Biodiversity and Conservation 6, 45±58 (1997)

Ectomycorrhizal fungi of the Philippines:

a preliminary survey and notes on the geographic
biodiversity of the Sclerodermatales
K. SIMS
Research School of Biosciences, University of Kent, Canterbury, Kent CT2 6NJ, UK

R. WATLING
Royal Botanic Gardens, Inverleith Row, Edinburgh, UK

R. DE LA CRUZ
BIOTECH, University of the Philippines at Los BanÄos, Laguna, The Philippines

P. JEFFRIES*
Research School of Biosciences, University of Kent, Canterbury, Kent CT2 6NJ, UK

Received: 21 September 1995; revised and accepted 20 November 1995

Basidiomata of putative ectomycorrhizal fungi have been collected from under pine or dipterocarp
stands in Central Luzon in the Philippines. Over 100 collections are reported. Among the material,
75 taxa have been recognised and assigned to known genera or at least placed close to already
documented European, Japanese or North American taxa. Formal identi®cations for 46 species are
given along with comments on previous collections from South-East Asia. Species collected from
under the native Pinus kesiya resemble the communities found under three-needled pines in North
America and the West Himalayas, whereas those from dipterocarp forests were similar to com-
munities found in Malaysia and Japan. Members of the Sclerodermatales were particularly common
and are probably the pioneer colonizers of young seedlings in these ecosystems.
Keywords: fungal biodiversity; ectomycorrhiza; Philippines; Sclerodermatales; biogeography.

Introduction
All forest trees depend on bene®cial symbioses with mycorrhizal soil fungi for the ecient
uptake of minerals. This symbiotic relationship is vital to tree growth in natural ecosys-
tems, and also in commercial forestry, both in the nursery and in the plantation. In the
tropics, for example, the South-East Asian rain forest is dominated by trees from the
Dipterocarpaceae, and ectomycorrhizal associations have been reported from at least 13 of
the 16 genera in this family (Lee, 1990). Many of the fungi that form ectomycorrhizal
associations with forest trees belong in the Basidiomycotina, a group which includes
mainly genera that produce large fruit bodies or basidiomata. Those genera which form
pu€balls, earthballs and earthstars are grouped in the gasteromycetes, in which the spores
mature within an enclosed basidiome which contains the basidia from which they are not
forcibly discharged. The most common examples of these fungi, especially in Australasia

*To whom correspondence should be addressed.

0960-3115 Ó 1997 Chapman & Hall
46 Sims et al.
and other warm, dry regions, are the Sclerodermatales. It is assumed that these are im-
portant symbionts of many tropical and temperate trees, including several of economic
importance, although their ectomycorrhizal connections have only been demonstrated
unequivocally for relatively few examples.
Few records have been collected of the ectomycorrhizal fungi in the Philippines.
Pampolina (1991) made a survey of ectomycorrhizal fungi associated with dipterocarps in
the Mount Makiling area, Southern Luzon, and seven genera were recognised, viz. Rus-
sula, Lactarius, Amanita, Boletus, Paxillus, Scleroderma, Cantharellus. Five and three
species, respectively, were recorded for the ®rst two genera, including Lactarius piperatus
(Scop.: Fr.) S.F. Gray which was found throughout the collecting period, but voucher
material was not available to compare these specimens with European collections (Wa-
tling, pers. obs.). Surveys of this nature are necessary since it is important to identify the
mycorrhizal fungi occurring in natural forests so that new plantations, or replantation
sites, can be assessed for the presence of appropriate e€ective mycorrhizal fungi. In their
absence, seedlings will not establish and rehabilitation schemes will fail (Je€ries and Dodd,
1991). In such circumstances inoculation of tree seedlings at the nursery stage is necessary,
and research is required to select bene®cial and competitive strains that possess the ap-
propriate physiological traits necessary for inoculation techniques in the ®eld.
As part of an EU-funded project relating to the use of mycorrhizal fungi in refor-
estation schemes, a preliminary survey of ectomycorrhizal fungi was undertaken during a
collecting trip in Central Luzon in 1993. Ninety-one collections from this survey are
reported here. Among the material, 46 taxa were recognised, of which half were assigned
to known species or at least placed close to already documented European, Japanese or
North American taxa. Members of other families (10 Russulaceae and 4 Cortinariaceae),
although documented, are not discussed in detail. For the ®rst time it is possible to relate
named species of South-East Asian pines and dipterocarps with named fungi, even if only
by association. Many if not all of the records appear to be new to the Philippines and do
not appear in Quimio and Capilit (1981) or Pampolina (1991), the most up-to-date lists of
Philippine fungi.

Materials and methods

Collection sites
All the collections were made in pine woods at Baguio and in pine woods near Bokod, and
dipterocarp woodland at Zambales, and in mixed growth at the Baguio Botanic Garden.
To these collections have been added material collected by K. Sims at Quezon National
Park, by P. Je€ries (University of Kent) on an earlier trip to Baguio, and by B. Dell and N.
Malajczuk (University of Murdoch, W. Australia and CSIRO, W. Australia respectively)
also from Baguio at a later date. Fresh basidiomata were photographed and described,
and if appropriate, sections were removed and transferred onto a Modi®ed Melin
Norkrans agar medium for culturing. Basidiomata were then placed in plastic bags con-
taining silica gel to dry them. The silica gel was changed frequently to ensure ecient
drying. All the material is deposited in the herbarium at the Royal Botanic Garden,
Edinburgh (E) and appropriate accession numbers are quoted in the records. Records of
Malaysian fungi lacking cited references are observations made by one of the authors
(RW) and Evelyn Turnbull. Any collections with K number codes refer to material col-
lected by K. Sims which is now deposited in the herbarium at the Royal Botanic Garden
Ectomycorrhizal fungi of the Philippines 47
Edinburgh (in E). Material given the code Sims 6 is the original name of a culture col-
lection which has no dry material stored in E.

Identi®cation
Smell, surface texture and colour of gills, stipe and pileus of basidiomata were noted, as
well as the type of gill attachment, presence of latex (coloured or not), presence or absence
of veils, consistency of the ¯esh, etc. following Henderson et al. (1969). The reaction of
iron II sulphate was determined by rubbing a crystal onto the stem of the basidiome and
noting the colour change: pink, salmon, bu€ or green (negative reaction). The treated area
was always removed before drying of basidiomata. Classi®cation in the main follows
Julich (1981) and Knudsen (1995).

Results
Despite the relatively small scale of the collection, a diverse range of basidiomata of
ectomycorrhizal fungi was found. Over 120 individual collections were made, representing
at least 75 taxa. Forty-six of these have been assigned to existing species, one is probably a
new species, and 21 more (all Russulaceae or Amanita species) as yet have not been
formally identi®ed. Further details are given below. The most frequently encountered
basidiomata were from the Sclerodermatales, especially Scleroderma dictyosporum Pat.
and S. verrucosum Pers. Both these fungi were also observed to be fruiting within the
plastic bags containing seedlings of Pinus kesiya Royle ex Gordon in the nurseries at the
University of Los BanÄos. These plants had been inoculated with `Mycogroe' tablets, a
local form of mycorrhizal inoculum containing spores of Pisolithus arhizus (Pers.) Raus.
and other gasteromycete basidiomata collected locally. In these particular cases it was
possible to gently remove the soil from the bags and observe mycelial strands extending
from the basidiomata to the tree roots. In the ®eld it is much more dicult to trace
mycorrhizal connections between roots and basidiomata, but in the case of Scleroderma
sinnamariense Mont. the bright yellow colouration of the mycelium enabled successful
connections to be traced between the basidiomata and roots of the associated dipter-
ocarps.

List of fungi found

BOLETALES
Boletaceae
1. Boletus aemelii Barbier in Bull. Soc. Mycol. Fr. 31,54 (1915)
Baguio, Roosevelt Plantation, with Pinus kesiya, legit B. Dell & N. Malajczuk, 23.6.93
Wat. 25599; Bokod, Bobok Comm., BMC forest, with Pinus kesiya, 17.6.93, Wat. 25588.
A collection by Je€ries from Baguio (27.6.92) probably also represents this bolete (Wat.
25183). There has been much discussion as to the correct name which should be used for
this fungus (Watling and Li, unpublished). Although described from Europe, it has been
found amongst dried collections from Australia (Watling and Li, unpublished).
2. B. destitutus Corner in Boletus in Malaysia 227 (1972)
Quezon Nat. Park, with Dipterocarpus grandi¯oris Blanco, and Anisoptera sp., legit K.
Sims, #7/41, 29.6.93, Wat. 25594. This small bolete was known previously only from
Peninsular Malaysia growing on the soil in woodland.
48 Sims et al.
3. B. discolor QueÂl. (= B. junquilleus s. Watling, 1969) in Fl. Champ. S. Fr. 382 (1909)
Bokod, Bobok Comm., BMC forest, with Pinus kesiya, on slope among needles, 17.6.93,
Wat. 25712. This is considered by some a variety of B. queletii Schulz., the yellow stipe
with yellow or orange but no red punctae is characteristic. Because of confusion its
distribution is poorly known.
4. B. cf. patouillardii Singer in Amer. Midl. Nat. 37, 55 (1947)
Labrador, South Zambales, under Shorea and Anisoptera, coastal zone, 24.6.93, legit B.
Dell & N. Malajczuk, Wat. 25644. This collection di€ers from the type (Cambodia, Petelot
222, FH) particularly in its larger size.
5. B. rubellus probably subsp. borneensis Corner in Boletus in Malaysia 120 (1972)
Baguio, Roosevelt Plantation, under Pinus, legit B. Dell & N. Malajczuk, 23.6.93, Wat.
25592. This is a member of a widely distributed complex which needs critical re-
examination.
6. B. subvelutipes Peck in Rept. N.Y. State Mus. 2, 142 (1889)
Baguio, Roosevelt Plantation, under Pinus kesiya, 23.6.93, legit B. Dell & N. Malajczuk,
Wat. 25713. This collection was ®rst thought to be the recently described B. horakii Lakh.
from India, but it lacks the prominent caulocystidia.
7. B. variipes Peck in Ann. Rept. N.Y. State Mus. 41, 76 (1808)
Baguio, Roosevelt Plantation, under Pinus kesiya, 16.6.93, Wat. 25597; ditto, legit B. Dell
& N. Malajczuk, 23.6.93, Wat. 25591; Bokod, Bobok Comm., BMC forest, 17.6.93, Wat.
25598. A very common widespread member of Sect. Edules. A collection of an immature
basidiome was made by P. Je€ries (Baguio, 27.6.92; Wat 25185).
8. Rubinoboletus balloui var. fuscatus (Corner) Heinem & Rammeloo in Bull. Jard. Bot.
Nat. Belg. 53, 296 (1983)
Photographed by K. Sims, Quezon. A collection from Baguio, Roosevelt Plantation,
under Pinus kesiya, 27.6.92, legit P. Je€ries, Wat. 25712 also belongs here. This is a variety
of a widely distributed bolete found in N. America and Australasia extending into Japan.
It is recorded from both Malaysia and Indonesia.

Paxillaceae
9. Paxillus rubicundulus P.D. Orton in Notes Roy. Bot. Gdn. Edinb. 29, 110 (1969)
Baguio Botanic Garden in mixed stand of Pinus and Alnus, legit B. Dell & N. Malajczuk,
23.6.93, Wat. 25589. P. rubicundulus was described from alder carrs in the British Isles
(Orton, 1969) but is apparently widespread elsewhere in Europe.
10. Phylloporus bellus (Mass). Corner in Nova Hedwig. 20, 798 (1970)
Baguio Botanic Garden, in mixed stand with Pinus kesiya, 18.6.93, Wat. 25614; in old
growth mixed dipterocarps mostly Dipterocarpus grandi¯oris and Anisoptera sp., legit. K.
Sims (#7/41), 29.6.93, Wat. 25594. This species was originally described from the Singa-
pore Garden Jungle but has subsequently been collected elsewhere in Pen. Malaysia, in
mixed plant communities; it is known from Australia (Queensland).
11. P. bogoriense (HoÈhnel) in Sitzber. Kaiserl. Acad. Wiss. Wien Math.-naturu. Kl. 123,
89 (1914)
Cobalt Mine Co., near Santa Cruz, Zambales, under Shorea and Anisoptera, 22.6.93, Wat.
25616. Originally described from Java, it is now known from N. America (Florida) and
from Queenland, Australia (Watling and Gregory, 1991). Smits (1994) recorded it from
Indonesia.
Ectomycorrhizal fungi of the Philippines 49
12. P. foliiporus (Murr.) Singer in Persoonia 9, 424 (1978)
Baguio Botanic Garden, several collections under Pinus kesiya, 18.6.93, Wat. 25602; Ba-
guio, Roosevelt Plantation, 16.6.93, Wat. 25595 & 96. The distribution of this species has
been extended recently by Watling and Gregory (1991) to include Australia (Queensland).
It was originally based on the North American Gomphidius foliiporus Murrill.
13. P. leucomycelinus Singer in Persoonia 9, 426 (1978)
Baguio Botanic Garden, in mixed stand of Pinus kesiya, 18.6.93, Wat. 2560l. The no-
menclature for this fungus is complex and has been outlined by Watling and Gregory
(1991). It is found in both North and Central America and in Australia, associated with a
variety of plant communities.

Gomphidiaceae
14. Gomphidius roseus (Fr.) Gill in Champignons de la France, 623 (1874)
Associated with Suillus bovinus (L.: Fr.) Kuntze, Baguio, Roosevelt Plantation, under
Pinus kesiya, 16.6.93, Wat. 25611. The relationship with Suillus is now well-known from
many parts of its range and has been recently documented microscopically (Agerer, 1991).

Strobilomycetaceae
15. Strobilomyces polypyramis Hook. f. apud Berk. in Hook. J. Bot. Kew Gdn. Misc.
3, 78 (1851)
Antimonan, Quezon National Park, second growth mostly Dipterocarpus grandi¯oris,
29.6.93, legit K. Sims, Wat. 25570. Known previously from India (type locality: Sikkim)
and from Java, it has recently been recorded by Smits (1994) and is also known from
Pen. Malaysia. It is well characterized by the pyramidal scales on the pileus.
16. S. velutipes Cke. & Mass. in Grevillea 18, 5 (1889)
Baguio, Roosevelt Plantation, under Pinus kesiya, legit B. Dell & N. Malajczuk, 23. 6.93,
Wat. 25569. Also among collections made by P. Je€ries, same locality, 27.6.92, Wat.
24988. S. velutipes was originally described from Australia but is known from India,
Africa and Pen. Malaysia including Singapore.
17. Suillus albidipes (Peck) Singer in Farlowia 2, 45 (1945)
Bokod, Bobok Comm., under Pinus kesiya, in troops, in needles and among grass, 18.6.93,
Wat. 25606. This fungus is widespread from New England to the Paci®c Coast of North
America. It is not known from Japan.
18. S. borealis Smith, Thiers and Miller in Lloydia 28, 123 (1965)
Bokod, Bobok Comm., under Pinus kesiya, in troops among needles and sparse grass,
18.6.93, Wat. 25605. Previously this bolete was only known from the Paci®c Northwest of
North America.
19. S. bovinus (L.: Fr.) O. Kuntze in Rev. Gen. Plantarum 3, 535 (1898)
Baguio, Roosevelt Plantation, with Pinus kesiya, in large troops, 16.6.93, Wat. 25566-68
inc.; also see Gomphidius roseus above. This is a common boreal bolete with Pinus syl-
vestris L. It is known from Japan and has been introduced into Australia.
20. S. granulatus (L.: Fr.) O. Kuntze in Rev. Gen. Plantarum 3, 535 (1898)
Bokod, Bobok Comm., BMC forest, in du€ under Pinus kesiya, 17.6.93, Wat. 25612; ditto,
14km from Kayapa, 18.6.93, Wat. 25615. This is probably one of the most widespread
species of Suillus having been introduced with exotic plantings of pine to Africa, South
America and Australia. There is a record but suprisingly not with pine in Pen. Malaysia
50 Sims et al.
(Corner, 1972), although it is known to occur there with planted Pinus merkusii Jungh. &
De Vr., P. kesiya, P. oocarpa Scheide ex Schlect. and P. massoniana Lamb ( Ivory, 1975).
21. S. subaureus (Peck) Snell in Lloydia 7, 30 (1944)
Near Chromite Mine, Santa Cruz, Zambales, under Pinus merkusii, 23.6.91, Wat. 25641;
ditto, Wat. 25642. This bolete is known from central and eastern United States and
southern Canada, and from Japan.
22. S. a€. subluteus (Peck) Snell in Lloydia 7, 34 (1944)
Baguio, Roosevelt Plantation, with Pinus kesiya, 16.6.93, Wat. 25604; Bokod, Bobok
Comm., BMC forest, with P. kesiya, 18.6.93, Wat. 25603. This was ®rst called S. co-
thurnatus Singer in the ®eld from the baggy ring but this soon collapses and is not as
persistent as it would be in S. cothurnatus. There has been much discussion as to the
delimitation of this bolete. The present collections di€er from S. cothurnatus also in the
spores which in size approach S. subluteus which is a fungus known from eastern and
central North America and Japan. It is probably what Singer referred to as `cothurnatus ±
northern form' when he visited Michigan. Also found in the ®rst locality on 27.6.92 by P.
Je€ries as Wat. 25181.

Tylopilus
23. `Boletus' cf. funerarius Mass. in Kew Bull. 207 (1909)
Baguio, Roosevelt Plantation, under Pinus kesiya, 27.6.92, legit P. Je€ries, Wat. 25176.
This is apparently a common species in Pen. Malaysia and has been interpreted here in the
sense of Corner (1972); Corner's earlier interpretation (Corner, 1947) should be referred to
another very di€erent bolete, B. phaeocephalus Pat. & Baker. Singer (1947) places B.
funerarius close to Tylopilus velutipes (Pat. & Baker) Singer, a bolete which should be
referred to Rubinoboletus Heinem. & Rammeloo.
24. `Boletus' nigerrimus Heim in ReÂv. Mycol. Paris 28, 281 (1963)
Baguio, Roosevelt Plantation, under Pinus kesiya, 27.6.92, legit P. Je€ries, Wat. 25186.
This is interpreted in the sense of Corner (1972) whose description di€ers from that of
Heim (1963) in the colour of the spore-print, the rufescent ¯esh and abundant cystidia.

Xerocomaceae
25. Boletellus chrysenteroides (Snell) Sing. in Mycologia 33, 422 (1941)
Baguio, Roosevelt Plantation, under Pinus kesiya, 16.6.93, Wat. 25563 & 64; Baguio
Botanic Garden, under Pinus kesiya, 18.6.93, Wat. 25565. A collection from the Roosevelt
Plantation was also made by B. Dell and N. Malajczuk (Wat. 25590, 23.6.93). It is known
from North America and Japan.
26. B. emodensis (Berk.) Singer in Annls. Mycol. 40, 18 (1942)
Baguio, Roosevelt Plantation, legit B. Dell & N. Malajczuk, 23.6.92, Wat. 25572; Bokod,
Bobok Comm., under Pinus kesiya, 17.6.92; unfortunately Wat. 25571 is immature, but
the taxon is very distinctive. It is known from Australia to India through Pen. Malaysia to
Japan, and Smits (1994) recorded it in Indonesia.
27.Boletellus sp. (nov. sp.)
Baguio, Roosevelt Plantation, under Pinus kesiya, 27.6.92, legit P. Je€ries, Wat. 25180.
This collection is very close to B. chrysenteroides but di€ers in the cross-striae on the
basidiospores: see above.
Xerocomus rubellus (Krombh.) Sing. see under Boletus.
Ectomycorrhizal fungi of the Philippines 51
Rhizopogonaceae
Rhizopogon is a hypogeous relative of the Boletaceae s.lato. Several collections of members
of this genus were made in Luzon but unfortunately all but one proved to be immature.
28. Rhizopogon roseolus (Corda) T.M. Fries in Svensk Bot. Tld. 3, 288 (1909)
Mindanao, Malaybalay, Bukidnun, legit Elsie B. Lorilla, 27.8.93, 93-P14.
29. R. vulgaris (Vittad.) M. Lge. in Dan. Hypog. Macromyc. 56 (1956)
Bokod, BMC forest, on bank under Pinus kesiya, (Sims 5) 18.6.93, Wat. 25673. R. vulgaris
is widespread under pines, although despite the epithet generally never as common at least
in Europe as R. luteolus f. apud Fr. & Nordh. and R. roseolus (Corda) T.M. Fries.

SCLERODERMATALES
Pisolithaceae
30. Pisolithus arhizus (Pers.) Raus. in Zeits. Pilzk. 25, 51 (1959)
Baguio, Roosevelt Plantation, on rocky slope under Pinus kesiya, 16.6.92 (same locality,
legit P. Je€ries, 27.6.93); Bokod, Bobok Comm., BMC forest, on side of bank, trail under
Pinus kesiya, 17.6.93, (Sims 6), Wat. 25672. Also with Pinus kesiya in pots, Los BanÄos,
Biotech, 24.6.93.

Sclerodermataceae
31. Scleroderma cepa Pers. in Syn. Meth. Fung. 155 (1801)
Malaybalay, Butiknun, Mindanao, 27.8.93, legit Elsie Lorilla, 93-P13. A widespread but
never common earthball and possibly southern in its distribution in Europe.
32. S. columnare Berk. & Br. in J. Linn. Soc. (Botany) 14, 80 (1875)
Cobalt Mining Co., near Santa Cruz, Zambales, in mixed dipterocarp grove with Ani-
soptera sp., 22.6.93, K18; ditto, with Dipterocarpus sp., Shorea polysperma (Blanco) Merr.
and Anisoptera sp., 23.6.93, K19-25 inclusive. Also known from pot cultures, Los BanÄos,
Biotech, legit P. Je€ries. Recorded from Pen. Malaysia.
33. S. dictyosporum Pat. in Bull. Soc. Mycol. Fr. 12, 135 (1896)
Los BanÄos, Makiling Botanic Garden, with Parashorea malaanonan (Blanco) Merr., 23.6.92,
legit P. Je€ries, Wat. 24987 & 24990; ditto, mixed dipterocarps, along trail, 23.6.93, K27;
Quezon National Park, with Dipterocarpus sp. and Shorea sp., 29.6.93, K28-30. Also known
in pot culture at Los BanÄos, Biotech, with Eucalyptus spp., 3.4.92, legit P. Je€ries; ditto, with
Pinus kesiya, K26. This is a pantropical earthball recognised in several important rain forest
communities e.g. recorded from Indonesia by Smits (1994).
34. S. leptopodium Har. & Pat. in Bull. Soc. Mycol. Fr. 15, 84 (1909)
Cobalt Mining Co., near Santa Cruz, Zambales, in mixed dipterocarp forest, 23.6.93, K32
& 33. This has only recently been recognised (GuzmaÂn, 1969) as being distinct from S.
columnare with which it was placed in the genus Veligaster. It di€ers particularly in the
spore size. Smits' record (1994) of S. columnare may be this species.
35. S. sinnamariense Mont. In Annls. Sci. Nat. 2 ser. 14, 331 (1840)
Cobalt Mining Co., near Santa Cruz, Zambales, in mixed dipterocarp forest with Ani-
soptera sp., Dipterocarpus grandi¯oris and Shorea polysperma, 22.6.93, K34, K35 and K36.
This is a widespread pantropical species and found in all the major tropical forests of the
Old and New World. Consequently it has been given many di€erent names. It is well-
known in Pen. Malaysia.
52 Sims et al.
36. S. verrucosum Pers. in Syn. Meth. Fung. 154 (1801)
Los BanÄos, Makiling Botanic Garden, with Parashorea malaanonan, 23.6.92, legit P.
Je€ries, Wat. 24983-86 inclusive; ditto, with Shorea polysperma, legit P. Je€ries; ditto, with
Hopea philippinensis Dyer, 30.6.93, K44; ditto, with Parashorea, 30.6.93, K45; ditto, with
Hopea foxworthi Elm., 30.6.93, K46. Also same locality, 30.6.93, K41 & 42. Bokod, Bobok
Comm., BMC forest, along forest trail near Pinus kesiya, 17.6.93, K38. Also known from
Biotech, Los BanÄos. Growing in pots with Eucalyptus spp., 3.4.92, legit P. Je€ries, Wat.
25174; with Pinus kesiya, 24.6.93, K39 & 40. This species although found in Europe and
North America has probably a more southerly distribution. It is quite widespread in the
tropics (GuzmaÂn, 1970).

Astraeaceae
37. Astraeus hygrometricus (Pers.) Morg. in J. Cincinnati Soc. Nat. Hist. 12, 20 (1889)
Los BanÄos, Makiling Botanic Garden, with Shorea contorta Vidal, 30.6.93, K59; ditto with
Hopea plagata (Blanco) Vid., 30.6.93, K60 & K61; ditto with Anisoptera thurifera (Blanco)
Blume, 30.6.93, K62 & K63; ditto with Dipterocarpus grandi¯oris, 30.6.93, K64. A
widespread fungus in Europe, Asia including the Indian subcontinent, North America,
Africa and Australia, it is easily confused with members of the Geastraceae.

GOMPHALES

The Gomphaceae and its coralloid members now placed in the Ramariaceae have been
related by some to the Boletales. A single representation of this family was made:
38. Ramaria zippelii (LeÂv.) Corner in Clavaria and allied genera 632 (1950)
Cobalt Mining Co., near Santa Cruz, Zambales, under Anisoptera and other mixed dip-
terocarps including Dipterocarpus grandi¯oris and Shorea polysperma, 22.6.93, Wat.
25679. Members of the genus Ramaria are thought to be mycorrhizal (Knudsen, 1995). R.
zippelii has been found in India, Sri Lanka, Java, Malaya, Borneo, New Guinea and N.
Caledonia. The present collection agrees with var. gracilis of Corner (1950).

THELEPHORALES

The Thelephoraceae has also been linked to the Boletales. A single representative of this
family was made:
39. Thelephora ramarioides Reid in Kew Bull. 227 (1958)
Cobalt Mining Co., near Santa Cruz, Zambales, under Casuarina equisetifolia Forst., on
bank accompanied by Lycoperdon polymorphum Vitt., 22.6.93, Wat. 25678. A widespread
fungus in Australasia (apparently speci®c to growing with Casuarina equisetifolia). It is
known in close association with several trees (e.g. Acacia mangium Willd.) from Australia,
Indonesia, Malaysia etc., but inoculation experiments indicate that it might not be my-
corrhizal under all conditions.

Non-boletoid records
TRICHOLOMATALES
Hydnangiaceae
40. Laccaria amethystea (Bull.) Murrill in N. American Flora 10, 1 (1914)
Baguio, Roosevelt Plantation, under Pinus kesiya legit P. Je€ries, 27.6.92, Wat. 24994. A
Ectomycorrhizal fungi of the Philippines 53
widespread mycorrhizal fungus found throughout Europe, North America and Northern
Asia. It is also recorded from Japan and probably occurs in Pen. Malaysia judging from
recent ®nds. In Britain it is usually con®ned to fagaceous woods with rather deep shade.

AMANITALES
Amanitaceae
41. Amanita gymnopus Bas & Corner in Persoonia 2, 259 (1962)
Cobalt Mining Co., Zambales, near Santa Cruz, under Anisoptera sp. with Shorea poly-
sperma, and Dipterocarpus grandi¯oris, 22.6.93, Wat. 25833. This was described from
Malaysia (Johore) and based on a single collection. Watling & Turnbull have several
collections from around Kepong, Pen. Malaysia (Watling, pers. obs.). The present ma-
terial extends the distribution.
42. A.hemibapha (Berk. & Br.) Sacc. in Syll. Fungorum 5, 13 (1887)
Bokod, BMC Co., under Pinus kesiya, 17.6.93, (immature); ditto, Baguio, Roosevelt
Plantation, 23.6.93, legit B. Dell & N. Malajczuk, Wat. 25834. Although originally de-
scribed from Sri Lanka, it is widespread in South-East Asia occurring in many forms. It is
also thought to occur in the Southern States of North America. The Philippine material
agrees with the subspecies javanica Corner & Bas.
43. A. pantherina (Fr.) Krombh. in Abbild 4 pl. 29 (1836)
Bokod, BMC Mining Co., under Pinus kesiya, 17.6.93, Wat. 25836; ditto, Baguio, Roo-
sevelt Plantation, 23.6.93, legit B. Dell & N. Malajczuk, Wat. 25835. This is a widespread
species known from Europe and N. America in addition to the Himalayas (see Watling
and Gregory, 1980).
Note:12 other collections of Amanita have been made but as yet have not been allocated to
speci®c groups: the material has been fully documented both macroscopically and mi-
croscopically.

RUSSULALES
Russula and Lactarius
Both these genera are important ectomycorrhizal genera (Thoen, 1993; Watling & Lee,
1995) and representatives of both have been collected (eight species of Russula; two of
Lactarius). Although fully documented only one, Russula virescens, is dealt with herein;
the two Lactarius spp. belong to Sect. Dapetes.
44. R. virescens (Schae€.) Fr. in Epicrisis Syst. Mycol. 355 (1838)
Bokod, Bobok Comm., BMC forest, with Pinus kesiya, 17.6.93; Baguio, Roosevelt
Plantation, with P. kesiya, legit B. Dell & N. Malajczuk, 23.6.93. This is a widespread
agaric in Europe, North America and Asia. It is widespread in Pen. Malaysia.

Aphyllophoralian fungi
HYMENOCHAETALES
Coltriciaceae
Coltricia is considered by some authorities to be mycorrhizal (Thoen, 1993). Two col-
lections of the same species were made during the present study.
45. C. cinnamomea (Pers.) Murr. in Bull. Torr. Bot. Club 31, 343 (1904)
Bokod, Bobok Comm., BMC forest, on sandy soil, under Pinus kesiya, 18.6.93, Wat.
54 Sims et al.
25680; near Santa Cruz, Zambales, with Pinus merkusii, on sandy soil, 23.6.93, Wat.
25677. This is a widespread member of the genus being recorded from Africa, Sierra Leone
to Kenya, South Africa, southern Europe and North America. It is widespread in Asia and
Australia where it has been called C. oblectans (Berk.) G.H. Cunn.; the Australian concept
agreed with the collection from Bokod.

CANTHARELLALES
Scutigeraceae
46. Albatrellus cristatus (Pers.: Fr.) Kotl. & Pouz. in Ceska Mykol. 11, 154 (1957)
Bokod, Bobok Comm., BMC forest, 14 km from Kayapa, under Pinus kesiya only as
immature basidiomes, on bank of red clay, 18.6.93, Wat. 25613. Albatrellus is a terrestrial,
rather ¯eshy, genus of polypores which have been considered mycorrhizal (Singer et al.,
1983). This species is known from Europe and Japan as well as eastern and mid-west
regions of the USA; it has recently been recorded from India. It is characterized by the
greening of the bruised or ageing basidiomes.

Discussion
Most of the specimens found were from the Boletales or related orders, for example the
Sclerodermatales. This order has been linked to the Boletales through common secondary
metabolites (Gill and Watling, 1986) and the fact that they play host, in parallel to
members of the family Paxillaceae, to the hyperparasite Sepedonium chrysospermum Tul.
There are several ®rst records from the Philippines. For example, Strobilomyces velu-
tipes and S. polypyramis form a closely related pair and can be considered together. The
Philippine collections suggest that this genus is widespread in the Old World tropics and
undoubtedly associated with a range of trees both angiosperms and gymnosperms. This
may indicate that Strobilomyces may not be mycorrhizal, or, if it is, then the species must
either be facultative mycorrhizal or capable of switching hosts rather dramatically. Bo-
letellus emodensis parallels S. velutipes in its mycogeography and probably biology.
On the other hand, Suillus is considered to be only associated with Pinaceae particularly
Pinus spp. Although Corner (1972) had noted a collection of S. granulatus in Malaysia
where no conifers were to be found, such records are rare, although the occurrence of
some Suillus species under hardwoods is also reported by some American authors (e.g.
Smith and Thiers, 1971). It has also become traditional to think that Suillus spp. are
restricted to trees of a speci®c section of the genus, e.g. two-needled, three-needled pines,
but this is not necessarily true and in tropical plantations both Suillus granulatus and S.
luteus form ectomycorrhizas with many di€erent Pinus spp. from all sections of the genus
(Ivory, pers. comm.). Switches are also known under arti®cial conditions, e.g. S. luteus
usually with P. sylvestris (two-needled) growing with P. radiata D.Don (three-needled) in
Australia (Watling and Gregory, 1989). Thus S. bovinus, a European species also found
with P. sylvestris, is found in the Philippines with P. kesiya and in addition in the exciting
combination with Gomphidius roseus, just as it is in Europe and in Japan. Pearson (1950)
was familiar with S. bovinus under P. radiata in South Africa but not with P. sylvestris.
These collections extend the range of hosts for S. bovinus, and the other species of Suillus
recorded, and their geographical distribution. Some Suillus spp. might be expected with P.
merkusii (two-needled).
Ectomycorrhizal fungi of the Philippines 55
The genus Boletus and its satellites are an interesting mixture of temperate European/
North American taxa and possible Malaysian elements. The number of species of Phyl-
loporus found in the present study supports the singularly rich Malaysian ¯ora described
by Corner (1970). The two species of Boletellus, other than B. emodensis, hint at a rich
¯ora here which also contrasts markedly with the European biomycota. Contrary to many
beliefs, hypogeous basidiomycetes are not absent from the tropics; there is no doubt
further collections will be added to the Rhizopogon spp. found in the short collecting trip.
The earthballs (Sclerodermatales), including the devils foot fungus' (Pisolithus), are
common with an interesting mixture of tropical and subtropical elements.
Ramaria zippelii is widespread in tropical rain forests, and although Thelephora ra-
marioides has only comparatively recently been described it seems to be widespread in
distribution associated with Acacia spp., Eucalyptus spp. and Casuarina equisetifolia, al-
though it has not yet been shown to form true ectomycorrhizal structures with the asso-
ciated roots. The Amanita spp. and Russula virescens, although not associated with
Casuarina, all have a wide distribution, possibly even wider than that of T. ramarioides.
Amanita hemibapha has been the subject of much debate because of confusion certainly in
the United States with the European A. caesarea (Scop. ex Fr.) QueÂl. Finally, Laccaria
amethystea is a rather unusual sighting, but only probably because of preconceived ideas
regarding its temperate distribution. These observations have indicated that there are a
variety of new host/fungus relationships to be found in the Philippines.
Quimio and Capilit (1981) compiled a very useful list of the fungi recorded for the
Philippines between the years 1936±77. Of the species listed, ®ve agarics could be con-
sidered potentially ectomycorrhizal: viz. Amanita (as Amanitopsis; Quimio, 1983; Quimio
and Suayan, 1975: in our survey three species were found); Lactarius sp. (Quimio and
Suayan, 1975: two species were collected in the present survey); Paxillus (Quimio, 1978:
see P. rubicundulus), Russula (Quimio and Suayan, 1975; Quimio, 1983: eight species
reported herein) and probably Tricholoma, depending on the taxon involved (Quimio and
Suayan, 1975; Quimio, 1983: no collections here). The two Cortinarius spp. would need to
be re-examined before this genus could be put onto the list (three Inocybe spp. were
currently found). Surprisingly no boletes were recorded; these made up the biggest haul of
taxa in our collection. Among the gasteromycetes, two species of Pisolithus and ®ve species
of Scleroderma were recorded, including S. ¯avocrocatum (as `favo') which is considered a
synonym of and is dealt with herein under S. sinnamariense (Sims et al., 1995); it is a
widespread pantropical species. In fact the authors listed above only scratched the surface
of what is a very rich macromycete ¯ora, and failed to hint at the incredible diversity of
agaricoid fungi, including potentially ectomycorrhizal taxa.
In the Philippines, the putative ectomycorrhizal fungi that we collected from the Pinus
kesiya communities resemble in their broad fungal components the forests of three-needled
pines in North America (North-West and Central/East) and the Western Himalayas. Some
European elements were also identi®ed unless these really turn out to be more cosmo-
politan elements and through ignorance they are at present assigned to Europe. Both
geographical and host speci®city details have been expanded. The general dipterocarp
forests were parallel in some ways to those of Malaysia which are themselves related to
communities found in Japan (Watling and Lee, 1995). Indeed, any furtherance of our
knowledge of the Philippine biomycota will be dependent on consulation of texts and
exsiccata from North America and Japan, and volumes on various groups published by
E.J.H. Corner and summarized in Corner (1993).
56 Sims et al.
In areas which are scheduled for reforestation with trees that are obligately ectomy-
corrhizal (as are many parts of the Philippines), it is necessary to ensure that inoculum of
appropriate fungi is present, and that these fungi are capable of persisting in the edaphic
conditions of the replanting site. It is thus essential to choose fast growing `early-stage'
(sensu Deacon et al., 1983) mycorrhizal fungi with a catholic host range as initial colo-
nizers of newly planted soils and members of the Sclerodermatales would appear to be
ideal candidates for warm forest ecosystems, especially in the early stages of establishment
when the soil is more prone to drying than when a full canopy has developed. Both
Pisolithus arhizus and Scleroderma sinnamariense are relatively fast-growing in culture
which may also be signi®cant in this context. In this study we collected six species of
Sclerodermatales (Pisolithus arhizus, Scleroderma cepa, S. columnare, S. dictyosporum, S.
leptopodium, S. sinnamariense) in the Philippines. In addition, we have found Scleroderma
columnare and S. leptopodium in Indonesia, and Scleroderma columnare, S. dictyosporum,
S. echinatum (Petri) GuzmaÂn, S. sinnamariense and S. verrucosum are known from Ma-
laysia (Watling, 1994). Within the genus Scleroderma, both restricted and widespread
distributions are known. Thus, S. sinnamariense and S. echinatum are restricted to rain
forest areas. The former under its many synonyms is widespread in both neo- and pan-
tropical regions, whereas S. echinatum is apparently restricted to South-East Asia (Rifai,
1987; Watling, 1994; Walting and Lee 1995). Other species generally prefer more arid
conditions (e.g. in Mediterranean, S. meridionale Dem. & Mal.) but on the whole soils of a
more mineral status are preferred, certainly for fruiting. Scleroderma verrucosum probably
increases southwards in Europe and is a constituent of tropical ¯oras. Scleroderma co-
lumnare, placed by GuzmaÂn (1969) in the new but super¯uous genus Veligaster (Sims et
al., 1995), is also probably more tropical than temperate.
The biogeography of ectomycorrhizal fungi seems to be determined by the distribution of
suitable hosts. From molecular analyses the major lineages within the ectomycorrhizal
fungi seem to have evolved later than those of the less host-speci®c arbuscular mycorrhizal
species, probably about 130 Myr ago (Berbee and Taylor, 1993) and consequently after the
origin of the angiosperms. The greater frequency of associations with the Pinales and
lower angiosperm families compared with the higher families of angiosperms may con-
tradict this, however, and suggest an evolutionary time scale closer to the evolution of the
early angiosperm lineages (i.e. 200±250 Myr ago). It has also been suggested that ecto-
mycorrhizal associations evolved from saprotrophic relationships, thus ectomycorrhizal
fungi would have become associated with their plant partners over a period of time rather
than being an inherent component during the evolution of the terrestrial ¯ora. The dis-
tribution of ectomycorrhizal fungi is consequently more complex than that of their ar-
buscular mycorrhizal counterparts, especially when complicated by host speci®city such
that some species are con®ned to the geographical range of their host plants. This con-
fusing situation has been exacerbated by the importation of exotic trees into di€erent areas
of the world, often associated with their ectomycorrhizal symbionts. The catastrophic
failure of some trees to establish in the absence of imported fungi (Mikola, 1983) has
demonstrated the heterogeneous distribution of these fungi across the world. It is also
clear that some fungi are more important than others as initial colonizers of young tree
seedlings and bene®t the plant in terms of aiding establishment in stressed ecosystems.
These fungi tend not to be host-speci®c and thus have a more widespread distribution than
the fungi associated with the later stages of tree growth. Thus it is common to ®nd species
of Laccaria, Hebeloma and Thelephora in most temperate ecosystems where young trees
Ectomycorrhizal fungi of the Philippines 57
are found. As discussed above, these tend to be replaced in warmer, drier ecosystems of
many regions of the world by the Sclerodermatales forming ectomycorrhizas with a large
range of trees.

Acknowledgements
We thank our many colleagues in the Philippines who helped us to make the collections,
especially Elsie Lorilla, Joy Zarate, and Sixto `Estoy' Tanyag. We thank B. Dell and N.
Malajczuk for supplying additional specimens. This work formed part of a project funded
by the EU STD programme contract CI1*-CT91-0904 entitled `The use of mycorrhizal
fungi in reforestation programmes in the Philippines'. R. Watling wishes to thank Eliza-
beth Watt who assisted in the identi®cations by documenting the microscopic characters of
the collections.

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