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 STALACTITts. THta SPECIES •• NOT
INCLUDED ON SACCARDo•s "SYLLOQE FUNGORUM" AND HAS NOT 8!£N
RECOGNtZEO AS A VALID SPECIES•
PACKARD (1879) COLLECTED SEVERAL FUNGI FROM MAMMOTH
CAVE WHICH WERE IDENTIFIED BY THE LATE PROFESSOR ROLAND -1-
THAXTER OF HARVARD UNIVERSITY. THESE INCLUDED AN ISOLATE
REFERRED TO AS 0ZONIUH AUBANTIACiUM ltNK, AND ISOLATES DESCRIBED A8 "A REDDISH BUTTON•SHAPED FUNQU81 A GREEN MOLD,
A LONG WHITE MOLD, AND A FUNGUS QROWtNQ ON THE HIND BODY OF
A CAVE CRICKETTe" IN THE SAME PAPER, PACKARD MENTtONEO AN
UNIDENTIFIED 8PECt!a OF PEZtZA AND A COLORLESS AGARIC FROM
WEYER'8 CAVE.
IN A SERIES OF TWO PAPERS CALL (t879 A, 1879 B) PU8Ll8H£D A LIST OF FUNGI COLLECTED IN MAMMOTH CAVE,
KENTUCKY. THESE WERE ALSO IDENTIFIED BY PROFESSOR THAXTER
OF HARVARD UNIVERSITY AND INCLUDED THE FOLLOWING SPECIES: COPRtNUS MICACEUS BULL., FOMES (POLYPORUS) APPLANATUS PERS., RH!ZOMORPHA MOLINARI& (?), MtCROASCUi b0NOIR08TRl8 ZUKAL,
LASMIDIUM CELLAR! FR., GYMNOASCUS SETOSUS EtDAM 1 !• UNCtNATU8
EIDAM 1 TWO ADDITIONAL SPECIES OF GYMNOA8CUS, A SMALL, REDOISH-
BROWN SPECIES OF P£ZIZA, ISAR!A (SPOROTRICHUM) OENSUM LINK,
THE LATTER GROWING ON THE DEAD BODIES OF CRICKETS (HAOENOA§CU8 ayBTERRANEU8), !• FL!VISSIMUM LINK, lABOULBENtA 8U@TERRANE6
ON ANOPHTHALMUS TELLKAMPFU, A NEW SPECIES OF PAPULOfPORA 1 A NEW SPECIES OF BOUDl£RA1 CARYCHIUM 8TVGJUM, AND MUCOR MUCEDO LINN.
FOLLOWING A VtStT TO THE UNITED STATE8 1 MAHEU (1926) PUBLISHED A LIST OF Hl8 EXTENSIVE COLLECTIONS FROM THE
KENTUCKY CAVERNS INCLUDtNG THE FOLLOWING CAVERNICOLOU8
FUNGI: COPRINU8 MICACEUI BULL., FLAMMULA 8P. 1 MVCENA sP., -1-
PAX!LLU8 PANUOIDES FR., CORlOLUS ELONQATUS BERK., RAOULUM
.!f!•• TRAMETES ODORATA (WULFe) FR., PEZIZA §!.•, ISARIA OENSA
ltNK GROWING ON THE CADAVERS OF H~DEHAECUS SUBTERRANEUS, AND
MYCELIAL STAGES BELONGING TO THE GENERA Svaaua, RHt%OMORPHA.
AND 0ZONtUM• MORE RECl?NTLV, BAILEY (1933) NOTED THE
PRESENCE OF M,aRASfOUfl flGRIPES SoHw., COPRINUS ATR!MENTARIUS
(BULL.) FR., A SPECIE& OF STEREUM• AND ONE OF THE BLACK
MOLD8 1 ALSO FROM THE CAVES OF KENTUCKYe
FROM HtS STUD1ES OF CAVE LlFE IN HUNQARV 1 GABHARDT
(1932) MADE REFERENCE TO ONLY TWO SPEC1ES OF FUNGI, THE
DISCOMYCETE, HYALINA FASCICULARE AND THE AGARIC 1 COPRINUS ATRAMENTARIUS (BULL.) FR.
WOLF (1938) S80LATED FROM SOIL SAMPLES, BAT EXCREMENT,
AND FRA88 FROM LEAF-CUTTING ANTS, COLLECTED FROM CAVES IN
YUCATAN, MEXICO, THE FOLLOWING FUNGI: AsPERGtLLUS CLAVATUa
DESM. AND PENICILLIUM 8Pe FROM BAT DUNQ 1 AND PyTNIUM SP.
FROM DAMP ROCK 8CRAPINQ8•
A LARGE CAVE SYSTEM tS USUALLV WELL VENTILATED ANO
AIR-BORNE SPORES OF FUNGI MAV BE CARRIED TO THE MOST REMOTE
AREAS OF THE CAVEe SUCH SPORES, IF DEPOSITED ON A SUITABLE
SUBSTRATUM, WILL GROW. THIS ACCOUNTS, PERHAPS, FOR THI ORfQIN OF THE FUNGUS OE&CRIBEO BY BtRO (1949) QROWINQ ON CANDLE IREA8E IN AN ENGLISH CAVE ANO tOENTIFIED AS
PENICILLIUM fTALICUM W£HM£R. AN EARLIER COLLECTION FROM ~-
THE 8AME CAVE AND FROM SIMILAR 8UB8TRATUM WAS MADE BY
WALTON (1944) WHO PROVED EXPERIMENTALLY THAT THIS FUNQU8
SERVED AS THE SOL! SOURCE OF FOOD FOR THE SPRINOTAIL
COLLEMBOLA• BtRD ALSO RECORDED FROM OTHER ENGLISH CAVES
A8PERGILLUS VER8ICOLOR (VUtLL.) TIRABOSCHI AND AN
UNIQENTIFIED SPECIES OF HELOTIUM. " -
HAZELTON AND GLENNIE (1953) AL80 MENTIONED THE
PRESENCE OF FUNGI IN BRITISH CAVES, ALTHOUGH THESE WERE
NOT IDENTIFIED. ONE TYPE WAS DESCRIBED AS "VERTICAL
QROWTH8 A FEW MILLIMETERS HIGH, ABOUT A CENTIMETER APART,
A FOREST OF MINUTE FUNGI QROWINQ ALL OYER A WIDE BENCH OF
AN OLD PA88AQE ABOUT FOUR FEET ABOVE THE PRESENT FLOOR,•
AND A SECOND ORGANISM, POSSIBLY A FUNGUS, WAS FOUND QROW-
IN8 ON BARE ROCK IN "SMALL CIRCULAR PATCHES, OFTEN WtTH
A YELLOWISH CENTER, SELDOM MORE THAN A CENTIMETER IN DIAMETER•" ANOTHER UNIDENTIFIED QROWTH, APPARENTLY THAT OF A WATER MOLD, WAS FREQUENTLY OBSERVED AT THE EDQE OF
RIM8TON£ POOLS.
Two 8PECfE8 OF CAVERNICOLOUS FUNGI REQUIRE SPECIAL MENTION HERE SINCE THEY HAVE SEEN R£PORTED ONLY FROM CAVES•
BOTH FUNGI WERE DESCRIBED BY PETCH (1934) AS PARASITES OF tN8£CT8• THEii SPECIES ARE HtRSUTELLA DIPTERICA PETCH,
PARASITIC ON TWE DtPTEROUI FLY, LERIA SERRATA, FROM CAVES
IN ENGLAND; AND STtBELLA KERVILLEI (QUEL.) ltNGELSH. 1 -9~
PARASITIC ON THE FIRST NAM£0 FUNGUS. BOTH FUNGI ARE
APPARENTLY THRESHOLD SPECIES, BUT AT TIMES MAY BE FOUND
IN THE tNTERtOR OF CAVES StNCE THE HOST FLV 1 NORMALLY A
THRESHOLD SPECIES, WILL FREQUENTLV MIGRATE DEEPER INTO
THE CAVE.
DR. Te Ce BARR, JR. HAS REPORTED (PERSONAL COMMUNI- CATION) SEVERAL COMMON SPECtES OF FUNGI ON WOOD AND DUNG
FROM CAVES IN TENNESSEE• THESE INCLUDE MUCOR MUC!DOa
PHVCOMVCES NITEN!, XVLARIA POLYMORPH~, A SPECIES OF ISARtA
FOUND ON THE DUNG OF DAVE RATS, AND ~TOMOPHTHORA ON THE
CAVE CRICKET HADENOAECUS $UBTERRANEUSe BARR ALSO NOTED
THAT THE REO BEETLE, PSEUDOAN~PHTHALMUS, MAV BECOME SEVERELY PARAllTIZED av ...... LABOULSENtA --SP. -10 ...
M.l~ TER L'\LS AND METHODS
WtTHttt GILES, CRAIG, ROANOKE, ANO MONTGOMERY COUNTIES
OVER 255 CAVE' HAVE BEEN FOUND. ALL OF THESE ARE IN THE
LIMESTONE BELT OF THE APPALACHIAN VALLEY. SUCH CAVES
OCCAStONALLY CONTAIN SMALL BODltS OF STANDING WATER OR
UNDERGROUND STREAMS AND, I~ NOT, THE S01L FOUND THEREIN
13 QUITt WET. THIS 13 'TRUE EVEN DURING THE SUMMER MONTHS
WHEN THESE CAVES WERE SAMPLES. ALTHOUGH MOST OF THESE
CAVES HAYE BEEN KNOWN FOR YEARS, THEY ARE VISITED ONLY
BV SPELEOLOGISTS INTERESTEO IN THE EXPLORATION AND
SCIENTIFIC STUDY OF SUBTERRANIAN HABITATS. SUPERSTITION,
FEAR, AND GENERAL LACK OF CURtOSITY HAV! KEPT THE LOCAL
HUMAN INHABITANTS FROM VENTURING VERY FAR INTO THE INTERIOR.
THESE CAVES ARE GENERALLY UNDISTURBED BY THE C~SUAL VISITOR
AND THE ORGANISMS FOUND THEREIN MAV BE REGARDED AS
NATURALLY OCCURRING FUNGI.
A NUMBER OF WELL ... KNOVN COMMERCIAL CAVERNS WERE At.SO
VISITED DURING THIS INVESTIGATION. THESE WERE LOCATED IN
THE AREAS BORDERING THE SHENANDOAH VALLEY• SINCE THESE
ARE FREQUENTED ev THE GENERAL PUBLIC, SUCH CAVERNS CONTAIN
ABUNDANT DISCARDED DEBRIS• THE ORGANISMS INHABITING SUCH
SUBSTRATA ARE NOT GENERALLY REGARDED AS NATURALLY OCCURRING
INHABITANTS OF THE CAVERNIOO~OUS ENVIRONMENT. IN COM-
MERCIAL CAVERNS CARE WAS TAKEN TO COLLECT MATERtAL FROM
THE LESS FREQUENTED AREAS IN ORDER TO LESSEN THE CHANCE OF
COLLECTING TROOLOXENIC CONTAMINANTS. -1 t-
SAMPLES OF SOIL AND WATER WERE COLLECTED FROM BOTH
COMMERCIAL CAVERNS AND NON-COMMERCIAL CAVES• SEVENTEEN
SOIL SAMPLES AND THIRTY-ONE WATER SAMPLES WERE OBTAINED
FROM FIVE COMMERCIAL CAVERNS LOCATED IN WIDELY SEPARATED
AREAS OF THE SHENANDOAH VALL!Ye NINETY-TWO SOIL SAMPLES
ANO THIRTEEN WATER SAMPLES WERE COLLECTED FROM THE CAVES
IN THE BLACKS&URQ AREAe
THE PROCEDURE USED IN THE COLLECTION AND ISOLATION
O~ OAVERNICOLOUS FUNQJ WAS ESSENTIALLY THE SAME AS THE
TECHNIQUES INITIATED BY BUTLER (1907) AND HARVEY (1925) 1 AND SINCE ADOPTED BY MOST STUDENTS OF THE SOIL AND WATER INHASITINQ PHVCOMYOETOUS FUNGle SOIL SAMPLES WERE COL-
LECTED IN STERILIZED 60 MLe GLASS BOTTLESe SUCH SAMPLES WERE OBTAINED BY SCRAPtNO THE SURFACE SOIL OR BY OIQQtN8
FRESH EXCAVATION8e WATER SAMPLES W!R! OBTAINED SY
PARTIALLY SUBMERGING STERILIZED 60 ML. QLA88 BOTTLES BELOW
THE SURFACE TO ALLOW THE WATER TO FLOW FREELY INTO THE CONTAINER. IN ALL CASES THE SAMPLES WIRE CAREFULLY LABELED AND RETURNED TO THE LABORATORY. THE TIME LAPSE BETWEEN
COLLECTION AND BAITING VARIED FROM SEVERAL HOURS TO
8£V~RAL DAYSe DURING THE LONGER DELAYS SAMPLES WERE
STORED IN A REFRIGERATED CONTAINER.
IN THE LABORATORY A ~ORTION OF EACH SOIL SAMPLE
SUFFICIENT TO COVER THE BOTTOM WAS TRANSFERRED TO A -12-
STERILE, DEEP PETRI-DISH. To EACH WAS ADDED APPROXtMAT£LY 75 cc. OF STERILE SOIL WATER ANO EACH DISH 80 PREPARED WAS
BAITED WITH HALVES OF SOILED HEMP SEED 1 SMALL PIECES OF
BOILED SNAKE 8KIN 1 SQUARES OF BOILED CELLOPHANE, A~D SMALL
FRAGMENT& OF STERILE, CHLOROPHYLL-EXTRACTED GRASS LEAVES. WATER SAMPLES WERE DISPENSED INTO STERtLE P!TRt-Dl8HES AND BAITED IN THE SAME MANNER• STERILE aGIL WATER WA8 OFTEN USED TO SUPPLEMENT THESE SAMPLES IF THE ORIGINAL MATERIAL WAS NOT SUFFICIENT TO FILL A 0£1P PETRI-DISH.
SAMPLES WERE MAtNTAINEO AT ROOM TEMPERATURE AND BAITS
EXAMINED MICROSCOPICALLY AT REGULAR INTERVALS• PR08A8L£
ANO POSITIVE l80LATE8 WERE TRANSFERRED TO FRE8H DISHES
AND REBAtTEO tN THE SAMI MANNER• SUCCEIStVE TRANSFERS
RESULTED IN UNtFUNQAL ISOLATES R£LATIYELY FREI OF
BACTERIAL CONTAMINATION. NO EFFORT WAS MADE TO OBTAIN
BACTERIAL-FREE CULTURES OF THE FUNGI STUDIED. THE
IDENTIFICATION AND MORPHOLOBtCAL OBSERVATtONS REPORTED IN
THE FOLLOWING lttltTION OF THIS PAPER WERE MADE FROM UNIFUNQAL MATERIAL MAINTAINED AS STOCK CULTURES• -13-
FUM G f COll ECT::::D
CH YTf-
t N I) t A 8 0 T • S l::R • , l: 14 f • f 9 07 •
MYCEL!UM COPIOUS, EXTRA-MATRICAL; HVPHAE MUCH BRANCHED,
IRREGULAR IN SfZE, MEASURING 3.3-5.0 p IN DIAMETER;
ZOOSPORANQIA CLAVATE TO SPHERICAL, TERMINAL OR INTER-
CALARY 1 M~ABURING 23.3-3f .6 µ tN WIDTH BY 63.3-19.9 µ
IN LCNQTH WITH A LONG APICAL PAPILLA OPENING BY AN
OPERCULUM MEASURING a.a µ IN OIAM~TER, APOPHYSATE, THE APOPHYSIS SPHERICAL TO CAMPANULATE ANO MEASURINQ to.o-
13.3 µ IN DIAMETER; ZOOOPORES ESCAPING FULLY OEVELOP£D
tN MASS, SPHERICAL, UNlFLAG£LLATE, CONTAINING A SINGLE
OIL DROP, MEASURING 6.7 µ tN DIAMETER; RESTING SPORES
NOT OBSERVED.
FOUNO ONCE ON A ROTTING STICK tN WATER, SHENANDOAH
CAVERNS, SHENANDOAH COUNTY, VtRGINIA•
RAMOS.A, WAS FIRST DESCRIBED ON ROTTING STEMS OF TRtTtCUM
VO!,OA,RE JN IHOIA (8UTLtrn 1 1907), AND WAS LATER REPORTED
OH VEGETABLE DEBRIS FROM HUNGARY (OOMJAN, 1936)1 ANO
FROM BRAZIL (KARLfNQ 1 1944). THE LATTER INVESTIGATOR HAS
FOUND THIS SPECIES ON SIMfLAR SUBSTRATA THROUGHOUT TH£
SOUTHERN UNtTED STATES (KARLING 1 1941 1 1942 A). SPARROW -14-
AND BARR (1955) HAVE RECENTLY FOUND THE SAME FUNGUS ON MORIBUND _...._..,.._CHARA BP. FROM NORTHERN MICHIGAN.
THE VtRQINIA ISOLATE AGREES IN MOST RESPECTS WITH
THE DESCRIPTIONS OF PREVIOUSLY DESCRIBED FORMS• RESTING
SPORES WERE NOT OBSERVED, NOT WERE THE PROLIFERATING
OELLS OF THE RHtZOMYCELIUM, AS NOTED BY BUTLER (1907).
NOWAKOWS_,l(f ELLA RAMOS A MAY BE 0 lSTINGU I SHED FROM OTHER
APOPHVSATE MEMBERS OF THE GENUS, BY THE POSSESS&OH OF
NON-PROLIFERATING ZOOSPORANQIA MEASURING 25-45 µ IN
DIAMETER, AND BY THE ABSENCE OF SETAE ON THE ZOOSPORANGIAL
WALL. IT ts OF INTEREST TO NOTE THAT, ALTHOUGH TH•s
ISOLATE MAY SE READILY CULTtVATEO, IT WILL GROW ONLY ON
WOOD CHIPS AUD FAILS TO DEVELOP ON OTHER BAITS USED IN
THIS STUDY.
2. ? RHIZOPHLYCTfS !!f.•
SOIL ANO WATER COLLECTIONS FROM A NUMBER OF CAVES
YIELDED A VARIETY OF MONOCENTRtC CHYTRIDS APPARENTLY
ALLIED TO RHIZOPHLYCTIS. SEVERAL CHARACTERISTIC TYPES
OF ZOOSPORANGIA WERE OBSERVED BUT TIME DID UOT ALLOW
FOR THE tNDUCTtON OF ZOOSPORE DISCHARGE. HENCE, IDENTI-
FICATION WAS NOT FEASIBLE. THESE ISOLATES APPEARED
PRIMARtLY ON SNAKE SKIN BAITS AND HAVE BEEN MAINTAINED
FOR FURTHER INVESTIGATION. OWING TO THE CONFUSED STATUS
OF TH£ SPECIES ANO TO OUR LACK OF UNOERSTANDINQ OF THE -as-
RHtZOPHLYTt8-COMPLEX 1 IDENTIFICATION ANO FURTHER STUDY OF THESE ISOLATES WOULD BE WELL WORTHWHILE.
BLASTOCLAD I ALES
3. A~LO~VCES CY8TOGENUS EMERSON, LLOYDIA, !,: 136. 1941.
HYPHA ABUNDANT, PSEUOOSEPTATE 1 MEASURING 23.3 p AT
TH! 8A8E TO 10.0 µ AT THE TIP; THIN-WALLEO Z008PORANGIA
TERMtNAL OR SUBTERMINAL IN A ROW, 08LONQ 1 MEASURING 26.6- 31.6 µ IN WIDTH AND 56.6-80.1 µ IN LENITH (AVERAGING 29.3 X 72.3 µ); RESTING SPORANQIA THICK-WALLED, PITTED, BROWN, DICIDUOU8, MEASURING 31.6-35.9 µ IN WIDTH AND 45.o-st.6 µ IN LENGTH (AVERAGING 33.3 X 49.3 p); GERMINATION OF RESTING SPORANOIA NOT OBSERVED; QAMETANGIAL PHASE LACKING.
l&OLATEO 5 TIMES FROM WET SOIL ON HEMP SEED FROM SMOKE HOLE ANO HOQ HOLE, GILES COUNTYJ AND FROM CLOVER HOLLOW, CRAIG COUNTY, VtRQtNtA. ALLOMYoga CV8TOGENU8
HA8 BEEN PREVIOUSLY OE8CRIB£D AS A SAPROPHYTE IN SOIL
FROM ARIZONA, VENEZULA, 8URMA 1 AND CHINA (EMERSON, 1941), AND 18 APPARENTLY WIDELY Qt8TRl8UTED IN SOILS•
THE CAVERNICOLOUS 180LATE8 AGREE WITH TM! DE9CRIPTtON OtVEN BY EMERSON (1941) FOR ALLOMYCE8 CY8TOGENU8• THE
MEASUREMENTS OF THE HYPHA£ 1 Z008PORANQIA 1 AND RESTINQ
SPORANQIA FALL WITHIN THE RANGE REPORTED FOR THAT SPECIES• FURTH£RMOR!1 !.• CVITOQENUS MAY BE DISTINQUfSHEO FROM -16-
OTHER SPECIES OF THE GENUS BY THE POS9ESStON OF D!CtOUOU8 RE$TINQ SPORANGIA,, BY TH£ COLORLESS CYTOPLASM,, AND BY A l.IF'E-CVCl.E WHICH CON8tST8 SOLELY OF AN ASEXUAL PHASE•
4. BLASIOC~ADtOPata PARVA (WHtFFEN) SAPRROW,, J. OF WA8H1NQTON ACAD. Set., iQ.1 sa. 1950~
THALLU8 UP TO 500 µ IN LENQTHJ HVPNAE DICHOTOMOUSLY 8RANCM£D1 OFTEN WITH ONE BRANCH 8UPPREl8EO, MEASURING
8.9-14.0 ~ IN DtAMETERJ ZOOSPORANGIA CLUB 8HAPED,
MEASURING 36.6-44.9 µ IN DIAMETER, WITH ONE APICAL AND
ONE AXILLARY PAP1LLAJ R£8TfNQ SPORES 8MOOTH-WALLED 1 NON-
PUNCTATE, SPHERICAL TO ovo10, MEASURING 25.0-33.3 ~ IN DIAMETER (AVERAQtNO 28.0 µ), LYING LOOIELY IN A SMOOTM-
WALLED CONTAINER, THI LATTER SPHERtC~L TO ELLIPSOIDAL M!A8URINQ 27.8-36.7 µ fN DIAMETER; GERMINATION OF THE RE8TINQ 8,0RE NOT 088£RV£D•
faOLATED ONCE ON HEMP 8££0 FROM WET SOIL, HOG HOLE•
GILES COUNT'Y1 VtRGINlA• BLA8TOOLADtOP818 PARYA HAS BEIN
PREYIOU8LV OS8CRt8ED AS A SAPROPHYTE lN SOIL FROM TEXAS (WHtFFEN1 1943) AND FROM CUBA (SPARROW, 1950). IT HAS
8EE.N REGARDED GENERALLY AS A FUNGUS RAt! IN tT8 OCCURRENCE.
THta l80LAT£ VAS OBTAINED ON HEMP 8EED, APPARENTLY AN UNUSUAL BAIT FOR tT8 t80LATION• PREVIOUS RECORDS OF THIS
8,101£8 INDICATE ITS ABtLITV TO GROW ON SNAKE SKIN ANO -17-
GRASS, BUT THErtE f S NO REPORT OF ITS OCCURR~NCE ON HEMP
SEED• THE FACT T~AT IT GROWi IN CULTURE OH HEMP SEED AND
THIS tSOLATE, AT LEAST, ts ALWAYS FOUND AMONG THE HEAVY
MYCELIAL QROWTH OF AN UNtDENTIFIEO IMPERFECT FUNGUS, MAY
WELL ACCOUNT FOR THE R£PORTtD RARlTV OF THIS FUNGUS.
THE MEASUREMENTS OF THtS ISOLATE ARE SLIGHTLY SMALLER
THAN THOSE CU Vim FOR 8.f.;,AS,TOCl.AD,IOP,S,l~ PARVA BY SAPRROW
( 1950). THIS SPEOIE8 es,, AT PRESE~lT, THE ONLY DESCRIBED
MEMBER OF THE GENUS. HOWEVER, SCOTT (PERSONAL COMMUNICA-
TION) OBTAINED A LARGE NUMBER OF ISOLATES FROM TROPICAL AND
SUBTROPICAL SOILS. THESE VARIED QREATLV WtTH RESPECT TO
SIZE AND NUMBER OF ZOOSPORANGIA AND NUMBER OF DISCHARGE
PAPtlLAE, AND IN TH~ StZE A~D COLORATION OF RESTtNQ
SPOr.Es. IT IS QUITE POSSIBLE, THEN, THAT THIO GENUS MAY
BE REPRESENTED ev SEVERAL SPECIES, AND THAT THE CAVERNl-
COLOUS ISOLATE MAY, INDEED, REPRESENT A NEW AND UNOESCRIBEO
MEMBER OF !LASTOCLADtOPStS.
THE GENUS BLASTOCLADIOPSIS MAY BE DtSTINQUt8H£0 FROM
THE CLOSELY RELATED GENUS BLASTOCLADfA ev POSSESSING NON-
PONCTATE RESTING SPORES WITH COMPLETELY SMOOTH WALLS AND
THESE NOT FILLING THE RESTING SPORANGIUM. FURTHERMORE,
THE MULTl-PAPILLATE NATURE OF THE ZOOSPORANGIA ANO THE
SUPRESSED DICHOTOMY OF THE THAt.t..US SERVES TO OtSTINGUISH
TH£Sl GENERA• -18-
SAPROLEGNtALES
5. ACHLYA DUSI! COKER, SAPROLEGNIACEAE1 P. 135. 1923.
HYPHAI STOUT, SPARINGLY BRANCHED, MEASURING 41.8- 78.5 µ tN DIAM!TERj QEMMAE ABUNDANT IN OLD CULTURES, IRREGULAR, TERMINAL OR fNTERCALARY; ZOOSPORANQIA ABUNDANT,
FUSIFORM, MEASURING 42.8-49.9 µ IN WIDTH BY 337.6-514.0 ~ IN LENGTH (AVERAGING 47.1 X 426.9 µ), RENcWED 8YMPODIALLV OR IN BASIPETAL SUCCESatoN; THRAU8TOTHECOID zooaPORE 018-
CHARGE FROM PRIMARY Z008PORANQIA 1 ACHLYOIO DtSCHARQE FROM SECONDARY ZOOSPORANGIAj ENCY.TED Z009PORE8 MEASURING 10.0 µ IN DIAMETER; OOGONIA ABUNDANT ON LATERAL BRANCHES, SPHERICAL
TO PYRtFORM 1 SMOOTH WALLED, PITTED ONLY UNDER ANTHERIDfAL ATTACHMENT, MEASURING 39.g_53.3 µ tN DIAMETER (AVERAGING 46.6 µ IN DIAMETER); 008PORE8 ECCENTRIC, SPHERICAL, 4-7 PER OOQONIUM, USUALLY 4, MEASURING JG.7-22.2 µ IN DIAMETER
{AVERAGING 19.3 µ IN DIAMETER), NOT ,ILLINQ THE OOGONIUM;
ANTHERtDIA MONOCLINOU8 1 ONE PER OOQONIUM 1 ANTHERIDIAL CELL
APPRE88£D; OOSPORE GERMINATION NOT OBSERVED.
ISOLATED ONCE ON HEMP SEED FROM WATER, SKY LINE
CAVERNS, WARREN COUNTY, VIRGINIA• ACHLYA DUBIA HAI BE£N 018CRl8ED PREVIOUSLY A8 A SAPROPHYTE IN SOIL FROM NORTH
CAROLINA, SOUTH CAROLINA, GERMANY, AND PUERTO RICO (JOHNSON, 1956 e). -19-
THE CAV!RNICOLOU8 l80LATE DIFFERS IN ANTHERlDlAL
ORIGIN FROM THOSE ISOLATES DESCRIBED BY JOHNSON, AND THE
Z008PORAN8lAL AND OOGONIAL MEASUREMENTS OF THIS ISOLATE
8UCIGEST A C.LOfl»R R£LATIO"f 8H tP TO ACHLYA CAROL IN tANA THAN
TO THAT OF ACHLYA DU8IA 1 ALTHOUGH THE THRAUITOTHECOIO
TYPE OF ZOOSPORAN81AL DISCHARGE 19 CHARACTERISTIC OF THE
LATTER SP£CJ!8• IT MAY WELL 8£1 THER£FORE1 THAT TfffS
ISOLATE 18 A FORM INTERMEDIATE BETWE£N THE TWO AFORE•
MENTIONED SPECIES. THE MOST DISTINCTIVE CHARACTERISTIC
OF A.• DU81A 18 ITS THRAUSTOTHECOID TYPE OF ZOOSPORE 018- 0HARGE FROM THE PRlMARY ZOOSPORANQIA, AND IT fS THIS
FEATURE WHICH DISTINGUISHES THIS SPECll8 FROM ALL OTHER
MEMBERS OF TH! GENUS•
6. APHANOMYCE8 LA£Vl8 DEBARY, JAHRB. wtss. BOT., £1 179. 1860.
HVPHAE SPARINGLY 8RANCH£0 1 UNDULATED, MEA8URING 5.0- 6.6 p IN DIAMETER; ZOOSPORANllA FtLAEMNTOUa, UNDIFFERENTIATED FROM YEOITATIVE HYPHAE, MEASURING 5.8 p IN WIDTH AND VARY•
INQ tN LENGTH; PRIMARY %008PORE8 ROD-SHAPED, MEASURING
16.7 ~ tN LENGTH; ENCYSTED ZOOSPORE8 MEASURING 7e5 p tN
OIAMETERJ OOQONIA TERMINAL ON SHORT LATERAL 8RANCH£9 1
SPHERICAL TO 8U88PHERtOAL, THIN WALLED SOMETIMES 8£COMING
IRREGULAR DUE TO THE DISINTEGRATION OF THE ATTENDANT
ANTHERIDIA, MEASURING 17.1-33.3 p IN DIAMETER (AVERAGINQ
22.6 ~); 008PORE8 8INQLE 1 HYALINE 1 CONTENTS GRANULAR, -20-
WtTH A LARQE CENTRAL OIL GLOBULE, MEASURING 15.7 µ IN
DIAMETER; ANTHERIDIA ONE TO SEVERAL, VERMIFORM, ANTHERIDIAL
STALK SIMPLE 1 OfCLINOUS OR MONOCLINOU8 1 FERTILIZATION TU8E8
ABSENT; OOSPORE GERMINATION NOT OB8ERVE0e
ISOLATED 5 TIMES ON SNAKE SKIN FROM SOIL AND WATER, STRALEY'• CAVE, GILES COUNTY; GRAND CAVERNS, AUGUSTACOUNTYj LURAY CAV!RUI• PAGE COUNTY; ANO SHENANDOAH CAVERNS, SHENANDOAH COUNTY; VtRQINIAe APHANOMYCES LAEVIS HAS BEIN PREVIOUSLY OE8CRIBEO AS BOTH A PARASITE ANO A SAPROPHYTE ON MANY DIVERS! SUBSTRATA FROM WIDELY SEPARATED ARCAS OP THE WORLD (SCOTT, t958). IT 18 APPARENTLY THE MOST
COMMON AND MOST WIDELY DISTRf8UTED MEMBER OF THE GENUS.
THE CHARACTERISTICS OF THtS CAVERNICOLOUS ISOLATE
AGREE WITH THE MODIFIED DE8CRtPTtON QIYEN IV SCOTT (1958).
APHANOMYCEa LAEYl8 MAY BE DISTINGUISHED FROM OTHER SPECIES
OF THE GENUS HAVING OOQONIA WITH SMOOTH WALLS OR WALLS.
MADE ROUQH BY THE DISINTEGRATION OF ATTENDANT ANTHERIDIAL
CELLS, BY POSSESStNQ NON-COILtNQ ANTHERIDIAL 8TALK8 1
SIMPLE Z008PORANllA1 AND BY THE SIZE OF OOGONIA AND OOIPORES• IT 18 OF INTEREST TO NOTE THAT, ALTHOUGH APHANOMYCES LAEVIS HAS NOT BEIN REPORTED PREVIOUSLY AS A CAVERNICOLOU8
SPECl£81 ANOTHER MEMBER OF THE QENUS 1 !.• 8TELLA!U8, HA8 BEEN ISOLATED FROM ORY SOIL COLLECTED IN AN ALEUTIAN BURIAL CAY! tN ALASKA (SCOTT, 1958). -2t-
7. SAPROLEQNIA FERAX (GRUtTH) THURET 1 ANN. Set. NAT. Ill, -r 4 : 2 r 4. r a50. HYPHAE ABUNDANT, IRREGULAR, SPARINGLY BRANCHED,
M£ASURING 18.3-42.8 p IN DIAMETER; ZOOSPORANOIA TAPERtNQ
TOWARD THE APEX, EXHIBITING INTERNAL PROLIFERATION, MEASURING 28.6-42.8 p IN WIDTH BY 371.3-470.8 p IN LENGTH
(AVERAGING 42.8 X 379.8 p); ZOOSPORES MEASURING 11.7-
13.3 p IN DIAMETER UPON ENCYSTMENT; OOGONIA SPHERtCAL 1 MEASURING 53.3-98.2 p IN DtAHETER (AVERAGING 73a9 p) SOME WITH HVPHAL-LIKE EXTENSIONS CONTAINING OOSPORES IN A
SINGLE ROW, OOGONIAL WALL PITTED; OOSPORES SPHERICAL TO
ELLIPSOIDAL, CENTRIC, NUMBERING 4-16 PER OOQONIUM
(AVERAGING 8), MEASURING 23.3-33.3 p IN DIAMETER (AVERAG-
ING 28.8 p); ANTHERIOIA PRESENT ON LESS THAN 10~ OF THE
OOQONIA 1 MONOCLINOUS, ONE PER OOGONIUM; OOSPORE
GERMINATION NOT OBSERVED.
ISOLATED ON HEMP SEED ANO SNAKE SKIN 3 TIMES FROM
WATER, GRAND CAV~RNSt AUGUSTA COUNTY 1 AND fNOLESS CAVERNS,
ROOKINGHAP.1 COUNTY, VtRCUNIA. SAPROLEGNIA F"~RAX HAS SEEN
C!SCRIBED PREVIOUSLY AS A SAPROPHYTE FROM WATER AND SOtL
f'ROM THE UtUTED STATES AND Eur~OPE (COKER ANO MATTHEWS,
1937} ANO IS APPARENTLY ONE OF THE MORE COMMON WATER MOLDS
TO BE FOUNO IN THESE HABITATS. -22-
As JOHNSON (1956 A) STATES, SAPROLEGNIA FERAX
(GRUITH) THURET, ANO THE FOLLOWJNG SPECIES, ~· MIXTA
OE8ARY 1 REPRESENT A COMPLEX OF TWO SPECIES WITH MANY
VAR1ANTS. THE PRINCf PAL OIFFERENCE BETWEEN THESE SPECIES fS APPARENTLY THE PRESENCE OF FUNCTIONAL OOOONIA WlTHIN THE
EMPTY ZOOSPORANGIA OF ~· ~~AX 1 AND THE LACK OF THESE IN !· MIXTA. JOHNSON (Loe. CIT.) REFERS TO ISOLATES, OTHER-
WISE SIMILAR TO !• MIXTA 1 WHICH PRODUCE OOGONfA WITHIN
DISCHARGED Z00SPORANGIA 1 AND ISOLATES CLEARLY CHARACTERtS- TIC OF -s. FERAX, WHICH LACKED SVCH OOGOHtA. OosPORE NUMBER IS MUCH TOO VARlABLE IN THESE SPECIES TO BE USED
AS A DISTINGUISHING FEATURE. SAPROLEGNIA FERAX, HOWEVER,
CAN BE Ot9TINCUISHED FROM OTHER RELATED SPECIES WHICH
LACK ANTHERIOIA ON MORE THAN HALF OF THE OOGONtA AND WHICH
HAVE OOGONIA THAT ARE NOT PRODUCED IN CHAINS, BY THE
P0$SESSION OF LARGE OOSPORES AND OOGONIA, THE LATTER WITH
EXTENSIONS CONTAINING OOSPORES ANO BY THE FORMATION OF
OOQONIA WITHIN OLD ZOOSPORANGIA• THE JSOLATES ASSIGNED TO -S. FERAX DURJNG THIS STUDY AGREE IN ALL RESPECTS WITH THE OESCRIPTION OF THE SPECIES AS MODIFIED BY COKER AND
MATTHEWS ( t 937) •
8. SAPROLEQNIA MIXTA DEBARY 1 BoT. ZEIT. 1 -41: 38, 54. t883.
HYPHAE ABUNDANT, IRR£0ULAR 1 SPARINGLY BRANCHED, MEASUR-
INQ 18.0-47.9 µ IN OtAMETER; ZOOSPORANGIA TAPERING TOWARD -23-
THI APEX, iXHIBITINQ INTERNAL PROLIFERATION, MEASURING
28.6-42.8 µ WIOE, 314.6-421.3 µ IN LENGTH (AVERAGING 42.5 X 352.7 p); Z008PORE8 MEASURING 11.7-13.3 p tN
DIAMETER UPON ENCYSTMENT; OOOONIA SPHERICAL, TERMINAL
OR INTERCALARV, NEVER FORMED WITHIN OLD ZOOSPORANGIA,
MEASURING 63e2 µ IN DIAMETER, OOQONIAL WALL PITT!OJ
009PORES SPHERICAL, CENTRIC, NUMBERINQ 6 PER OOQONIUM 1
NOT FILLING THE OOQONIUM, MEASURING 26.6-29.9 µ IN DIAMETER (AVERAGING 28.3 µ); ANTHERIDIA PRE8ENT ON ABOUT
35% OF THE OOQONIA, MONOOLINOU8 1 f PER OOQONIUMJ OOSPORE
QIRMINATION NOT 088ERYED•
f SOLATID ON HEMP SEED TWICE FROM WATER, NEV RtYIR
CAVE, GILES COUNTY, VIRGINIA• SAPROLIGNIA MIXTA HAS BEEN
DESCRIBED AS A SAPROPHYTE FROM WATER AND SOIL FROM THE
UNITED STATE81 GERMANY, ANO EUROPE (COKER ANO MATTHEV81 1937). Aa PREVfOU8LY 8TATEO (Pe 22) THE VALIDITY OF
THIS TAXON 18 QUESTIONABLE•
TH! CAVERNICOLOU8 ISOLATES AGREE WtTH THE MODIFIED
D£8CRIPTION QIV£N BY COKER ANO MATTHEWS (1937) FOR
SAPROL!QN!6 MIXTA IH ALL RESPECTS EXCEPT OOSPORE SIZE•
THE 008POR£8 OF THESE t80LATES AVERAGE SOMEWHAT LARGER
THAN THOSE OF PREVIOUSLY DESCRIBED FORMS•
IT IHOULO BE NOTED THAT THESE llOLATE8 WERE
PARA8tTIZ£D BV 0LPIDIOP818 8!PROL£QNIAEe THl8 MAY WELL
ACCOUNT FOR THE LACK OF OOQONIA WITHIN OLD ZOOSPORANGIAe THE StGNIFICANC! OF THE LACK OF OOGONIAL EXTEN810N8 AND TH! ABSENCE OF OOGONIA WITHIN THE OLD ZOOSPORANGIA MUST -24-
6£ CONSt0£'RE:D1 AT LEAST IN THESE ISOLATES,, SINCE tN EACH
CASE HAO THESE ACCESSORY STRUCTURES OEVELOPE0 1 THE FUNGUS
MIGHT WELL BE ASSfGN£D TO SAPROLEGNtA FERAX•
LEPTOMITALES
9. APODACHLYA MtNIMA COKER ANO lEtTN£R 1 J. ELISHA MITCHELL Sc r. Soc., M_: 313. 1938.
MVCELIUM FLACCID ON HEMP SEED; MAIN HYPHAE SLENDER,
SEGMENTED, BECOMING NARROWER TOWARD THE APEX, SEGMENTS
MEASURtNQ 6.1-10.0 µ tN DIAMETER av 84.3-166.5 µ IN
LENOTH 1 SEGMENTS OF OOQONIAL BRAMOH MEASURING 6.7 µ IN
DIAMETER SY 6.7-15.0 µ IN LENGTH; PROTOPLASM THSN EXCEPT
IN OOGONIAL BRANCHES; ZOOSPORANOIA NOT OBSERVED, ZOOSPOR-
ANQIA ARl$1NG FROM SINGLE LATERAL BRANCHES tN GLOMERULJ,
OVOID OR PVRIFORM 1 MEASURING AFTER DISCHARGE 12.8•17.6 µ IN
DIAMETER BY 17.6-26.2 µ tN LENQTH 1 (JOHNSON, 1955) ; OOGONIA
SPHERICAL WITH SMOOTH, THICK WALLS, MEASURING 12.9-15.7 µ fN DIAMETER (AVERAGJNG 14.9 µ); OOSPORE SINGLE, COMPLETELY
FILLINQ THE OOOONIUM 1 AT FfRST ECCENTRIC, BUT LATER THE
CONTENTS ROUND UP INTO A HYALINE BALL CLOSELY APPRESS£D TO
A SMALLER HYALtNE OIL DROPLET; ANTHERIDIUM ORIQINATtNQ
FORM A SUBOOQONIAL CELL AS AN OUTGROWTH, BECOMING EMPTY
BEFORE MATURATION OF THE OOSPORE•
ISOLATEO TWICE ON HEMP SEED, ONCE FROM WATER FROM
ENDLESS CAVERNS, ROCKltHlHAM COUNTY,, VIRQINIA1 AND ONCE -25-
FROM SOIL AROUND ROOTS OF DRYOPTERIS INTERMEDIA FROM LURAY CAVERNS, PAGE COUNTY, VIRGINIA• APODACHLYA MINIMA HA8 BEEN 0£8CRIBED PREVtOUSLY FROM A SLtBffTLY CONTAMINATED
STREAM FROM NORTH CAROLINA (COKER AND LEITNER, 1938) AND
FROM SOIL FROM PUERTO RICO (JOHN80N 1 1955).
THIS ISOLATE AGREES IN ALL RESP£0T9 WITH THE ORIQINAL
DESCRIPTION OF APODACHLYA MINIMA A8 GIVEN BY COKER AND . lEtTNER (1938). SPORANGIA, HOWEVER, WERE NOT OBSERVED IN
THIS ISOLATE• THEY HAYE BEEN OBSERVED AND DESCRIBED IN
DETAIL BY JOHNSON (1955) 1 WHO COMMENTS ON THE DIFFICULTY
OF OBTAINING Z008PORANGtA IN CULTURE AND ON THE OBSCURE NATURE OF THESE STRUCTURES. A• MINIMA MAY BE DtSTINGUtSHEO FROM RELATED SPECIES OF THE GENUS HAVING 8MOOTH•WALLED
008PORES AND OOQONIA BORN£ ON SHORT MONILIFORM SEGMENT& BY
THE P088E88ION OF ANTHERIDIA ORfQfNATING A8 LATERAL OUT-
GROWTHS OF THE 8UBOOGONIAL CELL AND BV THE RARITY OF
ZOOSPORANGtAL FORMATION.
LAGENIOIALES
10. 0LPIDIOP8!9 !PH6NOMYCl8 CORNU, ANNe Set. NAT., !§.l 148. 1872.
ZooaPORANGtA SMOOTH, SPHERICAL, OVAL TO ELONGATE, 1-4 PER SWOLLEN HYPHAL TIP, MEASURING 23.3-39.9 µ IN WIDTH AND 34.9•39.9 µ tN L£NQTH; WITH ONE OR TWO EJ1T PAPILLAE -26-
MEASURING UP TO 10.0 µ IN LENGTH AND EXTENDING THROUGH
THE HOST CELL WALL; RESTING SPORES NOT OBSERVED.
ISOLATED ONCE AS A PARASITE OF APHANOMYCES LAEYl8
FROM WATER, SKY LtNE CAVERNS, WARREN COUNTY, VIRGINIA•
0LPIDIOP8f8 APHANOMYCIS HAS BEEN DESCRIBED PREVIOUSLY AS A PARASITE OF APHANOMVCES !f.• FROM FRANCE, DENMARK, GERMANY, INOIA, AND THE UNITED STATES (KARLING, 1942 B)
ANO IS APPARENTLY OF COMMON OCCURRENCE ON THIS HOST•
THE CAVERNtCOLOUS ISOLATE AGREES WITH THE MODIFIED
DESCRIPTION GIVEN FOR TH! ISOLATE BY KARLINQ (1942 8)
A~THOUGH HE FAILED TO INCLUDE THE Z008PORANGIAL MEASURE-
M!NT8 OF HIS ISOLATE•
0LPIOtOP818 APHANOMVCIS MAY OTHERWISE BE Dt8TINGUISHED
F.RQ~ OTHER SPECIES OF THE GENUS BY THE HOST ON WHICH tT GROWS. WHETHER OR ;\!OT HOST SPECfFtCtTY rs A VALID CRI-
TERIGN FOR THE fDENTIFIOATION OF SPECIES ts QU~8TtONABLEe
U~Tf~ f~RTHER STUDIES ON MEMBERS OF THtS GENUS ARE lNJTtATED ANO MORPHOLOGICAL OfSTfNCTtONS CLARIFIED, MOST
SP!CJE8 tNCLUOING THIS AND THE FOLLOWfNG ISOLATE MUST
REMAIN PROBLEMATaCAL.
''· 0LPIDIOP81& SAPROLEQNIAE (BRAUN) CORNU, ANN. See. NAT., -151 145. 1872. ZooaPORANQIA SMOOTH, HYALINE, SPHERICAL TO ELONQAT!,
4-12 PER SWOLLEN HYPHAL TIP OF THE HOST, MEASURING 14.9-33.3 ~ -27-
IN DIAMETER WITH 1-2 EXfT PAP,LLAE EXTENOtNQ TO SURFACE OF
HOST CELL; RESTING SPORE NOT OBSERVED.
ISOLATED TWICE AS A PARASITE ON SAPROLEGNIA .. PP. FROM .-.....-.....;...... --...... - WATER, NEWCASTLE ~lURDER HOLE, CRAIG COUNTV 1 ANO FROM WATER,,
New RIVER CAVE, GILES COUNTY, VtRGINIA. OLPfOfOPSIS
SAP~OLECNIAE HAS BEEN DESCRIBED PREVtOUSLV AS A PARASITE OF SAPROLEGNIA AND ISOACHLYA FROM GERMANY, FRANCE, Rusa•A,,
DENMAfH<, UNITED STATES, JAPAN, ENGLMJ01 ANO RUMANJA
(SPARROW, 1943).
THE ZOOSPORANGtAL MEASUREMENTS OF THIS ISOLATE AR£
WITHIN THE RANGE GIVEN BY KARLJNQ (1942 B) ANO SPARROW
(1943) FOR 0LPIDIOPSJS SAPROLEGNIAE. fT 19 INCLUDED HERE
BECAUSE OF ITS REGULAR SMOOTH WALLED ZOOSPORANCSA AND
BECAUSE OF THE HOST ON WHICH IT GROWS. WITHOUT RESTING
SPORES, THE fSOLATE CANNOT BE DEFtNJTELV DISTINGUISHED FROM ft• fNCRASSATA. As MENTIONED PREVIOUSLY THE IDENTIFI- CATION OF SPECIES tN THIS QENUS BASED ON HOST SPECIFICITY
8 ~UESTIOMA8LE. THIS t$0LATE, THEN, MAV ONLV BE INCLUDED
HER£ TENTATIVELY• -28-
PERONOSPORALES
12. ?!1HIUM !FERTILE KANOUSE AND HVMPHREV, PAPER8 MtcH. AcAo., !r 127. 1927.
HVPHA! BRANCHING, MEASURINQ 3.3-f0.0 µ tN DIAMETERJ
ZOOSPORANGIA UNBRANCHED, TERMINAL OR INT£RCALARY, MEASUR-
ING 3.3-S.0 µ IN WIDTH BY 83.2-527.0 ~ IN LENGTHI
Z008PORE8 FORMED IN A VESICLE MEASURINQ 23.3-35.7 ~ IN
DIAMET!R 1 ZOOSPORE8 MONOPLANETIC 1 MEA8URINQ 8.3-10.0 ~J
SEXUAL REPRODUCTION UNKNOWNI CHLAMYOOSPORES NOT OB8ERVE0e
ISOLATED ON HEMP SEED, GRASS, AND SNAKE SKIN 6 TIME8 FROM WET SANDY SOIL, CLAY, AND WATER; Hoo HOLE, GILES
COUNTYI EARHART'S CAVE 1 MoNTQOMERY COUNTY; AND LURAY CAVERNS, PAQE COUNTY, VIRGINIA• PYTHIUN AYERTIL§ HAS BEEN OE8CRIB£0 PREVIOUSLY AS A PARASITE OF ...... GossVPtUH ___SP • FROM ANGLO-fQVPTIAN SUDAN, OF VAUCHERIA s~. FROM THE
UNITED STATES, AND OF VIOLA TRICOLOR FROM THE NETHERLANDS
(MIDDLETON, 1943). THIS t8 APPARENTLY THE FtRST REPORT
OF THl8 SPECIES OCCURRING AS A SAPROPHYTE FROM 801L AND
ALL CAV£RNICOLOU8 ISOLATES OF PvTH!UN WHICH POS8E8S
FILAMENTOUS ZOOSPORANGIA AND NO SEXUAL REPRODUCTION HAYE 8E£N tNCLUDED UNDER THE BINOMIAL PVTH!UM AFERT!LE• MIDDLETON (1943) STATES THAT ttf• !FERTILE MAY EITHER BE -29-
MAtHTAtNEO AS A DISTINCT SPECIES, LACKING A SEXUAL STA8E,
OR AB A 81ffOMIAL WHICH INCLUDES A VARIETY OF RELATED
SPECl£8•·•~••••; TRANSFERENCE TO ANOTHER BINOMIAL WOULD
BE MANDATORY ONCE THE SEXUAL STAGE APPEARED tN THE 8PICIE8
OF THE LATTER CATEGORY•" StNCE MOST SPECIES OF PYTHtUM
CAN BE D19TINQUISHED ONE FROM ANOTHER ONLY OH THE BASea OF
TH!IR SEXUAL REPRODUCTIVE CHARACTERISTICS AND SINCE THE ZOOSPORANGIA OF l!• !FERTILE 18 NOT MARKEDLY DIFFERENT FROM THAT OF OTHER FILAMENTOUS ZOOSPORANGIAL FORMS OF THE GENUS,
TNE AUTHOR CONSfOER9 THIS TAXON TO BE AN AGGREGATION OF
UNRELAT£D IMPERFECT ISOLATES.
THE TWO ISOLATES INCLUDED HERE WHICH VERE OBTAINED
FROM PAGE COUNTY, VtRQtNIA WERE THE ONLY ISOLATES OF
PYTHIUM AFERTILE WITH A LtMIT!D HYPHAL GROWTH• THEIR HYPHAE NEYER REACHED A LENGTH OF MORE THAN 500 p, WHEREAS, ALL OTHER ISOLATES OF f• AFERTILE EXHIBITED
EXTEN81VE HYPHAL DEVELOPMENTe THt8 18 THE ONLY MARKED
DIFFERINCE IN THE ISOLATES INCLUDED UNDER THIS BfNOMIALe
13. PVTHIUM CAROLINIANUM MATTHEWS, STUDIES ON TH! GENUS
PYTHtUM, P. 71. 1931 •
. HYPHAE 8MOOTH, MEASURING 3.3-s.o ~ IN OfAMETERJ ZOOSPORANttA 8AMERICAL TO 8UBSPHERICAL, SHOWING INTERNAL
PROLIFERATION, MEASURING 16.7-30.0 ~ IN DIAMETER (AVERAGING 24.7 ~)J ZOOIPOR£8 NUMEROUS FORMED IN A STALKED VE81CAL ARISING FROM A SHORT APICAL PAPILLA; SEXUAL REPRODUCTION UNKNOWN. -30-
ISOLATED ON HEMP SEED TWtCE FROM CLAY ANO WET SANDY
SOIL, VICKER ROAD CAVE, MONTGOMERY COUNTY, VIRGINIA.
PYTHtUM CAROLINfANUM HAS BEEN PRE-tOUSLV DESCRIBED AS A
PARAS I TE OF SP I tlOOVRA !.!!.• FROM NORTH CAROL INA (MATTHEWS, 1931) ANO AS A SAPROPHYTE FROM VEGETABLE DEBRIS IN THE
UNITED STATES (MIDDLETON, 1943).
THESE ISOLATES AGREE WITH THE MODIFIED DESCRIPTION OF
PYTHJUH CAROLINIANVM GIVEN BY MIDDLETON (Loe. CIT.) EXCEPT
THAT THE RANGE IN SIZE OF THE ZOOSPORANQIA IS GREATER IN
THES£ ISOLATES THAN IN THOSE DESCRIBED BY MIDDLETON. LtKt PVTHIUM !FERTILE, f.• C!ROLtNIANUM HAS NO KNOWN SEXUAL REPRODUCTIVE STRUCTURES. HOWEVER, THE LATTER
SPECIES HA9 A ZOOSPORANQIUM WHICH 18 MARKEDLY DIFFERENT
FROM ALL OTHER PVTHIUM SPECIES. FOR THIS REASON, IT IS
REQARD~O AS A DISTINCT SPECIES AND NOT AS A BINOMIAL WHICH IS APPLIED TO A VARIETY OF RELATED FORMS• f• CAROLINIANUM MAY BE READILY RECOGNIZED ANO DISTINGUISHED FROM OTHER
SPECl£8 OF PVTHlUM HAVING SPHERICAL, PROLIFERATING
ZOOSPORANGtA Af'JO NO KNOWN SEXUAL REPRODUCTfON BY THE
POSSESSION OF A STALKED VESICLE ARISING FROM THE SHORT
APICAL PAPILLA OF THE ZOOSPORANQIUM. 14. PVTHIUM IRREQULARE BUtSMAN, MED. PHYTOPATH• LAB., -IJ: 1-1 s. 1927.
HVPHAE LOBULATE, FREELY BRANCHED, MEASURING 2.9 µ IN OtAMtTER: ZOOSPORANGIA SPHERICAL TO ovoto, TERMINAL OR -31-
INTERCALARY, NOT PROLIFEROUS 1 MEASURtNQ 22.9-30.0 µ IN DtAMETER (AVERAGtNQ 26.0 µ); OOQONIA SPHERICAL TO IRREGULAR, MEASURING 17.1-21.4 µ IN DIAMETER {AVERAQING 19.S µ); 008,0R!S SINGLE, APLEROTtc, MEASURING s.6-17.1 µ IN DIAMETER (AVERAGING 13~2 µ), CONTAINING A SINGLE OtL
GLOBULE; ANTHERIDIA I PER OOGONIUM 1 MONOCLtNOUS 1 SMALL,
APPLIED APICALLY TO THE OOQONIUM; OOSPORE GERMINATION NOT
OBSERVED.
ISOLATED ON HEMP SEED AND SNAKE SKIN TWICE FROM VET
ROCKY 80tL1 CLOVER HOLLOW, CRAIG COUNTY, VIRQINIAe PYTHIUM
!RR§GULARE HAS BEEN DESCRIBED AS A PARASITE ON 43 GPECIEB
OF FLOWERING PLANTS FROM THE UNITED STATES, NETHERLANDS,
HAWAII, ENGLAND, AND THE UNtON OF SOUTH AFRICA (MtDOLETON, 1943).
THta ts APPARENTLY THE FIRST REPORT OF THE OCCURRENCE
OF PYTHIUM IRREQULARE AS A SAPROPHYTE FROM SOILe GILMAN
(1957) INCLUDES THIS SPECIES AS A SOIL INHABITING FUNGUS AND REFERS TO AN ISOLATE REPORTED EARLIER BY YOUNKIN (1938).
THE LATTER INVESTIGATOR REISOLATED FROM EXPERIMENTAL SOIL
PLOTS A SPECIES OF PVTHIUM WHICH HE TENTATIVELY A881QNED TO
THIS BINOMIAL• HAWEYER, THE IDENTITY OF THE ISOLATE IS
QUE8Tf0NA9LE SINCE YOUNKIN FAILED TO INCLUDE A DESCRIPTION
OF THE FUNQUS. FURTHERMORE THE FUNQU8 WAS ORIGINALLY
ISOLATED FROM PARA81TtZED ROOTS OF WATERMELLON 8EEDLINQ8 -32-
AND WAS NEVER 1 AS FAR AS CAN BE DETERMINED, ISOLATED IN
NATURE FROM SOIL.
THE CAVERNICOLOUS ISOLATES OF PvIHIUM IRREQULARI
DIFFER FROM THOSE DESCRIBED BY MIDDLETON (1943) IN THE
AVERAGE SIZE OF OOQONIA ANO ZOOSPORANGIA• MEASUREMENTS
OF THESE STRUCTURES ARE WITHIN THE RANGE GIVEN FOR fHE
SPECIES, BUT THE AVERAGE DIAMETERS ARE MARKEDLY LARGER•
THE AVERAGE DIAMETER OF 008PORE81 HOWEVER, ARE 80M~WHAT
SMALLER THAN THAT GIVEN FOR ~REVIOUSLY DESCRfBED ISOLATES.
PyTHtUM IRREGULARE 18 THE ONLY MEMBER OF THE GENUS
WITH APLEROTIC 008PORE8 ANO IRREGULAR OOQONIA 1 WHICH IN
THE LATTER CASE, ARE NOT MADE IRREIULAR BY THE FUSION
OF THE ANTHERIDIAL CELL WITH THE OOQONIAL WALL. THE
VARIATION IN SIZE BETWEEN THESE ISOLATES ANO OTHERS
ASSIGNED TO THIS SPECIES 18 NOT CONSIDERED SIQNIFICANT
ENOUGH TO WARRANT THE ERECTION OF A NEW TAXON.
15. PYTH!UM MON08PERMUM PRINQSHEIM 1 JAHRB. Vt88• BOT., -t : 284. 1858. HVPHAE MEASURING 2.a-s.o "' IN DIAMETER, WtTH SWOLLEN
BUD-LIKI LATERAL OUTQROWTH8 1 BRANCHING FREELY: Z008PORANGIA
BRANCHED OR UNBRANCHED, UNDIFFERENTIATED FROM THE HYPHA£ 1
DELIMITED BY SEPTA, TERMINAL OR INT£RCALARV 1 MEASURING
3.3-5.0 µ IN DIAMETER, 83.3-642.6 ~ IN LENGTH; ZOOSPOR£8 6-12 IN NUMBER, to.o-ta.o µ IN DJ4METER; OOQONIA SMOOTH, SPHERICAL, TERMINAL ON SHORT BRANCHES, 11.4-20.0 µIN -33-
DIAMETER (AVERAGING 16.1 µ); OOSPORES SINGLE, PLEROTtC, 8.6-15.7 µ tN DtAMETER (AVERAGING 14.5 µ), CYTOPLASM
GRANULAR GIVING THE APPEARANCE OF SMALL GLOBULES CLOSELY
APPRESSED; ANTHERIDIA ANOROGENOUS OR MONOCLtNOUS, I PER
OOQONlUM; OOSPORE GERMINATION NOT OBSERVED.
ISOLATED ON HEMP SEED 3 TIMES FROM WET ROCKY SOIL,
CLOVER HOLLOW, CRAIG COUNTY; AUNT NELLIE'S HOLE,
MONTGOMERY Co UN TV; AND NEWCASTLE MURDER HOLE, CRAIG
COUNTY, VtRQINIA, PYTHIUM MONOSPERMUM HAS BEEN REPORTED
PREVIOUSLY AS A SAPROPHYTE IN ANIMAL DEBRIS FROM GERMANY
AND FRANCE, ON PLANT DEBRIS FROM GERMANY, IN SOIL AND
WATER FROM fRELANO ANO THE UNITED STATES, ANO AS A
PARASITE ON HORDEUM ~ULGARE, L£P10$UM SATIVUM, ~tCOTIANA
TABACUM, 0RYZA !ATIVA, PERSEA AMERICANA, RtCHARDIA
AETH&OPICA• SACCHARUM OFFICINARUM, SPINAC•A OLERACEA,
ANO ZINGIBER OFFICINALE (MtODt..ETOt4 1 1943).
ALTHOUGH EACH OF THE THREE ISOLATES INCLUDED HERE VARY
MARKEDLY ONE FROM THE OTHER, ALL THREE MAV BE ASSIGNED TO
THIS TAXON WITHOUT QUESTION• THE ABOVE DESCRIPTION
JNCLUOES THE COMBINATIVE VARIATIONS OBSERVED AND RESULTS
tN A SOMEWHAT BROADER CONCEPT OF THE SPECIES. AMONG THE
ISOLATES OBSERVED, LITTLE VARIATION OCCURS IN THE HYPHAE 1
OOQONIA 1 AND OOSPORE CHARACTERISTICS AND THESE FEATURES
ARE QUITE StMILAR TO THOSE OF PREVIOUSLY DESCRtSED t80LATES. -34-
THE ANTHERCDI~ HOWEVER, DIFFER AS TO ORlGIN. Two ARE
ANDROGENOU8 1 THE THIRD IS MONOCLINOUS. PREVIOUS INYESTt-
GATORS FAtL TO DISTINGUISH BETWEEN ANDROGENOUS ANO MONO-
OLINOUS ANTHERIDIA WITH RESPECT TO PVTHtUM MONOSPERMUM•
THE ZOOSPORANQIA OF THESE ISOLATES DIFFER MARKEDLY FROM
ONE ANOTHER ANO FROM PREVfOUSLY DESCRIBED ISOLATES. THE
MEASUREMENTS QfVEN BY MtOOLETON (1943) FOR THE ZOOSPORANGIA
OF PYTHtUM MONOSPERMUM ARE 2-5 µ x 80-170 µ.. Two OF THE
ISOLATES MENTIONED HEREIN MEASURE 5.0 µ X 93.3 µ. ANO 3.3 µ. X
83.3 µ RESPECTIVELY. THESE MEASUREMENTS ARE WITHIN THE
RANGE DESCRIBED BY MIDDLETON ALTHOUGH O(FFERtNG FROM ONE
ANOTHERe THE THtRO CAVERNICOLOUS ISOLATE IS EVEN MORE
STRICKING IN ZOOSPORANGIAL CHARACTERISTICS, MEASURING
5.0 µ tN DIAMETER AND UP TO 642.6 µ. tN LENGTH. WITH
RESPECT TO THE MODIFIED SPECIES DESCRIPTtON GIVEN ABOVE•
THESE THREE MAY SE ASSIGNED WITH CONVICTION TO PVTHIUM
"ONOSPERMUM DESPITE THE MARKED VARIATIONS IN THE
ZOOSPORANGIAL AHO ANTHEltDIAL CHARACTERISTtcs.
16. PVTHIUM NACAll ITO AND TOKUNAG0 1 J. FAC. AGR• HOKKAIDO IMPEfh UNIV., -32: 201. 1933. HYPHAE MEASURING 3.3-4.2 µ IN DIAMETER; ZOOSPORANGIA
SPHERICAL, NUMEROUS, TERMINAL OR INTERCALARV 1 SHOWING INTERNAL PROLIFERATION, MEASURING 20.a-2s.o p IN DIAMETER (AVERAGING 23.0 µ); ZOOSPORES FORMED IN A SPHERICAL VESICLE -35-
BORNe ON A SHORT EVACUATION TUBE; OOQONIA SPHERICAL,
MEASURtNG 20.0 µ IN DIAMETERJ OOSPORE APLEROTIC, SINGLE, SPHERICAL, THIN WALLED (0.8 µ), MEASURING 15.7 µ IN
DIAMETER; ANTHERIOIA NOT OBSERVED; OOSPORE GERMINATION
NOT OBSERVED.
ISOLATED ON HEMP SEED ONCE FROM MUDDY SOIL, VICKER
ROAD CAVE, MoNTGOMERY COUNTY, VtRQINfA• PVTHIUM NAGAll
HAS BEEN REPORTED fREVIOUSLY AS A PARASITE OF 0RVZA SATIV!
FROM JAPAN ANO FRO~ THE UNITEO STATES (MIDDLETON, 1943).
THIS IS APPARENTLY THE F1RST REPORT OF THIS SPECIES
OCCURRING AS A SAPROPHYTE FROM SOIL.
THE CAVERNtCOLOUS ISOLATE DIFFERS BUT LITTLE FROM THE
DESCRIPTION OF PVTHIUM NAGAll GIVEN BY MIDDLETON (1943).
OOQCNtA WERE OF RARE OCCURRENCE IN CULTURES OF THIS FUNGUS•
WHEN THESE WERE OBSERVED ANTHERIDIA WERE FOUND TO BE LACK- ING. NEVERTHELESS, f• NAQAll CAN BE READILY DISTINGUISHED FROM ALL OTHER RELATED SPECIES HAVING SPHERtCAL PROLIFERAT-
INQ ZOOSPORANGIA AND APLEROTIC OOSPORES BY THE POSSESSION
OF CHARACTERISTICALLY THIN WALLED OOSPORESe
17. PVTHIUM PAPILLATUM MATTHEWS, J. ELISHA MITCHELL Set. Soc., ...... 43: 229. 1928 •
HYPHAE LOBULATE, MEASURING 2.7-6.7 µ IN DtAMETER;
ZOOSPORANQIA FILAMENTOUS UNDIFFERENTIATED FROM THE -36-
VEGETATIVE HYPHAE• TERMINAL OR INTERCALARY 1 MEASURING
6.7 µ WIDE X 183.2 µ LONG; ZOOSPORES 8-16 PIR VE8lCL£1
MEASURING 9.9 ~ IN DIAMETER; OOQONIA SPHER&CAL TO sua-
8PHERICAL, OCCASIONALLY CATENULATE, M£A8URtNQ 12.9-15.0 ~ IN DIAMETER (AVERAGING 14.3 µ); OOSPORE8 PLEROTIC EXC!PT
AT OOQONIAL NECK, WITH A StNQLE OIL GLOBULE, MEASURING
12.3-14.3 ~ IN DIAMETER (AVERAGING t3.6 µ); ANTHERIDIA
NOT FORMED; 008POR£ GERMINATION NOT OBSERVED.
ISOLATED ON HEMP SEED ONCE FROM VET ROCKY SOIL 1
NEWCASTLE MURDER HOLE, CRAIG COUNTY, VtRGfNIAe PYTH!UM
PAP!LLATUM HAS BEEN DESCRllED PREVIOUSLY AS A SAPROPHYTE
FROM THE SOIL ANO FROM THE ROOTS OF HbROEUM VUhGARE FROM
THE UNITED STATES (MIDDLETON, 1943).
THIS ISOLATE OF PYTHIUM PAPILLA!UM POSSESSES REPRO-
DUCTIVE STRUCTURES WHICH MIA8URE SOMEWHAT SMALLER THAN
THOSE RECORO~D FOR TH! 8PECll8 IN TH! MODIFIED O!SCRtP-
TION GIVEN BY MIDDLETON (Loe. CIT.). NEITHER DOES THIS
t80LATI FORM PAPILLATE OOQONtA. IT DOIS, HOWEVER, POasrsa
ALL OTH!R CHARACTERISTICS USED IN THE SEPARATION OF ...P• PAPILkATUM FROM CLOSELY RELATED SPECIES AND MAY BE 018-
TINQUISHID FROM TH08£ SPECIES HAVING UNDIFFERENTIATED
FILAMENTOUS ZOOSPORANGIA BY P088ESSINQ APANDROU8 1 PLEROTIC
OOIPORES• -37-
18• PYTHIUM PULCHRUM MINDEN, MYKOL• UNTERSUCHe BERe R. FALCK, J_: 146. 1916.
PYTHIUM EPIQYNUM J-t6HNK 1 MYCOLOQIA, ~I 505. 1932.
HYPHAE MEASURING s.o-6.7 µ IN DIAMETER; Z008PORAN~IA
SPHERICAL TO PYRIFORM, TERMINAL OR INTERCALARY 1 23.3-33e3 µ IN DIAMETER (AVERAGING 29e3 µ), OCCASIONALLY CANTENULATE
WITH 2-3 ZOOSPORANGIA tN A SERIES; zooa~RES ABOUT 24 tN NUMBER, 13.0 µ IN DIAMETER, FORMING IN A VESICLE WHtCH ARl8£8 FROM AN APICAL OR LATERAL EVACUATION TUBE; OOQONIA
SMOOTH, SPHERICAL, t8.6-24e3 µ IN DIAMETER (AVERAfUNQ
21.s ~); 008PORES SINGLE, APLEROTIC, ts.1-20.0 µ IN DIAMETER (AVERAGtNQ 18.4 µ), CONTAINING A SINGLE LARGE OIL ILOBULE AND A SMALL LAT!RAL REFRINQENT soov;
ANTHERIDIA HVPOGYNOU8 1 USUALLY I PER OOQONIUM; 008,0R!
QERMINATtON·NOT 088ERVEDe
ISOLATED ONCE ON HEMP SEED ~ROM WET CLAY SOIL,
VICKER ROAD CAVE, MoNTQOMERY COUNTY, VIRGINIA• PYTHIUM
PMLQHBUM HA& BEIN REPORTED PREVIOUSLY AS A SOIL SAPROPHYTE
FROM THE UNtTED STATES (MATTHEWS, 1931; ffiHNK 1 1932, 1933;
MEREDITH, 1938)1 ON ANIMAL DEBRIS FROM GERMANY (MINDEN,
1916)1 ON PLANT MATERIAL FROM THE UNITED STAT!8 (MtOOLETON,
1943)1 ANO AS A PARASITE (THROUQH INNOCULATION) ON PHA!EOLUS
VUL86Rlla P18UM SATIVUM, AND !§.!.t:1A!.l (H8HNK 1 1932), AND ON
MIPICAGO 8ATIYIA (BUCHHOLTZ ANO MEREOtTFt 1 1938) FROM THE UNITED STAT!a. -38-
THE ISOLATE REPORTED HERE DtFFERS FROM THE ORIQl~AL
OESCRtPTION IN THE P089E8810N OF SMALLER Z008PORANGIA1
OOIONIA, AND 008PORE8 1 AND IN THE ABSENCE OF BOTH MONO-
CLINOU8 AND DICLINOU8 ANTHERIDIA• IN THESE RESPECTS IT
AGR!EI WITH THE DESCRIPTION OF THE FUNGUS ISOLATED BV
HgHNK (1932) AND REFERRED TO BY HIM AS A NEW SPECIES,
PJTH!UM EPtQYNUM• HOWEVER, AS MIDDLETON (1943) HAS
8U8QESTED, THE DfFFERENCES IN SIZE OF THE REPRODUCTIVE
STRUCTURES 18 NOT SUFFICIENT TO ALLOW SPECtFtC 8EQREQA-
TION. FURTHERMORE, 80ME STRAINS OF f.• PULCHRUM FAIL TO PRODUCE ANTHERIDIA OTHER THAN THOSE OF THE HYPOQYNOU8
TYPE. StNCE IN ISOLATES OF BOTH 8PECIE8 1 THE Z008PORANGIA
AND OOQONIA ARE SIMILARLY IHAPED, THE Z008PORANOIA ARE
FREQUENTLY PRODUCED IN CHAINS, AND THE OOSPORES ARE
APLEROTtC, THERE 18 APPARENTLY LITTLE JUSTIFICATION FOR MAINTAINING -P. EPllYNUM AS A SPECIES DISTINCT FROM -P. PU~Ck"UM. THE PRESENT ISOLATE IS A88tQN£D THE LATTER
BINOMlAL AND MAY 8E 018TINQUl8HEO FROM ALL OTHER SPECIES
WITH NON-PROLIFERATING Z008PORANQIA AND WITH APL!ROTIO
008PORE8 IN P088E8SINQ HYPOQYNOUS ANTHERIDIA. IT MAY AL80 BE 8IPARATEO FROM f• R08TRATUM AND f• HYPOGYNUM, TWO CLOIELY RELATED SPECIE&, BY THE PRESENCE OF APLEROTIC
008POR!S• -39-
19. PVTHJUM ROSTRATUM 6UTLER 1 MtM. DEPT. ACRe fNOIA BOT., 1: t. t 907.
HYPHAE MEASU~l~G 3.3-5.0 µ t~ DtAMETER; ZOOSPORANQIA SPHERICAL, TERMINAL, NON-PROLtFEROUS, MEASURtNQ 13.3-f6.7 µ tN DIAMETER (AVEAAGINQ 15.7 p); ZOOSPORES 16 IN NUMBER, MEASURING 5.0 X 8.3 µ; OOGONtA SPHERICAL TO SUBSPH!RICAL, TERMINAL OR INTERCALARY, MEASURING 13.3-16.7 p IN DIAMETER
(AVERAGING 14.7 p)J OOSPORES SINGLE, PLEROTIC 1 WITH A
SINGLE OIL GLOBULE; ANTHERIDIA MONOCLINOUS 1 I PER OOQONIUM;
OOSPORE GERMINATION NOT OBSERVED.
ISOLATED ON HEMP SEED ONCE FROM CLAY SOIL, EARHART'S
CAVE, MONTGOMERY COUNTY, VIRGINIA• PYTHIUM ROS~R!TU~ HAS
BEEN PREVIOUSLY DESCRIBED AS A PARASITE ON 28 SPECIES OF
FLOWERINQ PLANTS FROM THE UNITED STATES, HAWAtt, ANO
GER~ANY ANO AS A SAPROPHYTE FROM BOIL FROM FRANCE ANO THE
UNITED STATES (MIDDLETON, 1943).
THE CAVERNICOLOUS ISOLATE VARIES FROM THE MODIFIED
DESCRIPTION GIVEN BY MIDDLETON (1943) FOR PVTHIUM RO$TRATUM
IN THE POSSESSION OF CONSISTANTLV SMALLER REPRODUCTIVE
STRUCTURES. FURTHERMORE, THE ANTHERIDIA WERE NEVER SESSILE
AS HAS BEEN FREQUENTLY REPORTED FOR THIS SPECIES. THE
ABOVE MENTIONED DIFFERENCES ARE NOT, HOWEVER, CONSIDERED
SIGNIFICANT ENOUGH FOR THE EXCLUSION OF THtS ISOLATE FROM
THE SPECIES. -40-
20. PYTHtUM V£XAN8 DEBARY, JOURe SOT., 142 tos. 1876.
HYPHAE MUCH BRANCHED, MEA8URINQ 2.9 p IN DIAMETER;
ZOOSPORAN&tA 8PHERICAL 1 TEPMINAL OR INTERCALARV,MEASUR- INQ 16.7-18.3 µ IN DIAMETER (AVERAGING 17.3 µ); ZOOSPORE8 MANY IN NUMBER, MEASURINQ 7.2 µ; OOGONIA SPHERICAL, TERMI-
NAL ON SHORT LATERAL 8RANCHE8 1 MEASURING 12e2-17t2 µ fN DIAMETER (AVERAGING 15e2 µ); OOSPORES APLEROTtc, WITH A llNQL! RESERVE GLOBULE, MEASURING 8.6-14.3 µ IN DtAM!TER (AVERAGING 12.3 µ); ANTHERl01A ANDROGENOus, I PER OOQONIUMJ OOSPORE GERMINATION NOT OBSERVEDe
ISOLATED ON HEMP SEED TWICE FROM WET SANDY SOIL,
TONY 1 8 CAVE, GILES COUNTY, AND CATAWABA MURDER HOLE,,
ROANOKE COUNTY, VIRGINIA• PYTHIUM V§XAN8 HAS BEEN FOUND GROWING AS A PARASITE ON A LARGE NUMBER OF PLANTS FROM
THE UNtTED STAT!&, HAWAII, THE UNtON OF SOUTH AFRICA,
MALAYA, INDO-CHINA,1 NETHERLANDS, JAVA, INDIA 1 ANO GERMANY•
IT HAS ALSO BEEN REPORT!D IN SOIL FROM ENGLAND, IRELAND 1 FRANCE, AND THE UNITED STATES (MIDDLETON, 1943).
TH! MEASUREMENTS OF REPRODUCTIVE STRUCTURES ~F THESE
CAVERNJCOLOUI 180LATE8 ARE CONSISTANTLV SMALLER THAN THOSE GIVEN IN MtDOLETON's MODIFIED DESCRIPTION OF PYTHIUM VEXAN8,
&UT IN AU. O'THCR RESPECTS THESE ISOLATES AGREE WITH THOSE
PREVIOUSLY DESCRIBED• -41-
PYTHf UM VEXANS MAV BE OISTfNSUISHEO FROM OTHER RELATED
SPECIES HAVING SPHERICAL NON-PROLtFIROUS IOOSPORANGIA ANO
APLEROTtC 008PORES BY THE POSSESSION OF ANDROQENOUS 1
STALKED ANTHERIDIA• -42-
SUMMOtRY
A PRELIMINARY SURVEY OF CAVES LOCATED IN THE LIMESTONE
BELT OF THE APPALACHIAN VALLEY OF VIRGINIA, HAS REVEALED
THE PRESENCE OF NUMEROUS SOIL AND WATER INHABITING SPECIES OF AQUATIC PHVCOMYCETE8• A REVIEW OF THE LITERATURE INDI- CATES A PAUCITY OF INFORMATION DEALING WITH THESE ORGANl8M8•
THE CAVE& SAMPLED AND THE METHODS USED IN THIS INVESTIGATION
ARE BRIEFLY DESCRl8EOa
FROM NINETEEN CAVES, 162 SOIL AND WATER 8AMPL£8 WERE
COLLICTED FROM WHICH 33 ISOLATES HAVI BEEN OBTAINEOa IN
ADDITION A NUMBER OF MONOCENTRIC CHYTRI09 WERE 088ERVED 1
BUT HAVE NOT BEEN FURTHER IDENTIFIED. EACH SPECIES
SSOLAT!O HA8 BEEN STUDIED AND DESCRIBED UNDER UNIFUNGAL
CULTURAL CONOITtONSe THE OCCURRENCE ANO Ot8TRf8UTION OF
EACH 8P£CIE8 18 CITED AND ANY VARIATtON8 IN MORPHOLOQY OR
DEVELOPMENT ARE DISCUS8£De QUESTIONS DEALING WtTH THE
TAXONOMY AND NOMENCLATURE ARE CONSIDERED IN !ACH CAIE•
THE CAYERNICOLOU8 ENVIRONMENT MAY BE REGARDED AS A
NEW STATION FOR EACH SPECIES REPORTED HEREIN• THREE 8P£CIE8
OP' PyTHIUM, !• AFERTU.E, f• IRREQUL!RE, AND f• NftOAI 1,, ARE RIPORTED FOR THE FIRST TIME AS SAPROPHYTIC INHABITANT8 OF
THE SOIL• -43-
818L IOGRAPHY
BAILEY, VERNON. 1933. CAVE LIFE IN KENTUCKY. AMER•
Mf OlANO NATURALIST, J.i! 385-635.
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YOUNKIN, S. G. 1938. PYTHfUM IRREQULARE AND DAMPING-OFF OF WATERMELLONa. PHYTOPATH., -2a: 596. The vita has been removed from the scanned document ABSTRACT
A PRELIMINARY SURVEY OF THE FUNGUS FLORA OF CAVES. I• AQUATIC PHYCOMVC!TES • ., MARIARET MlRIHALL HOLLAND
TM&•la IVIMITTIO 1'0 TM! 8RADUATC FACULTY OF TNC VtR8tMIA PoLYTCOffMIC INITITUTI t• CANDIDACY VOR THI DIQR£E O'
...
BIOLOGY, 8eTAHY OPTION
MAY 16, f95f
8UOKl8VR•1 VtRttNIA A~3STRACT
A PR£Lt~INARY 8URVlV OF CAV~a LOCATlO fN THe LtMe•
$TOH! B!LT e~ THE APPALACHIAN VAL~EY OF VIRO&NIA1 HAI
RCYEALEO TH£ PR£$£NC£ OP NUMEROU$ SOIL AHD WATER INHAGIT-
HU~ &PtCtES OF AQUATIC PHVCOMYe£n::s. A REVIEW or tHE LITERATURE tNOtCATE8 A PAUCITY OF INFORMATION OEALJNQ
WITH Tffll~ ORQANISMI. THE CAY'S 8A~PL~D ANO THE M!TffOOS
UBEO US THUi INVUJTIQATfO.M Alli BRU.Ft..Y Olt$CRt.8£fh
FROM NINETEEN CAY!tS 1$2 SOIL ANO WATER SAMPL.£8 W£R£
COLLECT£0 FROM WHICH 33 IGOlATtS MAV~ BEEN OBTAINED. IN
AO&ITtON A -~MStR OF MOWOCENTRtC CHYTRID8 WtAE OS$£RV£0•
BUT ffAVI HOT BEOt: ~URTHER IDtnTIFIEO. EACH IPECIC$
U:JOt..ATltD HAI <ltt 8TtUH £0 ANG OESOfH $1?0 UNtU'lR UN f F'UIUa,u,.
CULTURAL CONOITIONS• TH£ OCCURRENCE ANO OISTR•BUTIOM OF
[.i\CH $t'te•~· IS C1T£0 AND ANV YMaATIOJt$ IN MORP>.J-Ot..OQY
O~ D£Y!LOPMJ!NT AR£ IH8CUS8tD• QUIUSTIOttS D!AL fNQ WITH THE
TAXONOMY ANO NOMENCLATURE ARE CON8tDaREO ·~ ~ACK eA•«·
TH£ CAV£RNtCOLOUS ENVJRONNENT MAY QI R£QARDIO A8 A
MtW STATION FOR EACH SP!Cl!S ftltPORTEO 1UtfUtU4• TffRE:E
SPtCl~ll Oft PVTHUJM, !• AF'ERTH,.E,1 f.• l!rnt:&Ui..Afl£ 1 A.NO f• MAIA••· AR£ Rl,ORT£n FOR THE FtnST TIM£ AS $APROPHYTtC tNHABtTANT9 OF TM~ 80tL.