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Department of Biology, Utah State University, Logan, Utah 84322
P. Brandon Matheny
Department of Biology, Box 351330, University of Washington, Seattle, Washington 98195-5325
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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We present a morphological and phylogenetic study of Inocybe chelanensis and other North American species of Inocybe that have unusually elongated nodulose spores. Taxonomy and illustrations of these species are provided, along with a key to these and similar species found in Europe and North America. The species concept of I. chelanensis is broadened to include the range of variation occurring for the species in North America. Despite similar basidiospore morphologies, I. chelanensis and I. candidipes are not closely related. Inocybe chelanensis is related more closely to I. stellatospora and I. candidipes is related to I. glabrodisca based on RPB1 and nLSU-rDNA nucleotide sequences. Distal enlongation of Inocybe basidiospores was achieved independently in at least two separate lineages of Inocybe. Inocybe candidipes and I. sierraensis are described as new.
Key words: Agaricales, Basidiomycota, RPB1, species concepts, systematics
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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, Horak 1977, 1979, 1980, Kobayasi 1952, Kobayashi 2002, Kuyper 1986, Losa Quintana 1960, Perez-Silva 1967, Stangl 1989), much of the work done on the genus in North America still consists of scattered reports and species descriptions. An early monograph by Kauffman (1924), together with a series of papers by Smith and Stuntz (1950), Stuntz (1947, 1954) and Grund and Stuntz (1968, 1970, 1975, 1977, 1980, 1981, 1983, 1984), provide an important platform for work on the genus within the continent. In addition, recent papers focused on North American taxa have provided keys to species from California (Nishida 1989), species associated with Populus tremuloides Michx. (Cripps 1997) and species within the Inocybe lanuginosa (Bull. : Fr.) Kummer group (Matheny and Kropp 2001). In many parts of North America, the species of Inocybe remain poorly documented and it is likely that at least some new taxa will be found as these areas are studied. While working on species of Inocybe in western North America, we discovered several interesting specimens sharing similar characters with the Inocybe chelanensis Stuntz group. Members of this group either are uncommon or rare, often occur in montane to subalpine settings in association with Abies and Pinus and are characterized by having spores with an elongated apex and one to several basal nodules that give them a distinctive rocket shape. Within North America, three species that appear to belong to this group, Inocybe chelanensis, I. rainierensis Stuntz and I. texensis Thiers, have been described. Outside North America, only two other species that share affinities with the group have been reported thus far. These are I. dolichospora Malençon, a species associated with Cedrus in North Africa, and I. ortegae Esteve-Raventós, reported with Quercus from Spain. Inocybe rennyi (Berk. & Broome) Sacc. is another similar species with unusual spores that occurs in North America and Europe. Although its spores are elongated and angular to nodulose, they typically lack the basal nodules that impart the rocket shape of the other species mentioned above.
Specimens from Utah and Arizona appear to belong to the I. chelanensis group because of their unusual spores, but they seem unique because their spores and, to a lesser degree, their cheilocystidia differ in length from those reported for the holotype of I. chelanensis (Stuntz 1947); in addition, the stipe covering of the Arizona material is different than is typical for I. chelanensis. A preliminary examination of herbarium specimens representing I. chelanensis indicated that more variation occurs within this species than anticipated. Thus, to determine the status of the Utah and Arizona collections, it was necessary to examine the holotype of each member of the group and to document morphological and phylogenetic variation among existing collections.
Species concepts in Inocybe have relied strictly upon gross morphological and anatomical data (Kuyper 1986). However, Matheny et al (2002) demonstrated that sufficient variability exists at the species level within Inocybe at the RPB1 locus, a gene that encodes the largest subunit of RNA polymerase II, to separate species. Less variation among species of Inocybe occurs at the nLSU-rDNA (nLSU) locus, yet the region also appears suitable to separate many samples at the species level (Matheny et al 2002, Matheny data unpubl).
In this paper, we present the results of a systematic study of I. chelanensis and similar species in North America, including species descriptions, known geographical distributions, phylogenetic data and a key to species that are either phenotypically similar to I. chelanensis or might be confused with it.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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). Samples from all specimens were rehydrated and examined directly in 10% NH4OH (or 3% KOH for I. texensis). Microscopic measurements were made under oil immersion at 1000x, and line drawings were made with the aid of a drawing tube. When possible, measurements of basidiospores were done using spores that had been deposited on stipes to avoid measuring immature spores. Spore measurements are given as averages and ranges for length and width with outlying measurements indicated in parentheses. Dimensions for basidia and hymenial cystidia are given as ranges. Spore colors were taken from microscopic preparations in 10% NH4OH using the Munsell book of color (Munsell Color 1976). Color notes from fresh material were taken from several different color standards (Ridgway 1912, Kornerup and Wanscher 1967, Munsell Color 1976). For the purposes of this paper, all colors used in notes from fresh material or given in previously published species descriptions have been converted to the nearest equivalent in the Munsell book of color by direct comparison. The other color designations used are our own. When only herbarium specimens were available, colors and other macroscopic characters either were obtained directly from the specimen or from field notes enclosed with the specimen.
The sampling strategy includes only nodulose-spored taxa of subgenus Inocybe (Kuyper 1986) and the minimally angular to smooth-spored species I. lacera so that we could evaluate both the placement of collections of an unknown taxonomic status and test the monophyly of Inocybe species with apically elongated nodulose-basidiospores. The inclusion of I. lacera is warranted based on the suggestion of others (Kühner 1980, Kuyper 1986) that I. lacera might be related to some goniosporous species such as I. curvipes. The two do share similar cystidial characters and oblong basidiospores. Whereas the spores of I. lacera are oblong-smooth, subfusiform, to minimally angular, those of I. curvipes are oblong angular-nodulose, generally with few to several low obtuse nodules. In addition, several caulocystidiate taxa that lack a cortina (i.e., I. calospora, I. praetervisa and I. glabrodisca) were included to assess the taxonomic and phylogenetic utility of apically elongated basidiospores. Both Kuyper (1986) and Matheny et al (2002) have suggested the polyphyly of Inocybe species with angular-nodulose basidiospores, thus phylogenetic conclusions about nodulose-spored taxa in general are not addressed. We were unable to extract DNA successfully from the holotype of I. rainierensis, which was collected in 1948.
DNA was extracted from dried basidiomata (samples listed in TABLE I), and nucleotide sequences were obtained for both RPB1 and nLSU following Matheny et al (2002). Primers A forward, C reverse, and B reverse were used to directly sequence the 5' end of RPB1 that includes conserved domains A to C; primers 5.8SR and LR7 were used to amplify the 5' nLSU region with internal primers LR0R, LR3R, LR16 and LR5 used for direct sequencing. Sequences were aligned using ClustalX (Thompson et al 1997), adjusted by eye and analyzed with PAUP* (Swofford 2002). The alignment is available at TreeBASE (S917) or from the second author on request.
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); hence, the two datasets were combined for further analysis. We used heuristic searches to find the optimal (MP) trees and 1000 bootstrap replicates (Felsenstein 1985) to evaluate branch strength. All searches for optimal trees employed 30 random-addition sequence replicates holding one tree at each step during stepwise addition and the tree-bisection-reconnection (TBR) branch-swapping algorithm. Inocybe ayangannae, a neotropical nodulose-spored species with a cortina (Matheny, Aime and Henkel 2003), was used to root the trees. Combined analysis of RPB1, nLSU, and an additional locus, RPB2, suggests I. ayangannae and several newly described species from the neotropics form a clade sister of the remainder of subgenus Inocybe sensu Kuyper (Matheny unpubl data).
We generated unconstrained and constrained maximum-likelihood (ML) trees under a best-fit model selected by Modeltest 3.0 (Posada and Crandall 1998, 2001) to evaluate suboptimal trees using the Shimodaira-Hasegawa test (SH-test) (Shimodaira and Hasegawa 1999). This permits testing of alternative (suboptimal) phylogenetic hypotheses to determine if they are significantly worse than the optimal ML topology (
= 0.05) (Goldman, Anderson and Rodrigo 2000). A Tamura-Nei model was chosen as best fit to the combined RPB1 and nLSU data (base frequencies equal; a rate matrix of 5.1873 and 9.9843 for AG and CT transitions, respectively, and 1.0000 for all transversions; a 0.4351 proportion of invariable sites and a gamma distribution shape parameter of 1.3616). Under this model of DNA substitution, we then constrained the monophyly of taxa with a rocket-shaped basidospore outline (two I. chelanensis collections, I. sierraensis sp. nov. and I. candidipes sp. nov.) and of the taxa belonging to the I. lanuginosa group (I. lanuginosa, I. leptophylla, I. teraturgus and I. stellatospora). The SH-test then was used to determine if our constrained topologies significantly were worse than the optimal ML topology ( = 0.05).
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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and three exon regions. Intron 2 is composed of 536 sites and intron 3 has 52 sites, including gaps. No indels were present in the exon regions.
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Species with rocket-shape basidiospores cluster in two separate clades (FIG. 1). Inocybe chelanensis is sister with strong bootstrap support (85%) of I. sierraensis. Both species share similar basidiospore morphologies but differ in other morphological characters. However, I. candidipes, which has the distinctive rocket-shape basidiospore outline similar to I. chelanensis, occurs in a separate clade sister of the caulocystidiate species, I. glabrodisca (=I. oblectabilis Britz. f. decemgibbosa Kühn. & Bours.), with 100% bootstrap support. Like I. glabrodisca, I. candidipes appears to lack a cortina and has caulocystidia on the lower part of the stipe, although not in abundance. When we constrained the monophyly of taxa with this peculiar spore shape under maximum likelihood conditions, the ML constraint tree was rejected (P < 0.05) (T1 = 9881.7031; TBEST = 9771.7970). This further supports the homoplastic status of this spore outline. Nodulose- to spinosespored species with caulocystidiate stipes (and that lack a cortina) appear paraphyletic, although this result is weakly supported (<50% bootstrap support).
Inocybe lanuginosa, I. leptophylla and I. teraturgus are strongly supported as monophyletic (100% support). However, I. stellatospora (=I. longicystis Atk.), also thought to be a member of this group, is excluded (Matheny and Kropp 2001). The strongly supported (97%) sister position of I. stellatospora to the I. chelanensis group is unexpected because I. stellatospora has nodulose spores that are merely subangular to subelliptic. When we constrained the monophyly of members of this group, the constrained topology (T2 5 9798.4200; TBEST = 9771.7970) is statistically worse (P < 0.05) than the optimal topology using the SH-test. This result substantiates the strong bootstrap support for the sister position of I. stellatospora and the I. chelanensis group.
The sister relationship of I. curvipes and I. lacera also is supported strongly (100%), and this is the first phylogenetic work that supports a smooth-spored species as sister of an angular-nodulose-spored species. Although it should be said that I. lacera often has slightly angular spores and thus is somewhat atypical of the smooth-spored group.
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ARTIFICIAL KEY TO INOCYBE SPECIES WITH APICALLY ELONGATED NODULOSE BASIDIOSPORES |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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of stipe length, with dense sand-filled mycelium at the base; stipe lavender (or vinaceous, 7.5R 8/4) in all developmental stages. Basidiospores 12.0–13.8–15.5 (–16.5) x (4.0–)4.5–4.9–6.5(–9.5) µm, variable in outline from smooth and elongated to irregular, a relatively small proportion (approximately 10–15%) more-or-less rocket-shaped with 1–4 basal nodules and an elongated apex; spore color yellowish under the microscope (2.5Y 9/2–9/4). Basidia 29–39 x 8.4–12 µm, clavate, mostly 4-spored with a few 2-spored basidia present. Pleurocystidia 52–71 x 14–25 µm, walls mostly less than or equal to 0.5 µm, sometimes up to 1.0 µm thick, walls hyaline or pale yellowish, mostly fusiform-ventricose usually above a basal pedicel, occasionally nearly pyriform with a tapered apex, apices obtuse, often crystalliferous at the apex. Cheilocystidia 40–74 x 14–25 µm, similar to pleurocystidia, mostly fusiform-ventricose, occasionally subcylindrical, usually with a basal pedicel, apices obtuse, mostly crystalliferous at the apex, no paracystidia seen. Lamellar trama hyaline, parallel hyphae about 8 µm diam. Pileipellis a cutis of repent hyphae up to 22 µm diam that are lightly incrusted and golden brown, incrustations pale yellowish. Stipitipellis apex (upper –) covered by a superficial layer of hyphae 3–18 µm diam that often are lightly incrusted with pale yellowish material, many of the hyphae terminate in thin-walled cystidioid cells that tend to be clustered, cauloparacystidia not observed; the remainder of the stipe covered by a superficial layer of tangled hyphae similar to those at the apex but lacking cystidioid hyphal tips. Clamps present. Habit, habitat and distribution. – Solitary on sandy soil, associated with loblolly pine (Pinus taeda L.) The only known specimens were collected in eastern Texas during April.
Specimens examined. – USA. TEXAS. Walker County, Sam Houston National Forest, near Huntsville, 19 Apr 1952, leg. H.D. Thiers, H.D. Thiers 1508 (HOLOTYPE MICH).
Comments. – Inocybe texensis is known only from the holotype that consists of two single halves of different basidiomata. The location of the complementary halves of these specimens is unknown. One of the pieces appears to have partially decayed before being dried and, although its microscopic characters are identical to the remaining specimen, it is not useful for macroscopic study. The other piece is in good condition and all microscopic measurements presented above, except for the details on the lamellar trama, were taken from it. Details on the lamellar trama presented above were taken from Thiers’ protologue to minimize sampling from the holotype.
The macroscopic description given above is a modification of Thiers’ original description from fresh material with added notes from the holotype. Our microscopic observations agreed with those presented in the original description, with the exception of a few minor details. We observed a wider range in the length of the hymenial cystidia than was given by Thiers, and the pileus cuticle did not stain dark brown in KOH as originally described.
A relatively low 10–15% of the spores of I. texensis have the rocket-shape outline characteristic of I. chelanensis. The majority of spores are somewhat variable in outline, and many resemble the rather unusual spores of I. hotsoniana Stuntz (Stuntz 1947) by being elongated and more-or-less amygdaliform with a narrowed apex. However, I. hotsoniana differs from I. texensis macroscopically and by lacking rocket-shaped chelanensis-type spores. It also is characterized by having distinctive cap-like pleurocystidial apices, although we are uncertain how consistent this trait is. Inocybe texensis differs from I. chelanensis and I. dolichospora by having a lavender stipe along with a pileus that tends to become scaly. In addition, the pleurocystidia and cheilocystidia of I. texensis are thin-walled, unlike those of I. chelanensis. The hymenial cystidia of I. sierraensis are thin-walled, but this species has been found only soon after snowmelt in the Sierra Nevada. It differs sharply from I. texensis by its overall whitish to cream coloration and glabrous to finely fibrillose pileus surface. Inocybe rennyi and I. ortegae both have very polymorphic spores.
To our knowledge, no one has reported I. texensis aside from the holotype and it is probably quite rare. Examination of nearly 100 unidentified specimens of Inocybe from eastern Texas housed at the Field Museum of Natural History (F) failed to turn up any additional material of this species. Additional collections are needed to further document the variation and distribution of this fungus, but the holotype is distinctive enough that, until more material can be studied, its status as an autonomous species is warranted.
Inocybe sierraensis Kropp et Matheny sp. nov. FIG. 2b
Pileus 20–40 mm latus, convexus vel applanatus, siccus et glaber, albus interdum laete brunneus, marginem versus leviter fibrillosus et cremi-color. Lamellae albae vel pallide brunneae, adnatae vel basi rotundato-adnatae. Stipes 10–30 mm longus, 5–10 mm crassus, aequalis, albus, interdum laete cinero-brunneolus tinctus, leviter fibrillosus. Sporae 10.5–12.8–16.5(–17.5) x (5.0–)5.5–6.8–7(–8.0) µm amygdalifomes vel angulatae vel triangulares; nodulis basalibus 1–3 plerumque solitaribus. Basidia 37–49 x 9–14 µm, clavata. Pleurocystidia 51–99 x 9–20 µm, subventricosa vel clavata, interdum septata, tenuitunicata. Cheilocystidia 60–112 x 10–21 µm, subventricosa vel clavata, interdum septata, tenuitunicata. Caulocystidia nulla. HOLOTYPUS: DED 6101, SFSU.
Pileus 20–40 mm diam, broadly convex to nearly plane at maturity, margin incurved to straight and sometimes lobed; surface dry and dull, disk smooth and glabrous, usually white, becoming finely, radially fibrillose toward the margin, this region cream (10YR 9/4) with pale ochraceous to tan fibrils, velipellis absent, occasionally the pileus color may appear mostly light brown to medium yellow brown with whitish areas, in some cases adhering soil may obscure the ground color; context thick, soft, white; odor not distinctive. Lamellae close, bluntly adnate to very shallowly adnexed, white to pale grayish white becoming pale brown or grayish brown; light brown on dried specimens (10YR 7/6–8/6), margins entire and faintly whitish; lamellulae one tier. Stipe 10–30 x 5–10 mm, equal, cylindrical, solid, lacking a bulb, white mycelium at the base; surface lightly covered with white, longitudinally arranged fibrils, most specimens not pruinose anywhere, although some may have a scurfy appearance near the stipe apex; stipe color entirely white becoming pale grayish brown on some specimens; cortina fugacious. Basidiospores 10.5–12.8–16.5(–17.5) x (5.0–)5.5–6.8–7(–8.0) µm, varying from angular to smooth with a slightly amygdaliform outline, most spores spurred, many with a single nodule at the base giving the spores a blunt triangular appearance, the number of nodules varies from 1–3 but relatively few spores have more than one nodule; spore yellowish under the microscope (2.5Y 8.5/4–9/4). Basidia 37–49 x 9–14 µm, clavate, mostly 4-spored with a few 2-spored basidia present. Pleurocystidia 51–99 x 9–20 µm, sometimes septate, thin-walled, fusiform-ventricose, narrowly utriform, to clavate, some but not all above a basal pedicel, apices obtuse, often crystalliferous, walls pale yellowish mostly less than or equal to 0.5 µm thick, occasionally up to 1.0 µm. Cheilocystidia 60–112 x 10–21 µm, sometimes septate, walls mostly less than or equal to 0.5 µm, mostly clavate to narrowly utriform, some but not all with a basal pedicel, apices obtuse, often crystalliferous, no paracystidia observed. Lamellar trama hyaline, smooth, parallel hyphae, 4–15 µm diam. Pileipellis a fairly dense cutis of repent hyphae 2.5–6 µm diam with incrusting and adhering yellowish or brownish yellow material; velipellis not present in cross sections. Stipitipellis covered by a sparse superficial layer of yellowish hyphae 3.0–9.0 µm diam, this scant on some specimens; hyphae sometimes with adhering and incrusting material; some cystidioid hyphal tips present, these solitary or in clusters and most common apically, decreasing in abundance toward the base, sometimes absent at the base; cauloparacystidia nowhere observed. Clamps present.
Habit, habitat and distribution. – Gregarious, subhypogeous on soil soon after snowmelt, early June. Some basidiomata joined at the stipe base. Associated with Abies sp. or with mixed Ceonothus sp. and Abies sp. in montane fir-pine forests. Known only from the Sierra Nevada of California.
Specimens examined. – USA. CALIFORNIA. Sierra County, Haskell Peak Road about 11 km from junction with Gold Lake Road, 6 Jun 1994, leg. D.E. Desjardin, DED 6101 (as I. chelanensis) (HOLOTYPE, SFSU); Sierra County, Chapman Creek Campground on California 49, 3 Jun 1996, leg. D.E Desjardin, DED 6471 (as I. chelanensis) (SFSU); Sierra County, on California 49 at Yuba Pass, 4 Jun 1996, leg. D.E. Desjardin, DED 6483 (as I. chelanensis) (SFSU).
Comments. –
Inocybe sierraensis is a snowbank fungus (Cooke 1955) characterized by its unusual whitish basidiomata that have cream-to-tan fibrillose areas on the pileus, especially near the margin. The combination of thin-walled, sometimes septate, hymenial cystidia and spurred spores is both distinctive and characteristic of this species. Other than I. sierraensis, none of the species of Inocybe with both nodulose spores and whitish basidiomata occurs as a snowbank fungus and all differ, in one or more ways, microscopically from I. sierraensis.
Inocybe dolichospora is whitish but generally only during early development (Malençon and Bertault 1970). The spores of I. dolichospora are fairly similar in size to those of I. sierraensis but have basal nodules below an elongated apex that give them the more typical rocket-shaped appearance of members of the I. chelanensis group. The spores of I. sierraensis differ by having fewer nodules than those of either I. dolichospora or I. chelanensis. In many cases, its spores either are entirely smooth or have only a single nodule, which gives the spores a spurred or triangular appearance. In addition, the cystidia of I. dolichospora are thick-walled and the species is known only from North Africa, where it associates with Cedrus species (Malençon and Bertault 1970).
Phylogenetically, I. sierraensis is sister of I. chelanensis (FIG. 1) that, like I. sierraensis, frequently occurs around melting snow in montane environments in western North America. A second sample of I. sierraensis was sequenced for nLSU and found to match the holotype but is not included in the phylogenetic analysis due to the lack of an RPB1 sequence.
Inocybe rainierensis Stuntz, Mycologia 42:109. 1950. FIG. 3a
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– of the stipe, otherwise longitudinally silky-fibrillose becoming streaked, bulb with veil remnants present, stipe color at first uniformly light pinkish tan (7.5YR 8/4), later gradually darkening to dark reddish or yellowish brown (7.5YR 5/4, 4/6) below a pallid apex; context unchanging. Basidiospores (11.0–)11.5–12.7–14.0 (–14.5) x (4.5–)5.0–5.8–6.5(–7.0) µm, mostly rocket-shaped in outline with 2–4 basal nodules and an elongated apex, some spores vary in outline from the typical rocket shape by having one or two lateral nodules, occasional spores are nearly square due to lateral nodules, and a few are nearly smooth in outline or have an elongated triangular shape; spore color yellowish under the microscope (2.5Y 9/4–9/6). Basidia 29.0–41.0 x 10.0–12.0 µm, 4-spored. Pleurocystidia 53.0–75.0 x 14.0–23.0 µm, cell walls mostly 0.5–2.5 µm thick, and pale yellowish in 10% NH4OH, fusiform-ventricose above a short pedicel, apices obtuse and usually crystalliferous. Cheilocystidia 40.0–72.0 x 16.5–25.0 µm, similar to pleurocystidia, usually with a short basal pedicel, no paracystidia observed. Lamellar trama fairly dense trama, of parallel, mostly hyaline, smooth hyphae up to 10 µm diam. Pileipellis a cutis of compact, repent, smooth hyphae up to 5.0 µm diam that are reddish brown to yellowish brown; no velipellis seen. Stipitipellis apex covered with golden brown hyphae, having some incrusting material; caulocystidia present, these variable and usually but not always thin-walled, with or without apical crystals; middle of the stipe and stipe base covered with a superficial layer of golden brown hyphae with some incrusting material, up to 12 µm diam, cystidioid hyphal tips sometimes present. Clamps present. Habit, habitat and distribution. – Associated with conifers in late August. Known only from Mount Rainier NP in the Cascade Mountains of western Washington.
Specimens examined. – USA. WASHINGTON. Lewis County, Mount Rainier National Park, at Reflection Lake, 28 Aug 1948, leg. A.H. Smith, Stuntz 4200 (HOLOTYPE WTU); Lewis County, Eagle Peak, 26 Aug 1948, leg. A.H. Smith, Stuntz 4161 (WTU).
Comments. –
The gross morphological description above has been modified from Stuntz (1947). In his description of I. rainierensis, Stuntz mentions the similarities between this species and I. chelanensis. He also emphasized that, at the time, I. chelanensis was known only from the type collection. As a result, little could be said about the extent of its variability with respect to the characters that define I. rainierensis. Since then, enough additional collections of I. chelanensis have been made to allow a better assessment of its variability and relationship to I. rainierensis, and these two taxa do not appear to intergrade with one another. Thus, we consider Inocybe rainierensis an autonomous species.
Inocybe rainierensis is distinguished from I. chelanensis by having a uniformly dark brown pileus, a stipe that is darker brown at maturity and a bulbous stipe base. It also lacks a velipellis, while one is typically, although not always, present in the material representing I. chelanensis. In addition, all known material of Inocybe rainierensis was collected in late August, while I. chelanensis usually appears in the spring just after snowbanks have melted, although it sometimes can be found throughout the summer.
Inocybe rainierensis and I. chelanensis differ little in their microscopic characters such that the two taxa only can be separated macroscopically. The hymenial cystidia of I. rainierensis and I. chelanensis are virtually identical, and their spores are also indistinguishable in outline. The spores of the I. rainierensis holotype are relatively short, only 11.5–14 µm in length compared to 14–19.5 µm for the holotype of I. chelanensis. However, they fall within the relatively wide range of variation for I. chelanensis as circumscribed here.
It should be pointed out that one collection (Stz 6241, WTU) made by Dr. Stuntz subsequent to the publication of I. rainierensis appears to be somewhat intermediate between I. rainierensis and I. chelanensis in exhibiting darker colors close to those of I. rainierensis. However, this collection lacks a bulbous stipe base and has the velipellis that usually is found on I. chelanensis. Dr. Stuntz (unpublished) noted this but did not assign the collection to either taxon, and it is unknown how he intended to interpret the material. This collection is considered by us to be a color variant of I. chelanensis.
Inocybe rainierensis is rare and is known only from two collections from montane environments in western Washington. Little is known about its geographical distribution or ecology.
Inocybe candidipes Kropp et Matheny sp. nov. FIGS. 3b, 6c
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Pileus 10–34 mm diam, conic to convex or nearly plane with an umbo; surface dry, disk covered with a pallid velipellis, radially fibrillose toward the margin, not rimose, somewhat felty, occasionally with appressed squamules in age; color of areas beyond the velipellis medium grayish brown (10YR 8/4, 7/4, 7/6, 6/6, 4/4); context white; odor faint, spermatic to acidulous. Lamellae close, adnate to sinuate, ventricose, up to 4 mm broad; color whitish when young, becoming gray-brown at maturity (2.5Y 6/4, 10YR 7/6); margins entire, faintly white-fimbriate; lamellulae one tier. Stipe 27–40 x 3–6 mm, solid, equal to the base that usually has a slight nonmarginate bulb; surface appearing nearly glabrous but lightly pruinose at the apex and sometimes lightly pruinose over the entire stipe, cortina not observed; color white when young becoming somewhat yellowish brown (2.5Y 8.5/4) on some specimens in age, this at least partly due to spore deposits; context white in young material developing gray brown tints (2.5Y 8/4, 7/4) in age. Basidiospores (10.0–)11.5–12.8–14.0(–14.5) x 5.0–5.6–6.0(–6.5) µm, mostly rocket-shaped in outline with 1–4 basal nodules and an elongated apex; occasional spores vary in outline from the typical rocket shape by having an elongated triangular outline or one or two lateral nodules that sometimes give a rectangular outline; spore color yellowish under the microscope (2.5Y 8.5/4). Basidia 27–36 x 8–14 µm, clavate, 4-spored, sometimes 2-spored. Pleurocystidia 49–69 x 14–24 µm, slightly thick-walled, walls 1.0–2.5 µm thick, pale yellowish, fusiform-ventricose, occasionally ovate, usually above a short pedicel, apices obtuse and often crystalliferous. Cheilocystidia 44–68 x 16–26 µm, walls 0.5–3.0 µm, similar to pleurocystidia, mixed with smaller and variably shaped cells. Lamellar trama parallel, mostly hyaline, smooth hyphae up to 16 µm diam. Pileipellis a cutis of repent hyphae that are brown, incrusted, and up to 12 µm diam; velipellis visible in section, composed of hyaline, sometimes lightly incrusted hyphae up to 12 µm diam. Stipitipellis nearly glabrous, a very light superficial layer of hyaline to yellowish, sometimes incrusted hyphae, can be seen especially near the base; caulocystidia present at the apex, these similar to the cheilocystidia but variable and often clustered, thick or thin-walled, with or without apical crystals, caulocystidia scattered over the remaining length of the stipe but very infrequent on the lower portion, at times mixed with short cylindrical cells. Clamps present.
Habit, habitat and distribution. – Gregarious on soil and associated with Pinus ponderosa in montane environments. Thus far, known only from Arizona. Appearing in midsummer.
Specimens examined. – USA. ARIZONA. Apache County, 23 Jul 1999, leg. B.R. Kropp, BK 23-Jul-99-26 (HOLOTYPE UTC); Coconino County, 24 Jul 1999, leg. B.R Kropp, BK 24-Jul-99-7 (UTC).
Comments. – Inocybe candidipes thus far has been found along the Mogollon Rim of Arizona in association with Pinus ponderosa. The taxon is known only from the material cited here, but the collections were found at sites widely separated from one another. Extensive collecting of Inocybe species has not been done in the southwestern United States. However, it is likely that I. candidipes will be found throughout that region and perhaps in adjacent parts of Mexico. Inocybe chelanensis occasionally occurs in association with Pinus ponderosa in parts of Washington, but material corresponding to I. candidipes has not been confirmed in that area.
Although I. candidipes has a spore outline similar to I. chelanenis, it is rather distinctive in the field because of its sometimes lightly pruinose stipe, which is usually white especially when young, and the lack of a rimose pileus. The stipe usually has a weak non-marginate bulb. Brownish hues may be present on the stipe in age. The basidiospore length of I. candidipes also occurs at the lower end of the range for I. chelanensis as circumscribed here. Moreover, both RPB1 and nLSU sequence data confirm that I. candidipes is not closely related to I. chelanensis (FIG. 1), and that the rocket-shape outline of their basidio-spores has arisen more than once independently over the course of evolution within Inocybe.
Inocybe chelanensis Stuntz, Mycologia 39:26. 1947. FIGS. 4a, b and 6a, b
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–, below this covered with a pallid silky fibrillose layer that is usually thin but is sometimes thick enough to impart an overall whitish color to the stipe (FIGS. 6a, b), occasionally nearly glabrous; ground color usually yellowish brown, or dull pinkish brown, sometimes dark reddish or yellowish brown (10YR 7/4–6/4; 7.5YR 8/4, 8/2, 6/4, 5/4, 4/6; 5YR 7/4, 5/4), rarely white, often having light rose or grayish pink hues (2.5YR 8/6, 6/6; 5YR 5/4, 8/6) near the apex which also may be pallid; stipe base usually white; context pallid or with weak flesh tones, unchanging. Basidiospores (10.0–)11.0–14.1–17.0(–19.5) x (4.5–)5.0–5.7–6.5 (–7.5) µm, mostly rocket shaped in outline with 2–4 basal nodules and an elongated apex, some are nearly smooth in outline or have an elongated triangular outline, occasional spores vary in outline from the typical rocket shape by having one or two lateral nodules, rarely quite polymorphic; spore color yellowish under the microscope (2.5Y 9/4, 8.5/6, 8.5/4, 8.5/2). Basidia 25–47 x 8–14 µm, clavate, mostly 4-spored. Pleurocystidia 41–81 x 25–12 µm, slightly thick-walled to thick-walled, walls 1.0–2.5 µm thick, pale yellowish, fusiform-ventricose, sometimes narrowly so, occasionally ovate, usually above a short pedicel, apices obtuse and crystalliferous. Cheilocystidia 29–85 x 10–27 µm, similar to pleurocystidia, no paracystidia observed. Lamellar trama parallel, hyaline, mostly smooth hyphae 4–21 µm diam. Pileipellis a cutis of repent hyphae that are reddish brown or dark to golden brown, either incrusted or smooth, up to 12 µm diam; velipellis usually visible in section, composed of hyaline, smooth hyphae 2.5–11 µm diam, velipellis sometimes lacking. Stipitipellis apex usually with a light covering of hyaline or yellowish hyphae 2–9 µm diam, with or without light incrusting material; caulocystidia usually but not always present at the apex, versiform, usually thin-walled and often occurring in clusters, with or without apical crystals; the remainder of the stipe usually covered with a light superficial layer of hyphae, cystidioid hyphal tips often present. Clamps present. Habit, habitat and distribution. – Associated with Pinus and Abies in montane environments throughout western North America. Solitary to gregarious, sometimes caespitose on soil. Appearing from spring to mid-August and occasionally found near melting snowbanks at high elevations.
Specimens examined. – USA. CALIFORNIA. Sierra County at SFSU field campus, 4 Jun 2000, leg. D.E. Desjardin, DED 7165 (SFSU); Sierra County, SFSU field campus, 1 Jun 1997, leg. P.B. Matheny, PBM 491 (WTU); IDAHO. Idaho County, Seven Devils Mountains, 30 Jul 1954, leg. Stuntz, Smith, Bigelow, Kennedy, Stuntz 8804 (MICH); OREGON. Baker County, Wallowa-Whitman National Forest, 2 Jul 1995, leg. S. Albee, BK 2-Jul-95-1 (UTC); UTAH. Rich County, 27 Jul 1997, leg. B.R. Kropp, BK 27-Jul-97-21 (WTU); Rich County, 27 Jul 1997, leg. B.R. Kropp, BK 27-Jul-97-22 (WTU); Rich County, 15 Jul 1997, leg. B.R. Kropp, BK 15-Jul-97-16a (WTU); Rich County, 15 Jul 1997, leg. B.R. Kropp, BK 15-Jul-97-16b (WTU); WASHINGTON. Chelan County, Chesapeake Saddle, 8 Jul 1938, leg. D.E. Stuntz, Stuntz 659 (HOLOTYPE WTU); Chelan County, Wenatchee Lake State Park, 27 May 1951, leg. D.E. Stuntz, Stuntz 6241 (atypical) (WTU); Kittitas County, Table Mountain, Naneum Meadows, 15 Aug 1999, leg. P.B. Matheny, PBM 1583 (WTU); Yakima County, near confluence of American and Bumping rivers, 23 May 2000, leg. R. Halling, PBM 1870 (atypical) (WTU); Kittitas County, Table Mountain, 28 Jun 2000, leg. P.B. Matheny, PBM 1917 (WTU); Kittitas County, Table Mountain, 28 Jun 2000, leg. P.B. Matheny, PBM 1918 (WTU); Kittitas County, Table Mountain, 23 Jun 2000, leg. P.B. Matheny, PBM 1919 (WTU); Kittitas County, Table Mountain at Naneum Meadows, 15 Aug 1999, leg. P.B. Matheny, PBM 1583 (WTU); Kittitas County, Table Mountain, 12 Jul 2000, leg. P.B. Matheny, PBM 1930 (WTU); Chelan County, Fish Lake, 8 Jun 2002, leg. P.B. Matheny, PBM 2314 (WTU); Kittitas County, Table Mountain, Reecer Canyon Road, 30 Jun 2002; leg. P.B. Matheny, PBM 2329 (WTU).
Comments. – Of the species treated, Inocybe chelanensis is the most common and widespread. It is reported thus far from montane to subalpine environments in much of the western United States and probably also occurs in similar habitats in adjacent Canada and Mexico. To our surprize, both RPB1 and nLSU sequences (FIG. 1) reveal a close relationship between I. chelanensis and the umber brown scaly I. stellatospora, despite their markedly different outward appearances and differences in basidiospore and cystidia morphologies.
Most collections of I. chelanensis macroscopically fit the protologue of Stuntz (1947) quite well and are characterized by having a light yellowish brown to medium gray-brown pileus with a pallid velipellis over the disk. Typical collections have stipes with a brownish overall ground color and incarnate reflections at the apex. They are lightly pruinose at the apices and are covered by a relatively light whitish, silky-fibrillose superficial layer below. However, variation occurs for pileus and stipe color as well as for the degree to which the velipellis and stipe covering are developed. The velipellis of some specimens is entirely lacking while in others, it is exceptionally thick and imparts a whitish overall appearance to the pileus. The incarnate hues typically seen at the stipe apices also may be absent and, in this case, the ground color of the stipe is brownish. A few of these variants, if considered individually, are fairly distinctive in appearance but fit within I. chelanensis as circumscribed here when all characters are considered together.
Most microscopic characters of I. chelanensis exhibit relatively little variation and the structure of the pileipellis, lamellar trama, and stipe surface remain quite constant between collections. Our measurements of the basidiospores and hymenial cystidia from the holotype of I. chelanensis agree well with those given by Stuntz (1947). However, the dimensions of the basidiospores were found to vary markedly among the collections representing this taxon. The basidiospores from the holotype were the longest encountered, while spores from the specimens from Utah were among the smallest and were nearly nonoverlapping in length with those of the holotype. A comparison of the available collections shows that their basidiospore measurements thoroughly intergrade and there is no correlation between spore size and any of the other characters. None of the variants of I. chelanensis studied warrants formal taxonomic recognition.
The basidiospores of a number of the collections of I. chelanensis are somewhat polymorphic but with the typical rocket-shaped outline exhibiting basal nodules and an elongated apex clearly predominant. However, certain collections, (i.e., PBM 1870, WTU) produce basidiospores that are strongly polymorphic, exhibiting outlines that ranged from the typical rocket shape to ovate, amygdaliform, rectangular or very irregular with lateral and/or apical nodules present in addition to the typical basal nodules (FIG. 4b). This atypical material agrees in all other respects with I. chelanensis. It is interesting to note that quite a few of the basidia present were malformed and some produced two or three sterigmata rather than four (FIG. 4b). Spores still attached in microscopic mounts to these odd basidia sometimes were quite unusual in outline. The polymorphic spores from these collections probably resulted from abnormalities in basidial development. Because of the variation in spore outline, the range in spore size and differences in macroscopic characters among collections of I. chelanensis, variants of the species should be interpreted cautiously.
The spores of I. ortegae (Esteve-Raventós 2001) also are polymorphic, although their outlines differ somewhat from the atypical collections of I. chelanensis. Inocybe ortegae differs from I. chelanensis primarily in the outline of its spores, although it also differs somewhat in pileus color. In addition, it occurs with species of Quercus and Cistus rather than with conifers (Esteve-Raventós 2001). We have not studied I. ortegae, but it is only known from the type collection and additional material needs to be found before an assessment of its variability and relationship to I. chelanensis can be made.
Inocybe dolichospora is another species occurring outside North America that appears very similar to I. chelanensis. The only substantial differences between these two species are that I. dolichospora is associated with Cedrus in North Africa and that its basidiomata are reported to remain whitish or pale ochraceous until maturity (Malençon and Bertault 1970).
Inocybe rennyi (Berk. & Broome) Sacc., Sylloge Fungorum 5:788. 1887. FIGS. 5a, b
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Agaricus rennyi Berk. & Broome, Ann. Mag. Nat. Hist. Ser. 5, III, p 205. 1879. Pileus 6–25 mm, broadly convex to nearly plane at maturity, margins not inrolled, most specimens umbonate, sometimes acutely so when young, becoming broadly umbonate in age; surface coarsely radially fibrillose except disk, which is nearly glabrous; context thin; color of herbarium specimens dark, reddish brown to dark brown at disk (7.5 YR 3/2–7.5 YR 4/4) becoming lighter brown (7.5YR 5/4) toward the margin, some nearly uniform in color. Lamellae close, adnate to somewhat adnexed, ventricose; color rusty brown (7.5YR 5/6–4/6) as observed on herbarium specimens; margins fimbriate; lamellulae one tier. Stipe 14–30 x 0.5–2 mm (45–50 mm long for holotype), equal, base with matted mycelium; cortina present, longitudinally fibrillose along entire length, occasionally lightly pruinose at apex in herbarium specimens; nearly concolorous with the pileus for most specimens except at the apex which is light grayish on herbarium specimens. Basidiospores (7.5–)9.5–13.3–16.0(–17) x 4.5–5.3–6.5(–7.0) µm, (10.5–13.9–19.5 [–22.5] x 5.0–5.8–7.5 [–9.0] µm for holotype), oddly shaped, extremely polymorphic in size and outline, usually greatly elongated apically, with numerous irregularly formed low nodules and occasional large nodules present along the spore outline; spore color pale yellowish under the microscope (2.5Y 9/4). Basidia 24–29 x 7–9 µm (28–32 x 9–11 µm for holotype), mostly 4-spored, some producing two spores, poorly reviving and difficult to measure. Pleurocystidia 42–62 x 12–21 µm (53–58 x 16–24 µm for holotype), cell walls mostly 1.0–2.0 µm thick and pale yellowish, some thinner (0.5 µm) or thicker (3.0 µm), cellular contents of some also pale yellowish, ovate to fusiform-ventricose with apices that are somewhat papillate to obtuse and often crystalliferous, pedicels often long but sometimes short. Cheilocystidia 40–65 x 13–25 µm (50–62 x 14–23 µm for holotype), nearly identical to pleurocystidia, cell walls 0.5–1.5 µm thick, some up to 2.5 µm, often pale yellowish, some with yellowish contents, ovate to fusiform-ventricose with papillate to obtuse apices, often with crystals at the apex, pedicels either long or short. Lamellar trama of parallel hyphae, 7–15 µm diam, smooth and light yellow as revived in NH4OH, poorly reviving. Pileipellis a dense cutis of poorly reviving hyphae 2–4 µm diam (up to 8 µm for the holotype); hyphae yellowish brown to golden yellow with some light incrustations. Stipitipellis apex covered with loosely tangled, mostly hyaline smooth hyphae, 2–5 µm diam, clusters of versiform caulocystidia present at the apex of some specimens, these usually thin-walled, sometimes thick-walled, with or without apical crystals; the stipe middle and base are covered with fairly dense, tangled hyphae that are mostly hyaline, some with light incrustations. Clamps present.
Habit, habitat and distribution. – Known both from Europe and North America. Collected during spring and summer on the ground in woodlands.
Specimens examined. – ENGLAND. Herefordshire, leg. J. Renny, K 60274 (HOLOTYPE K). USA. MASSACHUSETTS. Suffolk County 10 Jul 1916, leg. S. Davis (MICH 00041358); Suffolk County 19 Jul 1918, leg. S. Davis (MICH 00041357), Suffolk County 16 Sep 1918, leg. S. Davis (MICH 00041356).
Comments. –
Inocybe rennyi is a rare species that is known to occur in Europe and is represented in North America by very few collections (Smith 1939). Except where indicated, the above description is based on North American material housed at MICH that agrees very closely with the holotype of I. rennyi.
Inocybe rennyi is an unusual fungus that is readily recognized by its polymorphic and oddly shaped spores. In addition to the distinctively elongated, nodulose shape of its spores, I. rennyi differs from I. chelanensis by its coarsely fibrillose pileus that consistently lacks a velipellis. It differs from I. texensis by lacking lilac hues on its stipe and by having thick-walled cystidia. Inocybe sierraensis differs from I. rennyi by having both whitish basidiomata and thin-walled cystidia. Little information is available about the habitat or hosts of I. rennyi because none of the specimens examined contained notes on habitat. Heim (1931) indicates that it might occur in montane environments or, in one instance, on the ground in "Dinedor Fir Wood", while Alessio and Rebaudengo (1980) list hardwoods as associated hosts.
We are aware of only five specimens from North America that have been identified as I. rennyi (Smith 1939, Stuntz 1947). Three of these, collected in Massachussetts, are housed in MICH, and these are the only North American specimens that agree accurately with the holotype for this species. Many of the basidiomata in the MICH collections have caulocystidia at the stipe apices. We could not confirm the presence of caulocystidia on the holotype because the specimens are glued to a cardboard card and the surface of the stipe apex has now almost disappeared from the specimen. However, caulocystidia were nicely illustrated by Stangl (1975) from holotype material. He indicates their presence on the upper third of the stipe, while on the MICH material they are present only on the extreme apex.
The other two North American specimens were reported by Stuntz from Washington (Stuntz 1947). These specimens both vary to differing degrees from the holotype of I. rennyi and they also are quite different from one another. One of them (Stz 610, WTU) has a dark brown, squarrose to lanuginose, convex pileus that, as was noted by Stuntz, strongly resembles that of I. lanuginosa (Bull. : Fr.) Kummer. Except for its spores, we do not believe that this collection fits the type-concept of I. rennyi and have excluded it from our description. It might be an aberrant developmental form of one of the species in the I. lanuginosa group. The other collection (Stz 438, WTU) more closely resembles the holotype of I. rennyi but differs by having longer hymenial cystidia, somewhat different colors and by being more robust.
The status of I. rennyi has been somewhat controversial. Most authors have retained it as an autonomous taxon while pointing to its strong similarity to I. cicatricata Ellis & Everh. or I. curvipes P. Karst. Heim (1931), for example, emphasized that I. rennyi differs from I. cicatricata only in its elongated spores. Stangl and Enderle (1983) also indicated that the two species can be separated only microscopically. Alessio and Rebaudengo (1980) go further and, although they treat I. rennyi as an autonomous species, they pose the question of whether I. rennyi, I. cicatricata and I. lanuginella Schroet. (=I. curvipes) all are synonyms. Esteve-Raventós (2001) more recently discussed the similarities between I. rennyi and I. curvipes, expressing the view that an elongation of I. curvipes spores would give rise to spores like those of I. rennyi. Esteve-Raventós also indicates that the smaller spores present in the holotype of I. rennyi resemble the spores of I. curvipes. Our observations of the I. rennyi holotype agree with those of Esteve-Raventós (2001). We concur that the presence of curvipes-type spores in specimens of I. rennyi, the macroscopic similarities between the two, and the similarities between the cystidia of I. rennyi and I. curvipes call the recognition of two taxa into question.
Two of the collections of I. rennyi housed at MICH (00041356, 00041358) are mixed and contain basidiomes that correspond to both I. rennyi and I. curvipes. Except for the topology of the spores, the basidiomes in these collections are indistinguishable. Under the assumption that the basidiomes in these mixed collections were produced from the same mycelium, the unusual mix of these particular species within two separate collections suggests once again that the two names could represent a single taxon.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: brkropp{at}biology.usu.edu
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSARTIFICIAL KEY TO INOCYBE...TAXONOMYLITERATURE CITED |
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0.5 µm); pileus whitish with cream areas to light brown; velipellis absent
0.5 µm); pileus yellowish brown, grayish brown, avellaneous, or dark brown; velipellis often present
