192 CHAPTER 6 Systematics of the Gomphales: The Genus Phaeoclavulina Brinkmann Admir J. Giachini1, Michael A. Castellano2, and Éfren Cázares1 1 Oregon State University Department of Forest Science Corvallis, Oregon 97331-5752 USA 2 United States Department of Agriculture Forest Service, Pacific Northwest Research Station Forestry Science Laboratory 3200 Jefferson Way SW Corvallis, Oregon 97330 USA Submitted to Mycotaxon 193 ABSTRACT The genus Phaeoclavulina has 41 species characterized macroscopically by pileate or ramarioid, greenish violaceous, brown, or orange-red basidiomata, and wrinkled to smooth hymenia. Microscopic features include monomitic hyphae, presence of clamp connections, and echinulate to verrucose, subreticulate or reticulate spores. The six pileate species are transfers from Gomphus sensu lato. These have funnel- to fan-shaped basidiomata and strictly tropical to subtropical distribution. The ramarioid section of Phaeoclavulina includes species transferred from Ramaria subgenus Echinoramaria. A remarkable characteristic of many Phaeoclavulina species is a cosmopolitan distribution. Phaeoclavulina species range from tropical to temperate regions, but are more abundant in the tropics and subtropics. A few species present a pantropical distribution, occurring in tropical places and with a refugium in the Appalachian Mountains of the USA. No Phaeoclavulina species has been confirmed as either mycorrhizal or saprotrophic, although the wood inhabiting tendency of many species indicate the genus may be saprotrophic. Additional studies on ecological and evolutionary aspects are necessary to better understand the placement and role of this diverse fungal genus. 194 INTRODUCTION The nomenclatural history of species in Phaeoclavulina, as for many others in the Gomphales, has been unsettled over the years. For the pileate species, it started when Patouillard (Doassans and Patouillard 1886) coined the name Nevrophyllum to accommodate N. clavatum (≡Cantharellus/Gomphus clavatus), later including N. viride (Patouillard 1888). The two species were grouped together because of their fan-shaped pilei. Nevrophyllum clavatum was separated from N. viride primarily by spore morphology: N. clavatum was claimed to have ‘smooth spores’, while N. viride had rough spores. Nevrophyllum viride differed from N. clavatum not only in general microscopic aspects but also by its tropical distribution. On the basis of Patouillard’s descriptions of N. clavatum and N. viride, Murrill (1910) transferred the latter to Chlorophyllum, with C. viride as the type. After discovering the name Chlorophyllum was preempted (Massee 1898), Murrill (1911) coined the name Chloroneuron to accommodate C. viride. Later species of Nevrophyllum and Chloroneuron were accommodated under Gomphus. The history of the ramarioid species of Phaeoclavulina had a similar trajectory, although with an earlier start than the pileate ones. Formal cognizance of a coralloid member of this group came when Persoon (1794) described Clavaria abietina, a name that, although popular for a time, was misapplied by Fries (1821). Reliance on the literature of Fries on the coral fungi persisted longer than for the agarics, except for the writings of Karsten (Petersen 1981). Over the years many taxonomists overlooked the works of Karsten. One problem recognized with the works of Karsten is the listing of species. Karsten’s (1882, 1889) use of asterisks preceding some but not all species names made it confusing whether he was referring to species or varieties when employing such symbols. After Fries the next taxonomic progress with the coral fungi came from Junghuhn (1830), who introduced the name C. ochraceo-virens as a substitute for Persoon’s C. abietina. According to Petersen (1981), Junghuhn’s species 195 represented a very different taxon from C. abietina. Later Brinkmann (1897) erected the genus Phaeoclavulina to include P. macrospora, which, according to him was to be related to Clavaria testaceoflava, C. spinulosa, and C. abietina. Brinkmann (1897) assigned Phaeoclavulina to the Clavariaceae, which also included Clavaria, Clavariella, Clavulina, and Typhula. Unable to recognize Fries’ concept of C. abietina, and aware that Persoon’s description antedated the newly imposed starting point, Cotton and Wakefield (1919) introduced a new name, Clavaria invalii, for the Friesian concept of C. abietina (Petersen 1981). Besides the genera mentioned above, species of Phaeoclavulina were also accommodated under Cladaria, Cnazonaria, Dendrocladium, Lachnocladium, Penicillaria, Phaeopterula, Pistillaria, Pterula, Ramaria and Thelephora. Others contributed to increase the list and knowledge of coralloid fungi, providing more in-depth clarification of the natural classification within the group. Among them are Léveillé, Patouillard, Hennings, Berkeley, Overeem, Cleland, Imai, Atkinson, and Spegazzini, just to name a few. However, one of the more important pieces on the direct relatives of Phaeoclavulina, the ‘Ramarias’, came in 1923 with Coker’s “The Clavarias of the United Sates and Canada.” Others (Corner, Fawcet, Overeem, Petersen) followed Coker’s compendium as a base for the identification and new descriptions to come. The next step on the historical nomenclature and taxonomy of coralloid fungi came with Corner’s (1950) book “A monograph of Clavaria and allied genera.” In his publication Corner recognized almost 30 ramarioid taxa, mostly from the tropics, widening the concept of ‘clavarioid’ to a new breadth. Doty (1944) published on the clavarioides of Oregon, subsequently (Doty 1948) providing a key to genera similar to that proposed by Corner. Over the following years new interest on clavarioid/coralloid/ramarioid fungi re-appeared around the world, especially on the genus Ramaria. Corner (1970) proposed to subdivide Ramaria into subgenera Ramaria, Echinoramaria, and Lentoramaria, each composed of several series nova. Marr and Stuntz (1973) proposed Laeticolora as a 4th subgenus of Ramaria. Marr (1968), Olexia (1968), 196 Dodd (1970), Marr and Stuntz (1973), and McAfee (1978) made important contributions on the North American taxa. Parmasto (1965) and Schild (1975, 1978) expanded on the European and Scandinavian taxa. Yokoyama and Sagara (1973) studied the Japanese species, and Singh (1977) and Rattan and Khurana (1978) circumscribed the Indian taxa. The literature for this group continued into the late 1900’s. In the 1970’s and 80’s Petersen produced fundamental compilations for the genus Ramaria. He produced monographs on Ramaria subgenus Lentoramaria (Petersen 1975) and subgenus Echinoramaria – to which Phaeoclavulina was a synonym (Petersen 1981). He also published several articles on species of Ramaria around the world, with an inclusive manuscript on the clavarioid fungi of New Zealand (Petersen 1988). However, even while the literature continued to add information it revealed the complexity of the group. As pointed out by Petersen (1981), detailed treatment of the group was still lacking for large members of taxa around the world; modern examination of many species and their phylogenetic relationships with other members of the Gomphales is still missing. Recently taxonomists have employed molecular methods to determine intergeneric and interspecific boundaries for taxa assigned to Ramaria and other allied genera in the Gomphales. Humpert et al. (2001) and others have shown the genus Ramaria to be non-monophyletic. Based on morphology and molecular sequences of three combined genomic regions, Giachini (chapter 2) showed, among other things, that a few species of the pileate Gomphus sensu lato formed a monophyletic group with the coralloid species of Ramaria subgenus Echinoramaria. Detailed morphological analyses of those species indicated distinctive combinations of morphological features, which resulted in the resurrection of the genus Phaeoclavulina Brinkmann (Brinkmann 1897). Phaeoclavulina was long recognized as a synonym of Ramaria subgenus Echinoramaria section Dendrocladium (Giachini, chapter 2). Two main basidiomata shapes are now recognized in Phaeoclavulina: pileate and ramarioid. 197 Pileate forms are shared by species formerly in Gomphus sensu lato. Those formerly in Ramaria subgenus Echinoramaria share the ramarioid shape. The pileate species of Phaeoclavulina are characterized by cantharelloidgomphoid8 and unipileate to merismatoid (composed of several pilei) basidiomata that display some green or violaceous color on pileus and hymenium, and by fan- to funnel-shaped pilei. Pilei are glabrous to subtomentose, and hymenia decurrent with hymenial folds rather than gills. Basidiomata vary in size, and species are known only from the tropics and subtropics. Micromorphological features include presence of clamp connections and echinulate, verrucose, subreticulate or reticulate spores. Known pileate species of Phaeoclavulina are P. africana (Borneo), P. grandis (Borneo), P. guadelupensis (Guadeloupe, in the Lesser Antilles), P. retispora (Malaysia and the Solomon Islands), P. subclaviformis (Brazil and Trinidad and Tobago), and P. viridis (Australia, Belize, French Guiana, and Martinique). Ramaria subgenus Echinoramaria was originally proposed to include species of spiny-spored Ramaria, which were also alleged to be saprotrophic (Corner 1970). Today, ramarioid species of Phaeoclavulina are recognized to have greenish violaceous, brown, or orange-red basidiomata and to occur on soil or wood, mainly in the tropics and subtropics but also in temperate regions. Micro morphological features include monomitic hyphae, presence of clamp connections, presence of gleoplerous hyphae, and ornamented spores. The ornamentation is either echinulate or verrucose, and the spore apiculus is generally prominent. Species of Ramaria subgenus Echinoramaria have long been alleged to be saprotrophic (Corner 1970, Petersen 1981), but the substrate affinity has not yet been verified by scientific investigation. The objective of this study is to describe important morphological features used to identify species of Phaeoclavulina and present a dichotomous key followed 8 Basidiomata with the similar shape of a Cantharellus species but with decurrent, wrinkled hymenia rather than straight gills. 198 by detailed species descriptions. For a complete discussion on the nomenclatural history of Phaeoclavulina see Giachini (chapter 2). MATERIAL AND METHODS Collections examined Collections of Phaeoclavulina examined in this study were obtained from the following herbaria: BPI, BR, CANB, DAR, FH, GH, H, K, LPS, M, NCU, NY, NYS, OSC, OULU, PERTH, PDD, S, TENN, and TNS (http://www.nybg.org/bsci/ih/ih.html). Holotypes of synonyms were examined whenever available. Habitat and fruiting patterns are based on the original descriptions and other relevant literature. Names of authorities are according to Brummit and Powell (1992). Macroscopic characters Macroscopic features are based on the original descriptions and notes accompanying herbarium collections supplemented by my own data on dried specimens. Features of the cap, hymenium, stipe, flesh, odor and taste were recorded whenever possible. Dimensions of basidiomata are given as the range (min-max) from all collections examined. Application of the various chemicals often used and referred to in the literature in describing species of Gomphus sensu lato provided little data of value and therefore were discontinued in this study. 199 Microscopic characters Features observed with a compound microscope were described from freehand sections mounted in Melzer’s reagent, 5% KOH, cotton blue, or water. Structures were measured with an optical micrometer at × 1000 magnification, mostly in 5% KOH, water, or Melzer’s reagent. The ranges for hypha measurements are thin walls ≤0.5 µm wide, thick walls >0.5 µm wide. The pileipellis and stipitipellis were examined in surface view. When material was available, dimensions of structures were recorded from at least 3 specimens from each of at least 5 collections; both mature and immature specimens were examined for developmental changes. Maturity was judged by the relative abundance of fully ornamented spores. Spore shape was determined as the length-width ratio of 20-30 randomly selected spores. Dimensions are given as: minlength-maxlength × minwidthmaxwidth, excluding ornamentation and apiculus. Immature or oversized spores were not included in the measurements, though variation was noted. SYSTEMATICS OF PHAEOCLAVULINA Key to pileate species of Phaeoclavulina 1. Base of stipe significantly expanded and bulbous below substrate; pileus creamy brown; uncommon, Uganda………...…...…….………...…. P. africana 1. Base of stipe not expanded or bulbous below substrate……………………….. 2 2. Pileus funnel-shaped; spores partially reticulate to reticulate……...…....… 3 200 2. Pileus fan-shaped; spores echinulate, verrucose or partially reticulate….… 4 3. Hymenial cystidia absent; basidiomata frequently merismatoid, on soil or humus; pileus pale sordid olivaceous, pale yellow-brown toward the margin…………………………………………………………….... P. retispora 3. Hymenial cystidia 90-140 × 8-10 µm long, thin-walled, cylindrical, hyaline, acyanophilic; basidiomata unipileate, on soil or wood; pileus dark sordid green, olive-green or gray-olive, becoming dark yellow-brown at the center of the pileus with age……………...………………..………..………………. P. viridis 4(2). Spores <13 µm long; basidiomata up to 9 cm tall; gleoplerous hyphae thin-walled, up to 6 µm wide; Brazil, Trinidad and Tobago.……………………………………………...….. P. subclaviformis 4. Spores >14 µm long; basidiomata >10 cm tall; gleoplerous hyphae inflated to 22 µm or more often absent; Borneo and the Caribbean...…... 5 5. Gleoplerous hyphae absent; spores (14-) 16-17.5 × 5.5-7 µm; basidiomata up to 30 × 22 cm; hymenium with tall wrinkled folds, pale olivaceous to violet in age, all parts bruising vinaceous; Borneo………………...…………....…. P. grandis 5. Gleoplerous hyphae rare, aseptate, cyanophilic, in some places inflated up to 22 µm; spores 15-18 (-25.5) × 7.5-10.5 (-12) µm; basidiomata up to 15 × 10 cm; hymenium smooth to wrinkled, orange to brown or slightly darker; Guadeloupe…………………………....…...………..………... P. guadelupensis 201 Key to ramarioid species of Phaeoclavulina 1. Spores ornamented with discrete spines at distal and proximal ends and mixed discrete spines and spines connected into circular to semi-circular ridges in median area, the ornamentation sometimes covering the entire surface of the spores………………………………………………………………… P. decolor 1. Spores echinulate or verrucose, not ornamented with spines connected into circular to semi-circular ridges in median area…………..………..…………... 2 2. Branch axils9 with narrow, sterile webbing10……..…………. P. articulotela 2. Branch axils without sterile webbing………………………...……………. 3 3. Crystals present in trama or adhered to rhizomorph hyphae………..…...…….. 4 3. Crystals absent in trama or rhizomorphs hyphae…………….……...………… 5 4. Basidiomata green to orange-green, bruising green……………………….. 6 4. Basidiomata not bruising green; apices not becoming greenish; spores without distinct oblique apiculus…………………………..……………… 7 5. Spores verrucose, ornamentation eventually connected into randomly located ridges…………………………………………………………………………... 8 5. Spores echinulate………………………………………………………………. 9 9 The area at the junction of two or more branches. Resembling the foot of a waterfowl. 10 202 6(4). Branch apices greenish; spores 5-9 × 2.5-5.5 µm, with a distinctly oblique apiculus; basidia ≤45 µm long………….………...… P. flaccida 6. Branch apices yellow to creamy yellow; spores 8-11.5 × 4-5.5 µm, without a distinct oblique apiculus, but often somewhat curved; basidia >50 µm long…..……………………………...…… P. glauco-aromatica 7. Crystal aggregates only present in trama; basidia ≥45 µm long..…. P. eumorpha 7. Crystal aggregates present in trama and/or rhizomorphs; basidia < 45 µm long………………………………………………………………………….... 33 8(5). Basidiomata bruising olive, green or blue-green on handling or weathering………………………………………..……………...….... 10 8. Basidiomata not bruising olive, green or blue-green on handling or weathering, but rather remaining yellow-orange to brown……...…… 11 9(5). Hyphae with H-connections………………………………..…………….. 14 9. Hyphae without H-connections………………………………..…………. 15 10(8). Spores 6-12 × (3-) 4-4.5 (-5.5) µm, ornamented with cyanophilic, fine to rough warts………………………..……...………… P. ochraceo-virens 10. Spores 8-12 × 6-8.5 µm, ornamented with cyanophilic, forked or lobed, stout, blunt or truncate warts and longitudinal ridges up to 2.5 µm tall…………………………………………..…………... P. pancaribbea 203 11(8). Spores nearly all longer than 9 µm………………………………………. 12 11. Spores <9 µm long………………………………………..……………… 13 12. Basidia <50 µm long; basidiomata up to 5 × 2.5 cm; stipe up to 3 mm thick; southern South America…………………….…………… P. campoi 12. Basidia ≥60 µm long; basidiomata up to 17 × 5.5 cm; stipe up to 15 mm thick; India……………….……………………………………. P. sikkimia 13(11). Spores 10.5-14 × 4.5-6 µm, ornamented with blunt, scattered, cyanophilic warts <1 µm tall; hymenium generally amphigenous11; India………………………………………………………. P. echinovirens Spores 8-11 × 6.5-8 µm, ornamented with cyanophilic blunt warts and 13. longitudinal ridges up to 3 µm tall; hymenium unilateral; New Zealand……………………………………………..………. P. zealandica 14(9). Spores 10-15.5 × 6-10 (-11) µm………………..…………….. P. zippelii 14. Spores 18-20 (-24) × 9-11 µm…………………...…...……. P. angustata 15(9). Basidiomata branches or tips with tints of olive or green when fresh or dry…………………………………………………..…………………….. 16 15. Basidiomata branches or tips without tints of olive or green when fresh or dry, but rather gray, yellow, orange, cinnamon, red or brown……..…….. 17 11 All around as opposed to unilateral (generally fertile on the outside facing side of the branch). 204 16. Spores >10 µm long…………………..…………………. P. cyanocephala 16. Spores <9 µm long…………………………………………...………… 18 17(15). Spores nearly all longer than 15 µm…………………...………………. 19 17. Spores <15 µm long………………………………..………..…………. 20 18(16). Branch apices about 1 mm long and wide, straight, digitate, creamy yellow-green to orange; basidiomata up to 16 cm tall; basidia 40-55 × 5-6 µm……………………….……………………...……. P. mutabilis 18. Branch apices waxy, dichotomous, awl-shaped, pale yellow; basidiomata up to 5.5 cm tall; basidia 30 × 7 µm………... P. ochracea 19(17). Basidia with 2 sterigmata; branch apices slightly thickened and shortly dentate or subcristate, intensely brick red to nearly scarlet; spores 15-18 × 8-10 µm; Brazil………………………………………………. P. apiahyna 19. Basidia with (2-) 4 sterigmata; branch apices orange to cinnamon; spores (12.5-) 14-20 (-22) × 4.5-8 µm; Europe, Japan, and the USA……………………………………………………….. P. macrospora 20(17). Branch apices orange to vivid red…..……………...………………. 21 20. Branch apices white to silvery, gray-yellow, gray-brown, dark oliveyellow, yellow to brown, cinnamon, orange vinaceous……...…….. 22 205 21. Spores 7-9.5 × 3-4.5 µm, ornamented with small, sparsely scattered, cyanophilic spines <0.5 µm tall; basidia <40 µm long; basidiomata up to 5.3 cm tall, delicate…………………………………………………… P. camellia 21. Spores 9-16 × 4-7.5 µm, ornamented with acute, hyaline, strongly cyanophilic spines 1-2 µm tall; basidia >45 µm long; basidiomata up to 13 cm tall………………………………………………………………….... P. cokeri 22(20). Basidiomata >11 cm tall……………………………………………. 23 22. Basidiomata ≤11 cm tall…………………………...……………….. 26 23. Basidiomata cauliflower-like; branches up to 3 cm long…...….. P. campestris 23. Basidiomata not cauliflower-like; branches >5 cm long…………..……….. 24 25. 24. Hymenium unilateral…………………………...……………… P. insignis 24. Hymenium amphigenous……………………………………………….. 25 Basidia >80 cm long; branch apices up to 20 mm long, slender, bifurcating with subacute axils, abruptly tapered, abruptly white; spores 7-10 (-14) × 4.56.5 (-8.5) µm………………………………………..…………….. P. gigantea 25. Basidia <60 cm long; branch apices up to 3 mm long, rounded-acute, golden yellow to orange; spores 9.5-11.5 × 4.5-6 µm, spore aberrations common, usually of spores up to 17 µm long and slender to a maximum of 4.5 µm wide, these usually much less highly ornamented than normal…. P. tropicalis 206 26(22). Spores 12-15 µm long………………...……………. P. griseobrunnea 26. Spores ≤11 µm long…………………………………..……………. 27 27. Hymenium amphigenous12……………..………………………………… 28 27. Hymenium unilateral……………………………………………..………… 31 29. 28. Hymenium surface velutinose-pruinose……………...……. P. cervicornis 28. Hymenium surface rugulose……………………………………………. 29 Branch apices minute, acute, irregular and congested when young, becoming flattened and irregularly digitate in age, awl-shaped, usually drying off-white, white to silvery…………………………………………...………. P. argentea 29. Branch apices dichotomous to trichotomous, not divergent, yellow, orange, orange vinaceous to brown………………….…………...….……………… 30 30. Spores 5-6 × 3-4 µm; odor pleasant, slightly of anise; taste acrid to almost salty; on the ground near bamboo in southeast Asia…...…….. P. capucina 30. Spores 5-9.5 × 2.5-5.5 µm; odor slightly earthy; taste more or less earthy, somewhat bitter; under Quercus ilex, Italy…….….…….. P. quercus-ilicis 31. Spores (3.5-) 4.5-6.5 (-8) µm long; basidia 25-36 µm long..………… P. curta 31. Spores 7-11 µm long; basidia 40-50 µm long……………...………………. 32 12 All around as opposed to unilateral (generally fertile on the outside facing side of the branch). 207 32. Branches not over 2 mm thick; stipe 20-30 × 1.5-2 mm; basidia with 2 stout, slightly curved sterigmata; Brazil, French Guiana, Guadeloupe, Jamaica, and Panama…………..………………………...… P. guyanensis 32. Branches up to 5 mm thick; stipe up to 70 × 7 mm; basidia with (2-3-) 4 stout, slightly incurved sterigmata; USA………...……….... P. longicaulis 33(7). Branch apices usually in two thin lash-like branchlets; basidiomata >6 cm tall………………………………………..…………………… P. vinaceipes 33(7). Branch apices without two thin lash-like branchlets, but rather dentate, acute to rounded; basidiomata <5 cm tall.…………..………………...… 34 34. Crystal aggregates 1.5-5.5 µm, more rarely connected into larger bodies of 8-14.5 µm; branches 3.5-7 mm thick above, subclavate, rarely 2-4branched; apices 3-7 dentate…………………………...…. P. clavarioides 34. Crystal aggregates abundant, minute; branches up to 2 mm thick, sparsely to repeatedly branched up to 4 ranks; apices acute to rounded... P. roellinii Descriptions Species are described in alphabetical order. Descriptions include citation of all material examined and the herbarium where the type and other collections are deposited. Phenology and biogeography are based on notes available with collections and the observations made in this study. 208 Phaeoclavulina Brinkmann, Jahresbericht des Westfälischen Provinzial-Vereins für Wissenschaft und Kunst 25:197. 1897, emend. Giachini, emend. prov. ≡Clavaria sect. Phaeoclavulina (Brinkmann) Killerm., in Engler and Prantl, Die Natürlichen Pflanzenfamilien 6:155. 1928 (not located, fide Petersen, Bibliotheca Mycologica 79:44. 1981). =Lachnocladium sect. Dendrocladium Pat., Journal de Botanique 3:26. 1889. ≡Dendrocladium (Pat.) Lloyd, Mycological Notes 60:870. 1919. ≡Ramaria subg. Echinoramaria sect. Dendrocladium (Pat.) R.H. Petersen, Bibliotheca Mycologica 79:44. 1981. =Chloroneuron Murrill, Mycologia 3:25. 1911. ≡Chlorophyllum (Pat.) Murrill, nom. illeg., North American Flora 9:172. 1910. =Clavaria sect. Clavariella subsect. Terrestres Killerm., in Engler and Prantl, Die Natürlichen Pflanzenfamilien 6:154. 1928 (not located, fide Petersen, Bibliotheca Mycologica 79:100. 1981). =Neurophyllum subg. Chloroneuron Heim., Revue Mycologique 19:51. 1954. =Ramaria subg. Echinoramaria Corner, Beihefte zur Nova Hedwigia 33:238. 1970. ≡Ramaria subg. Echinoramaria ser. Grandisporae Corner, Beihefte zur Nova Hedwigia 33:239. 1970. =Ramaria subg. Echinoramaria ser. Flaccidae Corner, Beihefte zur Nova Hedwigia 33:239. 1970. ≡Ramaria subg. Echinoramaria sect. Flaccidae (Corner) R.H. Petersen, Bibliotheca Mycologica 79:100. 1981. =Ramaria subg. Echinoramaria ser. Virescentes Corner, Beihefte zur Nova Hedwigia 33:239. 1970. =Gomphus subg. Phaeoclavulinoides R.H. Petersen, Mycologia 68:432. 1976a. 209 Type species: Phaeoclavulina macrospora Brinkmann, Jahresbericht des Westfälischen Provinzial-Vereins für Wissenschaft und Kunst 25:198. 1897. Macroscopic characters: Basidiomata up to 1.5-30 × 1.5-22 cm, solid, cantharelloid-gomphoid, unipileate, merismatoid or ramarioid. Pileate species pilei up to 22 cm wide, flat to depressed, fan- to funnel-shaped, glabrous to subtomentose or floccose-squamose (P. retispora), white, pale orange, orange to sordid olivaceous, sordid green to olive-green, yellow-brown, brown with age; margin incurved, crenate, lobed or inrolled when young, undulate to straight with age. Ramarioid species of wide branches below, cylindrical, subcylindrical to flattened or with small grooves or ridges, sometimes palmate, erect, crowded to openly spaced, sometimes spreading, stout to fragile, dichotomous to irregularly branched (up to 6 ranks), velutinous-pruinose (P. cervicornis) to rugulose, in general sterile and subtomentose on the inner surfaces and on the uppersides of the axils, forked or cristate near the apices, pale pink, gray, olivaceous, olive-green, blue-green, creamy yellow, orange, brown, cinnamon to red, in general bruising vinaceous brown to olive-green (P. ochraceo-virens); internodes short to generally elongated, especially at the primary axes13, length diminishing gradually distally, forked or single near the apices, blue-green, creamy white to pale yellow, yellowbrown to dark brown, golden yellow to orange, olive-orange, cinnamon, brick red; axils subacute, acute to rounded, in general U-shaped, branches subparallel to slightly divergent, with narrow, sterile ‘webbing’14 (P. articulotela), gray-yellow, brick red to reddish brown, cinnamon when dry (P. eumorpha); apices waxy, acute, subacute to rounded, dentate, digitate, subcristate, Y-shaped to awl-shaped in some species, composed of two thin lash-like branchlets in P. vinaceipes, white to various shades of gray, olive, green, yellow, brown or red. Hymenium: pileate species – smooth in some species, lamellate to sublamellate or irregularly wrinkled with decurrent hymenial folds in others, generally longitudinally oriented, 13 14 First branches observed from the stipe. Resembling the foot of a waterfowl. 210 dichotomously forked, anastomosing in some species, pale to dark orange, pale to sordid olivaceous, brown, sometimes violet in age (P. grandis) or bruising vinaceous; ramarioid species – amphigenous or unilateral, with regular decurrent folds sometimes extending to the substrate level, smooth to rugulose, with wrinkles forking regularly, waxy where fertile, minutely felty and ferruginous where sterile, but in general concolorous with branches. Stipe (2-) 30-90 (-150) × (0.6-) 6-30 (45) mm, firm, solid to partially hollow, slender or stout, simple to fused, cylindrical, subcylindrical to flattened, ridged or grooved, more or less equal to generally slightly tapered downward, significantly expanded (up to 25 mm tall and 15 mm thick) below the substrate level to a bulbous shape in P. africana, in general coriaceous to brittle, surface glabrous to tomentose, especially at the base, frequently covered with hymenium, in general concolorous with pileus and branches, white below the substrate level, sometimes bruising blue-green, vinaceous, lavender-brown, brick-red or red-brown. Context firm, tender to tough, generally pliable and moist when fresh, gelatinous, punky pliable to fibrous when dry, white, creamy yellow to orange, in some species bruising vinaceous to brighter/darker colors than original color. Odor none to anise-like, earthy, spicy and aromatic, sweet to farinaceous, or paracresol-like. Taste none to bitter, musty, sour, mildly earthy, starchy, occasionally astringent, acrid to almost salty in P. capucina. Spore print cinnamon brown, yellow-brown, ferruginous. Microscopic characters: Pileipellis up to 100 µm thick, of thin- to slightly thickwalled, compact to loosely parallel or compactly interwoven, undifferentiated, agglutinated or smooth, in general cyanophilic hyphae 1.5-16 µm wide, eventually with ampulliform swellings; clamp connections present; gleoplerous hyphae present or absent. Stipitipellis of thick-walled, parallel to compactly interwoven, branched, hyaline, moderately cyanophilic hyphae up to 14 µm wide, with ampulliform swelling in some species; clamp connections present; gleoplerous hyphae common. Context of thin- to slightly thick-walled, parallel to compactly interwoven hyphae 2-13.5 µm wide, inflated up to 15 µm in others, generally smooth, with adherent amorphous crystals (P. eumorpha), H-connections present in 211 P. angustata and P. zippelii; clamp connections present; gloeoplerous hyphae, either present, cyanophilic, not inflated or inflated to 25 µm, yellow. Subhymenial trama of irregular, more or less parallel to interwoven, generally narrow, indistinctly defined hyphae. Rhizomorphic strands, when present, more or less parallel, with hyphae between 1-3.5 µm thick, usually flattened, moderately pliable, easily broken when dry, white, in some species scattered with polymorph crystaloid elements of 1.5-14.5 µm. Basidia (19-) 40-100 (-140) × 4-16 µm, clavate, cyanophilic, with (1-) 2 (-3-) 4 stout, slightly divergent, slightly incurved to straight sterigmata up to 11 µm long, persisting or not after spore discharge; clamp connections generally present at base. Hymenial cystidia present only in P. viridis. Spores (3.5-) 8-17.5 (-25.5) × (2.5-) 3.5-9 (-12) µm, golden yellow, yellow-brown, orange-brown to ferruginous in mass, ellipsoid, ovoid, obovoid, subglobose, lacrimoid (tear-drop-shaped) to fusoid; ornamentation of cyanophilic warts and spines up to 3.5 µm tall, occasionally anastomosing at base of ornamentation to form subreticulate to reticulate ridges; apiculus prominent, eccentric. Habit, habitat and distribution: Epigeous, single to gregarious or sometimes caespitose, on soil or wood on forest floors or in gardens, from tropical to temperate regions. Discussion: Giachini (chapter 2) showed Ramaria subgenus Echinoramaria (monomitic hyphae and mostly echinulate spores) to group in a separate clade from other ramarioid species (non-monomitic hyphae and verrucose to smooth spores). A few gomphoid species were seen nested among the ramarioid species. This culminated with the resurrection of Phaeoclavulina. Forty-one species are known for the genus: 6 pileate and 35 ramarioid. The basidiomata of these species display a great breath of sizes, shapes, and colors. Common characteristics of pileate and ramarioid species of Phaeoclavulina include: (a) echinulate to verrucose (anastomosed into subreticulate to reticulate ridges in some species) basidiospores, (b) presence of clamp connections, and (c) terrestrial and/or lignicolous substrate affinity. No species in this genus has been 212 proved to be mycorrhizal, perhaps due to the lack of investigation, although some have been suggested to be saprotrophic. Both Corner (1970) and Petersen (1981) recognized several sections and series within Ramaria subgenus Echinoramaria. Because not all taxa could be molecularly sampled for this subgenus, Giachini (chapter 2) did not adopt Corner or Petersen’s concepts of sections and series for Phaeoclavulina. Even tough shape of the basidiomata wouldn’t suggest an evolutionary link between ramarioid and pileate lineages, molecular genetics shows them to be evolutionarily more closely related than had been suspected. In light of the recent discoveries, more collecting needs to be done, especially in tropical and subtropical regions, to increase the knowledge for these gomphoid-ramarioid lineages of Gomphales. This will enhance understanding of species distributions, diversity and substrate requirements. Pileate species Phaeoclavulina africana (R.H. Petersen) Giachini, sp. prov. Fig. 6.1 ≡Gomphus africanus R.H. Petersen, Mycologia 68:429. 1976a. Macroscopic characters: Basidiomata up to 4.5 cm tall, cantharelloid-gomphoid, unipileate (Figure 6.1). Pileus up to 2.5 cm wide, funnel-shaped, flat when young, depressed when mature, yellow-brown; margin thin and shiny when young, eventually crenate. Hymenium decurrent, with regular wrinkled hymenial folds extending to the substrate level, forking regularly, brown when dry. Stipe solid, significantly expanded with a subsurface bulb up to 25 mm tall and 15 mm thick, white, covered with a tomentum incorporating much sand (see figure 6.1). Data on context, odor, and taste not recorded. Spore print brown. 213 a. b. b. Figure 6.1. Phaeoclavulina africana (R.H. Petersen) Giachini, sp. prov.: (a) basidiomata (× 0.4); (b) spores (bar = 10 µm). Spore drawings modified from Petersen (1976b), with permission. 214 Microscopic characters: Pileipellis of thin-walled, compactly interwoven, agglutinated hyphae, yellow to a depth of 50 µm, often with adherent spores and minute debris; clamp connections present; gleoplerous hyphae often clavate, not inflated, strongly cyanophilic, yellow. Pileus context of compactly interwoven hyphae 2-4 µm wide, with adherent amorphous crystalline material; clamp connections present; gloeoplerous hyphae not inflated, yellow. Basidia 70-72 × 810 µm, clavate, gleoplerous, with agglutinated material, yellow, with 2 sterigmata that are stout below and abruptly narrow above, hyaline; clamp connections present at base. Hymenial cystidia absent. Spores 12.5-18 × 6.5-8 µm, thin-walled, ferruginous in mass, ellipsoid to obovoid, contents moderately cyanophilic; ornamentation of acute, moderately cyanophilic cones up to 1 µm tall, occasionally anastomosing at the base to form short ridges (Figure 6.1); apiculus rounded. Habit, habitat and distribution: The only known specimen was collected on grassland in Uganda (Petersen 1976a). Collections examined: UGANDA: Bukoto Co., Masal District, Bale, Lake Nabugabb, 17 May 1972, K. Lye (TENN 36291 – as Gomphus africanus – holotype). Discussion: As noted by Petersen (1971, 1976a), the morphological characters of Gomphus africanus (≡P. africana) are strikingly similar to those of some species formerly in Ramaria subgenus Echinoramaria, a condition corroborated by the molecular data (Giachini, chapter 2). Moreover, for those species previously in Ramaria subgenus Echinoramaria bearing heavily ornamented spores, sterigmata number is often 2, and basidia are normally gleoplerous and agglutinated, as observed for P. africana. The most obvious characters that link P. africana to other species of Phaeoclavulina are the size and ornamentation of the spores and presence of clamp connections. Indeed, it would be difficult to distinguish between a microscope mount of hymenium and spores of Ramaria zippelii (≡P. zippelii) and Gomphus africanus (≡P. africana) (Petersen 1976a). 215 Phaeoclavulina grandis (Corner) Giachini, sp. prov. Fig. 6.2 ≡Gomphus grandis Corner, Nova Hedwigia 18:812. 1969. Macroscopic characters: Basidiomata up to 30 × 22 cm, merismatoid (Figure 6.2). Pileus up to 22 cm wide, fan-shaped, glabrous, pale orange and then orange; margin blunt, inrolled when young, undulate with age. Hymenium decurrent, with tall, wrinkled hymenial folds running down the stipe to the substrate level, forked occasionally, less crowded and smooth to sublamellate toward the margin than in the center of the pileus, pale olivaceous to violet in age, all parts bruising vinaceous. Stipe up to 150 × 45 mm, tapering downward, fibrillose to tomentose, especially at the base, pale brown to concolorous with pileus. Context pliable and moist when fresh, gelatinous, off-white to creamy yellow; context of stipe stringy, pale orange. Odor strong, resembling that of Ganoderma (paracresol). Taste not recorded. Spore print color not recorded. Microscopic characters: Pileipellis 50-100 µm thick, of loosely interwoven hyphae 2-4 (-5) µm wide. Stipitipellis a stratum 100-500 µm thick, with hyphae 2-3 µm wide. Context of parallel, inflated hyphae 3-12 µm wide; clamp connections present; gleoplerous hyphae absent. Basidia 95-140 × 11-14 µm, clavate, with (3-) 4 sterigmata 7-8 µm long. Hymenial cystidia absent. Spores (14-) 16-17.5 × 5.5-7 µm, ellipsoid; ornamentation of irregular, cyanophilic warts up to 1.5 µm tall (Figure 6.2); apiculus prominent, straight. Habit, habitat and distribution: This large fungus is known from a single collection by Corner in a dense forest of Agathis and Fagaceae at the mid-slopes of Kinabalu, North Borneo. Collections examined: BORNEO: Kota Kinabalu, 26 April 1964, E.J.H. Corner RSBN 8448 (BR 079158-06 – as Gomphus grandis – holotype). Discussion: Corner (1969) stated that G. grandis is close to G. guadelupensis in possessing clamp connections and lacking gleoplerous hyphae. His pictures show a 216 a. b. b. Figure 6.2. Phaeoclavulina grandis (Corner) Giachini, sp. prov.: (a) basidiomata (× 1); (b) spores (bar = 10 µm). 217 resemblance to Gomphus clavatus, although overall size of the hymenial folds and size and shape of the spores immediately distinguish it from G. clavatus. Phaeoclavulina guadelupensis (Pat.) Giachini, sp. prov. Fig. 6.3 ≡Stereum guadelupense Pat., Bulletin de la Société Mycologique de France 15:201. 1899. ≡Gomphus guadelupensis (Pat.) D.A. Reid, Persoonia 2:134. 1962. =Nevrophyllum ochraceum Pat., in Duss, Flore Cryptogamique de Antilles Françaises, p. 38. 1903. ≡Cantharellus ochraceus (Pat.) Sacc. & D. Sacc., Sylloge Fungorum Omnium Hucusque Cognitorum 17:36. 1905. ≡Gomphus ochraceus (Pat.) Singer, Lloydia 8:140. 1945. Macroscopic characters: Basidiomata 10-15 cm tall, unipileate at first and then merismatoid. Pileus up to 10 cm wide, fan-shaped, lobed, flattened to subundulate, sometimes depressed at the center, with a tomentum of hyphae over the disc, velvety, orange when fresh, becoming brown with age, crenate at margin. Hymenium decurrent nearly to the base of the stipe, smooth to wrinkled, with low, shallow, crowded, simple or branched hymenial folds, commonly anastomosing downward to form irregular diamond patterns, concolorous with pileus or slightly darker. Stipe 30-70 × 30 mm, firm, glabrous, concolorous with pileus and hymenium. Context orange. Odor and taste not recorded. Spore print cinnamon brown. Microscopic characters: Pileipellis of thin- to slightly thick-walled, parallel to interwoven, undifferentiated hyphae 2-10 µm wide, inflated away from the surface of the pileus, stipe, and hymenium into the context; clamp connections present, large; gleoplerous hyphae absent. Stipitipellis of undifferentiated hyphae covered with abundant spores. Stipe context of generally parallel, agglutinated, slightly cyanophilic hyphae; gleoplerous hyphae rare, aseptate, cyanophilic, in some places inflated up to 22 µm. Basidia 65-110 × 14-16 µm, clavate, strongly cyanophilic, 218 Figure 6.3. Spores (bar = 10 µm) of Phaeoclavulina guadelupensis (Pat.) Giachini, sp. prov. Spore drawings modified from Petersen (1971b), with permission. 219 with 2-3-4 stout, slightly divergent, slightly incurved sterigmata up to 8 µm long; clamp connections present at base. Hymenial cystidia absent. Spores 15-18 (-25.5) × 7.5-10.5 (-12) µm, cinnamon in mass, ellipsoid to lacrimoid or fusoid; ornamentation of coarse, slightly cyanophilic warts up to 2.5 µm tall, partially reticulate (Figure 6.3); apiculus prominent, conical, eccentric. Habit, habitat and distribution: Solitary to gregarious or subcaespitose. Two specimens are known, both on wood from Guadeloupe in the Lesser Antilles. This species likely occurs on other Caribbean islands and on the nearby mainland of South and Central America (Petersen 1971). Collections examined: GUADELOUPE: Camp Jacob, February 1898, A. Duss 120 (FH – as Stereum guadelupense – holotype). Discussion: This remarkable species needs to be redescribed from living material (Corner 1966). It is evidently closely related to P. grandis, P. subclaviformis, and P. viridis on the basis of spore morphology, biogeography, and molecular data (Giachini, chapter 2). Its large basidioma size and coarse spore ornamentation resemble P. viridis (Petersen 1968, 1971). Petersen (1968) has also drawn analogies between Gomphus guadelupensis (≡P. guadelupensis) and the spinyspored species of Ramaria, especially between P. guadelupensis and P. zippellii. Phaeoclavulina retispora (Corner) Giachini, sp. prov. Fig. 6.4 ≡Gomphus retisporus Corner, Annals of Botany Memoir 2:124. 1966. Macroscopic characters: Basidiomata merismatoid (Figure 6.4). Pileus 7-11 cm wide, funnel-shaped, floccose-squamose to nearly glabrous, pale sordid olivaceous, pale yellow-brown toward the margin, in some specimens yellow toward the margin; margin incurved at first, then lobate-undulate, minutely subvillose. Hymenium decurrent, with distinct gill-folds up to 1 cm tall, dichotomous near the margin, transversely wrinkled in the interstices, sordid olivaceous, pale orange toward margin. Stipe 40-90 × 8-18 mm, concolorous, bruising vinaceous. Context 220 soft, paler concolorous with the surface, with watery zones in the pileus and watery-marbled zones in the stipe, unchanging color on exposure. Odor none. Taste not recorded. Spore print color not recorded. Microscopic characters: Pileipellis of distinctly thick-walled, compact, heavily agglutinated hyphae. Stipitipellis a tomentum 500 µm thick, composed of more or less entangled hyphae 4-7 µm wide. Other hyphae thin-walled, 17-25 µm wide, inflated; clamp connections present. Basidia 60-125 × (7-) 10-13 µm, clavate, with (2-) 3-4 straight sterigmata 8-11 µm long. Hymenial cystidia absent. Spores (11.5-) 14-17 (-20) × (5.5-) 6.5-7.5 µm, cinnamon in mass, ellipsoid to ovoid; ornamentation of irregularly oriented, strongly cyanophilic, subreticulate to reticulate ridges varying in height along their length up to 3 µm tall (Figure 6.4); notes on apiculus not recorded. Habit, habitat and distribution: Fruiting gregariously on soil and humus in Malaysia and the Solomon Islands (Corner 1966, 1969). Type (holotype): MALAYSIA. BATU BALAI: Pahang, Flum. Cheka, 13 November 1930, E.J.H. Corner (CGE – as Gomphus retisporus). Discussion: The present description was based on notes from Corner (1966, 1969) and Petersen (1971). Corner (1966) described the species from a collection obtained in Pahang, Malaysia. Later Corner (1969) mentioned two more collections from the Solomon Islands. Petersen (1971) examined the isotype deposited at the National Herbarium Nederland (L) and concluded that it has “subreticulate rather than reticulate spores, and that the ornamentation anastomosed, often giving the impression of a reticulum.” The species has affinities with Phaeoclavulina viridis, although closer in color and agglutination of hyphae to P. guadelupensis. No collection was available for analysis in this study. Loan of the type was denied due to the specimen’s fragile condition, and we were unable to obtain loan of the isotype deposited at L. 221 a. b. Figure 6.4. Phaeoclavulina retispora (Corner) Giachini, sp. prov.: (a) basidioma (× 1); (b) spores (bar = 10 µm). Basidioma photo from Corner (1969). 222 Phaeoclavulina subclaviformis (Berk.) Giachini, sp. prov. Fig. 6.5 ≡Thelephora subclavaeformis15 Berk., Bulletin of Miscellaneous Information 8:238. 1856. ≡Gomphus subclavaeformis (Berk.) Corner, Nova Hedwigia 18:815. 1969. =Gomphus cavipes Corner, Nova Hedwigia 18:808. 1969. Macroscopic characters: Basidiomata up to 9 cm tall, unipileate to merismatoid (Figure 6.5). Pileus around 5 cm wide, fan-shaped, often lobed and becoming concrescent around the stipe, finally appearing slightly funnel-shaped, glabrous, pale orange to white; margin straight. Hymenium initially smooth, then more or less radially and irregularly wrinkled with shallow hymenial folds of about 0.3 mm tall, pale orange when immature, dark orange with age. Stipe 40 × 2-9 mm, tapering downward, glabrous, concolorous. Context moist-fibrous, bruising vinaceous. Odor slightly sour. Taste not recorded. Spore print color not recorded. Microscopic characters: Pileipellis of compact hyphae up to 10 µm wide; clamp connections present. Stipitipellis of thick-walled, parallel, branched, slightly inflated hyphae 2-7 µm wide; clamp connections present; gleoplerous hyphae thinwalled, aseptate, up to 6 µm wide (Figure 6.5). Generative hyphae often coiled, rarely branched. Basidia 52-85 × 8-10 µm, clavate, with 4 straight sterigmata up to 8 µm long; clamp connections present at base (Figure 6.5). Hymenial cystidia absent. Spores 10.5-12.5 × 5.5-7 µm, cinnamon in mass, ellipsoid to fusoid; ornamentation of round to irregular, non-anastomosed, cyanophilic warts up to 1 µm tall (Figure 6.5); apiculus up to 1.5 µm tall, eccentric. Habit, habitat and distribution: Solitary or gregarious, not caespitose, on the ground in closed forest in Amazonas (Panuré) and Mato Grosso States, Brazil, and Arima, in Trinidad and Tobago (Corner 1969). Collections examined: BRAZIL. MATO GROSSO: Chavantina, 1 February 1966, E.J.H. Corner K-969 (BR 079159-07 – as Gomphus subclavaeformis – holotype). 15 The ‘ae’ connective is an orthographic error (International Code of Botanical Nomenclature, article 60G.1) to be replaced with an ‘i’ (Greuter et al. 2000). 223 a. b. c. b. d. Figure 6.5. Phaeoclavulina subclaviformis (Berk.) Giachini, sp. prov.: (a, b) basidiomata (× 1); (c) basidium and gleoplerous hyphae (× 1000); (d) spores (× 1000). Figures b, c, and d modified from Corner (1969). 224 Figure 6.5 (cont.). Spores (bar = 10 µm) of Phaeoclavulina subclaviformis (Berk.) Giachini, sp. prov. 225 Discussion: Berkeley (1856) described Thelephora subclavaeformis from Panuré, from the northern regions of Brazil. Later Corner (1969) elaborated on the correct identity of this species and transferred it to Gomphus. Even though only one collection was reported, he regarded this as a common species in the forests North of Chavantina, Mato Grosso, in Central-West Brazil. It also occurs in Arima, in Trinidad and Tobago. The basidiomata resemble G. clavatus, although the similarities end there. Giachini (chapter 2) studied Corner’s (1969) description of Gomphus cavipes from Trinidad and Tobago (the loan of the type from CGE was denied because of its fragile condition) and synonymized it with P. subclaviformis because of the overall morphology and spore shape and size. Fig. 6.6 Phaeoclavulina viridis (Pat.) Giachini, sp. prov. ≡Nevrophyllum viride Pat., Journal de Botanique 2:406. 1888. ≡Chlorophyllum viride (Pat.) Murrill, nom. illeg., North American Flora 9:172. 1910. ≡Gomphus viridis (Pat.) Singer, Lloydia 8:140. 1945. =Cantharellus patouillardii Sacc., Sylloge Fungorum Omnium Hucusque Cognitorum 9:65. 1891. Macroscopic characters: Basidiomata up to 12 cm tall, unipileate (Figure 6.6). Pileus flat to depressed, funnel-shaped, fleshy-coriaceous, minutely velvety or felty, dark sordid green, olive-green or gray-olive, becoming dark yellow-brown at the center of the pileus with age; margin deeply lobed to crenate. Hymenium with wrinkled, decurrent hymenial folds, generally longitudinally oriented, dichotomously forked toward the pileus margin, suppressed below on stipe, olivaceous. Stipe 6-9 mm thick, glabrous, tapering downward, concolorous with pileus, paler below substrate level. Data on context not recorded. Odor none. Taste sour. Spore print color not recorded. 226 a. b. Figure 6.6. Phaeoclavulina viridis (Pat.) Giachini: (a) basidiomata (× 0.8); (b) spores (bar = 10 µm). Figure a modified fromNeale Bougher. Drawings modified from Petersen (1971b), with permission. 227 Microscopic characters: Pileipellis of thin-walled, compactly interwoven, undifferentiated hyphae 4.5-6.5 µm wide; clamp connections present. Stipitipellis of hyphae identical to those of pileipellis, up to 14 µm wide, with ampulliform thinwalled swellings. Stipe context of thin- to slightly thick-walled, interwoven, closely packed, moderately cyanophilic hyphae, yellow-green in mass, 8.5-12.5 µm wide, with thin-walled ampulliform swellings up to 15 µm; clamp connections present; gleoplerous hyphae hyaline, strongly cyanophilic. Basidia 90-110 × 6-8 µm, clavate, with 2-4 stout, divergent, straight sterigmata; clamp connections present at base. Hymenial cystidia 90-140 × 8-10 µm long, thin-walled, cylindrical, hyaline, acyanophilic; clamp connections present at the base. Spores 13-16 (-21.5) × 6.5-8 (-12) µm, cinnamon in mass, ellipsoid to lacrimoid; ornamentation of irregularly oriented, strongly cyanophilic, subreticulate to reticulate ridges varying in height along their length from <0.5 to 3.5 µm tall (Figure 6.6); apiculus prominent, eccentric. Habit, habitat and distribution: Single or gregarious, occasionally caespitose, on soil or rotting wood under Agathis robusta, Allocasuarina littoralis, Angophora sp., and Eucalyptus grandis in Australia, and Coccoloba belizensis and Pinus sp. in Belize. Also known from French Guiana and Martinique. Collections examined. AUSTRALIA. QUEENSLAND: Kennedy Road to Kirrama, 28 April 1990, N.L. Bougher and M.A. Castellano E4302 and E4303 (PERTH). BELIZE: Mountain Pine Ridge, Lower Nature Trail, Five Sisters Lodge, 20 November 2001, K.K. Nakasare and D.J. Lodge (OSC 97708), and 23 November 2001, D.J. Lodge and B. Ortiz-Santana (OSC 97707). FRENCH GUIANA: no location, no date, N.T. Patouillard (FH – as Nevrophyllum viride – holotype). MARTINIQUE: Marne Larcher, Pres du Camp Balata’, August 1900, A. Duss 1853 (FH). Discussion: The vegetation at the type locality was not described. Corner (1966) reported the species fruiting on the ground in swampy woods in French Guiana (one collection – type) and Martinique (one collection). In 2001, Dr. Jean Lodge collected it twice in Belize under Coccoloba and Pinus sp. 228 This species has an unusual distribution in Australia, Belize, and Central and South America and has been never reported elsewhere. It occurs in mixed forests containing Araucariaceae (Agathis), Casuarinaceae (Allocasuarina), and Myrtaceae (Angophora and Eucalyptus) in Australia, and under species of the Pinaceae (Pinus) and Polygonaceae in Belize. As noted by Corner (1966), the species has affinities with P. retispora and is brownish olive rather than truly green as described by Patouillard (1888). Ramarioid species Phaeoclavulina angustata (Lév.) Giachini, sp. prov. Fig. 6.7 ≡Merisma angustatum Lév., Annales des Sciences Naturelles Botanique 3(5):158. 1846. ≡Thelephora angustata Fr., Nova Acta Regiae Societatis Scientiarum Upsaliensis 3:116. 1854. ≡Ramaria angustata (Fr.) Corner, Beihefte zur Nova Hedwigia 33:242. 1970. Macroscopic characters: Basidiomata 6-30 × 5-15 cm, ramarioid, repeatedly branched (Figure 6.7). Branches 5-15 mm wide below, cylindrical, erect, crowded, sometimes spreading, generally regularly dichotomous with successive branching more or less at right angles, the branching sometimes more irregular, some times 34-chotomous below, sterile and subtomentose on the adaxial surfaces and on the uppersides of the axils, pale pink, then more or less dark yellow, orange to brown, often tinged cinnamon to orange from the spores, all parts soon turning sordid vinaceous on bruising; internodes gradually more elongated distally, short and obscure proximally, 1.5-2.5 mm wide at the tips; tips subacute, intensely blue or blue-green, or in some specimens white or creamy white. Hymenium unilateral. 229 a. b. b. Figure 6.7. Phaeoclavulina angustata (R.H. Petersen) Giachini, sp. prov.: (a) basidiomata (× 0.4); (b) spores (bar = 10 µm). Basidioma drawings modified from Corner (1950). 230 Stipe 6-65 × 3-16 mm, cylindrical, relatively slender, slightly tapering downward, generally attenuate at the subvillose base, with white mycelium, concolorous elsewhere. Context rather dry but fibrous-brittle, breaking when bent, white or pale yellow, dark yellow near the hymenium, soon turning sordid vinaceous on exposure. Odor slight to none. Taste slightly bitter and astringent. Spore print brown. Microscopic characters: All hyphae parallel, compact, smooth; H-connections frequent, with pale yellow walls, dark yellow-brown and with blue tones at the tips, often rather unevenly inflated, 2.5-10 µm wide; clamp connections present. Basidia 55-75 × 9-11 µm, subclavate, generally constricted below the sterigmata, with a long narrow base 3 µm wide, with (1-) 2 (-3) sterigmata 6-8 (-10) µm long. Hymenial cystidia absent. Spores 18-20 (-24) × 9-11 µm, orange to golden yellow in mass, lacrimoid, rather densely echinulate with conical, acute, hyaline, cyanophilic spines 1-3.5 µm tall (Figure 6.7); data on apiculus not recorded. Habit, habitat and distribution: Gregarious or rarely caespitose, on the ground in forests of tropical America (Petersen 1981), Malaysia (Corner 1950), and perhaps tropical Africa. Collections examined: COSTA RICA: Location unknown, no date, A.D. Oersted (BPI – as Thelephora angustata – holotype). Discussion: Corner (1950) considered this the most common species of Ramaria subgenus Echinoramaria in Malaysia. Its large size plus distinctive dark yellowbrown branches with blue, or blue-green tips makes it one of the more striking ramarioid species in the genus. 231 Phaeoclavulina apiahyna (Speg.) Giachini, sp. prov. Fig. 6.8 ≡Clavaria apiahyna Speg., Boletin de la Academia Nacional de Ciencias de Córdoba 23:435. 1918. ≡Ramaria apiahyana (Speg.) Corner, Annals of Botany Memoir 1:555. 1950. Macroscopic characters: Basidiomata 5-8 cm tall, ramarioid. Branches subcylindrical, glabrous, white below, orange above; apices slightly thickened and shortly dentate or subcristate, intensely brick red to nearly scarlet. Data on hymenium configuration and features of the stipe not recorded. Context white. Odor and taste not recorded. Spore print ferruginous. Microscopic characters: Context hyphae with agglutinating material, yellowish; clamp connections present. Basidia with 2 sterigmata. Hymenial cystidia absent. Spores 15-18 × 8-10 µm, orange in mass, ellipsoid obovoid; ornamentation of densely and minutely dispersed, cyanophilic spines 1-3 µm tall, 1-2.5 µm wide at the base, often flattened and generally in longitudinal rows (Figure 6.8); apiculus blunt. Habit, habitat and distribution: On the ground in forests in Brazil (Petersen 1981). Collections examined: BRAZIL. SÃO PAULO: Apiaí, January 1891, Puiggari 307 (LPS 13259 – as Clavaria apiahyna – holotype). Discussion: The species is near P. macrospora but seems more brilliantly colored (Corner 1970). It is also similar to P. camellia, which is said to have similarly colored basidiomata, similar distribution, but much smaller spores.