A PRELIMINARY SURVEY of the FUNGUS FLORA of CAVES. I• AQUATIC PHYCOMVC!TES • ., MARIARET Mlrihall HOLLAND
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A PRELIMINARY SURVEY OF THE FUNGUS FLORA OF CAVES. " I. AQUATIC PHYCOMYCETES. BY MARGARET MARS'.1ALL HOLLAND /.I THESIS SUBMITTED TO THE GRADUATE FACULTY OF THE VIRGINIA POLYTECHNIC INSTITUTE tN CANDIOACY FOR THE DEGREE OF MASTER OF SCIENCE IN BIOLOGY, BOTANY OPTION APPROVED: APPROVED: DEAN OF AGRICULTURE ·~AJOR PROFESSOR MAY 15, 1958 BLACKSBURG, VtRGINIA I I PREFACE A SYSTEMATIC INVESTIQATION OF THE AQUATIC FUNQI OCCURRING IN THE SOILS OF VIRGINIA HAS NEVER BEEN MADE. TH~RE AR! A FEW RECORDS OF FUNGI COLLECTED HERE SY VISITING MYCOLOGISTS, BUT NO INFORMATION IS AVAILABLE CONCERNING TH£ HABITS, PREVALENCE, ANO DISTRIBUTION OF THESE ORGANISMS IN THE STATE. THE PRESENT CONTRIBUTION CONSIDERS BUT ONE AMONG MANY HABITATS OCCUPIED BY THE AQUATIC PHYCOMYCETO·us FUNGl 1 AND IS A PART OF A MORE COMPREHENSIVE INVESTIOA- TION, UNDER THE DIRECTION OF PROF• WtLLIAM W. SCOTT, OEALINQ WITH THE TAXONOMY, MORPHOLOGY, ANO DISTRIBUTION OF THESE FUNGI. THE WRITER HAS BEEN PARTtCULARLY fNTERESTEO IN THE CHYTRIDtACEOUS 1 BLASTOCLADIACEOUS 1 SAPROLEQNfACEOU8 1 AND PYTHIACEOUS SPECIES OCCURRING IN THE SOIL ANO WATER OF CAVES IN SOUTHWESTERN VIRGINIA• ALTHOUGH IT IS NOT POSSIBLE TO ACKNOWLEDGE THE NUMEROUS INDIVIDUALS WHO HAVE CONTRIBUTED TO THIS fNVESTtGATION, THE AUTHOR WISHES TO EXTEND HER SINCERE GRATITUDE TO ALL THOSE WHO HAVE ASSISTED IN ANY WAY TOWARD THE COMPLETION OF THIS STUDY. SINCERE APPRECIATION IS EXPRESSED TO THE MEMBERS OF THE COMMITTEE UNDER WHOSE DIRECTION THIS INVESTIGATION WAS CARRIED OUT. THESE INCLUDED PROF. W. W. SCOTT, CHAIRMAN; DR. t. 0. WILSON, I J I HEAD, DEPARTMENT OF BIOLOGY (RETIRED); OR. F. S. ORCUTT, HEAO, DEPARTMENT OF BIOLOGY; OR. A. B. MASSEY; OR. K. W. KING; AND DR. 0. E. NORBY. THE WRITER IS ALSO VERY GRATE- FUL TO MRr LARRY GRIFFIN FOR HIS ASSISTANCE IN THE COLLECTION OF SAMPLES, ANO TO THE MANAGEMENTS OF THE FOLLOWJNQ CAVERNS FOR PERMISSION AND AID IN COLLECTING MATEP.IALSt GRAND CAVERNS, SHENANDOAH CAVERNS, LURAY CAVERNS, SKY LINE CAVERNS, ANO ENDLESS CAVERNS. APPRECIATtON IS ALSO EXTENDED TO THE VtRGINIA ACADEMY OF SCIENCE FOR FINANCIAL ASSISTANCE, WHICH ENABLED THE WRITER TO !XTEND THE SCOPE OF THIS tNVESTIGATION. THE WRITER WISHES TO ACKNOWLEDGE THE ASSISTANCE OF OR• PAUL Me PATTERSON, HOLLINS COLL£G£ 1 FOR THE tDENTIFICATtON OF THE BRYOPHVTES AND DA. A. B. MASSEY FOR THE tOENTIFICATION OF THE FERNS• IV TABLE OF CONTENTS INTRODUCTION. • • • ••••• • •• • • • • • • • • • LITERATURE REVIEW • •• • •••• • • • • • • • • • • 5 MATERIALS ANO METHODS • • • • • • • • • • •• • • • • 10 FUNGI COLLECTED • • • •• • •• • • • •• • • • • • • t3 CHYTR llH ALES • • • • • • • • • • • • • • • • • • 13 I. NOWA~OWSKIELLA RAHOSA E. J. BUTLER. • • • • • f3 2. ? RHtZOPHLVCTIS SPP. • • • • • • • • • • • • 14 BLASTOCLADtALES• • • • • •••• • • • • • • • • 15 3, ALLOMVCES CYSTOQ!NUS EMERSON. • • • • • • • • 15 4. BLASTOCLAOIOPSIS PARVA (WHtFFEN) SPARROW. • • f6 SAPROLEGNtALES • • • • • • • • • • • • • • • • • JS 5. ACHLVA OUStA COKER. • • • • • • • • • • • • • 18 6. APHANOMYCES ~AEVIS DEBARY • • • • • • • • • • 19 7. SAPROL£QNIA FER~X (GRUtTH) THURET • • • • • • 2f 8. SAPROLEGNIA MIXT~ DEBARY. • • • • • Ill • • • • 22 LEPTOMITALES • • • • • • • • • • • • • • • • • • 24 9. APODACHLYA MtNtMA COK£ft AND lCtTNER • • • • • 24 lAGENIDIALES • 111 111 • • • • • • • • • • • • • • Ill 25 10. O~PtDtOPStS APHANOMYCIS CORNU. • • • • • • • 25 ti. 0LPIDtOPSIS SAPROLEQNIAE (BRAUN) CORNU • • • 26 PERONOSPORALES • • • • Ill • • • • • • • • • • •• 28 12. PYTHtUM AFERTILE KANOUSE AN~ HuMPHREVe • • • 28 • 13. PYTHIUM CAROLINIANUM MATTHEWS. • • • • • • • 29 14. PYTHf UM IRREOULARE 8Ul8MAN • • • • • • • • • 30 v 15. PYTHIUM MONOSPERMUM PRtNeSHEIM. • • • • • • • 32 16. PYTHIUM NA861t ITO AND TOKUNAQO • • • • • • • 34 17. PYTH!UM PAPILLATUM MATTHEWS • • • • • • • • • 35 f 8. PYTHIUM PULCNRUM MINDEN • • • • • • • • • • • 37 19. P'ITHIUM ROSTR6TUM BUTLER• • • • • • • • • • • 39 20. PYTHIUM VEXAN! OEBARY • • • • • • • • • • • • 40 SUMMARY ••• • •• • •••• • • • • • • • ••• • • • 42 BfBLIOGRAPHY • • • • • • • • • • • • • • • • • • • • • 43 VITA • • • • • • • • • • • • • • • • • • • • • • • •• 50 INTRODUCTION ALTHOUGH THE FAUNA OF CAVES HAS BEEN STUDIED tNTEN- 81VELY1 RELATIVELY LITTLE ATTENTION HAS BEIN QtVEN TO THE PLANT LIFE PRESENT IN SUCH HABITATS• THll DISCREPANCY 18 DUE, FOR THE MOST PART, TO TH£ DEPENDENCE OF CHLOROPHYLLOU8 PLANTS UPON LIQHT. StNCE LIQHT ta ESSENTIAL FOR THE QROWTH ANO DEVELOPMENT OF QREEN PLANT8 1 THE ABSENCE OY THESE FROM THE CAYE ENVIRONMENT MIQHT BE ANTICIPATED. A FEW MEADER REPORTS ARE FOUND IN THE LITERATURE, HOVEY£R 1 CONCERNING THE PRESENCE OF CERTAIN ALGAE, MOSSES, AND LlVERWORT8 F~UNO IN CAVE8e THESE GENERALLY GROW NEAR THE CAVE'S ENTRANCE OR IN THE VICINITY OF ARTIFICIAL ILLUMINATION, AND MAY NOT BE REGARDED AS TRUE CAYE INHA8fTANT8• MJ88 HARRING (1930), FOR EXAMPLE, HA8 REPORTED THE FOLLOWING BRYOPHYTES FOUND GROWING NEAR ELECTRIC LIGHTS IN H'WE CAVERNe: MARCHANTl6 POLYMORPHA, AMPHIDIUM MONGEOT!I, AMILY!TEJIUM JURALZKANUM, 8RACTYTHECIUM RUIABULUM, BRYUM C6E8PITICU~, 8RYU~ RAPILLARE, FUNARI! HYGROMETR!CA1 AND LIPTOBRYUM PYRIFORME• IN ADDITION, SCHl8T08TJGIA 08MUNDAC§A HA8 BECOME TH! CLASSICAL EXAMPLE OF A GREEN PLANT STRUCTURALLY ADAPTED FOR GROWTH ANO DEVELOPMENT IN THE 8UIDUED LIGHT OF CAVEi (VEROOORN 1 1932). THE HIGHER CON- CENTRATION OF CARBON DIOXIDE IN CAVES 18 APPARENTLY ONE OF THE FACTORS MAKING tT POSSIBLE FOR MOSSES TO 8ROW UHDIR THE8E CONDITION&. COINCIDENTAL WITH THE COLLECTION OF -2- MATERIAL FOR THIS STUDY, THE FOLLOWtNQ BRYOPHVTES WER! COLLECTED ANO IDENTtFIEDS FtSSIDEN! MINUTULUS SULL., LEPTODICIYUM RIPARIUM (HEDW.) WARNST., THUIDIUM VIRQINIANUM (BRIO.) LINOB., EURHVNCHIUM 8ERRUL!TUM (HEow.) KINDe •• ANO A&TOMUM MUHLENBERQIANUM (sw.) GROUT. ALSO FOUND. NIAR AN ELECTRIC LIGHT IN LURAY CAVERNS, WAI THE COMMON WOODLAND FERN 0RYOPTERl8 tNTERMEOIAe As MIGHT BE EXPECTED SUCH 8PECtMENS ARE GREATLY ETIOLATED AND POORLY DEVELOPED OFTEN TO THE EXTENT OF MAKINQ IOENTtFICATION DIFFICULT. SEED- ~tNG8 OF CERTAIN HIGHER PLANTS MAY SOMETIMES BE FOUND IN TH£ REMOTE AREA8 OF CAVES• THESE ARE REMARKABLE FOR THE EXTREME DEGREE OF ELONGATION EXHfBtTEO. SUCH SEEDLINGS DIVELOP APPARENTLY FROM QERMINATINQ 8EEDS CARRIED •NTO THE CAVES BY AIR CURRENTS OR BY ANIMALS, GROW AT THE EXPENSES OF THEIR STORED FOOD, THEN WITHER AND DIEe SINCE MANY OF THE FUNGI CAN COMPLETE THEIR DEVELOP- MENTAL CYCLES tN THE ABSENCE OF LIGHT, IT MIGHT BE EXPECTED THAT SUCH SPECIES WOULD THRIVE IN CAVE8• IN THE CASE OF CERTAIN OTHER FUNGI, HOWEVER, LtQHT EXERTS A DECIDED MORPHOQElltC OR A PHOTOSTATIC EFFECT OR, IN SOME CASE8 1 MAY BE REQUIRED FOR THE FORMATION OF REPRO- DUCTIVE STRUCTURE&. IT WOULD APPEAR LIKELY THAT SUCH SPECIES WOULD BE ENTIRELY ABSENT FROM THESE HABITATS OR 1 IF PRESENT, WOULD BE ABORTIVE OR STERILE. PREVIOUS tNVESTIQATORS HAVE NOTED SUCH ABORTIVE FRUITING STRUCTURES -3- AND OTHER REPORTS HAYE INDICATED THE FREQUENT PRESENCE IN CA¥E8 OF STERILE MYCELIA• FOR MANY SAPROPHYTIC FUNQt, HOWEVER, THE ENVIRONMENTAL CONDITIONS FOUND IN CAVES WOULD S!EM TO BE IDEAL• THE UNIFORMLY HIGH HUMIDITY, TEMPERATURE OF s2-sso F., AND THE NEUTRAL TO 8Ll8HT ALKALINITY OF THE SOIL AND WATER ARE ALL FAVORABLE CONDITIONS FOR THE DEVELOP- MENT OF MANY FUNQt. THE ABSENCE OF LIGHT AND THE RESTRICTED SUPPLY OF AVAILABLE NUTRIENTS MfQHT BE EXPECTED TO LIMIT THE VAAICTY OF FUNGI IN CAVES. NUMEROUS ACCOUNTS HAVE 8£~R PUBLISHED CONCERNING THE FUNGI FOUND IN MfNE8 1 TUNNELS, CELLARS, AND SIMILAR MAN- MADE HA8fTAT8• THE ORGANISMS ENCOUNTERED IN THESE HYPO•EAL ENVIRONMENTS DO NOT NEOE88ARILY REPRESENT THE KINDS OF FUN4t WHICH ONE WOULD EXPECT TO FIND IN NATURALLY OCCURRING CAVES. CAVERNICOLOU8 FUNGI MAY BE DIVIOED 1 FOR CONVENIENCE, INTO GROUPS. THOSE FUNGI, FOR EXAMPLE, WHICH OCCUR NORMALLY ta CAVES AND THRtVE THEREIN BECAUSE OF FAV0RA8LE ENVIRONMENTAL CONDITION8 MAY BE TERMED "TROQLO- PHfLE8• (TROQLO • CAVE8 1 PHILEO •LOVE). THE SPECJES WHICH ARE INTRODUCED INTO CAVES AND EXIST THERE ONLY TEMPORARILY MAY BE TERMED HTROQLOXEM!S" (TROGLO • CAVE, XENO! • GUl8T). THE LATTER ARE FOUND MOST FREQV!NTLY ON TtMBER, DISCARDED FOOD, AND OTHER REJECTAMENTA INTRODUCED BY EXPLORERS OR ~EFT BEHIND BY CASUAL VISITOR&. -4- THE PRESENT INVESTiCATtON IS CONCERNED PRIMARILY WITH THE TROQLOPHflttS 1 THOSE FUNGI OCCURRING HATURALLV tN THE SOIL AND WATER OF' CAV!!S. Tut WRITER IS PARTICULARLY fNTERESTED tN THE AQUATIC PHYCOMYC!TES 1 HENCE CONSfOERABLE ATTENTION HAS BEEN GIVl::N TO THESE FUNQl ounrnn THE PRESENT INVESTtGATION• -&- LITERATURE REVIEW OUR KNOWLEDGE OF CAYERNICOLOUS FUNGI IS LARGELY THE RESULT OF INCIDENTAL 098ERVATtON8 AND COLLECTIONS MAD£ BY QEOLOQleT8 AND ZOOLOGl8T8• THE POSITIVE tOENTIFtCATfON8 OF FUNGI FROM THE INTERIOR OF CAVES ARE FEW• Mo8T OF THE FUNGI WHICH HAVE BEEN REPORTED ARE 09VtOU8LV TROQLOXENE&. THESE INCLUDE WOOO-ROTTINQ FUNGt 1 COPROPHILOUS SPICIES 1 ANO OTHER SAPROPHYTIC 8PECtES QROWtNQ ON DISCARDED MATERIALS AND ORIQtNATtNG FROM AIR-BORNE SPORES• IT 18 PROBABLE, HOWEYER 1 THAT THE PARASITIC FUNGI FOUND tNF!CT- tNQ CAYE ANIMAL& ANO THOSE SAPROPHYTES OCCURRINQ IN THE 90tL AND WATER ARE TRULY TROILOPHILE8• AMONG THE EARLY RECORDS IN THE LITERATURE ARI SEVERAL WHICH APPEARED BEFORE THE TURN OF THE CENTURY• ACCORDING TO KYRLE (1923), FRtE8 DESCRl8~D IN 1874 FOUR ~UNQt FROM ·FALKEN8TEINERH8HLE. THESE INCLUDED "YELLOW-BROWN MYCELIA . OF RH!ZOMORPH!, A FELT-LIKE TUFT OF lg~TOMITUB LACT§U8, MMCOR MUCEOO QROWtNQ ON BAT DUlll, ANO THE FUNQU8 MXXOTRICHELLA SPELAEA•" HOVEY (1879) COLLECTED FUNQI FROM LURAY CAVERNS AND OESCRaBED ONE OF THE BLACK MOLDS A8 A NEW 8PECIE81 MUCOR