults recently been done by British and New Zealadd may cause considerable damage while emerg- workers on various biological and life history ing from infested timbers in homes by boring phases of several species of the Siricidae, through plaster walls, hardwood floors, and little is known about our North American spe- GO BULLETIN OF THE CALIFORNIA INSECT SURVEY cies. Most of our biological information con- channel of the ovipositor and not into the ovi- sists of scattered and fragmentary notes on duct. The egg becomes infected as it is laid, ovipositing or on emerged specimens. Except and the wood-destroying fungus penetrates the for the very fine and detailed work by several wood surrounding the larva as it feeds. Experi- British workers on the life history and para- ments have demonstrated that larvae can live sites of the introduced American species Sirex for at least three months on a pure culture of cyaneus Fabricius, not one single American the fungi. It has been theorized that the larvae siricid has been adequately studied. may be more or less mycetophagous and may Adults fly mostly in bright sunshine, and not need cellulose for their subsistance. Parkin females are usually more abundant than males. (1942) found the fungi associated with the lar- Males are reputed to resort to the treetops or vae and confirmed previous work which showed high ground where pairing takes place, and that the adult female becomes infected in the this may in part account for their relative scar- pupal chamber after shedding the pupal skin city in collections. Thar males are not'yet well before emergence. known is probably due to this lack of material A number of natural enemies prey upon the for study rather than through any intrinsic dif- horntails; of these, various Hymenoptera are ficulties in distinguishing them. The genitalia most important. Members of the cynipid genus seem of slight and doubtful significance for Ibalia (fig. 18) and the ichneumonid genera purposes of taxonomic distinction. Rhyssa and Megurhyssa (fig. 19) parasitize Females usually oviposit in trees weakened the larvae of Siricidae. or dying as a result of fire, disease, or other The life cycle of Zbalia requires at least injury. Occasionally they select a tree which two years. It oviposits in the young siricid to all appearance is still healthy. Ovipositing larva just before or more rarely just after it females insert their ovipositors deeply into the hatches, utilizing the oviposition bores of the wood and lay their eggs. Sometimes they are Sirex for this purpose. The parasite larva is unable to withdraw their ovipositors and die in endoparasitic. Parasitized host larvae confine this position. their borings largely to the outer parts of the The larvae bore into the trunk of the tree, tree trunk and usually make their final cell making perfectly cylindrical holes in the wood just under and approximately parallel to the and pack the frass from their borings in the surface. tunnels behind them. Tremex may complete Females of Rhyssa and Megurhyssa lay their larval growth and pupate within a year, but no eggs in the burrow of, and near to, the almost definite data are available. According to Hanson fully grown host larva or the pupa by piercing (1939), the life cycle of Sirex cyaneus normally the solid wood with their ovipositors. Feeding extends for a period of three years from egg to by the ichneumonid parasite is entirely ecto- adult, but development may ,be retarded and the parasitic and takes only a few weeks, during adult insects may not emerge from the timber which the host larva may burrow a little farther. until several additional years have elapsed. A The winter is passed by Rhyssa or Megarhyssa number of generations may be present in a as a resting larva. Pupation takes place in the single log at any one time, and this greatly spring, and the whole life cycle normally oc- complicates understanding the composition of cupies one year. the population. Woodpeckers, contrary to popular belief, Horntails are of additional interest to biol- consume few if any siricid larvae. Beal(19111, ogists because of the symbiotic relationship in his paper on the food of North American of Sirex, Urocerus, and Tremex with certain woodpeckers, fails to mention larvae in their wood-destroying fungi. The fungi are commonly diet. found apart from the siricids but with the ex- -As in previous publications in this series, ceptions of Xeris (Francke-Grosmann, 1939) all the locations of the specimens from which the female wood wasps examined have had asso- records have been taken are reported. The fol- ciated fungi. No fungi or fungal sacs have lowing abbreviations have been used for insti- been found in adult males. tutional and other collections: In the female, the fungus is contained in a California Insect Survey, University of Cal- pair of small invaginated intersegmental fungal ifornia, Berkeley (C.I.S.) sacs which project into the body cavity at the California Academy of Sciences (C.A.S.) basal end of the ovipositor, opening into the University of California, College of Agri- THE SIRICID WOOD WASPS OF CALIFORNIA 61 culture, Davis (U.C.D.) (fig. 5); eye distinctly longer than broad University of Oregon (U.O.) (fig. 17); hind tibia with two apical spurs. California State Department of Agriculture, ...... Urocerus (p. 66) Sacramento (C. S. D. A. ) California Forest and Range Experiment Station, Berkeley (C. F. R.E.S.)' Genus Sirex Linnaeus, 1761 Los Angeles County Museum (L.A.C.M.) San Jose State College (S. J.S.C.) Sirex is Holarctic in distribution with about University of Idaho (U.I.) 20 species described, 9 of which occur in I am indebted to the authoritjes of these North America; all but 3 of these are reported institutions, as well as to individuals whose from California. The females oviposit in the records are separately rsported, fo: the privi- trunks of various species of conifers in the lege of studying the specimens which have genera Sequoia, Cupressus, Libocedrus, formed the basis of this paper, and to Mrs. Pseudotsuga, Pinus, and Abies, and the larvae Celeste Green, Scientific Illustrator, for the are wood borers. Species in the genus Sirex drawings which accompany this study. lack a pale area behind the eyes dorsally; normally the cornus is not constricted; the forewing has at least the basal half of vein to Key the Subfamilies and Genera Cu, present. The California species may be of California Siricidae separated by means of the following key. Key to the California Species of 1. Antennae short, slightly swollen in middle, at most as long as head and thorax com- Sirex bined (fig. 4); cross veins CUI and 2 r-m absent; cell 2A of forewing with base of 1. Females; ovipositor present ...... 2 3A present (fig. 3); 'labial palpi 2-seg- Males 1; ovipositor absent (some species mented. Larvae bore in angiosperm trees not separable with any de~reeof certainty) T (Tremicinae) ...... I ...... Tremex colurnba (p. 70) Antennae long, filiform, much longer than Females head and thorax combined (fig. 2); cross vein 2 r-m present; cell 2A with base of 2. Ovipositor as long as the forewing, usually 3A absent (fig. 1); labial palpi +segment- longer...... 3 ed. Larvae bore in conifers (Siricinae). . 2 Ovipositor shorter than forewing .... 4 2. Head without pale areas above and behind 3. Legs with tibiae and tarsi red; cornus as the eyes; cornus of female usually not in fie. 8 ...... longicauda (p. 65) constricted in middle (except slightly in Legs entirely black or blue-black; cornus Sirex longicauda) (fig. 8); forewing with variagle, slightly mere triangular (fig. 13) at least basal half of Cu, present ...... areolatus (p. 62) ...... Sirex (p. 61) 4. Abdomen, except for basal segments, reddish Dorsal surface of head entirely pale or with brown; forewing with a transverse fuscous at least a pale spot behind each eye; cor- band beneath the stigma and a fuscous nus of female constricted itl middle (fig. apical margin; cornus mildly shouldered 15); forewing with vein Cu, usually com- (fis. 12); two basal abdominal segments pletely atrophied .....; .... 3 blue-black ...... behrensii (p. 63) 3. Head with a lateral carina behind the eye Abdomen entirely bluish black ..... 5 (fig. 2); ovipositor about half again as 5. Legs except coxae and trochanters reddish long as forewing (except Xeris macgilli- brown; wings usually nearly hyaline ex- vrayi); hindwing without a closed anal cell cept for infuscated apical margin; cornus (fig. 3.); eye almost round (fig. 16); hind not shouldered (fig. 10). cyanells (p. 64) tibia with only one apical spur ..... Legs black or dark feddish blue .....6 ...... Xeris(p. 68) 6. Forewing infuscated on apical margin and Head without a lateral carina; ovipositor at most scarcely longer than forewing; 1 TL males of californicus, longicauda, and hind wing usually with a closed anal cell obasus are unknown.

'Since going to press the name of this branch of the U.S.D.A. has been changed to Pacific South- west Forest and Range-. Experiment . Station. 62 BULLETIN OF THE CALIFORNIA INSECT SURVEY

across middle; cornus seen from above tri- Contra Costa Co.: Pleasant Hill, 9, IX- angular or shouldered. californicus (p. 63) 15-57 (T. A. Smith, C.I.S.). Wings uniformly dark violaceous; cornus Humboldt Co.: Eureka, 0,%, IX-29-49 ((2.1. distinctly shouldered (fig. 9); tarsal pads S.); 03 12-9-39 (1. Torio, C.S.D.A.); Arcata, ivory white in color, contrasting sharply 9, X-18-34(W. E. Peacock, C.S.D.A.). with the dark legs .... obesus @. 65) Marin Co.: Alpine Dam, 9, IX-1935 (Edward Ross, C.A.S):,, Muir Woods, 9, XI-15-51 (Alice Males and J. G. Edwards, Donald Burdick). 7. Head, thorax, legs, antennae, basal and Monterey Co.: Carmel 9, IX-7-30 (L. S. Slevio, C.A.S.); Monterey, XI-19-47, redwood apical abdominal segments metallic blue- $, black; abdominal segments 111-VII yellow- (W. K. Dayton, W.W.M.); 4 9, XI-18 to 19-47, redwood (W. K. Dayton, C.I.S.). ish orange areolatus (p. 62) ...... Orange Co.: Anaheim, IX-1929 (L.A. Head and thorax black or blue-black; legs 9, not above; antennae blue-black bas- C. M.) as or Sacramento Co.: N. Sacramento, 6,VI- ally reddish ...... 8 25-56 (Miller, C.S.D.A.). 8. Wings hyaline, a little smoky at the tip; fore- and midlegs except their coxae and San Francisco Co.: Cliff House Beach, trochanters reddish brown; basal segments ?, XI-26-22 (C. L. Fox, C.A.S.); San Fran- of antennae red; abdomen except basal cisco, 9, X-7-28 (E. Walther, C.A.S.); ?-, segments reddish ..... behrensii (p. 63) X-1-09 (F. X. Blaisdell, C.A.S.); 8,1X-22-08 Wings slightly yellow with a faint smoky (F. X. Williams, C.A.S.); ?, VIII-10-09 (F. band around the margins; posterior legs X. Williams, C. A.S.); d’, X-5-26 (E. Walther, bluish black except the distal segment of C.A.S.); 6,X-1-25 (F. M. Woods, C.A.S.). the trochanter; fore- and midlegs beyond San Mateo Co.: Palo Alto, 10 8,IX-8 to trochanters mostly reddish brown; t40rax XI-9-32, Cupressus macrocarpa (C.I.S.) 7 9, IX-8 to 25-32, Cupressus macrocarpa (C.I.S.); metallic blue or greenish; base of antennae reddish brown or black. cyaneus (p. 64) San Mateo, 8 $, X-18 to 25-25 (B. H. Murray, C.A.S.). Santa Clara Co.: Los Gatos, 8, X-19-16 (H. E. Burke, C.F.R.E.S.); g, IX-19-17 (F. B. Sirex areolatus (Cresson) Herbert); Palo Alto, ?, IX-15-30, Monterey cypress (H. E. Burke, C.F.R.E.S.); 52 g,20 (Western horntail) 9, IX to X-1932, Monterey cypress (H. E. Burke, C.F.R.E.S.). San Jose, P, X-30-53, in Urocerus areolatus Cresson, 1867. Trans. house (J. Wirth, Donald Burdick). Amer. Ent. Soc., 1:375. Type 9, New Mexico. Santa Cruz Co.: d, X-5-20 (E. A. Dodge, Sirex gracilis Westwood, 1874. Thesaurus Ent. (C.A.S.); Big Basin, %, VII-18-46 (Don Mead- Oxon., 114. 9. ows, L.A.C.M.); Santa Cruz, -, X-1923 (E. A. Urocerus caeruleus Cresson, 1880. Trans. Dodge, C.A.S.); % XI-9-20 (E. P. Van Duzee, Amer. Ent. Soc,, 834. 9. C.A.S.); 9, (C. Fuchs, C.A.S.). Sirex apicalis Kirby, 1882. List Hym. Brit. Sierra Co.: E. of Sierra Buttes near MUS., 1~377. d. Salmon Cr., 8, X-23-54 (C. Worthington, Geographic range: Arizona, British Columbia, Donald Burdick). California, Colorado, New Mexico, Oregon, Washington. Sonoma Co.: Mesa Grande, 2 d,VII-17-08 California records: (F. E. Blaisdell, C.A.S.); Cr, 9, X-1907,q Alameda Co.: Berkeley d‘, X-10-25 ovip. in redwood (Baumberger, C.A.S.). (C.I.S.): CY‘ M-23-15, Cupressus macrocarpa Tulare Co.: Paradise Creek, P, IX-19-30, (C.A.S.); $, IX-1933 (E. C. Zimmerman, Libocedrus decwens (J. M. Miller, C.F.R. C.A.S.); 3 ?-, XII-2-22 (S. E. Flanders, C.I.S.);, E.S.); Sequoia Nat’l. Park, d’, IX-8-18, Pinus ?, X-16-34 (C.I.S.); Leona Hts., 9, X-24-20 larnbertiana (R. D. Hartman, C.F.R.E.S.). (E. P. Van Duzee, C.A.S.); Oakland, ?, X-23- Yuba Co.: MaFysville, @, XI-1-34(Crane, 53 (J. V. Lonergan, C.S.D.A.). C. S. D. A. 1. THE SIRICID WOOD WASPS OF CALIFORNIA 63

Hosts: Specimens have been reared from, or (C.1.S.X 9, VI-1933 (C.I.S.). collected on, the following conifers: Se- Modoc Co.: Badger Township, 9, X-16-37 quoia sempemirens, coast redwood; Cupres- (K. A. Salman, C.F.R.E.S.); Crowder Flat, sus macrocarpa, Monterey cypress; Pinus 2 8, IX-1-27, Pinus ponderosa (F. P. Keen, contorta, lodgepole pine; Pinus Jeffreyi, C. F.R. E.S.). Jeffrey pine; Pinus lambertiana, sugar pine; San Diego Co.: Mt. Laguna, 2 ?, XI-26-40, Pinus radiata, Monterey pine; Li bocedrus Pinus Jeffreyi (D. DeLeon, C.F.R.E.S.); 1 d’; decurrens, incense cedar; Pseudotsuga 2 9, X-27-41, Pinus Jeffreyi (D. DeLeon, taxifolia, douglas fir; and on cedar. This C.F. R. E .S). species commonly attacks the coast red- Santa Clara Co.: 9, 1P28 (C. D. Duncan, wood, cypress, and cedars and is found less S.J.S.C.); Santa Clara 9, X-20-31 (Forest frequently on pines. Smith, S. J.S.C.). Parasite:*.* The parasite Ibalia ensiger was Santa Cruz Co.: Santa Cruz, 2 P (J. P. reared by Flanders from Berkeley speci- Strohbeen, C.A.S.). mens, XII-2-22. Sonoma Co.: Santa Rosa, 5 ?, XI-7-30, lap- Discussion: anese pine (H. E. Burke, C.F.R.E.S.); 6 ?, In California this siricid is predominantly XI-2-30, Monterey pine (S. Lockwood, C.S.D.A.). coastal in distribution. The steel blue body Stanislaus Co.: Ceres, 9,V-29-27 (D. Davis, and legs and the long ovipositor, which is as C.A.S.). long as the forewing, distinguish the female Hosts: Specimens have been reared from, or of this species. Our only other Sirex with a collected on, the following conifers: Pinus long ovipositor is S. longicauda Middlekauff Jeffreyi, Jeffrey pine; Pinus lambertiana, which has the tibiae and tarsi red rather than sugar pine; P. ponderosa, yellow pine; P. bluish-black. The cornus is fairly uniform in radiata, Monterey pine; Cupressus macro- shape but may, in some species, be slightly carpa, Monterey cypress; and Japanese more triangular than shown in figure 13. Sever- pine. al females from Palo Alto, reared from Monter- Parasite: A log of Pinus ponderosa from Oak- ey cypress, are extremely small, not exceeding hurst produced a number of s. behrensii as three-quarters of an inch from head to tip of well as two female specimens of Zbalia ovipositor. This phenomenon of size variation ensiger Nort. is rather common in the Siricidae. Discussion: Essig (1926) reports females ovipositing In spite of the fact that this species has in cured redwood lumber in the yards of Hum- not yet been recorded from Oregon, it is reason- bolt and Mendocino counties in California. able KO assume that it will be taken there even- tually. It has been collected both north and south of Oregon, and suitable hosts occur ex- tensively in the state. The reddish brown abdomen beyond segment I1 and the shape of the cornus (fig. 12) will Sirex behtensii (Cresson) serve to distinguish most of the females of this species. Several females have been seen Urocerus behrensii Cresson, 1880, Trans. with considerable black on the sides of the Amer. Ent. Soc., 8:35. Type ?, California. abdomen. Geographic range: California, Nevada, Wash- ington. California records: Sirex californicus (Ashmead) Alameda Co.: Berkeley, 9, IV-29-26 (E. C. Van Pyke, C.A.S.); XI-25-22, ovipositing in Pawwus californicus Ashmead, 1904, Canad. dead Monterey cypress, (S. E. Flanders, C.I.S.); Ent., 36:64. Type 9, California. 8,X-20-15, (E. P. Van Duzee, C.I.S.); Liver- Geographic range: British Columbia, Califor- more, ?, X-1909 (6.A.S). nia, New Mexico, Oregon, Washington. Madera Co.: Bass Lake 9, X-12-34, reared California records: from sugar pine (C.I.S.); 9, X-21-34 (C.I.S.); Alameda Co.: Berkeley, $, X-2-25, Monterey Oakhurst, 2 d’, 29, XI-1955, Pinus ponderosa pine (E. C. VanDyke, C.A.S.). (C. B. Eaton, W.W.M.). El Dorado Co.: Meyers Station, ?-, IX-21-16, Mariposa Co.: Yosemite Valley, 6 V-23-42 Pinus Jeffreyi (F. B. Herbert, C.F.R.E.S.); 64 BULLETIN OF THE CALIFORNIA INSECT SURVEY

?, VIII-31-15, Pinus mwrayana (F. B. Herbert, Newfoundland, and Hudson Bay, west to C.F.R.E.S.); 2 ?, IX-2-15, Pinus mwrayana Vancouver Island; south along the Rocky (F. B. Herbert, C.F.R.E.S.). Mountains to New Mexico; and in the Paci- Inyo Co.: Parcher's Camp, Bishop Cr., 9, fic coa_st states of Oregon, Washington, and IX- 10-39 (L. A. C. M. ). California. It has been established in Brit- Lassen Co.: Facht, 9, VII-20-21 (J. 0. Mar- ain where it is often imported in timber. tin, C.A.S.). California records: Modoc Co.: Warner Mts., Davis Creek, Alameda Co.: Oakland, 9 IX-15-48, pine 5,600-6,000' elev., 3 ?, VII-7 to 28-22 (A. W. lumber from Sierra (C.I.S.). Lindsey, C.A.S.); Crowder Flat, 2 9, IX-1-27, Amador Co.: Dew Drop Camp, 3 ?-,emerged Pinus. ponderosa (F. P. Keen, C.F.R.E.S.). 111-3 1-55, Abies concolor Hopkins No. 34-02OE Plumas Co.: Quincy, 4 mi. W., p, VII-7-49 (R. G. Struble, C. B. Eaton, C.F.R.E.S.). (W. H. Wade, C.I.S.). Contra Costa Co.: Lafayette, $, VII-6-50 San Bernadino Co.: Swartout Valley, 9, (E. 0. Essig, C.I.S.). IX-23-23, Pinus leffreyi (H. E. Burke, C.F.R. Los Angeles Co.: Alhambra, 9 (E. T. 01- E.S.). son, L.A.C.M.); Los Angeles, 9, IX-8-26, Tulare Co.: Camp Nelson, 4,700' elev., 2 spruce (Western Hardware Co., C.F.R.E.S.). 9, IX-18-14 (R. L. Beardsley, C.A.S. . Madera Co.: Miami Range Station, 2 ?, VI- Tuolumne Co.: Pinecrest, 3 d , XtlO to 1942 (C.I.S.). 16-33 (R. P. Allen, C.A.S.); Tenaya Lake, ?, Mendocino Co.: Mendocino, < XI-15-57 VIII-20- 17, Pinus mwrayana (J. E. Patterson, (J. R. Helfer, Don Burdick). C.F.R.E.S.); Yosemite, 9, VIII-1933 (W. G. Modoc Co.: Buck Creek Ranger Sta., 5 ?-, Scott, C.I.S.). 5 6,X-25-29, Pinus ponderosa (A. Wagner, Hosts: Specimens have been reared from, or C.F.R.E.S.); Crowder Flat, 9, IX-1-27, Pinus collected on, the following conifers: Pinus ponderosa (F. P. Keen, C.F.R.E.S.). Jeffreyi, Jeffery pine; Pinus rnurrayana San Francisco Co.: San Francisco, 9, XI-5- (= contorta), lodgepole pine; Pinus pon- 09 (C.A.S.); 9, VII-2-17 (E. P. Van Duee,C.A. derosa; yellow pine; Cupressus macrocarpa, S.); ?, IX-29-25 (R. M. Woods, C.A.S.); P, 910- Mont erey cypress; Pseudot suga rnenzies ii, 33 (C. E. Browning, C.S.D.A.).' Douglas fir. ' 'Siskiyou Co.: Yreka, 9 (H. E. Burke, C.F. Discussion: R.E.S.). This fairly common species is similar to Tuolumne Co.: Pinecrest, ?, IX-28-33 the less abundant obesus in having the body (C.I.S.). and appendages dark blue and the ovipositor Hosts: Specimens have been reared from, or short. It differs from obesus in having an in- collected on, the following conifers: Abies fuscated band around the apical margin of the balsamea, balsam fir; Abies concolor, white wings and one across the wing beneath the fir; douglas fir; pine; and spruce. stigma. The basal and central areas of the Parasite: Rhyssa lineolata (Kirby). wings are nearly hyaline. The male is unknown. Biology: lhe life history has been carefully studied Sitex cyaneus Fabricius by Chrystal (1928, 1930) and Chrystal and Meyers (1930). The following notes are ex- Sirex cyaneus Fabricius, 1781. Species Insect- tracted from their observations. torum, 1:419. Type ?, North America. The flight period seems to-be at its height Sirex duplex Shuckard, 1837. Mag. Nat. Hist. from July to October; some wasps have been (n.s.), 1:630. d', 9. taken as early as June and as late as Novem- Urocerus nitidus Harris, 1841. Rpt. Ins. Mass., ber. The adults fly in bright sunshine making a Injurious to veg., p. 391. 9. characteristic noisy buzz. As with other spe- Sirex varipes Walker, 1866. In Lord, Natura- cies of Siricidae males appear less frequently list in Vancouver Is. and B.C., 2:342. ?. in collections. Workers in Denmark observed Sirex hirsuus Kirby, 1882. List Hym. Brit. that siricids pair in the treetops, after which Mus., 1:380. 3. the females descend to the lower levels for Geographic range: This species is found in egg laying. the Hudsonian and Canadian Life Zones Females normally oviposit in warm sunny from Georgia north into New Brunswick, weather. Before the final site for oviposition THE SIRICID WOOD WASPS OF CALIFORNIA 65 is selected the female closely examines the Sirex longicauda Middlekauff bark with the antennae and the ovipositor. In a matter of less than ten minutes she begins to Sirex longicauda Middlekauff, Pan- oviposit and may lay from 1 to 7 eggs in the Pac. Ent., 24 (4):189-190. Tyi:*!: Cali- oviposition tunnel, spaced at intervals while fornia, withdrawing the ovipositor: 3 to 4 fusiform Geographic range: California. translucent eggs per tunnel seem to be average. California records: Sometimes the females are unable to withdraw Alameda Co.: Berkeley, 9, IV-1932, Abies the ovipositor and die in this position. The concolor (C.A.S.). Holotype. tunnel penetrates the wood at a right angle to Madera Co.: Miami Ranger Station, $, the axis of the trunk, or at a slant, to a depth VI- 1942 (C.F.R.E.S.). Paratype. of G to 20 mm. Chrystal (1928) found by dissec- Mariposa-Tuolumne cos.: Yosemite Nat’l. tion that females in England possessed 300 to Park, 8, IX-25-33 (C.I.S.); 5 8, K-28-33, white 400 eggs on the average. They are 1.25 to 1.5 fir (C.I.S.). mm. in length and are markedly constricted at San Bernardino: Hanna Flats, ?, IX-28-52 at the posterior end and bluntly rounded at the Menke, Jr., Arnold Menke); Silver Lake, other. , IX-1932 (H. Plank, C.S.D.A.). Host: Abies concolor, white fir. The incubation period is 3 to 4 weeks, after Discussion: which the young larva bites its way out of the This species closely resembles areolatus egg and begins to burrow into the wood at in body color and length of ovipositor, but right angles to the oGiposition tunnel. The may be readily separated on the basis of the larva tunnels in the outer sapwood at first; reddish tibiae and tarsi and the slightly more when it has attained the length of 8 to, 9 mm. constricted cornus. First described in 1948, it turns inward toward the heartwood. After it is still rare in collections; the male is un- reaching the heartwood the larva then curves known. round, returning toward the surface in prepara- tion for pupation and the emergence of the adult. Chrystal (1929) feels that 10 inches to Sirex obesus Bradley 1 foot is a fair estimate of the maximum length of the tunnel. The larva molts three to four Sirex obesus Bradley, 1913, Jour. Ent. Zool., times and packs the cast skins with the frass 5:12. Type?, Arizona. at intervals in the tunnel. The terminal spine Geographic range: California, Arizona. on the iarva is €or the purpose of packing the California recoras: frass or boring dust in its tunnel as well as Mari osa-Tuolumne cos.: Yosemite Nat’l. serving as a terminal support, being driven into Park, B , VIII-14-35, Pinus contorta, (G. R. the sides of the tunnel for this purpose. Struble, C.F.R.E.S.). Nevada Co.: Nevada City, 9, VIII-1306 (J. The pupal chamber is formed at the end of P. Baumberger, C.A.S.). the larval tunnel and in a direct line with the Host: Pinus contorts, lodgepole pine. outside. The adult cuts its way to freedom Disc us sion: through the remaining distance which is usually This relatively rare species is similar to less than three-quarters of an inch. Males are californicus in having the deep blue head, normally somewhat closer to the surface than body, and appendages and a short ovipositor females. The duration of the pupal stage is which seldom projects more than 2 to 3 mm. 5 to 6 weeks. The minimum period for the beyond the tip of the cornus. It may be dis- complete cycle is two years. tinguished by the dark, violaceous wings and white tarsal pads. The male is unknown. Discussion: A Holarctic species, generally believed Genus Urocerus Geoffroy, 1762 to be American in origin. Benson (1951) con- siders it to be adventitious in Great Britain. Members of this genus have been found in Eu- Some authors have confused this species rope, Palearctic Asia, Japan, North Africa, with the Palearctic juvencus (L.) or have de- and North America. About 20 species and sub- scribed it as a subspecies of juvencus. The species have been described, 5 of which occur systematics of the species need clarification. in North America and 3 in California. It con- 66 BULLETIN OF THE CALIFORNIA INSECT SURVEY

tains ow largest species, some members at- Urocerus albicornis (Fabricius) taining a body length (head to tip of cornus) of nearly 40 mm. Females oviposit in the trunks Sirex albicornis Fabricius, 1781, Species In- of various conifers in the genera Abies, Pinus, sectorum, 1:419. Type ?, North America. Tsuga, Picea, Pseudotsuga, and Libocedrus. Urocerus abdominalis Harris, 1841, d, Rpt. Species in this genus possess in common Ins. Injurious to Veg., p. . with Xeris a distinctive pale spot behind the Suez stephensi Kirby, 1882, $92, List Hym. Brit. eye, a constricted cornus and vein CUI in the Mus., 1:375. forewing which is usually completely absent. Geographic range: From -British Columbia, They may be distinguished from Xetis as fol- northern Ontario, Nova Scotia, and New- lows: head lacks a lateral carina, the ovipositor foundland south to Louisiana. South in the is usually shorter than the forewing,hind wing Rockies to New Mexico and in the Pacific possesses a closed anal cell, the eye is oval coast states to California. A common insect instead of nearly round, and the hind tibiae in the Canadian Life Zone extending into possess two apical spurs. the Hudsonian. California species may be separated by California records: meansof the following key. Humbolt Co.: Weott, 9,VII-13-29 (E. D. Van Dyke, C.A.S.). Madera Co.: Miami Ranger Station, ?, VI- 1942 (C.I.S.). Shasta Co.: Sweet Briar Camp, 8, (C. L. Key to the California Species of Fox, C.A.S.). Castella, VII- 1912 (Baumberger, 1915). Urocetus Hosts: Specimens have been reared from, or collected on, the following conifers: Abies 2. Females; ovipositor present ...... 2 balsamea, balsam fir; pine; hemlock; spruce; Males; ovipositor absent ...... 4 firs; Douglas fir; cedar. Parasites: Megarhyssa nortoni guebecensis (Prov.) and Rhyssa Zineolata (Kirby) have Females been reared from this species. Discussion: 2. Abdomen black with segments VII and VI11 This species is relatively uncommon in Cal- and sometimes 11, yellow ...... ifornia, and only a few locality records are ...... gigas flavicomis (p. 68) available. Abdomen usually entirely black or blue The female is reported to prefer freshly black...... 3 killed wood in which to oviposit. The life cy- 3. Wings golden yellow; flagellum of antennae cle is reported by Belyea (1952) to last t&o and basal half of tibiae and metatarsi years from egg to adult. Other authors state golden yellow; a large light-colored spot that the cycle may be completed in one year. behind eye ...... californicus (p. 66) The larval burrows run in all directions and Wings infuscated, brown; most of flagellum, are closely packed with a very fine dustlike a spot behind eyes, sometimes lateral frass. spots on the abdomen and the bases of the Our only other Urocetus with entirely black tibiae and basitarsi white .albicomis (p. 66) abdomen is californicus. The brownish infuscated wings .will serve ta separate albicornis Males females from the golden-winged females of californic us. 4. Abdomen reddish brown; wings golden yellow; legs and antennae reddish brown ...... californicus (p. 66) Abdomen with segments I11 to VI yellow, the Urocerus californicus Norton remainder black; wings nearly hyaline. . 5 5. Hind basitarsus 4.1 to 5.G times as long as Urocerus albicornis var. californicus Norton, broad ...... albicornis (p. 66) 1869, Trans. Amer. Ent. Soc., 2:360. Type Hind basitatsus 6.4'to 8.1 times as long as P, California. broad ...... gigas flavicornis (p. 68) Urocerus fulvus Cresson, 1880, 3, Trans. THE SIRICID WOOD WASPS OF CALIFORNIA 67 Amer. Ent. Soc., 8:35. Napa Co.: St. Helena, 9,VII-7-09 Urocerw fkrvipennis Kirby, 1882, ?-,List Hym. Nevada Co.: Sagehen, nr. Hobart Brit. Mus., 1:380. IX-5-57 (E. G. Linsley, C.I.S.); Donner Summit, Geographic range: Alpine regions in Arizona, 9, V-1940 (E. Swift, C.S.D.A.); Washington, 5 British Columbia, California, Colorado, Ida- mi. S., 9, VI-30-57 (H. W. Michalk,U.C.D.). ho, Mexico, Neva.da, New Mexico, Oregon, Placer Co.: Cisco, ?, VII-1924 (F. E. Blais- Utah, Washington. dell, C.A.S.); ?- VII-1911 (C. Van Geldern, Calif or nia records: C.A.S.); 6 8,9, VI -1910 IC. Van Geldern, Alameda Co.: Oakland, %, VIII-22-58(C.I.S.). C.A.S.); Mich. Bluff, 9 (E. R. Leach, C.A.S.); Calaveras Co.: Camp Wolfboro, 6,VIII-1- Middle Fork of American River, 9, VIII-15-07 39 (C.I.S.); DorTington, 2 ?-,VIII-7-33 (R. P. (C. L. Fox, C.A.S.); Whiskey Creek, 2?, XI-3- Allen, C.A.S.); 3 ?, IX-13 to 14-33 (R. P. 48 (Peter C. Ting); Carnelian Bay, Lake Tahoe, Allen, C.A.S.). ?, IX-24-56 (R. M. Bohart, U.C.D.). El Dorado Co.: China Flat, 2 $, VIII-2-49 Plumas Co.: Hartless, 9, IX-7-13 H. E. (J. W. MacSwain, C.I.S.1; Eagle King Mine, Burke, C.F.R.E.S.); McKinney Creek, b, IX- Grizzly Flats, ?, IX-8-38 E. P. Chace, L. A. 1914, (C.F.R.E.S.); Meadow Val., 5,000-6,000 ' C.M.). Fallen Leaf Lake, 6, VI-24-28 (L.A.C. elev., O, VII-20-39 (H. T. Reynolds, C.I.S.); M.); $,VI-23 to 30-1934 (Carl D. Duncan, S. J.S. 29, VII-27-24 (E. C. Van Dyke, C.A.S.); Onion C.); Pyramid Ranger Sta., 2 9, VIII-1-49 (J. W. Valley, 9, VIII-11-13 Abies concolor, (H. E, MacSwain, C.I.S.); 9, VIII-10-15 (F. B. Herbert, Burke, C.F.R.E.S.); 9, IX-15-13 (H. E. Burke, C.F.R.E.S.); 29, VIII-4 to 6-15 (C.F.R.E.S.); C.F.R.E.S.); Quincy, 9, IX-6-41 (W.W.M.); 10 9, VI11 to IX-1914 Abies concolor (H. E. Bucks Lake,$, VIII-29-56 (R. F. Wilkey, C.S. Burke, C.F.R.E.S.); $, IX-3-14, Libocedrw D. A. ). decurrens (H. E. Burke, C.F.R.E.S.); Snowline Riverside Co.: Marion Mt. Camp, San Jacin- Camp, 9, VI-29-48 (P. D. Hurd, C.1.S.). 3 ?, 6, to Mts., ?, VII-1-52 (1. W. MacSwain, C.I.S.). VII-14-48 (A. Bartel, C.I.S.); Vade, 9, Abies San BetnardinoCo.: Beak Lake, 9, VII-4-19 mugnifica (H. E. Burke, C.F.R.E.S.); Echo (J. D. Martin, C.A.S.); Hanna Flat, $, VI-20- Lake, 39, VIII-10-24 (E. 0. Essig, C.I.S.);?, 52 (Arnold Menke, Jr., Arnold Menke); San VIII-14-55 (H. R. Moffitt, U.C.D.). Lake Tahoe, Betnardino Mts., 7,50O'elev.,?, VII-2-17 (J. D. P, 28, IX-1929 (C.1.S.). Martin, C.A.S.); Bluff Lake, 2 $, July-August, Fresno Co.: Huntington Lake, 7,000' elev., 1948 (G. Chrisman, C.I.S.) Lake Arrowhead, 4 9, VII-lS-19(?) (E. P. Van Duzee, F. C. P, 6-8-40 (N. Gtatz, L.A.C.M.); g,VIII-13-40 Clark); 2 9, VII-21-30 (L.A.C.M.); ?-,VII-27- (N. Gratz, L.A.C.M.). 19(?) (E. P. Van Duzee, C.A.S.); S. Fork Kings Santa Clara Co.: Palo Alto, 9, VIII-9-54 (R. River Canyon, 5,000' elev., ?, VII-4-10 (E. C. Dixon, U.C.D.). Van Dyke, C.A.S.). Santa Cruz Co.: Big Basin, 2 ? VIII-7-56 Los Angeles Co.: Crystal Lake, ?, VII-1-52 (Don Meadows, L.A.C.M.); Felton, 9, IX-8-48 (L. A. Stange, L.A.C.M.). (Ralph Nix, Donald Burdick); Soquel, 9, VIII- Madera Co.: Bass Lake, P, VIII-31-34 (C.I. 11-32, white fir (C.F.R.E.S.). S.); Whiskey Creek, 9,VIII-3-31 (K. A. Salman, San Francisco: San Francisco, 9, VIII-27- C.F.R.E .S.). 58 (C.I.S.). Mariposa Co.: Pohono Trail, Yosemite Nat'l. Shasta Co.: 6 VII-6-21 (J. A. Kusche, C. Park, 3, VIII-8-33 (C.I.S.); Yosemite Valley, A.S.); Hat Creek,$, VIII-7-54 D. D. Linsdale, 9, VII-8-21 (E. C. Van Dyke, C.A.S.); 39, 3, C.I.S.); Lassen Nat'l. Park, k , IX-21-40 (T. (C.I.S.); Tuolumne Meadows, 8,500' G. Aitken, C.I.S.). , VII-1915 (C. L. Fox, C.A.S.). Sierra Co.: ?, X-1927 (C.A.S.); Calpine, 9, Mariposa-Tuolumne cos.: Yosemite Nat'l. VIII-27-48 (J. W. MacSwain, C.I.S.); Gold Lake, Park, 7,00017,750' elev., ?, VII-30-38 (C.I.S.); ?, VII-20-21 (C. L. Fox, C.A.S.); Sierra City, Yosemite, 49, IX-28-33 white fir, (C.F.R.E.S.); 9, X-1-36, cedar (A. Michie, C.S.D.A.). 9, IX-18-13 (J. J. Sullivan, C.F.R.E.S.); 9, Siskiyou Co.: Shasta Retreat, 2,416' elev., WII-1914 (1. J. Sullivan, C.F.R.E.S.); 9, VI- 9, VII-3-05 (F. E. Blaisdell, C.A.S.); Shasta 24-29 (J. Homing, springs, P VIII-1-20 (c. E. Masson, c.A.s.); Modoc Co.: , VII-10-22 (C. L. Big Flat, 9, Vm-4-31 (R. L. Usinger, C.I.S.); Fox, C.A.S.); Modoc Nat*l. Forest, ?-, XI-6-30, Yreka, 9,VII-20-40 (J. 0. McKitlney, C.S.D.A.); Abies concolor (C. A. Johnson, C.F.R.E.S.). Sonoma Co.: Santa Rosa, 9 (no date) (Lois 68 BULLETIN OF THE CALIFORNIA INSECT SURVEY

B. Stiles, Donald Bucdick). Sirex latifasciatus, Westwood, 1874, 6, The- Tehama Co.: Red Bluff, 9, VIII-1929 (L.A. saurus Ent. Oxon., p. 114. C.M.). Vrocerus riparius MacGillivray, 1893, ci’, Trinity Co.: Carrville, ?, VII-3-13 (E. C. Canad. Ent., 24:244. Van Dyke, C.A.S.). Geographic range: Northern coniferous belt of Tulare Co.: Giant Forest, 6 9, VIII-20-19, North America south along the Rocky Moun- Abies concolor (A. Wagner, C.F.R.E.S.); Min- tains to Arizona and New Mexico, Mexico eralking,?, IX-14‘-53 (R. W. Parker, L.A.C.M.); and in the Pacific coast states of Washing- Sequoia Nat’l. Park, 2 9, IX-10-18, Abies con- ton, Oregon, and California. Has been intro- color (R. D. Hartman, C.F.R.E.S.); ?, VII-26- duced into Britain in timber but is. not es- 39 (C.A.S.); Sequoia Nat’l. Park, Crystal Cave tablished. area,-more than 9,000 ‘elev., ?, VII-1940 (C.A. California records: S.); Three Rivers, ?, IX-10-52 (1. W. Hiner- El Dorado Co.: Lake Tahoe, 9, VII-1937 man, C.I.S.); 29, VII-I918 (A. Wagner, C.F.R. (C.I.S.). E.S.); Sherman Pk. National Forest, ?, X-6-37 Inyo Co.: Manigens Meadow, 9, VII-1-47 (Romain Young, C.S.D.A.). (C.I.S.). Tuolumne Co.: .?, JX-14-56, on dead pon- Hosts: Specimens have been reared from, or derosa (Roberta Hood, S.J.S.C.); Long Barn, 2 collected on, the following conifers: spruce, 9, 193 1, Abies concolor (R. G. Struble, C.F.R. pine, fir, Douglas fir. E.S.); ‘Pinehurst, 11 9, VI1 to VIII-1930, Abies Discussion: concolor (R. G. Struble, C. F.R.E.S.); 2 9, VI- A somewhat smaller species than californi- 24, 26-31 Abies concolor (R. G. Struble, C.F. cus. It is distinctively colored having abdom- R.E.S.); k, VII-12-27 (C.A.S.); 9, VII-10-29, inal segments VI1 and VI11 and sometimes 11, white fir (C.F.R.E.S.); 9, VIII-1936 (C.I.S.); yellow. Strawberry, 9, VII-8-51 (R. W. Morgan, C.I.S.). Hosts: Specimens have been reared from, or collected on, the following conifers: Abies Genus Xeris A. Costa 1894 balsamea, balsam fir; Abies concolor, white fir; Abies mgnifica, California red The genus Xeris is Holarctic in distribution fir; Libocedrus decurrens, incense cedar; with but four known species. Three species, Douglas fir; lodgepole pine. one of which is Holarctic, are recorded from Discussion: North America. A fourth spesies is confined to It is the largest and possibly the commonest the Himalayas. western siricid; the adult female may exceed The species of this genus possess a dis- an inch and a half in length from head to tip of tinctive carina behind the eye; the hind wing cornus. Although californicus may attack sev- is without a closed anal cell; the hind tibia has eral other genera of conifers it is primarily a but one apical spur; and the eye is nearly fir feeder, readily attacking the balsam, white round. Two of our species have the ovipositor and red firs. It is our only siricid with a post- longer than the body, and all possess a white ocular white spot, black abdomen, and golden- spot behind the eye. yellow-colored wings and antennae. The Bucks Lake female has a small yellow spot laterally above the spiracle on abdominal Key to the California Species segment VIII. of X eris

Urocerus gigas flavicornis 1. Females; ovipositor present ...... 2 Males; ovipositor absent ...... 4 (Fabricius) Females Sirex flavicornis Fabricius, 1781, Species In- sectorum, 1:418. Type ?, Labrador. 2. Cornus not constricted in middle or only Sirex bizonatus Stephens (n.n.), 1892, Syst. slightly so (fig. 14); tibiae and tarsi Cat, Brit. Ins. Mand., p. 342. usually considerably lighter than femora; Urocerus abdominalis Harris, 1841, 8, Rept. antennae basally darker than apical’ part Ins. Mass., p. 392. of flagellum ...... macgillivrayi (p. 69) THE SIRICID WOOD WASPS OF CALIFORNIA 69

Cornus distinctly constricted in middle Amer. Ent. Soc., b:25, 67. (fig. 15); legs mostly unicolorous reddish Urocerus indecisus MacGillivray, 1893, a”, ...... 3 Canad. Ent., 25:243. 3. Abdomen except first segment, reddish ... Geographic range: High coniferous forests of ...... morrisoni (p. 69) northwestern United States into Canada. Abdomen entirely black ...spectrum (p. -70) Arizona, British Columbia, California, Col- orado, Nevada, Oregon, Utah, Washington. California records: Alpine Co.: Lake Alpine, 9, VII-13-35 (R. Males P. Allen, C.A.S.). Calaveras Co.: Camp Wolfboro, P, VIII-1- 4. Abdomen black ...... spectrum (p. 70) 39 (C.I.S.); Dorrington, ?, VII-7-33 (R. P. Abdomen mostly reddish ...... 5 Allen, C.A.S.). 5. Postocellar area very rugose; median fur- El Dorado Co.: Echo Lake, 9, VII-13-31 row shallow, broadly rugose (fig. 6) .... (E. 0. Essig, C.I.S.); d’, VII-12-24 (P. 14...... macgillivrayi (p. 69) Wymore, C.A.S.); 9, 8, VII-16 to 18-33, red Postocellar area sparsely pitted not rugose, fir (C.I.S.); 79, 7 6,VII-10-24 ovipositing in polished, a distinct, narrowly rugose me- red fir (E. 0. Essig, C.I.S.); 6, VII-20-28 (E. dian furrow (fig. 7) .....morrisoni (p. 69) 0. Essig, C.I.S.); Fallen Leaf Lake, 9, VII- 17-15 Abies concolor (H. E. Burke, C.F.R.E. 9, VII-1915 (L. S. Rosenbaum, C.A.S.); Xetis macgillivrayi Bradley s$,VII-16-15, Abies co-ncolor (F. B. Herbert, C.F.R.E.S.); 7$, VIII-1931 (0. H. Swezey, C. Xeris macgillivrayi Bradley, 1913, Jour. Ent. A.S.); 9, VII-24-15 (E. C; Van Dyke, C.A.S.); Zool., 5:24. Type 9, Washington. Glen Alpine Creek, 3 ?, VII-16-20 Geographic range: California, Washington. Duzee, C.I.S.); Snowline Camp, California records: (0. E. Myers, Donald Burdick); 2?, 2 6, VI- Alameda Co.: Berkeley, 9, XI-25-22 (S. 0-48 (C. D. MacNeill, Donald Burdick); 14 Flanders, (C.I.S.); P, XX-25-22, Montetey cy- , 7 8, VI-25 to 30-48 (J. W. MacSwain, A. press (S. Flanders, C.I.S.). Bartel, L. W. Quate, D. Carter, P. D. Hurd, Modoc Co.: Hackamore, y, IX- 1930, attract- C.I.S.); 8 ?, 2 8, VU-3 to 4-48 (P. D. Hurd, ed to fire (Wagner, C.F.R.E.S.); Willow Ranch, C.I.S.); 4 9, 2 8, VII-7-48 (P. D. Hurd and S. 6 9, Cr, 11-1934, Libocedrus decurrens (K. A. A. Sher, C.I.S.). Salmon, C.F.R.E.S.). Fresno Co.: Huntington Lake, 3 9, 2 d’, San Mateo Co.: 5 9, X-21 to 25-25 (B. H. VII-12 to 18-19 (E. P. Van Duzee, C.A.S.). Murray, C.A.S.). Madera Co.! Soquel Meadow, Willow Creek, Santa Clara Co.: Palo Alto, 76 9, 114 8, ?, VII-1952 (W. C. Day, C.A.S.). VI11 to X, 1932, Monterey cypress (H. E. Mariposa-Tuolumne cos.: Yosemite Nat’l. Burke, C.F.R.E.S.). Park, 9, VII-10-13, Pinus contorta (J. J. Sul- Tuolumne Co.: Pinecrest, #, X-16-33 (R. livan, C.F.R.E.S.); 9, VIII-1-38 (C.I.S.). P. Allen, C.I.S.). Modoc Co.: Hackamore, 2 9, VII-2-50 (M. Hosts: Cupressus macrocarpa, Monterey cy- Wasbauer, Donald Burdick); 9, VII-5-50, Pinus press; Libocedrus decwrens, incense cedar. ponderosa (M. Wasbauer, Donald Burdick). Discussion: Napa Co.: Mt. St. Helena,?, V-22-26 (M. C. The unconstricted cornus of the female, the Van Duzee, C.A.S.). darker femora, and basal part of the antenna Nevada Co.: Sagehen, nr. Hobart Mills, 9, will distinguish macgillivrayi from our other VII-16-54 (R. H. Goodwin, C.I.S.). species. A female was reported attracted to Placer Co.: Cisco, 8, VII-1911 (C. Van fire. Geldern, C.A.S.); Towle, 9, VII-15-33 (C.I.S.). Plumas Co.: Bucks Lake, 8, VII-14-49 (J. W. MacSwain, C.I.S.); Meadow Valley, 3,500- Xeris morrisoni (Cresson) 4,000‘ dev., &’, VI-1G24 (E. C. Van Dyke, C.A.S.); Onion Valley, $, V-lFlG, Abies con- Urocerus morrisoni Cresson, 1880, Trans. color (H. E. Burke, C.F.R.E.S.). Amer. Ent. SOC., 8:35. Type 9, Colorado. Riverside Co.: Marion Mt. Camp, San Jacin- Urocerus tarsalis Cresson, 1880, 9, Trans. to Mts.,q7 VII-1-52(W. V. Garner, C1.S.). 70 BULLETIN OF THE CALIFORNIA INSECT SURVEY San Bernadino Co.: Falls Public Camp, 6, Turkestan, the Caucasus, Himalayas, and VII-2-56 (Arnold Menke, L.A.C.M.). Atlas Mountains. Introduced but not es- San Francisco Co.: San Francisco, 9, VII- tablished in Britain. In North America it 1908 (E. C. Van Dyke, C.A.S.). has been recorded from Alaska, British Co- Santa Cruz Co.: Santa Cruz Mts., 9, V-20-26 lumbia, California, Colorado, Maine, Ne- (J. A. Kusche, C.A.S.). vada, New Brunswick, New Hampshire, Shasta Co.: 3 mi. S.E. Mt. Lassen, 9, VII- Nova Scotia, Oregon, Utah, Washington. 18-53, Abies rnugnifica (H. W. MacSwain, C.I. California records: S.). El Dorado Co.: Echo Lake, 6 c?’, VII-10 to Siskiyou Co.: McCloud, ?, VII-27-14 (E. C. 24-24 (E. 0. Essig, C.I.S.); Snowline Camp, Van Dyke, C.A.S.); 2 d’, VII-2-14 (E. C. Van 9, VII-7-48 (J. W. MacSwain, C.I.S.); $, VIII- Dyke, C.A.S.). 3-48 (J. W. MacSwain, C.I.S.). Trinity Co.: 6, Pinus ponderosa (C.F.R.E. Lassen Co.: Blue Lake, 2 mi. N.W., 8, S. ). VII-19-47 (D. W. Adams, C.I.S.)‘ Tulare Co.: Sequoia Nat’l. Park, 9, summer Mariposa Co.: Lake Tenaya, Yosemite Nat’l. . 1937 (Jack Applegarth, S. J.S.C.). Park?, VII-24-18, Pinus murrayana (J. E. Pat- Tuolumne Co.: Mather, 6,VII-20-30 (E. C. terson, C.F. R.E.S.); Tuolupne Meadows, $, Zimmerman, C.A.S.); Pinecrest, 9, VI-28-31 VII-25 to VIII-5-15 (C. L. FOX,C.A.S.). . (C.F.R.E.S.); 29, VII-1 to 15-30 (G. B. Struble, Mariposa-Tuolumne cos.: Yosemite Nat’l. C.F.R.E.S.); 9, VII-7-29, Abies concolor (W. Park, 9, VII-10-13, Pinus contorta (J. J. Sulli- D. Edmonston, C.F.R.E.S.); 2 9, VI-26-29, van, C.F.R.E.S.). Abies concolor (H. E. Burke, C.F.R.E.S.); d’, Plumas Co.: Bucks Lake, 9, VI-23-49 (J. VII-15-29, Abies concolor (H. E. Burke, C.F. W. MacSwain, C.I.S.). R.E.S.); Strawberry Lake, 8,VII-1949 (Brown- Host: The lodgepole pine, Pinus contorta, is ing, C.S.D.A. ). probably the favorite host in California. Hosts: Abies concolor, white fir; Abies mag- Parasite: Rhyssa persuasoria L. nifica, red fir; Pinus contorta, lodgepole Discussion: pine; Pinus ponderosa, yellow pine. Females of this dark-colored species may Parasites: MacSwain and others took large be separated from morrisoni on the basis of numbers of Megarhyssa nortoni nortoni the entirely black abdomen. (Cresson) from trees where morrisoni had be en ovipositing. Discussion: Genus Tremex Jurine, 1807 This common Xeris is widely distributed in the mountains California where it attacks More than 20 species in this genus are known of from Europe, Palearctic and tropical Asia, mostly firs and lodgepole pine. Essig (1926) Borneo, New Guinea, Africa, and North Ameri- reports taking females ovipositing in dead and dying white fir and lodgepole pine. The reddish ca; some species are of doubtful validity. One species, with several phases, has been abdomen will serve to distinguish it from the color closely related spectrum. described from North America. The genus Tremex is so distinctive that it is placed in a separate subfamily. The antennae Xeris spectrum (L’innaeus) are short, more or less fusiform‘at most as- Ichneumon spectrum Ljnnaeus, 1758, Syst. long as head and thorax combined; the labial Nat. ed. 10, 1:560. Type%, Sweden. palpi are two-segmented; the forewings lack Urocerus nanus 0. F. Muller, 1776, 8,Zool. cross vein 2 r-m and possess an appendicular Dan. Prod. vein (3A); the hind wings lack an anal cell Sirex emrginatus Fabricus, 1793, 6,Ent. and the hind tibiae possess but a single api- syst., 2: 128. cal spur. Urocerus cadatus Cresson, 1865, 9, Proc. Ent. SOC.Philadelphia, 4:247. Tremex columha (Linn aeus) Sirex melancholicus Westwood, 1874, ci”, The- saurus Ent. Oxon., p. 116. (Pigeon tremex) ; Geographic range: Holarctic in northern coniferous forests, extending into mountains far- Sirex columba Linnaeus, 1763, Amoenitates ther south. North America, Eurasia, Japan, Academicae, 6:412. Type 9, “Amerika.” THE SIRICID WOOD WASPS OF CALIFORNIA 71

Sirex pennsylvanicus De Geer, 1773, Mem. a result of disease or other cause. Fattig Serv. Hist. des Ins., 3~593.?-. (1949) noted a female ovipositing near Atlanta, Sirex cinctus De Geer, 1773, Illus. Nat. Hist., Georgia at 6:OO A.M. It was the only one noted 2:72. P. from June, 1947, to June, 1248, despite many Sirex americana Christ, 1791, Naturg. Ins., p. observations of an infested tree. 412 P. The female deposits her eggs singly to a Tremex obsoletus Say, 1-823, West. Quart. depth of 10 to 12 mm. Rept., 2:73. dl The larvae, which obtain a length of 40 Trernez sericeus Say, 1823, West. Quart. Rept., mm., bore tunnels in the trunks and aid in the 2:73. 9. disintegration of the trees. They are of little Tremez seruillei Brull;, 1846, Hist. Nat. Ins., or no economic importance. It is stated that Hym., 4x545. 9. the larvae probably completes its growth and Tremex maurus Westwood, 1874, 6,Thesaurus transforms to a pupa within a year, but no def- Ent. Oxon., p. 116. 8. inite data are available. Fattig noted that Tremex columba aweus Bradley, 1913, Jour. when Tremex; emerged from the tree it dropped Ent. Zool., 5:25, 26. ?. to the ground and almost at once was off in a Geographic range: Widely distributed over the flash. United Staies and southern Canada, Arizona, Discussion: California, Colorado, Connecticut, Georgia, The adults of this species vary in color Illinois, Indiana, Kansas, Massachusetts, and markings, and three so-called races have Maryland, Michigan, Missouri, New Jersey, been recognized in the past, based upon these New Mexico, New York, Nova Scotia, On- variations. To a considerable extent these tario, Pennsylvania, Quebec, Tennessee, races are geographical with overlapping ranges. Utah. In the typical race, columba, the abdomen California records: is black with ocher-yellow bands and spots Riverside Co.: Neighbours, 6, V-1929 (L. A.C.M.). along the sides. It is most commonly found in northeastern United States and southeastern Hosts: Specimens have been reared from, or collected on, the following deciduous trees: Canada. In the race aweus the ground color of apple, beech, birch, box elder, elm, hack- the abdomen is yellow and the markings black. berry, hickory, maple, oak, pear, sycamore. The wings are golden yellow. This is the com- Parasites: The cynipid lbalia maculipennis mon form in Colorado, New Mexcio, Arizona, Haldeman and the ichneumonids Megarhys- and California. In sericeus the entire body is fulvous, the legs beyond the femora yellow, sa atrata (F.), Megarhyssa greenei greenei Viereck, and Megarhyssa macrurus macrwus and the wings dark reddish brown. This race (I..) have been recorded as parasites of the is found in the southeastern United States and pigeon tremex. An extensive bibliography as far north as Pennsylvania and west into on the biology of Megarhyssa spp. is given Colorado, New Mexico, Arizona, and California. by Townes and Townes (1951). A careful study in the light of modern sys- Biology: tematic thinking is needed to clarify the sta- Information on the biology of the pigeon tus of these color forms, to determine if they tremex is very scanty and confined entirely to are populations which are more or less isolated scattered short notes. No exhaustive studies as distinct geographical or ecological races have ever been made. Adults can be collected and hence deserve the status of subspecies. from June to September. They oviposit in dead The recent trend has been to consider them as or weakened deciduous trees or those dying as infrasubspecific categories. LITERATURE CITED

BAUMBERGER, G.P. FATTIG, P.W. 1915. Notes on the Siricidae of California. 1949. Some observations on Megarhyssa. Ent. News, 26(1):34. Ent. News, GO:@-71. BEAL, F.E.L. FRANCKE-GROSMANN, H. 1911. Food of the woodpeckers of the U.S.A. 1939. Ueber das zusammenleben von holz- U.S.D.A. Biol. Surv. Bull. no. 37, wespen mit Pilzen. Zeit. angew. Ent. pp. 1-64. Berlin, 25(4):647-680. BELYEA, R.M. HANSON, H.S. 1952. Death and deterioration of balsam fir 1939. Ecological notes on the Sirez wood- weakened by spruce budworm defolia- wasps and their parasites. Bull. Ent. tion in Ontario. Canad. Ent., 84(11): Res., 30:27-65. 325-335. HEDICKE, H. BENSON, ROBERT 1938. Siricidae. Hymen. Cat. pt. 6:l-36. 1943. Studies in Siricidae especially of Eu- Gravenhage. rope and Southern Asia. Bull. Ent. KEEN, F.P. Res., 34(1):27-51. 1952. Insect enemies of western forests. 1951. Handbook for the identification of U.S.D.A. Misc. Publ. 273. British insects. Roy. Ent. SOC.Lon- MANSOUR, K., and J. J. MANSOUR-BEK don, 6( 2a): 1-49. 1935. On the digestion of wood by insects. BRADLEY, J.C. Jour. Exp. Biol., 11:243-256. 1913. The Siricidae of North America. Jour. MIDDLEKAUFF, W. W. Ent. ZOO^., 5(1):1-35. 1948. A new species of Sirex from Califor- CHRYSTAL, R.N. nia. Pan-Pac. Em., 24(4): 189- 190. NORTON, EDWARD 1928. The Sirer woodwasps and their importance in forestry. Bull. Ent. Res., 1869. Catalogue of the Tenthredinidae and 19:2 19-247. Urocetidae of North America. Trans. Amer. Ent. Soc., 2:321-368. 1930. Studies of the Sirer parasites. Oxf. PARKIN, E.A. For. Men., 11: 1-63. 1942. Symbiosis and siricid woodwasps. CHRYSTAL, R. M., and J. G. MYERS Ann. Appl. Biol., 29(3):268-274. 1928. Natural enemies of Sirex cyaneus F., RIES, D.T. in England and their life history. 1951. Family Siricidae. In C.F.W. Muese- Bull. Ent. Res., 49:67-77, 1 pl., 2 beck, et aL, Hymenoptera of America figs. north of Mexico. Synoptic Catalogue, CRAIGHEAD, F.C. U.S.D.A. Mono., 2:83-85. 1950. Insect enemies of eastern forests. TOWNES, HENRY K.,'and MAfORIE TOWNES U.S.D.A. Misc. Publ. 657. 1951. Genus Megurhyssu. In C.F.W. Muese- ESSIG, E. 0. beck et al., Hymenoptera of America 1926. Insects of western North America. north of Mexico. Synoptic Catalogue. New York: MacMillan, 1035 pp U.S.D.A. Mono., 2:200-202. PLATES 74 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 4

Fig. 1. Fbrewing of Sirex. Fig. 2. Head and antenna of Xeris. Dorsal view. Fig. 3. Fore- and hindwing of Tremex. Fig. 4. Antenna of Tremex. Fig. 5. Anal region of hindwing, Urocerus. Fig. 6. Xeris macgillhayi, male: Dorsal view of head. Fig. 7. Xeris morrisoni, male. Dorsal view of head. Fig. 8. Sirex Zongicauda, female. Dorsal view of cornus and ovipositor. Fig. 9. Sirex obesus, female, Dorsal view of cornus and ovipositor. Fig. 10. Sirex cyaneus, female. Dorsal view of cornus and ovipositor. Fig. 21. Sirex californicus, female. Dorsal view of cornus and ovipositor. Fig. 12. Sirex behrensii, female. Dorsal view of cornus and ovipositor. Fig. 13. Sirex areohus, female. Dorsal view of cornus and ovipositor. Fig. 14. Xeris macgillivrayi, female. Dorsal view of cornus and ovipositor. Fig. 15. Xeris murrisoni, female. Dorsal view of cornus and ovipositor. Fig. 1G. Xeris morrisoni, female. Lateral view of head. Fig. 27. Urocerus californicus, female. Lateral view of head. THE SIRICID WOOD WASPS OF CALIFORNIA 75

-* cy- CARINA-'

--OVIPOSITOR

13 \ I

16 17 76 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 5

Fig. 18. Female Megarhyssa nortoni nortoni (Cresson), an ichneumonid parasite of larval woodwasps. Slightly larger than natural size.

Fig. 19. Female Ibalia ensiger Norton, a cynipid parasite reared from a log infested with Sirex behrensii (Cresson). Three times natural size.

Fig. 20. Section of redwood lumber showing female Sirex areolatus (Cresson) attempting to cut her way to freedom. About natural size. THE SIRICID WOOD WASPS OF CALIFORNIA 77