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Biological Species Concepts in Eastern North American Populations of Lentinellus Author(S): Andrew N

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Biological Species Concepts in Eastern North American Populations of Lentinellus Author(S): Andrew N Mycological Society of America Biological Species Concepts in Eastern North American Populations of Lentinellus Author(s): Andrew N. Miller and Andrew S. Methven Reviewed work(s): Source: Mycologia, Vol. 92, No. 4 (Jul. - Aug., 2000), pp. 792-800 Published by: Mycological Society of America Stable URL: http://www.jstor.org/stable/3761437 . Accessed: 12/11/2011 12:41 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Mycological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Mycologia. http://www.jstor.org Mycologia, 92(4), 2000, pp. 792-800. © 2000 by The Mycological Society of America, Lawrence, KS 66044-8897 Biological species concepts in eastern North American populations of Lentinellus Andrew N. Miller1 segregate taxa has led to considerable taxonomic Universityof Illinois at Chicago,Department of confusion as numerous taxa have been proposed BiologicalSciences, Chicago, Illinois 60607-7060, and only to later be reduced to synonymy (Miller and TheField Museum,Department of Botany,Chicago, Stewart 1971, 1983, 1996). Illinois 60605-2496 Pegler Segedin Among the most confusing species complex is the one which Andrew S. Methven includes L. ursinus (Fr.) Kiihner, L. castoreus (Fr.) EasternIllinois University,Department of Biological Romag., a European taxon, L. vulpinus (Sow.:Fr.) Sciences,Charleston, Illinois 61920 Kuhner & Maire, and several other morphospecies (Stalpers 1996) which have not been adequately dif- ferentiated biologically or phylogenetically from L. Abstract: As part of a biosystematic study of the Len- ursinus s.s. Although morphotaxonomy has been tinellus ursinus species complex in eastern North used to distinguish L. ursinus from L. vulpinus (Mill- America, field and herbarium specimens from North er and Stewart 1971), it has not accounted for addi- America, Central America, and Europe were exam- tional cryptic taxa within the L. ursinus species com- ined. Crossing single basidiospore isolates demon- plex and reveals nothing about reproductive com- strated tetrapolar (bifactorial) mating systems. Three patibility between morphospecies. Since preliminary biological species are recognized as L. ursinus, L. an- data suggested that more than one biological species gustifolius, and L. vulpinus. The production of gloeo- was represented in the L. ursinus species complex in cystidia and/or chlamydospores is considered taxo- eastern North America, intercollection matings were nomically significant for identifying somatic cultures proposed to identify sexually compatible populations of these species. All three taxa produced laccase, ty- (i.e., biological species) in the L. ursinus species com- rosinase, and peroxidase in culture and are white-rot plex. fungi. This is the first confirmed report of L. angus- The objectives of this study were: (i) to determine tifolius from North America. the mating systems operating in the L. ursinus com- Key Words: Auriscalpiaceae, Basidiomycetes, cul- plex; (ii) to identify sexually compatible and incom- ture mat analyses, Hericiales, mating systems, phen- patible populations; (iii) to determine the taxonomic oloxidase enzymes, systematics value of culture mat analyses and phenoloxidase re- actions, and (iv) to identify additional morphological characters which might be used to more readily seg- INTRODUCTION regate taxa. The genus Lentinellus Karsten (Hericiales, Auriscal- piaceae) includes a group of clitocyboid to pleuro- MATERIALSAND METHODS toid fungi most commonly collected in deciduous Basidiomatawere collected in six states in eastern North and coniferous forests throughout North temperate America from Aug through Nov, 1995 and 1996. Macro- regions. Although the genus Lentinellus is easily rec- morphologicalfeatures were recorded using standardtech- ognized by serrate lamellae and amyloid, ornament- niques (Largent 1977) with colors terms taken from Kor- ed basidiospores, many taxa are extremely variable nerup and Wanscher(1967). Basidiomatawere dried and and often difficult to distinguish macroscopically deposited in the herbariumat Eastern Illinois University Additional were obtained from herbaria and/or microscopically. The difficulty in segregating (EIU). specimens at the Botanical Gardens, (E), the Uni- taxa is due primarily to the relatively long life of the Royal Edinburgh versityof Tennessee (TENN), the Universityof Michigan basidiomata, which results in considerable morpho- (MICH), and Uppsala (UPS). Supplementaryspecimens variation in tomentum and logical especially pileus with spore prints were also provided by Thomas J. Volk of tramal and Stewart degree amyloidity (Miller (TJV).The names of the authorsare abbreviatedANM for 1971). The use of morphological characters alone to AndrewN. Millerand ASMfor AndrewS. Methven. Micromorphologicalanalyses of specimensfollowed stan- Accepted for publication February 14, 2000. dard practicesand terminology (Largentet al 1977). Sec- 1 Email: [email protected] tions were mounted in 10% NH4OH,Melzer's reagent, or 792 MILLER AND METHVEN: BIOLOGICAL SPECIES CONCEPTS IN LENTINELLUS 793 sulfobenzaldehyde and observed under bright field optics 11-14 d (28 d for RHP 7966), 5-mm-diam plugs were re- with a Nikon research microscope. A minimum of 10 basid- moved from the margin of the hyphal mat and transferred ia, 10 pseudocystidia, and 30 basidiospores were measured to 12 replicate plates each of MEA and Difco potato dex- for each collection. For basidiospore data, x ± SD and Q trose agar (PDA). Growth rates and macromorphological (spore length/width) were calculated for each species. and micromorphological features were recorded at 2-wk in- tervals according to Nobles (1948, 1965), Stalpers (1978), and iso- Monospore polyspore isolates.-Single basidiospore and Desjardin (1990). Color terms were taken from Kor- lates were collected one of the (SBIs) utilizing following nerup and Wanscher (1967). Nobles Species Codes (1965) methods: small sections from fresh basidiomata (i) pileus and Staplers Species Codes (1978) were determined for were from the lid of a 90-mm Petri suspended Pyrex plate each species. with and allowed to fall onto petroleum jelly basidiospores Spot tests for the phenoloxidase enzymes laccase, tyrosi- malt extract Difco (1.25% Difco) agar (2% Bacto) (MEA) nase, and peroxidase were conducted on culture mats at for 15-60 min; or from a were (ii) basidiospores spore print weeks 2 and 6 (Desjardin 1990). Syringaldazine (Harkin affixed to a sterilized and streaked onto moistened, loop and Obst 1973, Harkin et al 1974) and ot-naphthol (Kiaarik MEA in a 90-mm Petri Pyrex plate. Basidiospore germina- 1965) were used to test for laccase activity, L-tyrosineand p- tion occurred after 3-6 d of incubation in the dark at 23 cresol for tyrosinase (Marr 1984, Marr et al 1986), and py- C. Individual were transferred to MEA in germlings initially rogallol plus H202 for peroxidase (Taylor 1974, Staplers 90-mm Petri and then transferred Pyrex plates subsequently 1978). Distilled water and 95% ethanol were used as nega- to individual screw cap test tubes containing MEA after the tive controls. colonies reached ca 1 cm diam. Polyspore isolates were ob- tained by transferring a 5-mm2 block of MEA containing numerous germlings to MEA in screw cap test tubes which RESULTS were then stored in the dark at 23 C. In the case of ANM 321 and RHP 8768, polyspore isolates were created by self- Lentinellus ursinus (Fr.) Kfihner, Le Botaniste 17:99, crossing the mating types identified in each collection. Ad- 1926. ditional sets of SBIs, along with accompanying polyspore Basidiomata morphology. Miller and Stewart (1971) isolates and voucher specimens, were obtained from R. H. adequately described this taxon and provided pho- Petersen (RHP) (TENN). Ancillary sets of SBIs and voucher tographs and line drawings of salient microscopic fea- from Austria and Costa Rica were obtained from specimens tures. Although minor variations in some characters I. Krisai-Greilhuber and R. H. Petersen, respectively. All were found in this study, none were significantly dif- SBIs (including tester strains) and polyspore isolates uti- ferent from those reported by Miller and Stewart lized in this study were deposited in the University of Ten- (1971) for the of oleiferous nessee Fungus Culture Collection (TENN). except presence hyphae which were: (i) abundant throughout pileipellis, at Intra- and intercollection matings.-Intra- and intercollec- times terminating as solitary, cylindric to narrowly tion matings followed Petersen and Cifuentes (1994) with clavate, thin-walled pileocystidia; (ii) infrequent to minor modifications. connections were observed in Clamp common in trama; (iii) abundant in subhymenium, a 1:1 mixture of 2% phloxine and 2% congo red at X400 terminating as clavate pseudocystidia, and (iv) thin- under bright field illumination using a Nikon research mi- walled, unclamped, unbranched, inamyloid, yellow- croscope. Intra- and intercollection matings were scored on ish in NH40H, always with resinous contents, staining a grid for the presence or absence of clamp connections
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