Southeast Asian Myxomycetes. I. Thailand and Burma' DON R. REYNOLDS2 and CONSTANTINE J. ALEXOPOULOS2

TROPICAL SOUTHEAST ASIA includes the Phillip­ Overeem, 1922 ; Penzig, 1898; Raciborski, 1884; pines, the Indo-Malay Archipelago and Penin­ Zollinger, 1844). Chip (1921) and Sanderson sula, Eastern Indochina, and parts of Thailand (1922) published from Singapore and the lower and Burma (Richards, 1952). Europeans initi­ Malay Peninsula." Other collections were studied ated the modern phase of botanical exploration abroad (Emoto, 1931 b; Lister, 1931; Saccardo in this region. The floristics were done either and Paoletti, 1888). In the Philippines, though locally by resident foreign botanists or by spe­ some mycological work has been done, most of cialists in their native country, working with the plant taxonomists who have worked there contributed materials. That which the early resi­ have known little about the fungi. The Myxo­ dents could not competently identify was sent mycetes of Indochina are completely unknown largely to European and American specialists. in the literature. The specimens, of necessity, had to be dried or The present collections are being treated in otherwise preserved for a long sea journey. As a two parts. This paper deals with the material consequence many prominent mycologists pub­ from Thailand and Burma; the second part con­ lished on material they knew only from her­ cerns collections from the Philippines and will barium specimens. In spite of the disadvantages be submitted for publication to the Philippine of possible misinterpretation and duplication of Agriculturist. work, it is fortunate that this procedure became Heim (1962) refers briefly to an abundance prevalent; the duplicates now in the herbaria of of Myxomycetes in Thailand. The only Thai Europe, South Africa, and America are the only specimens of which we are aware are those col­ representatives of a large amount of pre-World lected in April-June 1967 by Dr. Emory Sim­ War II work. mons and deposited by him in the U.S. National Of the early Southeast Asian botanical centers, Fungus Collections (BPI) . These specimens, only the Bogor collections remain largely intact. which we have not seen, were determined by The extensive collections in the Philippines Na­ Dr. Marie 1. Farr who kindly supplied us with tional Herbarium in Manila and the Baker her list of identifications. They are cited under Fungus Collection in the University of the Phil­ the collector's (EGS) numbers at the appropriate ippines, College of Agriculture, were completely places. No citations have been found from destroyed during the Japanese and American in­ Burma. vasions of World War II. Only a few fungus specimens, mostly Baker duplicates, remain in MATERIALS AND METHODS the Singapore Botanical Garden. Collections were made during the summer of Within Southeast Asia, Java, Borneo, and Sumatra were among the better collected areas. 1967 by the senior author. Field specimens and leaf and bark materials for moist chamber cul­ Collections from these places were examined by ture were mailed from Thailand to the Univer­ many foreign workers (Berkeley and Broome, sity of Texas under USDA labels. No actual 1876; Boedijn, 1927, 1928, 1940; Cesati, 1874; specimens were collected in Burma; the moist Emoto, 1931 a; Fischer, 1907; Hohnel, 1909; chamber material was hand-carried back. Junghuhn, 1838; Moritzi, 1845; Overeem and

3 We examined 34 Myxomycetes collected in Singa­ 1 Supported by National Science Foundation grant pore and the lower Malay Peninsula wh ich are now in GB·68I2x to C. ] . Alexopoulos and by a grant-in-aid the Singapore Botanical Garden (SING) and found from the Society of the Sigma Xi to D. R. Reynolds. them to be dilapidated. Many of these are dated, anno­ Manus cript received May 15, 1970. tated, or both, so that they could have been the basis 2 Department of Botany, The University of Texas at of some of the Malayan citations or includ ed in early Austin, Aust in, Texas 78712. monographs. 33 34 PACIFIC SCIENCE, Vol. 25, January 1971

The nomenclature to be used in this report tion, The reticulate pattern of the peridium is follows Martin and Alexopoulos (1969). Un­ evident with magnification of 50X. It is not so less otherwise noted, the Thailand localities are distinct as the original illustration might suggest referred to in the following manner: A, Khao for the recessions between the anastomosing Yai National Park; B, Khaopang, Namtok; C, ridges scarcely show any shade of difference in Namtok, Tabkawg, Gankoy, Srabury. The Bur­ color. In a lactophenol mount these ridges flatten mese material was all taken from Dalhouse Park, out, suggesting that they were formed by a Royal Lake, Rangoon. shrinkage of the peridium. The spores of the The specimens examined are cited after each Thai material are smaller (9-1°11) than those locality in which they were collected by the col­ of the Indian type (13 .5-1511)' Our specimen lection number of Don R. Reynolds (DRR). An thus approaches L. operculate, but because of a asterisk following a number indicates that the complete lack of operculum we tentatively assign specimen was developed in moist chamber cul­ it to L. erecta. ture according to Gilbert and Martin (1933). Family Representative specimens are deposited in the University of Texas Myxomycete Collection, BPI Genus A dans. Fam. PI. 2:7.1763 and NYBG. 1. Lycogala exigtmtn Morgan. Thailand. A, 1258 ; C, 1209 Class MYXOMYCETES The sporangia in the Khao Yai collection are Subclass 1. CERATIOMYXOMYCETIDAE 1-2 mm; the spores are reticulate, 4.5-:-5.411; the Order CERATIOMYXALES capillitium densely spiny without inflations at Family CERATIOMYXACEAE the ends or at intervals as in L. epidendrum. The Genus Schroeter in Engler and external appearance of the Namtok collection is Prantl, Nat. Pfi. 1(1) :16. 1899 more nearly like that of L. epidendrum. How­ 1. Ceratiom yxa [ruticulos« (Miill.) Macbr. ever, in both collections the sporangia bear Thailand. A, 1200, 1221, 1231, 1234, blackish scales that are internally divided into 1235, 1237, 1170, 1247; B, 1266; C, numerous chambers. 1172, 1204, 1207 Martin (1967) writes that, "L. exigunm does The spores in some collections are 9-11 X suggest a small dark phase of L. epidendrum:" 5.4-6.311. The normal range is 10-13 X 6-711. He concludes that the smaller, darker fruit body Ceratiomyxa sphaerosperma Boedijn known containing slender often hyphalike pseudocapil­ from Indonesia and the neotropics was not litium and smaller, less strongly reticulate found. spores in addition to the distinct but variably tesselate warts, separates the former species from Subclass 2. L. epidendrum. Order Family LICEACEAE Family Cribrariaceae Genus Schrad. Nov. Gen. PI. 16. 1797 Genus Pers. Neues Mag. Bot. 1: 91. Other than the citation by Penzig (1898) for 1794 Licea variabilis Schrad. in Bogor, there are no 1. Cribraria microcarpa (Schrad .) Pers. Thai­ prior records of Licea from tropical Southeast land . A, 1251, 1260 ; B, 1174,1175,1205, Asia. Moist chamber work should yield many 1213 more records. In two collections, the cup is lacking. The 1. Licea biforis Morgan. Thailand. C, 1312* stalk of DRR 1175 is about 6/7 of the total 2. Licea erecta Thind and Dillon. Thailand. height, the dictydine granules are pulvinate and C, 1345* pale ; the cup is completely lacking. This collec­ This species was described in 1967 from the tion exhibits characteristics common to C. lan­ foothills of the Eastern Himalayas. Our collec­ gtlescens, C. pachydictyon, and C. microcarpa. tion comes from a forested area in central Thai­ Some sporangia of DRR 1213 have no peridial land. The wood on which the sporangia cups; others have definite, but small, cups re­ developed was not bamboo, as in the type collec- sembling C. languescens. Southeast Asian Myxomycetes-REYNOLDS AND ALEXOPOU LOS 35

2. Cribraria violacea Rex. Thailand. A, 1259; 3. Hemitricbia stipitata (Massee) Macbr. B , 1297* Thailand. A, 1220, 1230; B, 1267; C, 1173, 1202 Genus Dictydium Schrad. Nov. Gen. PI. 11. H emitricbia clavata (Pers.) Rost. is included 1797 in the Indonesian and Malayan lists. Martin 1. Dictydium cancellatum (Batsch) Macbr. (1949) states, "reported from the tropics, but Thailand. C, 1180 all tropical collections examined have proved to be H. stipitata:" This was our experience with Order these collections and other specimens from the Family ECHINOSTELIACEAE vicinity of Kuala Lumpur (DRR 1149, DRR 1145, Genus Ecbinosteliam de Bary, in Rost. Versuch DRR 1160) as well as with recent collections in 7. 1873 the same area by Dr. A. Nawawi of the Univer­ 1. Ecbinostelis m minutum de Bary. Thailand. sity of Malaya. A , 1296*; C, 1345* Genus Pericbaena Fries, Symb. Gast. 11. 1817 This is the first known collection of any 1. Pericbaena chrysosperma (Currey) Lister. echinosteliaceous species from tropical Asia. E. Thail and. C, 1295* minutum is known from the neotropics and from Australia as well as Europe and North Order STEMONITALES America. We also have a specimen in our collec­ Family STEMONITACEAE tions (UTMC-2102) collected in Malaya by Dr. Genus Clastoderma Blytt, Bot. Zeit. 38:343. A. N awawi in 1969. 1880 1. Clastoderma debaryanu m Blytt. Thailand. Order C, 1346* Family TRICHIACEAE Genus Comatricha Preuss, Linnaea 24: 140. Genus Wiggers. Prim. Fl. Holsat. 109. 1851 1780 1. Comatricha elegans (Racib.) G. Lister. 1. Arcyria cinerea (Bull.) Pers. Th ailand. A , Thailand. A, 1227 1228*, 1232, 1257, 1308*; C, 1181, 2. Comatricha laxa Rost. Thailand. A, 1225 1203; Pak Thong Chai, EGSDF 13. Burma. 3. Comatricha longa Peck. Thailand. A , 1226; 1309* Pak Thong Chai, EGS 117 Both the very minute globose form with 4.Comatricha tenerrimn (M. A. Curtis) G. sporangia only 0.1 mm in diameter and the typi­ Lister. Thailand. A, 1256 cally tropical digitate form are present among The only records from Asia of this species are these collections. from Japan. 2. Arcyria dent/data (L)W ettst. Thailand. 5.Comatricha typhoides (Bull.) Rost. Thai­ A, 1244; C, 1171, 1183, 1184; Pak land. A, 1223, 1235; C, 1217 Thong Chai, EGS GL 9 DRR 1235 is variety beterospors Rex. The 3. Arcyria insignis Kalch. and Cooke. Thai­ spores measure 5.5-6 !l and are finely reticulate land. C, 1200 in addition to exhibiting the characteristic wart 4. A rcyria magna Rex. Thailand. C, 1199 clusters. This and our collection from the Philippines 6. Comatricha sp. Thailand. C, 1179 (DRR 886) constitute the only records outside Th is specimen approaches C. subcaespitosa in the western hemisphere of this uncommon spe­ its general morphol ogy. The stalk is % to Y2 the cies. total height ; the capillitium is intricately branched and forms an almost complete net at Genus H emitricbia Rost. Versuch 14. 1873 the surface; the columella reaches almost to the 1. H emitricbia intorta (A . Lister) A. Lister. summit. Th e spores, however, measure uni­ Burma. 1352* formly 6-6.5 !l diameter which is much too 2. H emitricbia serpul« (Scop.) Rost. Thai­ small for that species and more like those of land. A, 1224 C. typh oides in size. They are, however, darker 36 PACI FIC SCIENCE, Vol. 25, January 1971

and uniformly punctate, lacking the clusters of Genus Physarella Peck. Bull. Torrey Club 9 :61. warts characteristic of that species. 1882 Genus Diacbea Fries, Syst. Orbis Veg. 143. 1. Physarella oblonga (Berk. and Curt.) Morgan . Thailand. C, 1313* 1825 1. Diacbea radiata Lister and Petch. Thailand. Genus Physarum Pers., Neues Mag. Bot. 1:88. A, 1242 1794 9-10 The spores are fl and appear distinctly 1. Physarum cinereum (Batsch) Pers.Burma. warted under the oil immersion lens. The perid­ 1348* ium is bronze on some sporangia and com­ The spores are 8-9 fl and appear distinctly pletely iridescent blue on others. The stalk and and closely warted under the oil immersion lens. columella are of calcareous nodules enclosed in The warts are dark brownish, rather than clear a hyaline membrane layer. The capillitium and violaceous. The calcereous nodes are small, peridium originate from this layer. When the roundish. The peridium is a calcareous, bluish stalk is broken, the calcareous nodules spill out. grey, rugul ose layer. Fruit bodies range from There is no continuous calcareous hypothallus in scattered individual sporangia 2.5 mm in diam­ the collection as recorded by Martin and Alex­ eter to plasmodiocarps 1-3 mm in length , these opoulos (1969) from a Panamanian collection, often flattened into sheets. but rather there is a circular shaped hyaline ex­ tension of the stalk's outer sheath. It must be noted that if not matured properly fructifications of several physaraceous species Genus Lamproderma Rost. Versuch 7. 1873 produced in culture tend to resemble closely 1. Lamproderma arcyrionema Rost. Thailand. those of P. cinereum and that, unless several A, 1236, 1239; Pak Thong Chai, EGS DF generations are cultured, there is always danger 20, 108 of misidentification. 2. Physarum compressum Alb. and Schw. Genus Stemonitls Roth. Mag. Bot. Romer and Thailand. A, 1278* Usteri 1(2) :25 . 1787 Most spores are 9-10.8 fl, but some are ovoid, 1. Stemonitis fusca Roth. Thailand. A, 1240 ; 9 XII.7 The spore wall appears to be double C, 1217 fl. as indicated by a distinct line in the approximate 2. Stemonitis herbatica Peck. Thailand. A , middle of the I-u-thick wall. The spore warts 1195 This species occurs on bark instead of the are quite evident and evenly distributed. usual habitat of living herbaceous plants. It 3.Physam m flavicomum Berk. Thailand. A, could possibly be referred to S. flavogenita. 1249, 1347* ; C, 1189 3. Stem onitis splendens Rost. Thailand.Boon­ 4. Pbysaram globulifertlm (Bull.) Pers. song Lekagul Clinic, 6 Custom House Burma. 1273* Lane, Bangkok, 1262 5. Physamm melleum (Berk. and Br.) Mas­ see. Thailand. A, 1276*; C, 1350*. Order Burma. 1349 * Family PHYS ARACEAE 6. Pbysarum nucieatum Rex. Thailand. C, Genus Erionem a Penzig. Myxom. Buitenz. 36. 1215 1898 7. Physamm nutans Pers. Thailand. A , 1249; 1. Erionema aureum Penzig. Thailand. C, C, 1185 1219 8. Physamm oblatnm Macbr. Thailand. 1322* This fruiting was from a large yellow phan­ The color of the peridium and the nodes is a eroplasmodium. The yellow elongate sporangia creamy coffee rather than yellow. This collection were massed on top of a large fallen log. The has sporangia that have a flattened central lime elastic capillitium has very few nodes; the spores knot as mentioned by Lister (1925) . The stalk measure 6-7 fl and, in addition to being mi­ is darker than the species description suggests. nutely punctate, bear distinctive warts in loose However, specimens on hand determined by clusters. G. W . Martin have a black furrowed stalk. Southeast Asian Myxomycetes-REYNOLDS AND ALEXOPOULOS 37

9. Physamm rigidum G. Lister. Thailand. Pak land. Acknowledgment is made to the Burmese Thong Chai , EGS National Airline for courtesies during an ex­ 10. Pbysarum tenerum Rex. Thailand. C, tended stay in Rangoon. 1176 11. Pbysarum oiride (Bull.) Pers. Thailand. LITERATURE CITED B, 1263; C, 1176, 1188, 1189, 1192, 1197, 1198 BERKELEY, M. J., and E. C. BROOME. 1876. Supplement to the enumeration of the fungi Family of Ceylon, Java. Transactions of the Linnean Genus Diderma Pers. Neues Mag. Bot. 1 :89. Society of London (Botany), vol. 15, pp. 82­ 1794 86. 1. Diderma bemispbericum (Bull.) Hornem. BOEDIJN, K. B. 1927. von Sumatra. Burma. 1272* Miscellanea Zoologica Sumatrana, vol. 17, In some fruitings the sporangia were sessile pp. 1-3. and confluent into plasmodiocarplike forma­ --- 1928. Verzeichnis der von Sumatra be­ tions. kannten Mycetozoa. Annales mycologici, Ber­ lin, vol. 26, p. 450. Genus Didymium Schrad. Von. Gen. PI. 20. --- 1940. The Mycetozoa, fungi and lichens 1797 of the Krakatau group. Bulletin du Jardin 1. Didymium clavus (Alb. and Schw.) Racib. botanique de Buitenzorg, series III, vol. 16, Thailand. A, 1241; C, 1210 2. Didymium iridis (Ditmar) Fries. Thailand. pp. 358-429. Pak Thong Chai , EGS DF 13. Burma. CESATI, V. 1874. Mycetum in itinere Borneensi. 1319*, 1353* Hedwigia, vol. 13, p. 186 (abs.). DRR 1319 has a black stalk. CHIP, T. F. 1921. A list of the fungi of the 3. Didymium squamulosum (Alb. and Schw.) Malay Peninsula. Gardens' Bulletin, Straits Fries. Thailand. A, 1314*, 1317* Settlements, Singapore, January 7, vol, 2, pp. 311-418. Genus Physarina von Hohnel, Sitz. Akad. Wien. EMOTO, Y. 1931 a. Javanische Myxomyceten. 118:431. 1909 Bulletin du Jardin botanique de Bruitenzorg, 1. Physarina ecbinocepbala Hohnel. Thailand. series II, pp. 161-164. A, 1351 * --- 1931 b. The Malayan Myxomycetes. The fruit body is pure white. The spores are Journal of Botany, British and Foreign, Lon­ 9!l and minutely but distinctly warted under don, vol. 69, pp. 38-42. 1,000X. This species was heretofore known FISCHER, E. 1907 . Ueber einige von Herrn. only from Java. Prof. E. Kissling in Sumatra gesammelte Pilze, Mittheilunge der Naturforschenden Ge­ SUMMARY sellschaft in Bern, 1906, pp. 109-123. GILBERT, H. and G. W. MARTIN. 1933. In this first part of a report of Southeast Asian c., Myxomycetes found on the bark of living Myxomycetes, 42 species from Thailand and 7 trees. Studies in Natural History, Iowa Uni­ from Burma are listed. Bcbinostelium minutum versity, vol. 15, pp. 1-8. is the first member of the Echinosteliaceae to be reported in print from the Asian tropics . Phy­ HElM, R. 1962. Contribution ala flore rnycolo­ sarina ecbinocepbala and Licea erecta were pre­ gique de la Thailande. 1. Revue de Mycologie, viously known only from the type localities. vol. 27, pp. 124-158. HOHNEL, F. VON. 1909. Fragmente zur Mykol­ ogie. Javanische Myxomyceten. Sitzungs­ ACKNOWLEDGMENTS berichte der kaiserl. Akadamie der Wissen­ Appreciation is expressed to Thep Lekagul schaften in Wien. mathem.-naturw. Classe, and the other members of Dr. Boonsong Leka­ vol. 118, pp. 427-442. gul's family for invaluable assistance in Thai- JUNGHUHN, F. W. 1838. Praemissa in Floram 38 PACIFIC SCIENCE, Vol. 25, January 1971

Cryptogamicam Javae insulae. Fascicules 1. landisch Ost-Indien bis dem jahre 1920 ge­ Java, pp. 1-86. fundenen Myxomycetes, Fungi und Lichenes. LISTER, A. 1925. A monogr aph of the Myceto­ Bulletin du Jardin botanique der Buitenzorg, zoa. 3rd ed, revised by G. Lister. London, series III, vol. 4, pp. 1-146. British Museum (Natural History). xxxii + PENZIG, O. 1898. Die Myxomyceten der Flora 296 pp. , 222 pIs. von Buitenzorg. Flore de Buitenzorg, series II, LISTER, G. 1931. Notes on Malayan Mycetozoa. Leiden, E. J. Brill, pp. 1-83. Journal of Botany, British and Foreign, Lon­ RACIBORSKI, M. 1884. Ueber die javanischen don, vol. 69, pp. 42-43. Schleimpilze. Hedwigia, vol. 37, pp. 50-55. MARTIN, G. W. 1949. Myxomycetes. North RICHARDS, P. W . 1952. The tropical rain forest; American Flora, vol. 1, no. 1, pp. 1-190. an ecological study. N ew York, Cambridge - - - 1967. Lycogala exiguum. Mycologia, University Press. vol. 59, pp . 155-160. SACCARDO, P. A., and G. PAOLETTI. 1888. My­ MARTIN, G. W. , and C. J. ALEXOPOULOS. 1969. cetes Malacenses. Atti del reale Instituto Ve­ The Myxomycetes. Iowa City, University of neto di Scienze, series 6, vol. 6, pp. 387-428. Iowa Press. 560 pp ., 41 col. pls, SANDERSON, A. R. 1922. Notes on Mal ayan MORITZI, A. 1845. Systematisches Verzeichniss Mycetozoa. Transactions of the British Myco­ der von H. Zollinger in den Jahren 1842­ logical Society, vol. 7, pp. 239-256. 1844 auf Java gesammalten Pflanzen vebst ZOLLINGER, H . 1844. Observationes phyto­ einer kurzen Beschreibung der neuen Gattun ­ graphicae praecipue genera et species nova zen und Arten. Solothu rn. nonnull a respicientes. N atuur-en geneeskun­ OVEREEM DE HASS, V., and c: VAN OVEREEM dig. Archief voor Nederlandsch -Indie, vol. 1, DE HASS. 1922. Verzeichnis der in N ieder- pp. 372-405.