Mycologia, 90(5), 1998, pp. 743-753. ~ 1998 by The New York Botanical Garden, Bronx, NY 10458-5126 Issued I October 1998 Cordyceps variabilis and the genus Syngliocladium Kathie T. Hodgel species were briefly described based on one or a few fruiting bodies and have never been fully character- . Department of Plant Pathology, Cornell University, Ithaca, New York 14853 ized. Mycologists are not necessarily good entomologists, and host records for insect pathogens are of- Richard A. Humber ten inaccurate or limited to general categories. Given the deficit in teleomorph studies, it is hardly surpris- Plant Protection Research Unit USDA-AR, Tower Rd., Ithaca, New York 14853' ing that the anamorphs of Cordyceps species have only Chris A. Wozniak2 rarely been characterized, despite their potential utility in elucidating infrageneric relationships. Known anamorphs of Cordyceps species belong to NCSL, USDA-AR, P.O. Box 5677, State University Station, Fargo, North Dakota 58105 hyphomycetous genera including Akanthomyces, " 60605 Abstract: Cordyceps variabilis has long been errone- ously reported to be a pathogen of beetle larvae; studies of newly collected and herbarium specimens )7 Mariannaea, Metarhizium, Nomuraea, Paraisaria, Paecilomyces, Toiypocladium and Verticillium (Evans and Samson, 1984, 1987; Gams, 1971a; Hodge et aI., 1996; revealed that C. variabilis attacks fly larvae, particu- larly those of the family Xylophagidae (Diptera). An Liang, 1991; Liang et aI., 1991; emended description of C. variabilis is provided and a lectotye designated. The anamorph of C. variabilis '56 Beauvera, Desmidiospora, Hirsutella, Hymenostilbe, is reported here for the first time, and is attributed to the hyphomycetous genus Syngliocladium. An allied species, S. tetanopsis sp. novo is reported as a Mains, 1950; Petch, 1932b, 1935; Samson, 1974; Samson and Brady, 1983; Shimazu et aI., 1988). Mycoparasites growing on Cor- dyceps stromata have sometimes been mistaken for anamorphs. This seems to have been the case for putative anamorphs recorded as Stilbella species by Ko- pathogen of larvae of Tetanops myopaeformis, the sug- bayasi (1941); many of these have been recognized 4 ar beet root maggot (Diptera: Otitidae). The taon- as species of Polycephalomyces Kobayasi by Seifert omy of the genus Syngliocladium is briefly reviewed, (1985). We believe that some putative anamorphs referred to Sporothrix may also be mycoparasites grow- '9) and the generic concept emended. Key Words: Clavicipitaceae, entomopathogen, ing on Cordyceps stromata. Hypocreales, Sorosporella, teleomorph 55108 In this paper, the anamorph of C. variabilis Petch is characterized for the first time, a correction is made as to the nature of the host, and an emended description of C. variabilis is presented. The ana- l9) -1999) INTRODUCTION 95 9) morph is assigned to the genus Syngliocladium Petch, Species of Cordyceps Fr. are obligate pathogens of ar- which is emended herein, and a similar fungus oc- thropods (insects, spiders), hypogeous fungi (Elapho- curring as a pathogen of the sugar beet root maggot, Tetanops myopaeformis Röder (Diptera: Otitidae), is described as a new species. myces spp.), or other clavicipitaceous fungi (Cordyceps 00) K 75962 and Claviceps spp.). Well over 400 species have been igy (199 described in Cordyceps, some of which can be expected to be eventually relegated to syonymy, but it is also apparent that many species have yet to be de- l091 ~ MATERIS AND METHODS scribed. Kobayasi, the foremost modern expert on Fresh specimens of C. variabilis were collected from rotting wood at various sites in New York. Refer to the genus, accepted some 280 species (1982). Many I) ,iear for: t) Accepted for publication April 21, 1998. 1 Email: khll(Qcornell.edu 2 Cu;rent address: 751lW U.S. EPA, Biopesticide and Pollution Pre~ention Division, 401 M Street S.w., Washington, D.C. 20460. Mention of a trademark or proprietary product does not consti- JÜlnica1(; posiage¡i and does not imply its approval to the exclusion of other products W705-15!J JhOLOCO~ ~incnt. ,end addJ 5126l!S.l Specimens examined for locations and accession infor- mation. Larvae of T. myopaeformis infected with an unknown fungus were collected during the course of field screening for biocontrol agents of T. myopaefor- mis by CAW starting in the summer of 1994. Color terms are those of Kornerup and Wanscher (1967), tute an endorsement or a guarantee of the product by the USDA that may also be suitable. and size ranges represent averages that are in some 743 ~ -t ~ ~ ~ ñ 744 MYCOLOGIA cases circumscribed by mInimum and maximum fied where possible. A slide made from the holotype measuremen ts. of S. aranearum Petch (ex Araneida, St. Leonard's Fungal cultures.-Cultures were obtained from su- Forest, Horsham, Sussex, u.K. 31 May 1931) was ex- perficial mycelium and from hyphal bodies within cadavers of T. myopaeformis, and from superficial my- celium, hyphal bodies, and part ascospores of C. variabilis. Observations of growth and morphology were taken from cultures on oatmeal agar (OA: a filtered aqueous solution of 30 g/L rolled oats cooked for 30 min, 1.5% agar), modified oatmeal agar (mOA: an aqueous suspension of 30 g/L rolled oats cooked for 30 min, 1.2% agar, supplemented after autoclaving with 3 mL/L extra virgin olive oil and 30 mg/L cho- amined through the kindness of H. C. Evans and the keeper of K; the tye specimen of S. intricatum Petch (on larva of Phyllopaga anxia, Apple Hil, Ontario, Canada. 1934) was also borrowed from K. Pathogenicity assay.-The ability of AREF 5497 and AREF 4080 to infect T myopaeformis larvae was tested in a bioassay. Ten third ins tar larvae of T. myopae- formis from field collections were placed in 125 g of autoclaved coarse sand in a 9-cm petri dish. A saline lesterol), malt extract agar (MEA, Difco3), and Sabourauds dextrose agar with yeast extract (SDAY: (0.85% w/v) spore suspension collected from mOA plate cultures of AREF 5497 or AREF 4080 was add- 40g/L dextrose; 20 g/L bacto-peptone; 20 g/L yeast extract; 1.5% agar). All cultures have been deposited in the USDA-AR Collection of Entomopathogenic Fungal Cultures (AREF; Plant Protection Research based on hemocytometer counts and fluorescein diacetate staining. Saline solution was added to 7 mL Unit, U.S. Plant, Soil, and Nutrition Laboratory, Tower Rd, Ithaca, New York 14853) and are referred to da at 24 C, larvae were removed from the sand, hereafter by their AREF accession numbers. Cultures of the Syngliocladium anamorph of C. variabilis included in the present study are AREF 4080 (from mycelium on surface of host bearing stroma of C. variabilis. NEW YORK Hamilton Co., Raquette L., Long Point. Xylo phagus sp. larva (Diptera: Xylophagidae) in wood, 29 July 1993 K. T. Hodge kth13); AREF 5365 and 5366 (from mycelium on surface of host bearing stroma of C. variabilis. NEW YORK Danby, Michigan Hollow State Forest. Xylophagus sp. larva (Diptera: Xylophagidae) in wood, 3 Aug. 1996 K.T. Hodge kth50a, kth50b (CUP 63882)); AREF 5367 (from mycelium on surface of host bearing stroma of C. variabilis. NEW YORK Danby, Michigan Hollow State Forest. Xylophagus sp. larva (Diptera: Xylophagidae) in wood, 3 Aug. 1996 K.T. Hodge kth51a (CUP 63883)); and AREF 5414-5429 (single ascospore isolates from C. variabilis. NEW YORK Danby, Michigan Hollow State Forest. Xylophagus sp. larva (Diptera: Xylophagidae) in wood, 3 Aug. 1996 K. T. Hodge kth50a, (CUP 63882)). ed to the sand to yield 3 X 105 viable spores per dish, total per dish; saline alone was used as a control for estimation of natural background infection. Mter 7 rinsed in sterile distiled water, and placed on 9 cm Whatman (Kent, UK) no. 1 filter discs in glass petri dishes wrapped with Parafilm M (American Can Co., Neenah, Wisconsin, USA). Discs were moistened weekly with sterile distiled water to prevent desicca- tion of the insects. Larvae showing external hyphal growth were transferred to separate dishes so that infection rates do not represent secondary infections among larvae. Forty larvae were used per treatment (5497, 4080, and control), and the experiment was performed three times. Cadavers were inspected for the development of externally sporulating mycelium. RESULTS FIGS. 1-1 = 2 mm. stroma arI: Cultural characterstics.-Cultures isolated from hy- 4. Small st: inside cadaver of T. myopaeformis larva. NORTH DAKOTA: Red River Valley, 6june 1995. c.A.'Wozniak (CUP 64915)); phal bodies, mycelium from the host surface, and single part ascospores of C. variabilis yielded the same hyphomycetous fungus in culture. Cultures isolated from T myopaeformis cadavers yielded a similar hyphomycetous fungus which differed in key characters AREF 5497 (ex tye culture (FSt1A8-2294; NRR 21854), as outlined below. from 3rd instar larva of T. myopaeformis. NORTH DAKOTA: Pembina Co., St. Thomas. 1 Aug. 1994. C.A. Wozniak (CUP Pathogenicity to T. myopaeformis.-Strains isolated Cultures of Syngliocladium tetanopsis sp. novo included in the present study were AREF 4972 (from hyphal bodies 64913)); and AREF 5577 (from hyphal bodies inside cadaver of 3rd instar larva ofT. myopaeformis. (FHilAll-295; NRR 21853). NORTH DAKOTA: Red River Valley, 6june 1995. C.A. Wozniak). Specimens.-Herbarium specimens of C. variabilis were obtained on loan from North American herbaria (CORT, CUP, DAOM, FH, MICH, TRTC). Speci- mens were examined for the presence of the ana- morph on the host body, and the hosts were identi- 10169), ba Cordyceps 44. 193 from T. myopaeformis were observed to be infective to 3rd instar larvae of T. myopaeformis; 85% of inoc- ulated larva were kiled by the fungus 30 da after inoculation. Infected larvae tyically died within 21 da, developed an orange coloration and eventually produced synnemata similar to those that developed = Cordy Dee.) = Cordy 1986. =? Cm'd in axenic culture on OA and mOA. The potential of Sci. lv these strains as biological control agents of T. my- =? Cord opaeformis will be explored in future investigations. Sci M HODGE ET AL.: C. VARIABIUS AND SYNGUOCLAlUM 745 FIGS. 1-6. Cordyceps variabilis. 1. Typical stroma arising from anterior end of larva of Xylophagus sp. (CUP 47800), bar = 2 mm. 2. Capitate stroma arising from anterior end of larva of Xylophagus sp. (CORT 7736), bar = 2 mm. 3. Robust stroma arising near midpoint of larva of Xylohagus sp. (the anterior portion has broken off) (TRTC 4344), bar = 2 mm. 4. Small stromata on unidentified dipteran larvae (hosts removed) (FH 9861), bar = 1 mm. 5. (inset) Section of stroma (FH 10l69), bar = 50 fJm. 6. (inset) Ascus containing part ascospores. (CUP 63882), bar = 10 fJm. TAXONOMY Cordyceps variabilis Petch, Trans. Br. MycoL. SoC. 21: LECTOTYPE. NEW YORK Ithaca, on larva of Xylophagus sp., 25 July, 1916, G.H. Duff (FH 6132A) 44. 1937 (25 Oct.) emehd. K. T. Hodge & Humber. FIGS. 1-6 = Cordyceps viperna Mains, Mycologia 29: 674. 1937 (1 Dec.). = Cordyceps ithacensis Balazy & Bujak., Mycotaxon 25: 11. 1986. =? Cordyceps ferTuginosa Kobayasi & Shimizu, Bull. Natn. Sci. Mus., Ser. B (Bot.) 6: 139. 1980. =? Cordyceps asyuensis Kobayasi & Shimizu, Bull. Natn. Sci Mus., Ser. B (Bot.) 6: 138. 1980. Anamorph. Syngliocladium sp. FIGS. 7-9, 14-15 Parasitic on larvae of Diptera, particularly of wood- inhabiting Xylophagidae; host discolored burnt sienna (7-8(D8) J as a result of fungal infection. Host cadavers packed with cream-colored hyphal bodies. Hyphal bodies frequently peanut-shaped, 10-35 J.m long, sometimes irregularly subglobose, not particularly thick-walled, and containing large, refractive lip- id droplets. Host body often bearing orange (6B8) ~ ~ ~ ~ ñ §. 746 MYCOLOGIA fertile cushions in which perithecioid ascomata are immersed, and which are composed of textura intricata with a pseudoparenchymous cortex. Cortex pig- n mented, 30-45 i-m thick, composed of an inner layer of thick-walled parallel hyphae and an outer one of inflated, thick-walled irregular cells with free ends measuring 4.5-7.5 X 3.5-4.5 i-m. Perithecia obpyri- form, 350-590 X 200-370 i-m, immersed perpendicular to the surface. Ostioles darker orange (6C8), protruding slightly above stromal surface and giving the fertile cushion a verrucose appearance. Perithecial walls 12-18 i-m thick, with an inner layer of waxy cells about 1 i-m wide and an outer one of thickwalled brick-like cells about 1.5 i-m wide. Ostiolar canal periphysate, periphyses 11 X 0.5 i-m. Paraphyses not observed. Asci cylindric, 210-330 i-m long by 6.0 i-m wide, with an apical cap 4.5 i-m deep, packed with eight filiform ascospores each extending almost the full length of the ascus. Ascospores becoming multiseptate, dividing at maturity into many cylindric part ascospores (5.0-) 8.5 (-10.0) X (1.5-) 2.0 (-3.0) i-m, with thick, refractive walls. Distal ends of terminal part ascospores tapered. Anamorph sometimes produced in sparse patches or tufts on the surface of the host or on immature stromata. Conidiogenous cells in monoverticilate 14 whorls or pairs. Conidiogenous cells enteroblastic, (10.0-) 14.0 (-19.2) X 2.5 i-m, with subcylindric base and abruptly narrowing to form a short, tapering neck without a conspicuous periclinal thickening or collarette. Conidiogenous cells often strongly hooked or bent; producing ameroconidia in copious slime. Orange granules often present on the hyphal cell FIGs. walls. Conidia subcylindric, on average (8.0-) 10.2 (- dadium an 12.4) X (1.9-) 2.6 (-3.1) i-m (n = 100), often slightly constricted in the middle and sometimes slightly curved. Ascospores germinating after one mo on FIGS. 7-13. Syngliocladium anamorph of C. variabilis (left) and Syngliocladium tetanopsis sp. novo (right). 7-9. Cultural morphology of Syngliocladium anamorph of C. variabilis (AREF 4080). 7. One-mo-old colony on SDAY, bar = 5 mm. 8. Conidiogenous cells, bar = 20 f.m. 9. Conidia, bar = 20 f.m. 10-12. Cultural morphology of S. tetanopsis (AREF 4972). 10. Six-wk-01d colony on MEA (AREF 4972), bar = 5 mm. 11. Conidiogenous cells, bar = 20 f.m. 12. Conidia, bar = 20 f.m. 13. S. tetanopsis on wild-collected larva of Tetanops myopaeformis (CUP 64914), bar = 1 mm. patches of mononematous Syngliocladium anamorph, especially at the intersegmental membranes. One or few stromata arising from body of host in rotten wood, varying widely in size (2-24 mm). Stipe round in cross-section, equal, 0.2-3 mm diam, MEA. Colonies developing very slowly in culture on OA, mOA, SDAY, and MEA, reaching only 7 mm diam after 2 wk on SDAY Colonies on OA composed 14-17 an Conidiophon Co., Larva ( 1970, C. H. ~ paign Co., Bi larvae (Dipte (ARSEF); MJ of zones of white mycelium which become orange as copious phialoconidia are produced in slime balls 2843*, syntyc that unite to form large macroscopically visible drop- York, Chris S lets. On OA and mOA occasionally producing blunt variabilis Pete orange synnemata (1 X 8 mm) with copious conidiogenesis and slime production. Not forming chlamydospores in the host nor on artificial media. Known distribution. Belize, Canada (Nova Scotia, Ontario), China, USA (Ilinois, Maine, New York, North Carolina, Pennsylvania, Tennessee, Washington, Wisconsin) (Liang et aI., 1995; Mains, 1937, 1939, 1958). 1920, R. Tha: (Diptera: Xyl 6l32A, hereIi Hamilton Co. (Diptera: Xyle (CORT 7736'" ger Lakes Tra in decorticate Hodge kth50 5366, 54l4-5' Finger Lakes chrome yellow (5A7), furfuraceous, with sterile apex, Specimens examined. (Specimens that include the ana- bearing one or more lateral, or occasionally terminal morph are marked with an asterisk) ILLINOIS: McLean dae) in decor ,. HODGE ET AL.: C. VARABIUS AND SYNGUOCLAIUM 747 nOuOOO 15 ~ 14 17 0 U 'C ~ ". '" ':' ~ ~. ~ " , ,." o G °000 FIGS. 14-17. Syngliocladium anamorph of C. variabilis (left) and Syngliocladium tetanopsis sp. novo (right). 14-15. Synglio- cladium anamorph of C. variabilis (AREF 4080). 14. Conidiophores. 15. Conidia. 16-17. S. tetanopsis (AREF 4972). 16. Conidiophores. 17. Conidia. Bar = 10 ILm. Co., Larva (Diptera: Xylophagidae) in wood, july-Aug. 1970, C. H. Runde (K. A. Harrison 9944, MICH); Champaign Co., Brownfield Woods. Rhacicerus nitidus johnson larvae (Diptera: Xylophagidae), 3 july 1973, D. W Webb (AREF); MANE: Kittery Point, 1895, R Thaxter (FH 2843*, syntye of C. variabilis Petch); Kittery Point, 20 Aug. 1920, R Thaxter (FH 6138*, syntye of C. variabilis Petch); York, Chris Swamp, R. Thaxter (FH 6146*, syntye of C. variabilis Petch); NEW YORK Ithaça, on Xylohagus larva (Diptera: Xylophagidae), 25 july, 1916, C.H. Duff (FH 6132A, herein designated LECTOTYPE of C. variabilis); Hamilton Co., Raquette L., Antler's Camp. Xylohagus larva (Diptera: Xylophagidae) in wood, 1 Aug. 1995, T.j Baroni (CORT 7736*); Danby, Michigan Hollow State Forest, Finger Lakes TraiL. Xylohagus larva (Diptera: Xylophagidae) in decorticated wood of Tilia amercana, 3 Aug. 1996, K. T. Hodge kth50 (CUP 63882;* Living cultures AREF 5365, 5366, 5414-5429); Danby, Michigan Hollow State Forest, Finger Lakes TraiL. Xylohagus larva (Diptera: Xylophagi- dae) in decorticated wood of Tilia amercana, 3 Aug. 1996, K. T. Hodge kth51 (CUP 63883;* Living culture AREF 5367); Danby, Michigan Hollow State Forest, Finger Lakes TraiL. Xylohagus larva (Diptera: Xylophagidae) immature stromata in decaying wood, 3 Aug. 1996, K. T. Hodge kth52 (CUP 63884*); Alpine, Lost Gorge. Xylohagus larva (Dip- tera: Xylophagidae) in wood, 15 july 1964, T. Plowman (CUP 47800); Danby. Xylophagus larva (Diptera: Xylophagidae) in wood, 8 Aug. 1971, Paul E. Powell (CUP 52353); Ithaca, Coy Glen. Xylohagus larva (Diptera: Xylophagidae) in wood, 30 Sept. 1982, john (fin-sheng) Hu (CUP 60118); Enfield, Connecticut Hil Wildlife Management Area. Xylophagus larva (Diptera: Xylophagidae) in wood, 10 july 1994, David Price (CUP 63314); Essex-Hamilton Co., Catlin L. Larva (Diptera: Xylophagidae) in wood, 19 Aug. 1934, A. H. Smith 387 (MICH,* holotye of C. viperna Mains); NORTH CAOLINA: Swain Co., Great Smoky Mountains National Park, Indian Creek. Larva (Diptera) in wood, 2 Sept. 1939, A. H. Smith 10834 (MICH); Swain Co., Great Smoky Mountains National Park, Indian Creek. Larva (Dip- tera) in wood, 5 Sept. 1937, A. H. Smith 7393 (MICH); I,' it'. . I .'11. ~, I ,i C" ': 748 MYCOLOGIA Swain Co., Great Smoky Mountains National Park, Indian selected as lectotye his syntype specimen FH 6132A resembling Creek. Larva (Diptera) in wood, 14 Aug. 1938, A. H. Smith (listed incorrectly in his manuscript as FH 613A). All of the many specimens examined in this study, cushions oj 10169 (MICH); PENNSYLVANIA: Bucks Co., State Game Land #157. Larva (Diptera: Xylophagidae) in rotten wood, 1 Aug. 1997, Judith Fuchs (CUP 64917*); TENNESSEE: Bur- bank, 7 Aug. 1886, R. Thaxter (FH 6144: syntye of C. variabiZis Petch); Sevier Co., Great Smoky Mountains National Park, Coon's Cove. Larva (Diptera) in wood, 8 Aug. 1938, A. H. Smith 9861 (MICH); WASHINGTON: Whatcom Co., Baber L. Larva (Diptera: Xylophagidae) in wood, 2 Sept. 1941, E. B. Mains (MICH*); Skagit Co., Rocks Port, Maple Creek Forest Camp. Larva (Diptera: Xylophagidae) in wood, 15 Aug. 1941, A. H. Smith 16166 (MICH*);Jefferson Co., Hoh R. Larva (Diptera: Xylophagidae) in wood, 30 June 1939, A. H. Smith 14687 (MICH); WISCONSIN: Blue Mounds, 8 July 1905, R. A. Harper (FH*, syntye of C. variabilis Petch); CAADA. ONTARO: Haliburton Co., University of Toronto Forest. Larva (Diptera: Xylophagidae) in wood, 13 Sept. 1968, D. Malloch (DAOM 136432); Haliburton Co., University of Toronto Forest. Xylophagus larva (Diptera: Xylophagidae) in poplar wood, 13 Sept. 1968, D. Malloch (TRTC 45496); Temagami F.R., Round Lake. Xylo phagus larva (Diptera: Xylophagidae) in moss, 2 Sept. 1932, H. S. Jackson (TRTC 4344); Lake Temagami. Larva (Diptera: Xylophagidae) in wood, 26 Aug. 1936, A. H. Smith 4185 (TRTC, paratye of C. viperna Mains); NOVA SCOTIA: Colchester Co., Earlton Rd. Larva (Diptera: Xylophagidae) in wood, 26 Aug. 1931, L. E. Wehmryer 1213a (MICH, paratye of C. viperna Mains); BELIZE. El Cayo District, Cohune Ridge. Larva (Diptera: Xylophagidae) in wood, 13 July 1936, E. B. Mains 3837 (MICH, paratye of C. viperna Mains) . Commentary. The epithet given by Petch (1937) to this species aptly reflects its extreme variability in size and in the aspect of the fertile portion of the stro- mata (FIGS. 1-4). Among the specimens examined, the height of the stroma varied from 2-24 mm, and the perithecial "cushion" formed one or more lateral patches on the stipe, exposing a sterile apex, or less frequently assumed a capitate form (FIG. 2). Cordyceps viperna was described by Mains (1937) the same year, and was later recognized by him to be a synonym of C. variabilis (Mains, 1958), a conclusion supported by our own observatio.ns of tye materiaL Both species were based on North American materiaL Cordyceps variabilis has also been reported from China by Liang et aI. (1995), and from the Democratic Republic of Congo by Moureau (1961); those specimens were not examined in the present study. Petch (1937) recorded the hosts of C. variabilis as coleopteran larvae or specifically as members of the coleopteran family Elateridae, despite annotations on mata (Kobe including type specimens of both C. variabilis and C. thors list tl viperna, were found to occur on larvae of Diptera. their figure Many of these can be readily identified as species of 23B, 1983b the genus Xyiophagus (Xylophagidae), others are too et aL. (198E small or degraded to determine to family. Mter death, hosts assume an orange-brown coloration that Moreover, t dipteran la confers some resemblance to elaterid larvae, and this tle from N may be the source of confusion. Xylophagid larvae lack legs, however, and have a sclerotized and variously lobed anal plate that Mains (1937) apparently Type mater regarding t confused with a beetle head. The larval head cap- species can necessary Sj sules of Xyiophagus spp. (FIGS. 1-3 (top) J are darkly melanized and conical with a narrow longitudinal slit to C. varia through which the mouthparts can sometimes be scribed as a seen to project minutely on the ventral side; the bod- (see Kobaye ma Kobaya~ ies are elongate (Stehr, 1991). We have not encoun- Moureau tered specimens of C. variabilis on hosts other than larval Diptera. iabilis on be Allies and synonyms of C. variabilis. Cordyceps cor- of Congo. 1viperna as pan allomyces Møller (1901) was described from Brazil on pseudo a "Kellerassel" (sowbug, Crustacea: Isopoda). It is similar to C. variabilis in its part ascospore size, red- (1937) in ( (1937) in i orange coloration and lateral perithecial cushion found no with a distinct cortex. Petch (1935) considered C. cor- North Amei allomyces to be a synonym of C. ainictos Møller. Ko- Petch's type bayasi (1941) reported C. corallomyces from enchymous iabilis. Unfortunately Møller's specimens were appar- changed his perina was if Japan on a dipteran larva; his description is referable to C. var- myces a nomen dubium. In the future, specimens col- went on to c Republic of differed fro) doparenchyi lected near the tye locality may help elucidate the of C. inconsj ently destroyed in Berlin during World War II (B. Hein, pers. comm.). Given the different host and lack of authentic specimens, we consider C. corallo- relationship or possible synonymy between it and C. variabilis. Cordyceps ithacensis Balazy & Bujakiewicz (1986) was described from a single collection on a woodinhabiting dipteran larva in the family Erinnidae ( = Xylophagidae fide Borror et aL., 1989) in the vicinity of Ithaca, New York. It was described as having a small capitate stroma with orange peri the cia and an almost furfuraceous cortical layer of cells. Al- iabilis were i Democratic comm.). Wit dress his ch, regarding tr c. variabilis of Congo. A number mata with lai though the holotye could not be obtained from included by POZN, the ilustrations and thorough description provided by the authors, the host family and stage, and the locality of collection indicate that it is cer- Eucordyceps i tainly a synonym of C. variabilis. Cordyceps ithacensis cludes C. mi ther of the (Greuter et , some FH labels by Roland Thaxter indicating that the is based on a specimen in which the fertile region occurs terminally on the stroma, a condition that we hosts were dipteran larvae. Mains (1937, 1958) de- have occasionally observed in C. variabilis (FIG. 2). sect. Latera! scribed the hosts as larvae of Coleoptera. Because Both C. ferruginosa Kobayasi & Shimizu and C. asyuensis Kobayasi & Shimizu were described as species of section. C Petch (1937) did not designate a holotye, we have therefore mi dyceps subg. i (1941), but' HODGE ET AL.: C. VARlABILlS AND SYNGLlOCLAlUM resembling C. variabilis in that they produce lateral cushions of perithecia on acicular, furfuraceous stromata (Kobayasi and Shimizu, 1980). Although the au- bayasi and Shimizu, 1980) was described from 749 Japan on larvae of Elateridae (Coleoptera). It differs from C. variabilis in its purplish coloration, longer part thors list the hosts for both as coleopteran larvae, ascospores, and smooth stipe. Cordyceps clavata Ko- their figures (Kobayasi and Shimizu, 1980, Fig. 23A, 23B, 1983b, PI. 4, Fig. 3) and also those of Imazeki et aL. (1988, p. 583) suggest that the hosts might be bayasi & Shimizu (1980) differs from C. variabilis in having perithecia that are partially free and in pro- dipteran larvae, as reported here for C. variabilis. Moreover, the descriptions and ilustrations differ little from North American material of C. variabilis. Cordyceps nigrpoda Kobayasi & Shimizu (1982) differs in its fuscous coloration, lack of a furfuraceous cortex, nondisarticulating ascospores, and its habit Type material must be examined before conclusions regarding the syonymy and relationship of these species can be reached; we were unable to obtain the necessary specimens from TNS. Cordyceps minutissi- ma Kobayasi & Shimizu also bears a close similarity to C. variabilis, and the small host (which was described as a beetle larva) may also be a dipteran larva (see Kobayasi and Shimizu, 1983b, PI. 11, Fig. 3). Moureau (1961) reported on a collection of C. variabilis on beetle larvae from the Democratic Republic of Congo. He rejected Mains' (1958) treatment of C. viperna as a synonym of C. variabilis because the pseudoparenchymous cortex described by Mains (1937) in C. viperna was not mentioned by Petch (1937) in his discussion of C. variabilis. We have found no evidence that Moureau examined any North American specimens; our own examination of Petch's tye specimens indicates that a pseudopar- enchymous cortex is present. In 1962, Moureau changed his views on this issue, deciding that C. viperna was indeed a synonym of C. variabilis. He then went on to erect a new species from the Democratic Republic of Congo, C. inconspicua Moureau, which differed from C. variabilis only in its lack of a pseudoparenchymous cortex.. Unfortunately the holotye of C. inconspicua and Moureau's collections of C. variabilis were inaccessible due to political unrest in the Democratic Republic of Congo (V. Demoulin, pers. comm.) . Without his specimens we are unable to address his changing arguments and remain uncertain regarding the status of C. inconspicua and whether C. variabilis might occur in the Democratic Republic of Congo. A number of other Cordyceps species produce stromata with lateral, pulvinate fertile portions. They are included by Kobayasi (1982) in Cordyceps subgenus Eucordyceps Kobayasi, section Laterales Kobayasi. Nei- ducing multiple stromata on a coleopterous host. on coleopterous larvae. Cordyceps pentatomae Koval occurs on adult Hemiptera and lacks a furfuraceous cortex (Kobayasi and Shimizu, 1978). Cordyceps ra- mosopulvinata Kobayasi & Shimizu differs in its branching stromata, sh0rter part ascospores, and its habit on larvae of Cicadidae (Kobayasi and Shimizu, 1 983a). Cordyceps unilateralis differs in its tyically fi- liform, dark stromata, nondisarticulating ascospores, and its habit on ants (Hymenoptera: Formicidae) (Kobayasi, 1939). Cordyceps baumanniana Henn. and C. interrupta Höhn. are poorly known species from Lepidoptera and Coleoptera, respectively. Petch (1932b) considered C. baumanniana to be an aberrant specimen of C. australis, a capitate species with obliquely embedded perithecia. Cordyceps interrupta produces branching, brown, filiform stromata which bear multiple discrete perithecial cushions (Höhnel, 1909) . Cordyceps michiganensis Mains (1934) is macroscop- ically similar to C. variabilis in its production of chrome yellow, acicular stromata with sterile apices and a furfuraceous layer of cortical cells. In C. michiganensis the peri the cia are superficial and free, seldom united in groups of more than one or two, and many stromata arise from a single host. No anamorph was observed in specimens present at MICH, and the host in all cases is a coleopteran larva (the host of one specimen (A. H. Smith 63538, MICH) was iden- tified by Dr. J. K. Liebherr (Cornell University) as Carabidae: Pterostichus sp.). Although C. michiganensis and C. variabilis are macroscopically similar and exhibit overlapping distributions, the two species are distinct. The anamorph of C. variabilis. Petch (1937) reported the presence of "acervuli" on the stromata of specimens from FH on which he found conidia matching in description those figured here (FIGS. 9, ther of these taxa is nomenclaturally acceptable 15). He believed, however, that they were produced (Greuter et aI., 1994): Cordyceps subg. Eucordyceps includes C. militaris, the tye species of Cordyceps, and by a hyperparasitic fungus, and asserted that the anamorph of C. variabilis was a Hirsutella species which he did not describe. Having examined the same specimens on which Petch based his conclusions, we be- therefore must be referred to by the autonym Cordyceps subg. Cordyceps (Art. 22.1, 21.3). Cordyceps sub- sect. Laterales was validly described by Kobayasi (1941), but was never validly transferred to the rank of section. Cordyceps purpureostromata Kobayasi (Ko- lieve that the conidia described by him are indeed those of the anamorph of C. variabilis and that no hyperparasite is present. Furthermore, the conidi- 750 MYCOLOGIA ogenous cells of the anamorph described below are similar in shape to the tyically subulate phialides of Hirsutella. In our studies, cultures isolated from hyphal bodies, surface mycelium on the host, and from part ascospores all yielded the same slow-growing hy- phomycetous fungus, which is described above as the discolored yellow-tan with darkened cuticular lesions. When held in a moist chamber, white to yellOw-or_ ange tufts of sporulating mycelium erupted through the cuticle, and synnemata ultimately developed after 4-5 wk at 25 C. The fungus differs from the ana- morph of C. variabilis in its smaller conidia, host of Syngliocladium anamorph of C. variabilis. origin, and smaller conidiogenous cells. Synglioc1adium tetaopsIs K. T. Hodge, R A. Hum- Syngliocladium Petch, emend. K. T. Hodge & RA. Humber ber et C. A. Wozniak, sp. novo FIGS. 10-13, 16-17 Habitus entomogenus, mononematosus vel synnemato- sus, conidiophoris super exosceleto hospitis vel in synnebus crassis efferentibus. Cellulae conidiogenae plerumque monoverticillatae vel geminatae, (10.0-) 12.2 (_ 13.7) X 2.0 ILm, basibus subcylindricis, in collum curtum abrupte decrescentibus; apicibus saepe valde uncinatibus mati flexibusve; ameroconidiis in muco copioso enteroblastice formantibus. Granula aurantiaca superficiaribus in hyphis et cellulis conidiogenis saepe praesentia. Conidia aseptata subcylindrica, (4.3-) 6.8 (-9.3) X (1.9-) 2.0 (-2.5) ILm (n = 100). Chlamydosporae non vidunt. Status teleomorphosus ignotus. HOLOTYPUS. CUP 64913 (cultura desiccata ex AREF 5497). Entomogenous, mononematous or synnematous, Synnematous or mononematous; synnemata either loosely fasciculate and bearing laterally projecting conidiophores with apical clusters of Gliocladium-like conidiogenous cells and tending to form upon germination of clusters of chlamydospores (Sorosporella synanamorphic state) or stout and comprised of relatively densely packed parallel hyphae, arising direct- ly from stromatic tissues either inside an infected host or from in vitro cultures. Conidiogenous cells (phialides) occurring in clusters, often with a Glio- cladium-like appearance, or singly; with a swollen or flask-like base narrowing to a usually obvious and of- ten prominent neck that may be straight, hooked (uncinate), or bent out of the axis of the cell base. Conidia one-celled, ovoid to cylindric with rounded ends, hyaline, released successively into apical mu- conidiophores produced on the surface of the host and on stout synemata. Conidiogenous cells frequently in monoverticilate whorls or pairs, (10.0-) ing with those on adjacent phialides into larger mu- 12.2 (-13.7) ¡.m X 2.0 ¡.m, with subcylindric base Chlamydosporic synanamorphic state: species of So- coid droplets, with mucoid droplets often aggregatcoid drops or masses. abruptly narrowing to a short, tapering neck; often strongly hooked or bent; producing ameroconidia enteroblastically in copious slime. Orange granules known): species of Cordyceps Fr. often present on the hyphae and conidiogenous discussed here present some problems in generic rosporella Sorokin. Teleomorphic state (where Commentary. Although the two anamorph species cells. Conidia one-celled, subcylindric, (4.3-) 6.8 (_ placement, we feel they are best accommodated in 9.3) X (1.9-) 2.0 (-2.5) ¡.m (n = 100). Chlamydo- spores not observed. Teleomorph unknown. the poorly known genus Syngliocladium, as emended above. Their affinities with Toiypocladium W. Gams PARTYPES. CUP 64915, Laboratory-infected T. myopaeformis larvae. 6 June 1995. C.A. Wozniak (source of living (1974), and Paraisaria R A. Samson & B. L. Brady culture AREF 4972); CUP 64914, T. myopaeforis, larvae. NORTH DAKOTA: Red River Valley, summer 1995; CUP 64916, T. myopaeformis, larvae. NORTH DAKOTA: Red Riv- er Valley near Fargo, summer 1994. Commentary. Syngliocladium tetanopsis was isolated from larvae of T. myopaeformis (sugar beet root mag- gots), an economically important pest of sugar beets. Fungus-infected T. myopaeformis larvae were collected from sugar beets in untreated border rows planted as buffer zones in a study of the efficacy of entomo- pathogenic nematodes as biocontrol agents. The studies were performed in Pembina Co., North Dakota, by CAW beginning in summer 1994. Dead or dying third ins tar larvae collected in the field were (1971b), Culicinomyces Couch, Romney & Rao (1983) are discussed below. The anamorph genus Syngliocladium was described by Petch for S. aranearum Petch on a spider from England (Petch, 1 932a). He described the genus as producing Gliocladium-like conidiophores arising laterally from synnemata on the host. The conidia are borne on subulate phialides in copious slime. Petch (1942) later added the species S. intricatum Petch (from a coleopterous larva) and S. cleoni (Wize) Petch. The status of Syngliocladium has been considered dubious by some researchers (Samson and Bra- dy, 1983; Shah and Evans, 1997; Carmichael et aI., 1980). Despite Samson and Brady's (1983) assertion that S. intricatum is a stilbellaceous fungus not closely allied with other species later included in the genus, HODGE ET AL.: C. VARABIIJS AND SYNGLIOCLAlUM our examination of the tye (K; FIG. 19) led us to conclude that Petch's concept of Syngliocladium was cohesive. No teleomorphs have previously been re- 18 ported for Syngliocladium, although its slimy phialoconidia, synnematous tendency, and insect pathogen- f 751 000 o () 0 ic habit suggest the now confirmed affinity to the entomopathogenic members of the Clavicipitaceae as noted by Speare (1920). o 0 Q o Sorosporella uvella (Krass.) Giard, characterized by the brick-red resting spores that fill the cadavers of its insect hosts, has been shown by Speare (1917, r ).e 1920) to have an unnamed synanamorph attributable to Syngliocladium. Not all Syngliocladium species have Sorosporella synanamorphs; we prefer to maintain the (a two genera as distinct for descriptive purposes and because of their disparate modes of spore ontogeny i- (Cams, 1982). Sorosporella infections of orthopteran r- td insects that have been by reported by Pendland and Boucias (1987), Shah (1993), Shah et aI. (1994), Is Shah and Evans (1997), and Welling et aI. (1995), 0- may represent one or more un )r described species, but the simple morphology of Sorosporella resting spores )f- makes the taonomy of the genus problematic. Most ~d Sorosporella species that have been studied in culture ,e. produce Syngliocladium synanamorphs, and these ~d uH- u~o- consideration that the Sorosporella state appears to be FIGS. 18, 19. Morphology of Syngliocladium spp. 18. Conidiophores and conidia of S. aranearum (holotye, K). 19. more commonly encountered in nature. Studies of Conidiophores and conidia of S. intricatum (holotye, K). Syngliocladium state is complicated by the practical Syngliocladium tetanopsis and the Syngliocladium anamorph of C. variabilis are morphologically similar and clearly congeneric. Syngliocladium tetanopsis differs from the anamorph of C. variabilis in its smaller istics. Whereas S. tetanopsis killed more than 85% of inoculated third instar larvae of T. myopaeformis (its thoptera: Gryllotalpidae). In culture, the Synglioclad- conidia and phialides, host, and cultural character- )m of S. tetanopsis against the host of the Syngliocladium , as lat- cause cultures of xylophagids were not available. kaaL., .ion sely ius, plex conidiophores as well as simple, frequently hooked phialides were observed (FIG. 19). Similar phialides were ilustrated by Speare (1917,1920) for the Syngliocladium syanamorph of Sorosporella uvella, and by Pendland and Boucias (1987) for an undescribed Syngliocladium sp. from mole crickets (Or- ied ze) sid- Bar = 5 ¡Lm. elucidate species in this long-neglected group. host of origin), the Syngliocladium anamorph of C. variabilis kiled fewer than 8%. Reciprocal bioassays are tch tch - That these fungi might best be characterized by their Les ms .ao idy o 00 0 diagnostic for taonomic purposes (Gams, 1982). host range and cultural morphology are needed to in ed 19 000 0 more highly differentiated states are likely to be more re ric o 0 0 0 C) anamorph of C. variabilis were not performed be- A slide taken from the holotye of S. aranearum Petch, the tye species of Syngliocladium, was gener- ously loaned by H. C. Evans and the curator of K. Because of its fragile condition (H. C. Evans, pers. comm.), the tye specimen itself was not inspected. Compound, Gliocladium-like conidiogenous cells were clearly visible; no simple conidiogenous cells were observed in the small synnema fragment mounted on the slide (FIG. 18). The tye specimen of S. intricatum Petch (K) was examined, and com- ium synanamorphs of undetermined Sorosporella spp. (AREF 1123, 1856) produced both simple phialides and complex conidiophores. Gams (1982) suggests that synanamorphs that differ in conidiophore complexity but not in conidial ontogeny need not be assigned different generic names. In the above cases the name Syngliocladium can be applied to both com- plex and simple conidiophores. We therefore treat the two taxa presented here as closely allied species of Syngliocladium on the basis of their simple, hooked, conidiogenous cells, conidia borne in copious slime, synnema-forming abilty, and entomopathogenic habit. Similar genera. Toiypocladium resembles Synglio- cladium in its frequently hooked conidiogenous cells 752 MYCOLOGIA and conidia borne in copious slime. The tye species, concept of Gliocladium advanced by Gams (1982), in T. inflatum, has been shown to have a Cordyceps sub- which similar conidia are produced by both simple and complex conidiophores. sessilis teleomorph (Hodge et aI., 1996). Toiypocladium cylindrosporum in particular bears a close resem- blance to the anamorphs described here. Toiypoclad- ACKNOWLEDGMENTS ium differs from Syngliocladium in its more complex, dendroid conidiophores. Culicinomyces also produces frequently hooked phialides and is pathogenic to larvae of Diptera. No teleomorph has been reported. Culicinomyces clavisporus and Cu. bisporalis (but not Cu. parasiticus) pro- duce two morphologically different tyes of conidiophores. The first (tye A) comprises short-necked, bottle-shaped phialides that are closely appressed to the upright axis of the conidiophore and occur singly or in whorls. The second (tye B) comprises subulate phialides with slender necks and larger, more cylin- dric conidia (Couch et aI., 1974; Goettel et aI., 1984; We are deeply indebted to Jennifer Altre for preliminary identification of the hosts of some C. variabilis specimens. We are grateful to the curators of FH, CORT, CUP, DAOM, K, MICH and TRTC for the loan of specimens, and to V. Demoulin for information concerning Moureau's collections. KTH thanks the Friends of the Farlow for generously supporting a visit to FH, and Claudia Olivier for her translation of parts of Møller's work. Harry Evans graciously shared his thoughts on taxonomic problems in Synglioclad- ium and made available his slide taken from the holotye of S. aranearum. Keith Seifert's comments and those of anonymous reviewers greatly improved the manuscript. Sigler et aI., 1987). This latter state is similar to Syn- gliocladium. Culicinomyces differs in its habit on aquatic larvae of flies of the more primitive suborder Nematocera (the species described here are known from Brachycera), in its failure to produce synnemata in culture, and in its production of distinctive tye A conidiophores. LITERATURE CITED Balazy, S., and A. Bujakiewicz. 1986. A new entomogenous species of Cordyceps: Cordyceps ithacensis sp. novo Mycotaxon 25: 11-14. Borror, D.j., C. A. Triplehorn, and N. F.johnson. 1989. An introduction to the study of insects, 6th ed. Saunders Col- anamorph of C. gracilis (Samson and Brady, 1983). lege, Philadelphia. 875 pp. Carmichael, j. W., W. B. Kendrick, 1. L. Conners, and L. Paraisaria has close affinities with Syngliocladium: it Sigler. 1980. Genera of hyphomycetes. University of Al- with a narrow neck, and is chiefly differentiated by berta Press, Edmonton. 386 pp. Couch,j. N., S. V. Romney, and B. Rao. 1974. A new fungus Paraisaria is a mono tyic genus described for the produces conidia in slime from subulate phialides the tendency of its conidiogenous cells to become polyphialidic. Microhilum Yip & Rath is also some- which attacks mosquitoes and related Diptera. Mycolo- what similar (Wright and Patel, 1992; Yip and Rath, Evans, H. C., and R. A. Samson. 1984. Cordyceps species and 1989), although the denticulate conidiogenous cells and dry conidia of Microhilum more closely recall Beauvera. Conclusion.-We have included S. tetanopsis and the Syngliocladium anamorph of C. variabilis in Synglio- cladium although they might alternately have been. accommodated in a new genus. We recognize their similarities with Toiypocladium, Culicinomyces, and Paraisaria, but, exclude them from these genera for reasons discussed above. We feel that these two species are likely to be closely related to other Synglio- cladium species, and that creation of superfluous anamorph genera is to be avoided. In emending the concept of Syngliocladium, we recognize the polymorphic nature of some existing species, which may form both simple and complex conidiophores producing conidia of the same ontogenetic tye. The two anamorph taxa described here produce only the simpler of the two tyes of conidiophores, but we believe they are best included within an expanded concept of Syn- gliocladium. 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