Conservation and Manageinent of Native Plants and _Fungi

Transcription

1 Conservation and Manageinent of Native Plants and _Fungi Proceedings from a C 1 onference of the Native Plant Society of Oregon Edited by Thomas N. Kaye, Aaron Liston, Rhoda M. Love, DanielL. Luoma, Robert J. Meinke, and Mark V. Wilson Foreword by Reed F. Noss

2 Laura A. Wilson 3930 NW Witham Hill #103, Corvallis, Oregon Abstract: Specimens were examined at the Oregon State University Herbarium and my personal collection in order to establish which species of Peltigera lichens are present in Oregon, their abundance, ecology, and distribution. A species list and key were constructed for the North American species of Peltigera including notes, photos, maps, and a glossary. Six taxa are reported for the first time from Oregon. Some species that have previously been reported from Oregon were not seen in the collections. Key words: Peltigera, taxonomy, key, lichens Pages T.N. Kaye, A. Liston, R.M. Love, D.L. Luoma, R.J. Meinke, and M.V. Wilson, editors. Conservation and Management of Native Plants and Fungi. Native Plant Society of Oregon, Corvallis, Oregon INTRODUCTION Peltigera lichens are easily recognized by their ecorticate, veined lower surface. Identification to species, however, is often difficult and frustrating. Nomenclatural changes, variation within species, and morphological characters that are difficult are some causes for confusion. Because of this, specimens in herbaria often have outdated or incorrect names. This makes it difficult to know which species are present in Oregon, how abundant they are, what habitats they occupy, and where they occur. Since lichens of the genus Peltigera contain the nitrogen-fixing algae Nostoc, they are an important source of nitrogen in forests, and could become candidates for future monitoring, management, and conservation. In order to update our knowledge of Peltigera in Oregon, specimens at the Oregon State University Herbarium and my personal collection were studied. METHODS AND MATERIALS Approximately 450 specimens at the Oregon State University Herbarium (OSC) and about 580 specimens from my personal collection were examined. Specimens were examined under a dissecting scope for morphological features; thin layer chromatography was not conducted. Maps were constructed showing distribution of species by county. Abundance, substrate, and habitat information were compiled from field observations. A key was constructed for all the Peltigera species listed in the Esslinger and Egan (1995) checklist using information from the keys and descriptions found in Goffinet and Hastings (1994), Goward et al. (1994), Goward et al. (1995), Holtan-Hartwig (1993), Vitikainen (1994), and personal observations. 272 RESULTS AND DISCUSSION Distribution and Ecology of Peltigera in Oregon Of the 450 specimens examined at OSC, approximately 200 specimens required name corrections. A species list was compiled, showing those species known from Oregon and those not seen from OSC or my collections. To date, 21 species and varieties of Peltigera and 2 phototypes have been documented from Oregon, and more taxa may be expected with additional field collecting. Maps were compiled for species present in the state. New finds for Oregon included Peltigera hymenina, P. cinnamomea, P. scabrosa, P. didactyla var. extenuata, possibly P. scabrosella, and the bluegreen phototype of P. britannica. Species that have previously been reported from Oregon but were not seen in the specimens examined include P. elisabethae, P. horizontalis, and P. polydactylon. Many packets labeled P. canina and P. polydactylon were found at OSC, but almost all were transferred to other species. Peltigera canina appears to be a rare species, and P. polydactylon is rare or nonexistent in Oregon. Most of the species had an exclusively western distribution. The largest number of species were present in the western Cascade Mountains. Typical Peltigera habitats included rocky outcrops, trail and road-cuts, areas close to lakes and rivers, and moist coniferous and mixed forests in general. Most species were collected on substrates such as moss, soil, rocks, and logs. Peltigera collina and P. neckeri were unusual in their preference for tree bark and twigs as a substrate, though they were occasionally found on soil. Common species of the coastal dunes and lake areas included P. hymenina, P. neopolydactyla, P. membranacea, P. britannica, and P. leucophlebia. Peltigera didactyla (both varieties), P. praetextata, P. scabrosa, and the blue-green photo- type of P. britannica were also be found at the coast but were not as common. In the Willamette Valley, P. collina and P. neckeri were found on trees; P. collina was one of our most abundant species. Though much of the Willamette Valley is agricultural or grassland, there are some grassy, rocky meadows, and disturbed areas that supported a small community of species, including P. membranacea, P. rufescens, P. ponojensis, P. didactyla var. didactyla, P. venosa and the light veined variant of P. kristinssonii. Only P. rufescens seemed able to thrive in the more arid central and eastern regions of Oregon, typically on so1~or rocks. A few species, such asp. didactyla var. extenuata, P. scabrosa, and P. cinnamomea, may be at the southernmost part of their range in Oregon. It is not known if the scarcity of collections in the central, southern, and eastern portions of Oregon reflects a true lack of diversity or a lack of collecting. The key includes all species listed for North America because it is possible that, with additional collecting in the northeastern corner of the state and the Mt. Hood area, some of the species not seen so far might be found. New Records for Oregon It is sometimes difficult to know if a species has been reported from Oregon, especially with all the nomenclatural changes that have occurred. Peltigera hymenina is reported for the first time for Oregon; it was found almost exclusively in coastal dune areas. It had been known asp. lactucifolia (now considered a synonym of P. hymenina) from Europe for some time, and was recently reported from coastal regions in British Columbia (Goffinet et al., 1995). Peltigera cinnamomea, a recently described species (Goward et al., 1995), is also a first report for Oregon; specimens were collected on mossy rocks along a trail cut near Opal Creek, Marion County, and on soil at the lake edge at Detroit Reservoir, Linn County. The thallus at Detroit Reservoir measured almost a meter in diameter! Specimens of P. cinnamomea were also seen at OSC from Latah County, Idaho and Winsor County, Vermont; these were also first reports for those states. Another recently named taxon, P. didactyla var. extenuata, was seen at OSC from Saddle Mountain in Clatsop County for the first time from Oregon. Several specimens of Peltigera scabrosa were collected at Cascade Head, Tillamook County, on a rotting log. Additional specimens collected a year later from the same log were appeared different; they were scabrous, but only at the margins, the upper surface was lighter in color, and the...,.thalli were smaller. These later collections may fit the scabrous form of P. neopolydactyla mentioned by Goffinet and Hastings (1994) found in western coastal areas. Peltigera scabrosella may also be present in Oregon. One specimen that fits the description of that species was collected on the north side of 273 Detroit Reservoir; however, chromatography is needed for verification. The specimen may have a tiny amount of tomentum near the lobe tips, andperhaps should be referred to the light veined variant discussed under P. kristinssonii. The blue-green phototype of Peltigera britannica is another new report. All the blue-green phototype specimens seen so far from Oregon appeared to be of P. britannica or P. venosa' types. Phototypes of P. britannica were seen from four local-. ities: Woahink Lake and Clear Lake near Florence, Lane County; Menagerie Wilderness, Linn County; and Opal Creek. In each case, the thalli were locally abundant on disturbed, steeply sloping soil along a trail or road cut. These thalli often had both green and blue-green lobes, and were easy to overlook because of their small size. Larger thalli that appear to lack any green lobules were rarely collected; they were strongly maculate and had diffuse dark brown veins on the lower surface. They bore a very close resemblance to P. frippii except they were tomentose on the upper surface. It is thought that the blue-green phototypes of the green Peltigera are thalli that grow out from cephalodia. They do not have their own species names. This is because the nomenclatural rules for lichens specify that the epithet applies to the fungal species of the lichen (which is the same in both the green and blue-green phototypes; only the primary alga is different). The blue-green phototype of P. venosa was also seen on soil under and around P. venosa lobes in most of the specimens examined. It is similar to Leptogium in appearance. Reports on distribution of the blue-green phototypes have been sketchy and tentative in part because of their rarity, but also because it is not always possible to determine which green Peltigera they represent. This is especially true of herbarium specimens where material of green Peltigera that usually grow in association with the phototype was not collected. Based on evaluation of herbarium specimens, we now know that at least the P. britannica and P. venosa phototypes occur in Oregon. Difficult Taxa Several species were especially difficult to separate, had a large degree of variation in their morphology, were unidentifiable, or merit further discussion for other reasons. The species of green Peltigera, P. aphthosa, P. britannica and P. leucophlebia, have characteristics that intergrade somewhat. It was common to find thalli that had both peltate and flat appressed cephalodia. Also, anomalous thalli with peltate cephalodia and patchy-corticate backed apothecia have been found. Peltigera collina varied somewhat in thallus morphology. Some forms were thick and robust, with prolific soredia. Other forms had a thin, delicate thallus with only sparse soredia and very narrow delicate dark brown veins. Laminal soredia were occasionally found.

3 Ct " ' ': -~ f a d e FIG. 1. Rhizine types: a. simple, threadlike, b. apically branched, c. tomentose, d. abundantly branched, e. fibri~lose, f-i. fascic~late, j. penicillate. FIG. 2. Textures of the upper surface: a. tomentum, b. scabers, ~ pruina, d. maculae. Fr~. 3. Apothecial types: a. honzontal, b. vertical, saddle-shaped, c. vertical, fingernail-shaped. FIG. 4. Cephalodia types: a. flat, b. wart-hke, c. peltate.. ~ c. It was often difficult to differentiate Peltigera canina and P. membranacea; intermediate forms were common in our area. Some specimens had the P. canina-like characteristics of a thick thallus, tomentum over most of the upper surface, and small to medium lobe width, and the P. membranacea-like characteristics of tomentose rhizines. Specimens that had both tomentose and apically to densely branched rhizines were also encountered. Peltigera membranacea was much more common in our area than P. canina, though some specimens appeared to be hybrids of the two taxa. ;~ Peltigera neopolydactyla '" was an extremely variable species. The undersurface of this species had very pale, indistinct veins, or brownish (sometimes very dark brown) distinct veins all the way to the margins, or a non-veined mostly brown lower surface with a pale margin. The rhizines were short or long, simple or fasciculate. Field color of thalli ranged from brown to gray to almost black or green. Specimens of these morphotypes seemed very different, yet intergradations existed between the extremes, and all had the common feature of large glabrous lobes. Distinguishing young or small coastal specimens from P. hymenina was sometimes difficult. It would be interesting to study specimens of Nostoc algae in the color variants of this species. However, if different species were discovered it would result in more nomenclatural problems for the genus. Peltigera kristinssonii appeared to be rare in Oregon. Typical specimens had broad, very dark veins that were flat or slightly raised, with short, fasciculate, dark rhizines. Several specimens collected in the Willamette Valley and the western Cascades had scabers and tomentum but did not fit the other characters of P. kristinssonii (see Figure 30). They had ropy, pale to light brown veins, and long (about 1 em), mostly dark brown, simple to delicately fasciculate rhizines. Specimens collected from one locality were all distinctly green when wet; other specimens did not have their field color noted. Some specimens also had a very strong, disagreeable odor (similar to Sticta odor) especially when wet but still very discernable when dry. Thin layer chromatography is needed to sort out the relationship of these specimens to P. kristinssonii, P. ponojensis and P. scabrosella. Another group of specimens did not key convincingly to any of the known species for North America. The specimens are similar to Peltigera degenii and P. pacifica but did not completely fit the descriptions for either species (see Figure 31). The thalli were glabrous and dull, the lobes averaged em in width, lobules were sometimes pres~t and occasionally abundant; the apothecia were small and reddish brown on short lobes. The veins were narrow, flat to ropy, and typically dark brown (though sometimes becoming pale in places); the rhizines were dark brown, long (to 1 em), simple or delicately fasciculate, sometimes becoming a bit fibrillose at the tips, but always remaining separate. Peltigera degenii differed in having pale veins and rhizines, a shiny upper surface, and pale apothecia. Peltigera pacifica was also close but had short fasciculate rhizines and broad veins. Chromatography is needed for a more complete evaluation of this species. Using the Key This key is designed with a sorting strategy in mind. I have found that comparison of specimens is critical to learning the species of this genus. I use boxes labeled according to the sections of the key, then within each box I sort according to species. After drying and packeting specimens, I put tentative species names in pencil on the packets. Later, when I have time, I try to work through all packets of that species in one sitting; that way, a specimen that looks different stands out from the rest. Colors described in the key refer to dried specimens (unless otherwise noted). Veins and rhizines are often paler and different near the margin than at the center. I recommend cleaning debris off the lower surface and evaluating veins and rhizines at least two centimeters in from the margin and a few different places on the specimen to determine forms and colors. Considerable variation in color and texture of veins and rhizines occurs in some species, and this is important to note (see Figure 1). Cleaning off debris is often easier once the specimen is dried and pressed well; a fresh specimen will tend to tear when dirt or moss is removed. Textures on the upper surface, especially near the lobe tips, are also important to note (see Figure 2). A good hand lens or dissecting scope is essential to see scabers, pruina, and maculae. Scabers and pruina are often hard to distinguish. Pruina consist of calcium oxalate crystals that form a white crust on the upper surface (usually just near the lobe tips) that resembles the whitish coating on a plum. Under a dissecting scope pruina look like salt crystals on top of the thallus. Scabers may look superficially similar to pruina, yet are not crystalline at all but rather a texture of the upper cortex. Under a hand lens scabers look like raised polygon-shaped mounds of hyphae; often minute cracking is present between the mounds. A scabrous thallus has a matt or velvety appearance, and is usually not shiny or glossy. Tomentum consists of brownish fuzzy or felt-like hairs that are fairly easy to see even without a hand lens. A glabrous upper surface is one that lacks tomentum and scabers, and is often shiny. Orientation of the apothecia (horizontal or vertical) is also important for distinguishing species (see Figure 3). Most species have brown to reddish-brown apothecia; but in a few cases apothecial color can be helpful to note. Peltigera coltina and P. neckeri are the only two species that have black apothecia. Pale-buff colored apothecia may be found on P J

4 degenii and sometimes also on P. membranacea. Green Peltigera have a green Coccomyxa algae in their thallus and blue-green Nostoc algae in cephalodia (tiny wart-like thalli attached to the upper surface). Cephalodia may take several P. aphthosa (L.) Willd. P. britannica (Gyel.) Holtan-Hartw. and Tonsb. P. canina (L.) Willd. P. cinnamomea Goward P. collina (Ach.) Schrader P. degenii Gyel. P. didactyla var. didactyla (With.) Laundon P. didactyla var. extenuata (Nyl. ex Vainio) Goffinet and Hastings P. elisabethae Gyel. * P. evansiana Gyel. * P. horizontalis (Huds.) Baumg. * P. h;ymenina (Ach.) Del. in Duby P. kristinssonii Vitik. P. lepidophora (Nyl. ex Vainio) Bitter* LAURA A. WILSON List of Peltigera Species in Key forms and are important for distinguishing some green Peltigera species; they can appear flat or pancake-like, wart-like, or peltate-squamulose (Figure 4). P. leucophlebia (Nyl.) Gyel. P. malacea (Ach.) Funck* P. membranacea (Ach.) Nyl. P. neckeri Hepp ex Mull. Arg. P. neopolydactyla (Gyel.) Gyel. P. pacifica Vitik. P. polydactylon (Necker) Hoffm. * P. ponojensis Gyel. P. praetextata (Florke ex Sommerf.) Zopf. P. retifoveata Vitik. * P. rujescens (Weis) Humb. P. scabrosa Th. Fr. P. scabrosella Holt.-Hartw. P. venosa (L.) Hoffm. blue-green phototype of P. aphthosa*, P. britannica, P. leucophlebia*, and P. venosa *no specimens were seen from Oregon. This list includes all species of Peltigera listed in Esslinger and Egan (1995). It also includes some species not on the list. KEY TO PELTIGERA GROUPS la. Thallus with a green photobiont, grass green when wet, greenish gray or brownish green when dry a. Apothecia!aminal..... Solorina (not treated here) 2b. Apothecia terminal on lobe tips... GREEN PELTS (Sect. A) lb. Thallus gray, brown, or yellowish when dry, not green when wet (occasionally blue-green when wet) a. Thallus sorediate... SOREDIATE PELTS (Sect. B) 3b. Thallus lacking soredia a. Apothecia oriented horizontally on lobe tips (see Figure 3a)... HORIZONTAL PELTS (Sect. C) 4b. Apothecia oriented vertically on lobe tips (see Figures 3b and c) a. Upper surface scabrous (check near lobe tips, see Figure 2b)... SCABROUS PELTS (Sect. D) 5b. Upper surface not scabrous a. Upper surface tomentose (check near lobe tips, see Figure 2a)... TOMENTOSE PELTS (Sect. E) 6b. Upper surface glabrous (lacking tomentum)... GLABROUS PELTS (Sect. F) SECTION A: GREEN PELTS Ia. Cephalodia absent from upper surface a. Thallus small; lobes fan-shaped; apothecia oriented horizontally; veins dark brown, distinct; occasional on soil, Willamette Valley to E. Cascades.... P. venosa (L.) Hoffm. (Figure 29. The blue-green phototype is often found with the green thallus. Look for small dark Leptogium-like lobules around the base of the thallus) 2b. Thallus not small and fan-shaped; apothecia oriented vertically; lower surface mostly black, veins lacking; thallus thick; similar top. aphthosa in appearance; no OR specimens seen (reported from Canada to CO).... P. malacea (Ach.) Funck. (Figure 19. Though this species does not have a green photobiont, it can appear green when wet and be mistaken for a green pelt) lb. Cephalodia present on the upper surface a. Cephalodia peltate-lobulate, attached to the upper cortex at the center but not at the margins, (see Figure 4c), easy to flick off with tweezers (check large ones); lower surface brown to black with a pale marginal zone, veins diffuse to lacking; apothecia with greencorticate undersides (check larger ones); common on soil, moss, rocks, coast tow. Cascades P. britannica (Gyel.) Holtan-Hartw. and Tonsb. (Figure 6) 3b. Cephalodia wartlike, flat, or cerebriform, the margins firmly attached to the upper surface of the thallus, not easy to tlick off the upper cortex (taking a chunk of upper cortex when flicked) (Figure 4a and b) a. Lower surface brown to black with a pale marginal zone, with few or no veins; apothecia with green-corticate undersides (check larger ones); rare on soil or moss, W. Cascades.... P. aphtlwsa (L.) Will d. (Figure 5) 4b. Lower surface with brown veins; undersides of apothecia with greenish corticate bumps (not continuously corticate) or ecorticate; common on soil, moss, rocks, coast tow. Cascades...?. leucophlebia (Nyl.) Gyel. (Figure 18) SECTION B: SOREDIATE PELTS la. Soredia marginal; very common on trees and moss, occasionally on rocks and soil, coast to E. Cascades..... P. collina (Ach.) Schrader (Figure 10. Note: if veins are lacking, check Nephroma parile, Erioderma and Leioderma, they closely resemble P. collina) lb. Soredia lamina!, in circular scar-like patches a. Thallus small, tomentose, consisting of small, scattered, erect, mostly fertile lobes; veins dark near center; rhizines simple to sparsely branched; occasional on soil, coast to E. Cascades.... P. didactyla var. didactyla (With.) Laundon (Figure 12) 2b. Thallus medium sized (up to 8 em); veins pale near center; rhizines densely fibrillose, forming a mat on underside; rare on soil, coastal..... P. didactyla var. extenuata (Nyl. ex Vain.) Goffinet and Hastings (Figure 13) la. lb. SECTION C: HORIZONTAL PELTS Lobules present on cracks and lobe margins; lower surface brown, with a pale marginal zone; veins absent; no OR specimens seen (reported from Canada to OR).... P. elisabethae Gyel. (Figure 14) Lobules absent; veins dark brown, distinct; no OR specimens seen (reported from Canada to OR)..... P. lwrizontalis (Huds.) Baumg. SECTION D: SCABROUS PELTS la. Upper surface tomentose (although sometimes only slightly so); thallus often yellowish; veins distinct, dark brown, flat or slightly raised; rhizines fasciculate or tomentose, dark brown; rare on soil, moss and rocks, W. Cascades..... P. kristinssonii Vitik. (Figure 16) lb. Upper surface lacking tomentum a. Lobes large (3-5 em), broadly rounded a. Rhizines short, dark brown, densely branched; scabrous over most of upper surface; veins pale to dark brown, diffuse; dried specimens with an unusual light greenish color; rare on rotting logs, coastal P. scab rosa Th. Fr. (Figure 27) 3b. Rhizines long, simple; upper surface irregularly scabrous; rare? (reported from western coastal areas) P. neopolydactyla (Gyel.) Gyel. 2b. Lobes smaller (up to 2.5 em) a. Veins dark brown, distinct; soredia present (check carefully with hand lens); very common on trees, moss, occasionally on rocks and soil, coast to E. Cascades..... P. collina (Ach.) Schrader (Figure 10) 4b. Veins pale, soredia absent a. Thallus with dark marginal and!aminal isidia and lobules; sparse hairs present near the lobe tips; no OR specimens seen (reported from Alberta, BC and eastern USA).... P. evansiana Gyel. 5b. Thallus lacking dark lobules and hairs; veins pale brown, diffuse; rhizines pale brown to white, slender, simple; rare on soil?, W. Cascades?.... P. scabrosella Holt.-Hartw. (Figure 28) SECTION E: TOMENTOSE PELTS I a. Isidia and/or lobules present and abundant a. Lobules present on stress cracks and/or lobe margins a. Thallus with both gray and green lobes, or attached to a green Peltigera; marginal lobules present; upper surface maculate; tomentum of sparse erect hairs near margin; lower surface with pale to brown, tlat, diffuse veins that merge near the center; rare (but may be locally abundant) on soil (esp. road and trail cuts), coast tow. Cascades... blue-green phototype of P. britannica (Figure 7) 3b. Thallus lacking green lobes and maculae; lobules wejj developed on cracks and lobe margins; tomentum sparse, restricted to lobe margins; veins distinct, pale to brown, flat to slightly raised; common on soil, moss and rocks, coast to E. Cascades P. praetextata (Florke ex. Sommerf.) Zopf. (Figure 25) 2b. Isidia/lobules!aminal (at least in part) a. Isidia flattened, pel tate, lobulate to squamulose; thallus small ( < 3 em); veins narrow, flat, brown; rhizines short (2 mm), pale, simple; no OR specimens seen (reported from BC to CO)... P. lepidoplwra (Nyl. ex Vainio) Bitter (Figure 17)

5 4b. Isidia granular to cylindrical, becoming squamulose on older lobes, often darker colored than the upper surface; thallus medium sized; veins narrow, low, pale to brown; rhizines pale to brown, simple; lobe margins sometimes with pruina and/or scabers; no OR specimens seen (reported from Alberta, BC and eastern USA).... P. evansiana Gyel. lb. Isidia or lobules absent, or occasionally present.... S Sa. Veins absent; lower surface brown to black with a whitish marginal zone, or veins diffuse and very indistinct a. Thallus with both gray and green lobes, or attached to a green Peltigera; upper surface maculate; tomentum of sparse erect hairs near lobe margins; lower surface with pale to brown, flat, diffuse veins that merge near the center; rare on soil (esp. road and trail cuts), coastal tow. Cascades... blue-green phototype P. aphthosa, P. britannica or P. leucophlebia (Figure 7. So far, those seen from OR appear to be all P. britannica type) 6b. Green lobes absent; maculae absent; thallus thick, green when wet; tomentum thick; lobe margins upturned; pruina sometimes present; lower surface black to brown, veinless, with a pale marginal zone; no OR specimens seen (reported from Alberta and BC to CO.... P. malacea (Ach.) Funck. (Figure 19) Sb. Lower surface with distinct veins a. Rhizines tomentose, simple, pale to dark brown; veins strongly raised and threadlike; lobe tips curling downward; very common on soil, moss, and logs, coast to E. Cascades.... P. membranacea (Ach.) Nyl. (Figure 20) 7b. Not with the combined characters of tomentose rhizines and narrow, raised veins... 8 Sa. Veins rusty-cinnamon colored (at least near center), flat to somewhat raised; thallus large; lobes up to 3 em wide; rhizines simple, pale to light brown; rare on soil, moss and rocks, W. Cascades.... P. cinnamomea Goward (Figure 9) 8b. Veins not rusty colored a. Lobe tips curling downward (check lobes without apothecia); rhizines abundantly branched, pale to dark; thallus thick; tomentose over most of upper surface; veins raised to tlattened; rare on soil, moss and rocks, Willamette Valley to central OR..... P. can ina (L.) Willd. (Figure 8) 9b. Lobe tips plane or curling upwards loa. Lower surface distinctly cottony, with deep, pitted interstices between broad, tomentose, pale to dark veins; thallus and lobes large; rhizines pale to dark brown, simple, fasciculate, or squarrose branched; no OR specimens seen (reported from AK to WA)...,.... P. retifoveata Vitik. lob. Lower surface not cottony and pitted lla. Thallus consisting of small, scattered, erect, mostly fertile lobes; upper surface with circular, scar-like areas that have soredia (check "scar" edges, easy to overlook); occasional on soil, coast to E. Cascades P. didactyla var. didactyla (With.) Laundon (Figure 12) llb. Thallus not of scattered, sorediate lobes a. Veins pale, raised; rhizines simple, long, pale to brownish; thallus small to medium sized; apothecia rare, very large (up to 1.3 em in diam.); occasional on soil, Willamette Valley to E. Cascades..... P. ponojensis Gyel. (Figure 24) 12b. Veins dark brown, flat or only slightly raised; rhizines dark brown, never simple a. Rhizines densely branched, fusing together to form a brown cottony mat over the lower surface; lobe margins wavy; tomentose over most of upper surface; thallus brown to dark brown, often with a crusty white coating; often difficult or impossible to remove substrate from lower surface, occasional on soil and rocks, coast to eastern OR... P. rufescens (Weis.) Humb. (Figure 26) 13b. Rhizines short, fasciculate or tomentose, separate, not fusing to form a mat; thallus often yellowish; upper surface scabrous, at least near lobe tips (but this is sometimes sparse and difficult to see); rare on soil and rocks, W. Cascades to eastern OR... P. kristinssonii Vitik. (Figure 16) SECTION F: GLABROUS PELTS la. Upper surface with isidia and/or lobules a. Lower surface dark brown with a pale marginal zone, essentially veinless; lobules present on lobe margins and cracks; rhizines dark brown, short, fasciculate; apothecia (when present) horizontally oriented; no OR specimens seen (reported from NWT to NM).... P. elisabethae Gyel (Figure 14) 2b. Lower surface not dark and veinless a. Isidia lamina!, granular, cylindrical, to squamulose; veins narrow, low, pale to brown; rhizines simple, pale to brown; lobe margins sometimes with pruina, scabers and sparse tomentum; no OR specimens seen (reported from BC and Alberta but distribution mostly eastern USA).... P. evansiana Gyel. 3b. Lobules/isidia mostly marginal, lamina! isidia occasionally also present a. Veins broad, dark, tlat to slightly raised; rhizines short, brown, fasciculate; lobules well developed;!aminal isidia sometimes also present; rare on moss, soil, rocks and tree bases, Willamette Valley tow. Cascades.... P. pacifica Vitik. (Figure 23) 4b. Veins narrow, pale, raised; rhizines simple, pale, long; lobules poorly developed to absent; rare on soil and moss, W. Cascades... P. degenii Gyel. (Figure 11) lb. Thallus lacking isidia and/or lobules, (occasionally with poorly developed lobules)..... S Sa. Veins narrow and raised, threadlike to ropy a. Rhizines tomentose, simple, pale to dark brown; lobe tips curling downward; this glabrous form is occasionally found on soil, moss, and logs, coastal..... P. membranacea (Ach.) Nyl. (Figure 20) 6b. Rhizines not tomentose, simple, pale; lobules poorly developed to absent; rare on soil and moss, W. Cascades P. degenii Gyel. (Figure 11. Note: the glabrous form of P. ponojensis may also key here) Sb. Veins not narrow and raised a. Apothecia black, fingernail shaped... 8 Sa. Soredia present; veins narrow, brown, flat, distinct over most of lower surface; very common, on trees and moss, occasionally on rocks and soil; coast to E. Cascades... P. col! ina (Ach.) Schrader (Figure 10) 8b. Soredia lacking; lower surface mostly brown with a pale margin; veins broad, brown, diffuse near margin (if present at all); upper surface often pruinose near lobe tips; occasional on trees and moss, Willamette Valley tow. Cascades P. neckeri Hepp. ex Mull. Arg. (Figure 21) 7b. Apothecia brown or lacking a. Lobes broa<hoften > 2.S em wide); veins extremely variable, slightly raised or flat, pale to brown, distinct or diffuse and merging; rhizines dark brown, fasciculate or simple; very common on moss, soil, rotting logs, coast to W. Cascades P. neopolydactyla (Gyel.) Gyel. (Figure 22) 9b. Lobes small to medium sized ( < 3 em wide).,... : 10 loa. Veins dark brown, flat, distinct over most or all of lower surface, some interstices present throughout lower surface lla. Rhizines short, fasciculate, near the lobe tips forming concentric rows; upper surface of thallus dimpled over outer rhizines; apothecia (when present) horizontal; no OR specimens seen (reported from Canada to NM).... P. horizontalis (Huds.) Baumg. llb. Outer rhizines not in concentric rows; upper surface lacking dimples; thallus often yellowish, shiny, with wavy margins; veins flat, distinct to margin; apothecia vertical, on elongate lobes; no OR specimens seen (reported from Canada to NM) P. polydactylon (Neck.) Hoffm. (Note: P. collina may key here, check carefully for soredia) lob. Lower surface with diffuse veins, or lower surface mostly brown and veinless a. Lower surface brown, veinless (some diffuse veins may be present near lobe margins); upper surface often pruinose near lobe tips; apothecia (when present) black, fingernail shaped; occasional on trees and moss, Willamette Valley tow. Cascades P. neckeri Hepp. ex Mull. Arg. (Figure 21) 12b. Lower surface with pale to brown diffuse veins, apothecia (when present) brown a. Lobes broad and rounded; veins extremely variable, slightly raised or tlat, pale to brown, distinct to diffuse and merging; rhizines dark brown, fasciculate or simple; very common on moss, soil, rotting logs, coast tow. Cascades P. neopolydactyla (Gyel.) Gyel. (Figure 22) 13b. Lobes narrow and acute; rhizines white to brown, simple to fasciculate; veins broad, diffuse, pale to ochraceous; occasional on sand or soil, coast to Willamette Valley..... P. hymenina (Ach.) Delise (Figure ls) ACKNOWLEDGMENTS I would like to thank Rick Meyer for computer assistance and for driving and accompanying me on numerous collecting trips. I also want to thank Tom Kaye for editorial assistance and for providing the Oregon map. And finally, thanks to the curatorial staff at OSC for allowing me access to the lichen collection, and for use of equipment there. LITERATURE CITED Esslinger, T.L. and R.S. Egan A sixth checklist of the lichen-forming, lichenicolous, and allied fungi of the continental United States and Canada. Bryologist 98: Goffinet, B. and R.I. Hastings The lichen genus Peltigera (lichenized Ascomycetes) in Alberta. Provincial Museum of Alberta Natural ~story Occasional Paper no. 21:1-54. Goward, T., B. Goffinet and 0. Vitikainen Synopsis of the genus Peltigera (lichenized Ascomycetes) in British Columbia, with a key to the North American species. Canadian J. Botany 73: Goward, T., B. McCune and D. Meidinger The lichens of British Columbia. Illustrated keys. Part. 1. Foliose and squamulose species. Ministry of Forests Research Program. Victoria, B.C. Holtan-Hartwig, J The lichen genus Peltigera, exclusive of the P. canina group, in Norway. Sommerfeltia 15:1-77. Vitikainen, Taxonomic revision of Peltigera (lichenized Ascomycotina) in Europe. Acta Botanica Fennica 152:1-96. GLOSSARY acute- becoming narrower near the tip. apothecia - reproductive structures of ascomycete fungi. In Peltigera, they are brownish, flat, circular structures located at the tips of the lobes. appressed - firmly attached to substrate. blue-green algae - a primitive type of algae (sometimes referred to as cyanobacteria) present in the body of the thallus or in cephalodia. They possess primitive pigments that give them a dark color, and have the ability to fix nitrogen

6 blue-green phototype - the thallus or lobes that contain blue-green algae as its primary photobiont; usually found growing on or in association with a thallus that contains a green algae as its primary photobiont. broad/rounded -broadening or expanding in a semicircular shape near the tip. cephalodia- tiny wartlike or lobe-like thalli containing blue-green algae, found on the upper or lower surface of green Peltigera (see Figure 4). chimeroid - lichens that have different looking lobes on the same thallus (both green and blue-green lobes). clavate- club or baseball bat-shaped. Coccomyxa - a unicellular green algae found in the green Peltigera as the primary photobiont. concentric - in semi-circular rows. coralloid- richly branched, similar in shape to coral fungi. cortex - the outer "skin" of lichen, consisting of tightly packed hypha! threads. corticate - having a cortex, with a smooth, skin-like or leathery texture (Peltigera are corticate on the upper surface of the thallus). cyanobacteria - blue-green algae. diffuse veins - veins that are not sharply or distinctly delimited from the interstices. dimples - small depressions in the upper cortex, usually above rhizines. ecorticate- lacking a cortex, cottony textured (Peltigera are ecorticate on the lower surface). erect - having an upright, or vertical habit. fasciculate- rhizines composed of brushlike bundles or groups of hyphae, (see Figure 1). fibrillose - rhizines that are richly branched and cottony. glabrous- smooth, without tomentum. globular - irregular shaped. granular- small, bump-like, with a grainy appearance. green algae - Coccomyxa algae present in the thallus of green Peltigera that contain chlorophylls a and b (green pigments); a more advanced type of algae than blue-green algae, but not able to fix nitrogen. horizontal apothecia - apothecia held parallel to the lobes, not on vertical, erect lobes (see Figure 3a). hyphae- thread-like fungal filaments. interstices - flat or pit-shaped areas between veins on the underside of the thallus. isidia - finger-like, corticate bumps found on the upper cortex, thought to function as vegetative propagules. isidiate -having isidia.!aminal - found on the upper surface of the thallus (as opposed to marginal). lobes - the flattened leaflike structures that make up the body of the lichen. lobe width (how to measure) - measure lobe width not at the tips of lobes but back in from the tip where the width is somewhat more uniform. lobulate - possessing lobules. lobules - tiny lobes, often crowded together on lobe margins or cracks in the upper cortex. maculae - small whitish spots in the upper cortex, caused by uneven distribution of algae near the surface, (see Figure 2d). maculate - with maculae. 280 margin- the edges of a lobe. marginal -present on lobe margins. medulla - the inner (usually whitish) part of the thallus, composed mainly of fungal hyphae. mycobiont - the fungal component of a lichen thallus, which is thought to provide the photobiont with moisture and anchorage. Nostoc - a filamentous blue-green algae, present in the thallus of a lichen or in cephalodia. peltate a flattened or umbrella-shaped structure attached to the upper cortex at its center, but not at its edges (see Figure 4c) photobiont - the algal component of a lichen thallus, which is thought to provide the mycobiont with carbohydrates. primary photobiont- the main algal component of a lichen thallus. pruina - crystals (calcium oxalate) which form a thin, whitish crusty layer on the upper cortex of some lichens, (see Figure 2c). pruinose- having pruina, (see Figure 2c). rhizines - root-like hypha! strands found on the lower surface of a lichen that hold it onto soil, rock, or bark. They may be stt inglike, fuzzy, or branched in a variety of ways, (see Figure 1). scabers - small polygonal shaped mounds of hyphae on the upper surface which give the upper cortex a roughened, matte appearance. Most often present near lobe tips, (see Figure 2b). scabrid- having scabers, (see Figure 2b). scabrous- having scabers, (see Figure 2b). so redia - powdery, granular, ecorticate clumps of hyphae and algae which emerge from the upper surface of the thallus or lobe margins. They are thought to function as vegetative propagules. sorediate - possessing so redia. squamulose- isidia that become lobe-like and resemble squamules. substrate - the medium a lichen grows on (soil, rock, moss, etc.). terminal - occurring at the end or tip of a structure (such as a lobe tip). thallus- the "body" of the lichen. tomentose - having tomentum. tomentum fuzzy, felt-like, usually brownish hyphae present on the upper surface of lichens, especially near lobe tips, (see Figure 2a), or on rhizines (see Figure lc). veins - vein-like structures present on the lower surface of the thallus, they may be raised and rope-like, or they may be flat. They vary in color from white to very dark brown. vertical apothecia - apothecia held upright on vertical lobes, (see Figure 3). FIG. 5. Peltigera aphthosa (Harris 1200, herb. Wilson). FIG. 6. Peltigera britannica (L.A. Wilson 1177, 1180). FIG. 7. blue-green phototype of P. britannica (L.A. Wilson 2004). FIG. 8. Peltigera canina (L.A. Wilson 2356, 2355). 281 j

7 FIG. 9. Peltigera cinnamomea (L.A. Wilson2256a, 2257b). FIG. 10. Peltigera collina (L.A. Wilson1438, 1420). FIG. 11. Peltigera degenii (L.A. Wilson 2260). FIG. 12. Peltigera didactyla var. didactyla (L.A. Wilson1470, 2015). 282 FIG. 13. Peltigera didactyla var. extenuata (L.A. Wilson 2363, 2364). FIG. 14. Peltigera elisabethae (Goffinet 3083, herb. Wilson). FIG. 15. Peltigera hymenina (L.A. Wilson 1630, 2124). FIG. 16. Peltigera kristinssonii (L.A. Wilson1635, 1640). 283

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9 FIG. 25. Peltigerapraetextata (L.A. Wilson1632). FIG. 26. Peltigera rufescens (L.A. Wilson 2412). FIG. 27. Peltigera scabrosa (L.A. Wilson1633). FIG. 28. Peltigera scabrosella (L.A. Wilson1634). 286 FIG. 29. Peltigera venosa (~.A. Wilson1406). FIG. 30. Unknown light, ropy veined, scabrous tomentose species (LA Wilson 2296) FIG. 31. Unknown dark vemed, glabrous species (L.A. Wilson 2083, 2086). ' 287