Mycorrhizal fungi of exotic forest plantations

Peitsa Mikola

Department of Silviculture University of Helsinki

Introduction South Africa 0.92 mill. ha, and Chile 0.35 mill. ha. In all these countries the main It is a well-known fact that Suillus grevillei species of plantations are pines (mainly Pinus (Boletus elegans) is to be found growing radiata of California, P. elliottii of Florida, under larch (Larix spp.) and nowhere else. and P. patula of Mexico), although the In late summer, sporophores of Suillus grevil­ indigenous floras of these areas include no lei are almost invariably found in all larch species belonging to the Pinaceae family. forests and plantations, and even under soli­ Australian eucalypts (Eucalyptus spp.) which tary trees. The great mycologist Elias Fries have ectotrophic mycorrhizae are also exten­ already wrote: " Ubi Larix, ibi Boletus sively grown outside their natural range, in elegans". Africa, Asia, and South America. Larch is an exotic tree species in Finland. The aim of this article is to review the Consequently, Suillus grevillei cannot belong fungal flora of exotic forest plantations and to the native Finnish flora but must have to discuss the possible modes of immigration 1 arrived there with or after its hosts, Larix of exotic mycorrhizal fungi. ) spp. The same applies to some other mycor­ rhizal fungi of larch, such as Boletinus asiati­ Mycorrhizal fungi of exotic plantations cus, B. cavipes, and Tricholoma psammopus, Local lists of fungi fruiting in exotic coni­ as well as to all the other areas where Larix ferous plantations have been published by spp. are grown as exotics. Likewise, mycor­ Birch (1937) and Rawlings (1950) for New rhizal fungi of other forest trees have followed Zealand, Purnell ( 195 7) for Victoria ( Aust­ along with their host species when these have ralia), Spegazzini (see references in Singer been cultivated outside their natural range. 1950) for South America, and others. Some Exotic forest trees are today grown exten­ data from these lists, supplemented with sively in many parts of the world, particularly personal communications and my own field in the tropics and subtropics. Systematic observations, are summarized in Table 1. In trials with exotic species have been conducted addition to the fungi listed, a great number since the last century (see Streets 1962) , and of species of Cortinarius, lnocybe, Lactarius, large-scale afforestations were started in some Russula, Tricho.Zoma, and other genera have countries, such as New Zealand and Chile, been described from exotic plantations (Sin­ as long ago as 1920's. According to the FAO ger 1950, and others). The fungal flora of Secretariat (1967 ), there are today some 80 million hectares of forest plantations in the 1) The review is based partly on ~he literature, world, a great many consisting of exotic partly on personal observations .and discussioil!S species. New Zealand, for instance, has 0.46 during a s·tudy tour round tihe world in 1967. mill. ha of plantations, Australia 0.3 mill. ha, The tour was sponsored by FAO.·

169 such plantations may also include indigenous Suillus variegatus is apparently quite a dif­ species. Anderson ( 1966), for instance, found ferent type and is not adaptable to warm numerous European mycorrhizal fungi in climates. Although S. luteus and S. varie­ Eucalyptus plantations in Italy. The majority gatus are equally common in the boreal pine of fungi, however, are usually exotic and, forests, the latter species can hardly be found thus, the fungal population of exotic planta­ anywhere outside its natural range.1 tions differs greatly from that of the indige­ Amanita is another mycorrhizal genus, nous forests in the same area. several species of which have spread with The most conspicuous and widely distrib­ exotic plantations. The most conspicuous uted mycorrhizal fungi of exotic coniferous species, A . muscaria, which forms mycorrhizae plantations are several species of Suillus, S. with both conifers and broad-leaved trees is grevillei being a typical example. Singer today common in New Zealand, Austraha, (1964) lists five species of Suillus for South South Africa, South America and elsewhere. America, and McNabb (1968 ) se en species It seems to avoid the most tropical elimates, for New Zealand, all of which have been however, and is unknown in East Africa introduced with exotic conifers. No species whereas it already grows in Rhodesia (I. A: of Suillus belong to the native flora of the S. Gibson, personal communication). Accord­ southern hemisphere. ing to field observations, it can probably form mycorrhizae with some indigenous trees too Suillus granulatus has been desribed at least with Nothofagus spp. in South almost wherever any species of Pinus has Arne~ rica and New Zealand, just as some indi­ been planted. It seems to be quite an adapt­ genous fungi may be able to form mycorrhi­ able species, growing in both cool boreal zae with introduced trees. forests and subtropical plantations. In regard Several Gasteromycetes (spp. of Rhizopo­ to soil pH, too, it has a wide range; it is gon, Scleroderma, and Pisolithus ) are also indigenous, for instance, in the Pinus hale­ common mycorrhizal associates of exotic pen.~is forests on limestone rocks of the l.Vfediterranean area and grows as an exotic forest plantations. The frequent occurrence under the same species on alkaline soils in of sprorophores of Rhizopogon in pine the La Pampa Province of Argentina. This nurseries was observed quite early (Kessell is somewhat surprising, because, according 1927 ), and in plantations they are often to pure culture experiments (Melin 1925, found as soon as one or two years after plant­ Modess 1941 ), the optimum pH of Suillus ing. Some species of Scleroderma are known granulatus is between 5 and 6 and it does or suspected to be mycorrhizal with eucalypts, not grow at all at a pH above 7.5. Probably sporophores being common under exotic Suillus granulatus is a collective species, eucalypt plantations (Pryor 1956; Levisohn including several subspecies or even separate 1958; Bakshi 1966 ) ; nevertheless, the fungi species with different ecological requirements. may also be indigenous in the area. Pisolithus Reichert ( 1940), for instance, has described tinctorius has been described as an indige­ four closely related species in Israel, none of nous mycorrhizal fungus of eucalypts in which is probably the true Suillus granulatus. Australia (Smith & Pope 1934) and grow­ One of these basophilous Mediterranean ing as an exotic under eucalypt plantations species, S. bellini, grows as an exotic in Cape in both Israel (Reichert & Avizohar-Her­ Province of South Africa (Stephens & Kidd shenzon 1959 ) and South Africa (Stephens 1953a). & Kidd 1953 b ), whereas, according to other Suillus luteus is another widely distributed sources, it is also indigenous in North Ame­ mycorrhizal associate of exotic pine plan­ rica (Schramm 1966), where it forms tations. Although it is the dominant species mycorrhizae with pines and has been intro­ under pine plantations in the East African duced into pine plantations of South Ame­ Highlands (Gibson 1963), for instance, it rica (van Suchtelen 1962 ). seems to avoid very warm climates. Thus, As a whole, exotic tree plantations have according to Singer ( 1963), S. luteus is the greatly enriched the fungal flora of many most common species under pines in south­ 1) According to a recent personal communica­ ern Chile and Argentina, whereas in the tion from Dr. E. Horak, Suillus variegatus is found warmer area of North Argentina it is under pine plantations in the South Island of replaced by S. granulatus. New Zealand.

170 tropical and subtropical countries. Thus, in South Africa, Australia, New Zealand, and South America there are no indigenous South America as long as two or three hun­ species of Amanita, Cantharellus, Hygro­ dred years ago. When, for instance, Dutch phorus or Gomphidius, there is one indige­ settlers planted European oaks around their nous species of Lactarius, and Tricholoma farms in South Africa, they could later ob­ only occurs in the extreme south (Singer serve the deadly poisonous death-cap ( Ama­ 1950). Today all these genera are represent­ nita phalloides) fruiting under these trees. ed by numerous species growing in asso­ Mycorrhizal fungi may also have arrived ciation with introduced trees, Gomphidius, through botanical gardens. The first spe­ for instance, by three North American cimens of many European and North Ame­ species (Singer 1964). rican trees were imported to botanical gar­ Not all the exotic fungi of forest plan­ dens as potted plants. Pinus radiata, for tations are necessarily mycorrhizal. Sapro­ instance, was first introduced into Australia phytic fungi, which in nature grow on such in 1857, when potted seedlings were brought substrates as pine needle litter, may have from Kew Gardens to the botanical gardens found a favourable environment under exot­ of Sydney and Melbourne (Fielding 195 7). ic pine plantations, too, and become natural­ Kew Gardens, in particular, has provided ized. botanical gardens throughout the British Introduced mycorrhizal fungi may some­ Commonwealth with exotic plants. times be of a considerable economic impor­ Likewise commercial nurseries, to which tance, improving the economy of exotic some living plants were also imported, were plantations. Many of the most common intro­ founded quite early, - in South Africa, for duced species, such as Boletus edulis, Suillus instance, in the seventeenth century (Donald spp. and Lactarius delicimus, are edible, and 1965). In those days there were no plant where mushrooms grow in masses in planta­ quarantine regulations restricting the import tions, they constitute important raw material of living plants. for canning industry. Their vah.1e is increased Because most of the immigrants, as well as by the fact that evidently the parasites of the the tree seedlings for botanical gardens, came mushrooms have not always been introduced from Europe, the introduced mycorrhizal with the hosts. Thus, for instance, in Austra­ fungi must be mainly of European origin, lia and South America the sporophores of alhough the tree species of coniferous plan­ Lactarius deliciosus are always clean, whereas tations are usually North American (Pinus in Europe they are almost invariably infest­ radiata, P. elliottii, P. patula, Pseudotsuga ed with insect larvae. Mushrooms, in partic­ menzzem, etc.). Several originally North ular species of Boletus and Suillus, are also American fungi, however, now also occur in a favourite food of many game animals. the .southern hemisphere, examples being Unfortunately, some extremely poisonous Suillus lakei and S. brevipes in New Zealand fungi have also followed their mycorrhizal (McNabb 1968 ) and S. brevipes in Argen­ hosts to new areas. Thus, Amanita phalloi­ tina (Singer 1963). des is a common associate of oaks in both Mycorrhizal infection may also have Argentina (Singer 1950) and South Africa arrived a.s spores attached to imported seed. (Stephens & Kidd 1953b). Contamination of seed with fungal spores may explain, for instance, the occurrence of Modes of migration of exotic fungi Suillus grevillei under larches in some remote localities where trees have been raised from In most countries where pines or other seed and no introduction af living seedlings ectotrophic trees are not indigenous, mycor­ is known (Stahl 1968). Spores o( mycorrhi­ rhizal fungi probably first arrived in the roots zal fungi may also spread with air currents. of living tree seedlings. According to old So far, very little is known about the viabili­ records, early settlers often brought trees ty and germination of the spores of mycorrhi­ from their home countries and planted them zal fungi. Although spores of many mycorrhi­ around their new homes (see Stephens & zal species are very difficult or even impos­ Kidd 1953a; Pryor 1958). Thus, large num­ sibe to germinate on synthetic media in the bers of mycorrhizal seedlings of pine, oak, laboratory, numerous experiences indicate and other European trees were planted in the great ease with which mycorrhizal in-

171 fection spreads (Robertson 1954) . In fact, of these experiments appear to have met with special precautions are usually necessary to success, the practicability of these methods protect experimentally grown non-mycorrhi­ on a field scale is still questionable. zal seedlings from air-borne infection. Into many countries, however, mycorrhi­ Some conclusions zal fungi have been imported intentionally. Numerous examples are known in which The history of the arrival of new fungal attempts to introduce exotic pines con­ species in various countries is not well docu­ sistently failed, until in one way or another mented. Potted tree seedlings were often appropriate mycorrhizal fungi were brought transported from one country to another in (see Mikola 1969). Usually, such impor­ hundreds of years ago and consequently the tation has taken place by bringing soil from most common mycorrhizal fungi were in­ a natural stand or established plantation of troduced to many countries long before any­ the respective tree species. The first intention­ thing was known about mycorrhizal symbios­ al importation of mycorrhizal soil and arti­ is. It may even be difficult to decide whether ficial inoculation of tree seedlings probably a species is native or introduced. Some orig­ took place in 1910, when, at the suggestion inally introduced fungi, saprophytic species of Kew Gardens, soil was brought from a and facultative mycorrhiza-formers in partic­ pine plantation in South Africa to Kenya ular, today grow outside exotic plantations as and, by application of this inoculum, the well. Paxillus involutus, for instance, is quite first successful pine plantation of East Af­ common in the natural forests of South Ame­ rica was established (Gibson 1963 ) . Later on, rica but may still belong to the introduced mycorrhizal soil has been imported, for flora (Singer 1964) . instance, to :Malawi (Clements 1941 ), Nige­ Even the present fungal flora of exotic ria (Madu 1967 ), Trinidad (Lamb 1956 ), forest plantations is very poorly known in Surinam (van Suchtelen 1962), and Puerto many countries. Afforestation activity is Rico (Briscoe 1959) . Since nowadays plant most intensive in developing countries, where quarantine regulations often prohibit the there is a great shortage of trained myco­ importation of living plants or unsterilized logists. Local lists and scattered observations soil, there has sometimes been great difficulty on mushrooms of plantations are usually in obtaining mycorrhizal inoculum. In fact, made by practising foresters, whereas the mycorrhizal soil has been smuggled to some few mycologists, if there are any, are usually countries illegally and so, of course, its im­ fully occupied with research on forest disea­ portation is not well documented. ses. Soil inoculum contains an indiscriminate In countries where planting of exotic trees mixture of all the fungi of the stand from dates back for a century or more, it may be which it is taken, both mycorrhizal and impossible to trace the early migrational saprophytic and even parasitic species. Thus, history of mycorrhizal fungi, and only the exotic saprophytes of forest plantations may present situation can be recorded. On the have arrived with soil inoculum too. Because other hand, in countries where afforestation of the risk of introducing pathogens, the use with exotics is recent or just beginning, it is of soil inoculum has been strongly criticized. eminently desirable that records of the Recently, several attempts have been made migration of exotic fungi should be kept up to introduce known species of mycorrhizal to date. Such a study is particularly recom­ fungi in the form of spore powder, dried mended for botany departments of univer­ sporophores, or pure cultures. Although some sities in these countries.

172 Table 1. Mycorrhizal fungi of exotic tree plantations (p. c. personal communication; P. M. p. o. = personal observation by the author) .

Fungus Country and reference Host species

Amanita muscaria Australia: Purnell ( 195 7) ; Pinus spp. W. Stahl, J. F. Titze (p. c. ) New Zealand: Birch (1937) Pinus, Larix, Pseudo­ Rawlings ( 1950, 1958) tsuga, Betula South Africa: Stephens & Kidd ( 1953b) Introduced trees P.M. p. o. Pinus spp. Rhodesia: Gibson (p. c.) « Argentina: E. A. Takacs (p. c. ) Cedrus deodara

Amanita phalloides South Africa: Stephens & Quercus spp. Kidd ( 1953b) Argentina : Singer (1950), « Takacs (1961)

Boletus fdulis South Africa: Stephens & Kidd Pinus spp. 1953a) P.M. p. o. « Uruguay: Singer (1964) «

Suillus bovinus Australia: W. Stahl (p. c.), P.M. p. o. Pinus spp. South Africa: Stephens & Kidd ( 1953a) « Argentina: E. A. Takacs (p. c.) Pinus elliottii

Suillus bre vipes New Zealand: McNabb (1968) Pinus spp. Argentina: Singer ( 1963 ) «

Suillus gr.anulatus Australia: Purnell (1957), Pinus spp. W. Stahl, J. F. Titze (p. c.), P. M. p. o. New Zealand: McNabb (1968 ) Pinus spp. South Africa: P. M. p. o. « Argentina: R. Piterbarg (p. c. ) Pinus halepensis Singer (1964), Takacs (1967) Pinus spp. Brazil: L. Cardoso (p. c. ), « P.M. p. o. Chile: Singer (1963), P.M. p. o. Pinus radiata

Suillus grevillei Australia : Stahl ( 1968) , Larix leptolepis P.M. p. o. New Zealand: Rawlings ( 1950, Larix spp. 1958 ), McNabb (1968) Europe: Kreisel (1963), « and others South America: R. Singer, « E. A. Takacs, H. Butin (p. c.)

173 Suillus lakei New Zealand: Rawlings (1958 ) Pseudotsuga menzt- (S. amabilis) McNabb ( 1968) estt Central Europe: Moser ( 1967 ) «

Suillus luteus Australia: Purnell ( 195 7), Pinus spp. W. Stahl (p. c.), P. M. p. o., New Zealand: Birch ( 193 7), Pinus spp. Rawlings (1950, 1958), Mc­ Nabb (1968) South Africa: P. M. p. o. Pinus spp. Kenya: Gibson ( 1963 ) « Uganda: Brown (1963 ) « Chile : P. M . p. o. « Argentina: Singer (1963, 1964) « Uruguay: Takacs (1967) Pinus elliottii

Suillus piperatus New Zealand: Rawlings (1950, Pinus spp. 1958 ), McNabb (1968 ) Pseudotsuga South Africa: P. M. p. o. Pinus spp. Trinidad: Singer ( 1964) «

Lcccinum durius­ South Africa: Stephens & Kidd Populus spp. culum ( 1953a)

Leccinum scabrum New Zealand: Rawlings ( 1950, Betula spp. 1958 ), McNabb (1968)

Hebeloma crustuli­ South Africa: Stephens & Kidd Pinus spp. niforme (1953b), P.M. p. o. Kenya : Gibson (1963) « Argentina : Takacs ( 196 7) «

Lactarius deliciosus Australia: Purnell (1957) Pinus s-prp. W. Stahl, J. F. Titze (p. c.) P. M. p. o. Pinus radiata South Africa : Stephens & Kidd Pinus spp. ( 1953a) Chile : P. M . p. o. Pinus radiata

Pisolithus tincto­ South Africa: Stephens & Kidd Eucalyptus spp. nus (1953b), Zambia: N. E. Cooling (p. c.) « Israel : Reichert & Avizohar­ « Hershenzon ( 1959 ) Puerto Rico: P. M.p.o. Pinus caribaea Surinam: v. Suchtelen (1962 ) «

Rhizopogon luteolus Australia: Purnell (1957 ) Pinus spp. W. Stahl, J. F. Titze (p. c. ) « New Zealand: Rawlings (1950) Pinus radiata

174 Rhizopogon roseolus Australia: W. Stahl, J. F. Titze Pinus spp. (p. c.) New Zealand: Birch (1937) Pinus radiata Rawlings (1950) Pinus spp. South Africa: P. M. p. o. Pinus spp. Nigeria: P. M. p. o. « Trinidad: P. M. p. o. «

Scleroderma spp. New Zealand: Birch (1937) Pinus spp., Pseudotsuga ( S. bovista, S. vulga­ Rawlings (1950) Pinus spp., Eucal,yptus re, S. verrucosum, spp. S. flavidum) Kenya: Gibson (1963) Pinus radiata Argentina, Uruguay: Takacs Pinus spp. ( 1967) Brazil: L. Cardoso (p. c.) Pinus spp., Eucalyptus spp.

REFER!ENOE·S

ANDERSON, J, 1966. lndagini sulla micorrizia in LEVISOHN, I. 1958. Mycorrhizal infection in Eu­ alcune specie di eucalitto nell'I talia Centra­ calyptus. - Emp. For. Rev. 37: 237-241. le. - Pubbl. Centro &per. Agr. For. VIII : McNABB, R. F . R : 1968. The Boletaceae of New 2519-274. Zealand. - N. Z. Jour. Bot. 6: 137- 176. BAKSHI, B. K. 19·66. Mycorrhiza in Eucalyptus in MADu, M . 1967. The biology of ectotrophic rny­ India. - Ind. For. 9'2: 19-20. corrhiza with reference to the growth o-f BIRCH, T. C. B. 1937. A synopsis of forest fungi pines in Nigeria. - Obeche, Jour. of Tree of significance in New Zealand. - N. Z. Glub, Univ. of Ihadan 1 (3): 9-18. Jour. For. 4: 1-09-125. MELIN, E. 192-5. Untersuchungen tiber die Bedeu­ BRISCOE, C, B. 1959. Early results of mycorrhizal tung der Baummykorrhiza. - G. Fischer, inoculation of pine in Puerto Rico. - Jena. Carib. For. 20: 73-77. MIKOLA, P. 1969. Mycorrhizal inoculation in BROWN, J. L. 1963. Working plan for Mafuga afforestation. - Intern. Rev. For. Res. 3. Central Forest Reserve. - Fo-restry Dept., Academic Press, New York (In print) . Uganda. (Stencil.) MoDESS, 0. 1941. Zur Kenntnis der Mykorrhiza­ CLEMENTS, J. B. 1941. The introduction of pines bildner von Kiefer und Fichte. - Symb. into Nyasaland. - Nyasaland Agr. Quart. Bot. Upsal. 5( 1) . Jour. 1 ( 4) : 5-15. MosER, M . 1967. Die ektotrophe Ernahrungsweise DoNALD, D . G. M. 1965. A study of the history, an der Wa1dgrenze. - Mitt. d. Forst!. practice and economics of forest nurseries Bundesvers.-anst. Wien, 75: 357-38.0. in South Africa. - Ann. Univ. v. Stellen­ PRYOR, L. D. 1956. Chlorosis and lack of vigour bosch 40A, 1. in seedlings of renantherous species of Eu­ F AO Secretariat. 196 7. Actual and potential role wlyptus caused by lack of mycorrhiza. - of man-made forests in the changing world Proc. Linn. Soc. N. S. W. 811: 9·1-96. pattern of wood consumption. - FAO PRYOR, L. D. 1958. How important is mycorrhiza World Symposium on Man-made Forests, to introduced conifers? - Austr. Jour. Sc. Documents 1: 4--50. 20: 215-216. FIELDING, J . M. 1957. Introduction of Pinus ra­ PuRNELL, H. 195 7. Notes on fungi found in Victo­ diata to Australia. - Austr. For. 21 : 15- rian plantations. - Plantat. Techn. Pap., 16. Foe. Comm. Victoria, 3: 9~ 13 . GIBSON, I. A. S. 196·3. Eine Mitteilung tiber die RAWLINGS, G. B. 1950. The mycorrhizas of trees in Kiefernmykorrhiza in den Waldern Kenias. New Zealand forests. - For. Res. Notes, Mykorrhiza, Intern. Mykorrhizasym­ N. Z. For. Service 1: 15-17. posium, Weimar 1960. 49~51. RAWLINGS , G. B. 1958. Some practical aspects KESSELL, S. L. 192 7. Soil organisms. The depend­ of forest mycology. - Proc. N. Z. Soc. ence of certain pine species on a biolo­ Soil Sci. 3: 41-44. gical soil factor. - Emp. For. Jour. 6: REICHERT, I. 1940. Studies on mushrooms and 70-74. other fungi of the forests of Palestine. I. KREISEL , H. 1963. Discussion of Singer 1963. - Boletus Boulieri Que!. and B. Bellini In­ Mykorrhiza, Intern. Mykorrhizasymposium, genza. - Palest. Jour. Bot. 3: 209-224. Weimar 1960. 229-230. REICHERT, I. & AviZOHAR-HERSHENZON, Z. 1959. A LAMB, A. F. A. 1956. Exotic forest trees in Tri­ contribution to the knowledge of higher nidad and Tobago.- Gov. Print., Trinidad furigi of Israel. - Bull. Res. Council Is­ and Tobago. rael 7D: 222-247.

175 ROBERTSON, N . .f . 1954. Studies of the mycorrhiza STAHL, W. 1968. Mycorrhiza fungi may be seed of Pinus silvestris. I. The pattern of develop­ borne. - Inst. Forest. Austr. New3letter 9·: ment of mycorrhizal roots and its signi­ 11. ficance for experimental studies. - The STEPHENS, E. L. & Krnn, M. M . 1953a. Some New Phytol. 53: 253-283. South African edible fungi. - Longmans, ScHRAMM, J. R. 196·6. Plant colonization studies Green & Co. Cape Town. on black wastes from anthracite mining in STEPHENS, E. L. & Krnn, M . M . 1953b. Some Pennsylvania. - Trans. Amer. Phil. Soc. South African poisonous fungi. - Long­ 56(1 ). mans, Green & Co. Cape Town. SINGER, R. 1950. Die hoheren Pilze Argentiniens. STREETS, R . ]. 1962. Exotic forest trees in the - Schw. Zeitschr. f. Pilzkunde 28 : 181- British Commonwealth. Clarendon 196. Press, Oxford. SINGER, R. 1963. Der Ektotroph, seine Definition, VAN SucHTELEN, M. J. 1962. Mykorrhiza bij Pinus geographische Verbreitung und Bedeutung spp. in de tropen. Med. Landb. in der Forstokologie. - Mykorrhiza, In­ Hogesc'h. Gent 27: 1104- 1106. tern. Mykorrhizasymposium, Weimar 1960. TAKACS , E. A. 1'961. Algunas especies de hongos 223-231. forma:dores de micorrizas en arboles fo­ SINGER, R . 1964 Boletes and related groups in restales cultivados en Ia Argentina. - Rev. South America. - Nova Hedwigia 7: 93 - For. Ar.gentina 5: S.0-82. 132. TAKACS, E. A. 1967. Produccion de cultivos puros SMITH, N. ]. G. & PoPE, F. B. 1934. The associa­ de hongos micorriz6genos en el Centro Na­ tion between the gasteromycete Polysacchum cional de Investigaciones Agropecuarias, and Eucalyptus roots. - Trans. Brit. Myc. Castelar. - Idia, Suppl. For. 4 : 83-87. Soc. 19: 95 . Received B. 2. 1969 Printed 29. 10. 1969

176