H.h. '. II BULLETIN OF THE NATIONAL SPELEOLOGICAL SOCIETY

VOLUME 31 NUMBER 3

Contents

PARADISE ICE CAVES, WASHINGTON

MICROECOSYSTEMS IN LEHMAN CAVE, NEVADA

JULY 1969 IL H. Hobbs Ill Information for [ontrihutors BULLETIN To the Bulletin of THE NATIONAL SPELEDLDGil:AL SOl:IETY Papers in any discipline of speleology are considered for publication in the BULLETIN. The upper limit for length is about 10,000 words or approximately 40 pages of manuscript, typed double-spaced. At least one copy but preferably two VOLUME 31, NUMBER 3 JULY 1969 copies of the manuscript (typed, double-spac~d) should be submitted to the Editor, CONTENTS Jerry D. Vineyard, Missouri Geological Survey, Box 250, Rolla, Missouri 65401. Photographs and line drawings, if required, should be submitted with the manuscript. THE PARADISE ICE CAVES, WASHINGTON : AN EXTENSIVE In general, prints and line drawings will be photo-reduced to the size necessary for GLACIER CAVE SYSTEM ...... Charles H. Anderson and William R. Halliday 55 use in the BULLETIN. MICROECOSYSTEMS IN LEHMAN CAVE, NEVADA ...... N. Stark 73 For general style, see papers in this BULLETIN. Abstracts are required for all papers. Abstracts should be brief and informative. Captions are required for all Published quarterly by the NATIONAL SPELEOLOGICAL SOCIETY illustrations, and all unusual symbols used should be explained. References to the literature should be by author and date, with specific pages Suhscription rate in ef feet January l. J Q6Q : $6.00. where desirable. Literature cited should be listed in an end bibliography with entries arranged alphabetically by author's last names. For book , give total pages; for Office Addre$: journal papers, inclusive pages. Consult bibliographies in this BULLETIN for general THE NATIONAL SPELEOLOGICAL SOCIETY format. 2318 N. KENMORE ST. Photographs must be sharp, with high contrast. Because of cost, only photo­ ARLINGTON, VIRGINIA 22201 graphs essential to the presentation should be included.

All l~ne drawin~ should ~ submitted in final .form, ready for photographSI DISCUSSION of papers published in the Bulletin is invited. Discussions should be ~roduction. That !s, all lettermg should be don~ with lettering instruments, strip prmter or other satisfactory means. Typed lettermg, unless done with an electric ~000 .W?rds or less in length with not more than 3 illustrations. Discussions of papers m this is.sue should be forwarded to the editor by October 15, 1969. !ypewriter with carbon ri~bon, is. not ordinarily satisfactory. Captions will be set in type and added to the 11lustrat1ons. All drawings must be inked with India ink or a satisfactory substitute. No drafting services are provided by the BULLETIN EDITOR ~taff. ~ecause of cost! maps and other drawings should not exceed 24" x 24" except m special cases. Agam, consult the Editor in case of doubt. JERRY D. VINEYARD Missouri Geological Sllf'Vey . P~pers of tim~ly importance! reports on new techniques and procedures, or Rolla, Missouri 65401 d1sc~1ons of published papers, 1f less than 10 manuscript pages in length are considered for the Shorter Contributions Section of the BULLETIN. ' ASSOCIATE EDITORS

Before public.ation, all paper~ will be reviewed by one or more authorities in THOMAS C. BARR, j R. THOMAS L. POULSON }OHN V. THRAILKILL the appropriate ~1eld~. Aft~r review, papers will be returned to the authors for Department of Zoology Department of Zoology Department of Geology approval and action 1f required. University of Kentucky Yale University Univers;ty of Kentucky Lexington, Kentucky New Haven, Connecticut Lexington, Kentucky 't SepaTrahtes may be ord~red at the time galleys are returned by the authors to the Edi or. ese separates will be furnished at cost. RANE L. CURL RICHARD J . REARDON WILLIAM B. WHITE Department of Chem. Eng. Sawyer College Department of Geochemistrv A page charge of $20.00 per printed page will be requested from institutions University of Michigan Box 347 Pennsylvania State Universih· sponsormg research reported in articles publi' .,h d · th B 11 t' p t f the h f n · bl' . ., e m e u e m. aymen o Ann Arbor, Michigan Arcadia, California University Parle, Pennsylvania page c arge 0 owmg pu •cation will entitle th · ft f t SO f · t If an institution is unable to th .e m.s i u ion o ree reprm s. the charge will not be a co ~~f ef page charge it will ·not be made, and payment of Copyright 1969 by the National Speleological Society, Inc. n 1 ion or acceptance of the manuscript. The BULLETIN is published . . Material to be included in . quarterly m January, April, J uly and October· to the first of the month oaf givbel~ n~mber must be submitted at least 90 days prior pu 1cation. NATIONAL SPELEDLOGil:AL SOl:IETY

THE NATIONAi. SPELEOLOGICAL Soc1ETY is a non-profit organization de­ voted to the study of caves, karst and allied phenomena. It was founded in 1940 and is chartered under the law of the District of Columbia. The Society is associated with the American Association for the Advancement of Science. The Paradise l1:e [aves, Washington: THE Soc1ETY serYes as a central a~ency for the collection, preservation, An Extensive lila1:ier [ave System and publication of information relating to speleology. It also seeks the preserva­ tion of the fauna, minerals, and natural beauty of caverns through proper By Charles H. Anderson and William R. Halliday conservation practices. THE AFFAIRS of the Society are controlled by an elected Board of Gov­ ernors. The Board appoints National Officers. Technical affairs of the Society are administered by a Research Committee of specialists in the fields that relate to speleology. INTRODUCTION PUBLICATIONS of the Society include the BULLETIN published quarterly. the NEWS appearing monthly, and the OCCASIONAL PAPERS. All members re­ The existence of caves in glaciers has long ard text (Cotton, 1945). Further, in de­ ceive the BULLETIN and NEWS. been known. The specific cave system dis­ glaciated areas, long sinuous ridges of gravel cussed in this report has been a tourist at­ A LIBRARY on speleological subjects is maintained by the Society. Mate­ and silt ( eskers) have long been interpreted traction for decades (fig. 1 ) . One of us has as deposits in extensive subglacial water­ rial is available to Society members at a nominal 'Charge to defray the cost discussed briefly pseudokarstic surface fea­ courses; in retrospect it seems surprising of handling and to others through inter-library loan. tures of glaciers (Halliday, 1960, p. 112), that little consideration has been given to and these are mentioned in at least one stand- their potential speleological implication with

OFFICERS 1969 - 1970 }ACK A. STELLMACK, Presid£-nt ALLAN p. HAARR State College, Pennsylvania Exec. Vice President DAVID R. MCCLURG Delmont, Pennsylvania Admin. Vice President DONALD N . COURNOYER , Treasurer Me11lo Park, California Arlinxtnn, Virginia

CAVE FILES ALICE R. STEINKE, Custodian 1829 M iller Drive lizwrence, Kansas 66044

LIBRARY National Speleological Society Library 21 William Street Figure 1. Old postcard view of Paradise Ice Cave tour party during 1920s, showing Closter, N ew Jersey 07624 entrance to Pillar Passage and Water Passage at that time. The cave system has retreated about 100 feet in this area since that time.

NATL. SPELEOL. Soc. BuLL., VoL. 31, No. 3, JULY 1969 55 / I\ I I ''--, PARADISE GLACIER --~~ I. !~ (,' 2.8

I '

l I ~. l.' Figure 2. fj,, PARAl)ISE AREA The Water Passage at moderately high water in August, 1967. I 'I Photo by Dave Mischke. c:u='lR? 8J 0(6 S3 ( Figure 4. Trails and roads in the Paradise Ice Caves area.

some exceptions (i.e., Russell, 1893). In son, Dave Mischke, Eric. Nelson, Bill Petty, general, it has been assumed that glacier caves Ronald Pflum, Rick Rigg and Ken Severa.­ are simple structures of insignificant length Jerry Frahm periodically maintained base and content. It has been believed that these communications at the Paradise Valley road­ form as small caverns melted annually within head. glacier snouts, largely or entirely closing in With subglacial torrents stilled by winter, winter as a result of glacial flow. exploration and study were found easier than Recent studies by the Cascade Grotto of in summer, once the caves were reached. By the National Speleological Society indicate midwinter of 1967-68, almost 1~-mile of that this is not true for the Paradise Ice Cave passage had been mapped. The existence of system of Mt. Rainier, Washington. recurrent seasonal speleothems had been dis­ PROJECT FORMAT covered, and new concepts of glaciospeleo­ genesis were fast evolving. Although access The "tourist section" of the Paradise Ice was hazardous (Halliday, 1968). the Cascade Caves was visited by the Cascade Grotto as Grotto did much of its work during that win­ early as 1961. Only in preliminary studies ter. The winter of 1968-69 saw record snow during the autumn of 1967, however, did the depths, forcing the project to halt tempor­ Figure 3. extent and complexity of the caves become arily. The longest single passage in the cave Snout of Paradise lobe of the Stevens Glacier in late August, 1968. The apparent. Participating repeatedly in the is still unmapped because of the volume of large entrances at left formed after July, 1967. project were Charles Anderson, Edith Ander- streamflow in summer (fig. 2). 56 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31, NUMBER 3, }ULY 1969 !Qi

ENTRANCE

PARADISE ICE CAVES PARADISE GLACIER MT. RAINIER NAT'L. PARK, WASH. SUICIDE PASSAGE SURVEY llY CASCADE GROTTO. NATIONAL SPELEOLOGICAL SOCIETY OCT 1967 COMPASS & CHAIN II. RIGG E &C . ANOEllSON 8. PETTY

- GENERAL - Figure 5. PASSAGES IN ICE WITH STREAM COBBLE FLOOR EXCEPT AS Entrance to the Pillar Passage and Water l NOTED. PASSAGE SIZE, LOCATION OF Passage at the beginning of the tourist ~ ENTRANCE OPENINGS TO SURFACE, ANO PASSAGE ACCESSIBILITY season in June, 1967. SUBJECT TO CHANGE . I 0 .0

Water Passage

fNTllANCf ENTRANCE ··-~ ', • J MAIN ENTRANCE t::.:."19... ~,.!•,i•:.,

SCAll \IOO

I • I ! ...... ~­ •cw•tlDll ,ttt

Figure 7. The Paradise Ice Cave system in October, 1967. A- J PARADISE ICE CAVES SUICIDE PASSAGE PARADISE GLACIER MT . RAINIER NAT'L. PARK, WASH.

SUllV!Y llY CASCADE GROTTO, ~~!~OA~~L& S~~~~NOLOGICAL SOCIETY JULY 196 7 0 II . RIGG f.&C. ANDERSON B. PETTY LOCATION contacted well in advance, to ascertain current conditions and regulations. The Paradise Ice Caves are in the Para­ - GENERAL - From the Paradise R_anger Station, an PASSAGES IN ICE WITH STREAM dise-Stevens Glacier on the southeast side of COBBLE FLOOR EXCEPT AS Mt. Rainier, a dormant 14,41()-foot volcano excellent trail leads ~-mile eastward to NOTED. Myrtle Falls. Thence, ·the Mazama Ridge PASSAGE SIZE, LOCATION OF in the Cascade Mountains of Washington OPENINGS TO SURFACE, ANO (fig. 3). The altitude of the caves is approx­ trail continues across the Paradise River just t PASSAGE ACCESSIBILITY imately 6,500 to 6,900 feet. below Sluiskin Falls and up Mazama Ridge SUBJECT TO CHANGE. to a junction with the Skyline Trail at a I A few decades ago, the Paradise and Water Passage point 1,34-mile from the ranger station. At Stevens Glaciers were distinct entities with a this junction, visitors proceed left along the common source. The Paradise Glacier has Skyline ·Trail, up Mazama Ridge, for about .:::;:UNSUIM"'t'lDll'ASUGI receded rapid}y, however, and some consider ~-mile. At this point, a rustic sign "Ice - lltOCK WAll r;;:9 LAROI N'.ICM it to have disappeared entirely so that the Caves" points right. Beyond this point, the • ICl FLOOR ~f.MA caves now should be considered to lie entirely (~;:1--· TO SIJMllC[ route is informal ·-(fig. 5) and often fogbound . ! ' • within a southwest lobe of the Stevens Glacier. In early summer, flags are emplaced in the The caves are approached from the Para­ snow to mark the route. Figure 6. dise Valley visitor center roadhead (fig. 4). The winter route requires special permis­ The Paradise Ice Cave system in July, 1967. The summer route requires no special arrange­ sion and parties must sign in and out at the ments if only tourist areas are to be visited, ranger station. It involves a more direct but otherwise the chief park ranger should be much steeper ascent on snow to and .across 58 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31, NUMBER 3, }ULY 1969 59 ENTRANCE ~-..

trii>Dllim~1... _ ..... /. Breakdown I , 1 ~ _ _,ENTRANCE ! ' J,

SUICIDE PASSAGE

PARADISE ICE CAVES PILLAR PASSAGE PARADISE GLACIER MT. RAINIER NAT'L. PARK, WASH. - GENERAL - PASSAGES IN ICE WITH STREAM SURVEY BY CASCADE GROTTO ~ NATIONAL SPELEOLOGICAL SOCIETY OCT . 1968 COBBLE FLOOR EXCEPT AS COMPASS & CHAIN NOTED. R. RIGG E & C. ANDERSON 8 . PE TTY PASSAGE SIZE, LOCATION OF OPENINGS TO SURFACE, Mm PASSAGE ACCESSIBILITY SUBJECT TO CHANGE.

, ""• '.. MAIN ENTRANCE ::::: Ufri!SUAVEYED PASSAGE Figure 9. JJJJJll ROCK WAll E!:J' LARGE ROCK

.IClflOOll ~JIUM SCALE I 100 Recently collapsed section of the Big Room in August, 1968. , ...... OHM ... TO SURfAC( \i".1 \. .

Figure 8. •; The Paradise Ice Cave system in October, 1968. the Skyline Trail via the "Golden Gate" areas, especially where constricted, but no above Myrtle Falls, thence down the steep air flow studies have been made. Rudimen­ wall of Paradise Canyon a few hundred yards tary temperature studies have been made at from the caves. Travel is routinely by com­ three points: 100 feet inside the cave in the pass in thick fog and blowing snow. Water Passage, at the Great Pillar and in Suicide Passage near its confluence with the DESCRIPTION Water Passage. Recorded air and water tem­ peratures have varied seasonally from 32°F As presently known. the Paradise Ice Caves to 36°F, with water temperatures one or two inchj~e at least 12 ,000 feet of passage in a degrees higer than ?..ir temperatures except branchwork pattern. Passages totalling about near the entrance u11 October 7, 1967, when 7,500 feet have been mapped to date, with the water and air temperatures respectively most of the remainder in the Water Passage were 36°F and 32°F. Only one water tem­ Figure 10. (figs. 6, 7 and 8). Collapse has segmented perature reading of 3 2 °F has been obtained. Now-collapsed section between the Flute the system during the course of this study Room and the entrance of the Pillar Pas­ (figs. 9 and 1O). While some parts of the Streams of various sizes are present in most of the passages in summer (fig.2). Marked sage in October, 1967 at an intermediate caves are little more than crawlways, most stage of segmental collapse. consist of glorious corridors a dozen or more variation of streamflow occurs, with some feet in diameter (fig. 11). Locally, the active freezing within the cave, but at no width and height are much more (fig. 12). time has all streamflow been found halted. Atypical eskers-in-development are present in Strong air currents are perceptible in many the stream beds; most include large stream

60 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31. NUMBER 3, }ULY 1969 61 Figure 11. Scalloped walls and ceiling in the Pillar Passage.

THE NATIONAL SPELEOLOGICAL SOCIETY 62 BULLETIN VOLUME 31, NUMBER 3, JULY 1969 63 Figure 14. Boulders and drifted snow near rear entrance of the Big Room in midsummer 1967. Figure 12. The Big Room in July, 1967.

boulders and cobbles in addition to finer sedi­ ments (figs. 13 and 14). Most of the system occurs beneath the glacier but some sections are floored by gla­ cial ice. Although far from homogeneous, the glacier ice is largely glistening white or blue­ white. Bedding and some impurities (fig. 15) can be observed. Speleogens and speleothems are well developed. These features and the effects of local illumination through the glacier render the caves entrancingly beauti­ ful. The play of color and light on the scalloped ice walls ranges from delicate green to deep ultramarine. The most prominent speleogens are large scallops or flutes (figs. 1, Figure 13. 2, 11, 12, 13 and 14), often more than a meter in width. Superficially resembling on Misery Crawl area. Note banding of ice. a grand scale the stream flutes of limestone Stream cobbles form a small island. caverns, they are probably aerogenic. Locally but not univer$ally, their shape and size Figure 15. appear modified by impurities or intraglacial bedding and related features. Fallen flakes at rear of Big Room in April 1968. Note banding (impurities) in ice at left rear. 64 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31, NUMBER 3, }ULY 1969 Figure 17. Entrance to the Pillar Passage in August 1968. Entrance to the Water Passage is also visible on the extreme right. Some upper entrances are present in the berg- schrund at the upper left.

Figure 16.

Medium-sized flake (right) near complex column in the Bi~ Room. The snow in the foreground drifted into the cave from the lower entrance. Another flake shows in the center distance. A small frozen waterfall can be seen along the surface of the complex column near the left edge of the picture.

Coalescence of scallops in irregular pat­ types of orifices connect the caves and the terns sometimes produces deeply undercut, surface of the glacier. Several of the rounded, hanging "flakes" of ice centimeters or meters domepit-like swallets termed moulins inter­ long (fig. 16), often weighing many tons. sect the caves (fig. 20) A crevasse about 4 Collapse of these flakes is a specific hazard feet wide and an estimated 500 feet long of exploration; some cannot be discerned intersects the Water Passage. As indicated, until one has passed beyond them and rela­ newly-formed collapse sinks have separated tively minor vibrations sometimes cause them the system into multiple caves during the to fall. course of this study; most occurred between Entrances to these caves are located at the August 19 and Spetember 30, 196 7. Figure 18. snout of the ice lobe and also in the bergs­ Large columns of glare ice form season­ Entrance to the Water Passage in early September 1967. The Tatoosh Range chrund (figs. 1 7, 18 and 19). Three other ally at the base of active moulins, and below is in the middle distance and Mt. Adams in the background.

66 THE NATIONAL SPELEOLOGICAL SOCIETY

BULLETIN VoLUME 31, NuMBER 3, JuLY 1969 67 Figure 21. The Great Pillar in winter, with drifted snow at the base.

l J Figure 22. The Great Pillar in late June.

Figure 19. Some view as Figure 18, but in early March 1968.

Figure 20. Looking up a moulin above the Great Pillar. The glacier is about 20 feet thick at this point.

68 Tl:IE NATION AL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31, NUMBER 3, JULY 1969 69 Figure 23. Figure 24. Water dripping through moulin at site of the Great Pillar. A small flake is Column in the Flute Room in March ] 968. visible in the foreground. A larger flake barely discernible in the middle distance is about 300 feet long. Tourists are seen in the entrance of the Pillar Passage in the distance. other drip points to a lesser degree. Most of compacting snow causes slow plastic flow are largely or entirely melted by mid-June of toward the wastage area. This process even­ most years (figs. 21, 2 2, 23, 24). Tapered tually delivers all parts of the glacier and its stalactites are fairly common, and participate cavities to the glacier's melting snout whether in column formation. Tubular stalactites have the snout is advancing, retreating or station­ been observed in the Water Passage. Ribbon ary (fig. 26). draperies form locally but are quite transient. The Paradise lobe of the Stevens Glacier Curious ice helictites are even more evanes­ is in a state of old age, and is in rapid cent. They have been observed only at the retreat. Almost all the seasonal snow which lower entrance of the Water Passage (fig. mantles it melts annually. The firn limit is 25) and on the surface of one column. almost at the upper end of the glacier. Com­ paratively little glacial flow thus appears to GLACIOSPELEOGENESIS be present to counteract two forces which enlarge glacier caves: stream action and Figure 25. Whether a glacier advances, retreats or circulation of warm air. Transient helictites in the Water Passage remains stationary depends on increases or As in the case of many limestone caves, entrance in April 1968. decreases in snowfall in the area of accumula­ the size of the cavern passages appears dis­ tion, with regional temperature changes also proportionate to the present streams on their a factor. floors, and some are presently stream-free. Unless virtually all the snow cover melts Some major passages show no evidence of from the surface of the glacier (in which case stream action on the ice walls, while others the glacier will soon disappear), the weight show typical vadose speleogens. Despite

70 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOI.UME 31, NUMBER 3, JULY 1969 71 Microecosystems in Lehman [ave, Nevada

By N. Stark

ABSTRACT Annual Surplus of Accumulation Over 70 species of plants and animals have been found and most of these identified from Lehman Cave near Baker in eastern Nevada. These organisms play Annual Wastage parts in 12 microecosystems, some of which are "pseudo" or lacking producer organ­ of Ice Figure 26. isms, and "terminal" or destined to cease to exist once the organic food supply left Diagrammatic side view of glacier. by former trogloxenes is exhausted. The microecosystems differ in degree of moist­ ness, dept of "soil" development, the presence or absence of light, and the numbers noteworthy fluctuations in streamflow, it staff of Mt. Rainier National Park. Partic­ and types of organisms they support. A strong controlling factor in an otherwise seems doubtful that tl:ie larger passages could ular thanks are due to Superintendent John quite stable environment is the extremes of temperature, air relative humidity, air be filled either by present floodwater invasion A. Townsley, Chief Park Naturalist Norman movement, and the drying power of the air caused by turning lights on or off in or by greater past runoff - or by flow when Bishop and Chief Ranger Joseph L. Orr. We various reflecting positions. Lights boxed on three or four sides produce conditions the passage was largely filled with esker. also wish to express our inestimable obliga­ too severe at 0.1 m distance for plant growth, but deep reflectored or open bulbs Rather, it appears that each passage was tion to all the members of the National Park ( 100 w) produce cool, moist conditions, favoring moss while shallow reflectored initiated by running water, but that initial Service, Mountain Rescue Council, Cascade bulbs favor algae on moist sites at 0.1 m. streamways admitted enough circulating air Grotto of the National Speleological Society to cause melting which produced most of the . and others who assisted during the 1968 res­ present volume of the cave. Some warming cue operation near the "Golden Gate." INTRODUCTION existence. The influence of the more than of circulating air by heat transfer from sub­ Simple ecosystems are prized by ecologists 50,000 visitors to this National Monument glacial streams may be a significant factor, because they are easy to study and still each year cannot be overlooked. but tentatively it is felt that most of the heat REFERENCES retain many of the factors and forces at work The Cave extends into the base of Wheeler transfer results from ingress of seasonal air in more complex ecosystems. Lehman Cave Cotton, C. A. Peak for a lateral distance under one mile above the freezing point. 194 5 Geomorphology: Whitcomb & in eastern Nevada at the base of Wheeler (fig. 1, Mohr and Sloane, 1955). The vege­ Photographs of the past few decades indi­ Tombs, Ltd., Christchurch, New Zea­ Peak ( 13,063 ft.) is a limestone cave com­ tation consists of Pinyon-Juniper, sagebrush cate that retreat of the glacier has caused land, 4th Ed. plex with a variety of simple microecosystems. ( Artemisia tridentata), rabbitbrush (Chryso­ loss of co :1siderable length of the cave. Plas­ The Cave is 10 miles west of Baker, thamnus nauseosus), and a few abundant tic flow also probably serves to reduce the Halliday, William R. Nevada at about 6,800 ft. elevation (Lat. herbs such as Penstemon palmeri and grasses. size of the caves. Yet direct observation dur­ 1960 Pseudokarst in the United States: 39 ° N., Long. 114° 50' W.) . These caves Natl. Speleol. Soc. Bull., v. 22, pt. ing this study suggests that collapse from are particularly interesting because they have ACKNOWLEDGMENTS 2,P. 112. vestiges of earlier microecosystems which over-enlargement is currently more important I thank the National Park Service at in destruction of the caves. 1968 The mountains don't care: Natl. were active before Lehman Cave was lighted, SpeleoL Soc. News, v. 26, n. 10, p. and a different group of organisms which Lehman Cave for supporting this research. I am grateful to Dr. Harold Humm of the 152, October. exist because of the lights. The flora and ACKNOWLEDGMENTS fauna of the Cave is not considered to be University of South Florida for verification This study was possible only because of Russell, Israel C. typical of caves in general and there are no of algal identifications, and to Dr. L. E. special permission and assistance rendered by 1893 Malaspina Glacier: Jour. Geology, troglobites, but it has a potpourri of organ­ Anderson of Duke University for moss ident­ the National Park Service and especially the v. 1, pp. 219-245. isms normally living elsewhere and generally ifications. Drs. George C. and Jeanette showing no specific adaptations to caves Wheeler identified the animals. The work 1117 36th Ave. East (Mohr and Poulson, 1966). This paper de­ was conducted from the mobile laboratories Seattle, Washington 98102 scribes the types of microecosystems which (Went, 1968) of the Desert Research Insti­ exist in Lehman Cave and some of the tute which made it possible to live near the • Manuscript received by the editor problem in the field. 5 March 1969 microclimatic factors which govern their NATL. SPELEOL. Soc. BULL., VoL. 31 , No. 3, }uLY 1969 73 72 THE NATIONAL SPELEOLOGICAL SOCIETY 0 M 0 ci c-.i N

0 METHODS percent of the lights had rock as a part of c-.i the substrate, while soil occurred at 25.0o/o M Careful observations ·and collections of and organic debris at 58.0o/o of the lights. G; 0 0 living material for identification were made The substrates were graded objectively as wet .J:2 ci at 200 of the existing lights along the .trails. to moist ( 48.0%), slightly moist ( 32 .0%) E ""'1:1' Q) c Dark areas were also studied. The survey and dry to damp ( 20.0%). > a u 0 ll') 0 of lights included: z0 0> ·- cxi ci 0 It') "() a) light wattage, position, and reflectance The Cave was divided into the entrance iA area which is dry to damp with the least Q) b) algae, mosses, ferns, other plants and >a algae, few mosses, no flow dusti and the most ll') 0 their conditions u old dung; the mid-section which has the most c ci c) visible evidence of fungi moisture and dripping, abundant algae and a N E mosses; and the final section which has less ,...ll')ll')Oll')Oll')Or.....oMO d) flies, other insects, larvae, spiders, ..c " 0 dripping, few mosses or algae, and the least ....Q) M oO M 0 M 0 M 0 M 0 N 0 cxi Annelids, mammals (signs) 00 0 0 0 0 0 "() recent animal signs (Table 1, fig. 1). .: e) the nature of the substrates i. e., part or Ill The entrance area is dry because the inflow Cl all rock, soil, wood, dung f of cold, relatively dry air during winter a f) the relative moistness of the substratum makes part of the entrance a variable temper­ ~.. g) the minimum and maximum distance ature zone. The middle section is protected a 0 0 from dry air movement and has the most "'CS from the lights to plant life It') active dripping and the best development of N I h) debris brought in by man. Cl) Ill microecosystems. These first two sections "'CS c"'D 0 0 0 0 c C·- N Data were compiled to show what percent­ were lighted in 1939-40 and have the health­ a 111(- ci age of the lights had various organisms and iest plant life. The last section (Talus Room) :? ..c conditions. Tentative food chains were dia­ was lighted in 1960. The amount of plant grammed for the various microecosystems. life which has become established in eight ~ 0 Five different representative light positions years is so small that most visitors are not 0 N were studied for microclimate with the lights aware that there are any plants there. on and off to better understand the effect of Collembola are scarce in the Talus Room ~ because they require an area of concentrated .. microclimate on the organisms. Air tempera­ c lft M N 0 tures, relative humidity, the evaporative plant life of about 4 x 4 cm. They do occur s -.! "() a ii: cxi power of the air, air movement, and light where there are fungi and no green plants. :0 intensity were studied at different distances a Dark areas with d_ust and lint, support fungi, Ill with lights on and off. c ea 0 "()""'1:1'0 00 00 M It') M 0 and occasionally Protozoans. Plant life in 0 e»- ·~ 1§ J cxi ci cxi ci c-.i this Cave is likely to remain in small areas =ij u N ""'1:1' N M SPECIES AND ENVIRONMENT c near lights. In 28 years, the largest plant to 0 Of the 200 lights studied, 77.5'}'o had algae, invade the Cave and stay is a smalll fern "' 0 0 ll') 0 "'CS It') ~ N O 20.5% had mosses, 15.0o/o had visible evi­ (Cystopteris jragilis var. fragilis). Limited c amounts of soil, low light intensity, and a " 0 dence of fungi, 30.5% had Collembola, and Ill possibly a deficiency of nitrogen and other E 31.5 % had various types of dead or live flies Ill (Table 1). A material which I called "flow elements is preventing the establishment of ·c: ll') 0 0 higher plants near lights. The energy level a ci dust" (fig. 2) appears as rivulets of mud or O> N dust on moist or dry stalagmites. Flow dust for the Cave ecosystem as a whole is ex­ .. tremely low. This may, in part, explain why 0 on moist lighted sites was composed of dia­ .. Cl) 0 a It') 0 0 toms, minerals and organic debris, and was the permanent organisms in the Cave are so O> small. All plants living in the Cave are >. found at 24.0% of the lights. This material a.. " N transient mammals bringing in seeds has not "' .... Ill as clayish-slimy vermiculations. An oligo­ Cl) chaete worm which appears to feed on the resulted in the establishment of any higher ..: Cl 0 diatoms occurred at only 10% of the lights. plants, possibly because the shortage of food 0 zo zo 0 0 0 Ill O~ 'cP. 0~Z.,,pZ .....0oz~o Gi Many ecological niches are not occupied in during the spring when young mammals are ~ tJ .iJ tJ U' .iJ ,..._ I 0 - > N N M M {!_ -c( the Cave after over 28 years. Ninety-eight born causes all usable food to be eaten. ~ BULLETIN VOLUME 31, NUMBER 3, jULY 1969 7'j 74 THE NATIONAL SPELEOLOGICAL SOCIETY ORGANISMS FROM LEHMAN C.WE from chemical reactions in the absence "'f light. Leptothrix sp. - iron bacteria. A. Producers-Autotrophs synthesizing carbo­ hydrates from C0 using light ener1cy. C. Primary Consumers - utilize plant food 2 for energy Algae Class Myxophyceae - blue green algae Members of the Phyla Flagellata, Sarco­ Anacystis montana (Lightfoot) dina, Ciliophora, Rotifera, Annelida, and Arthropo·'a were identified by the Drs. Drouet and Daily forma montana Wheeler. Oribatid mites, Collembola (three (Drouet and Daily) species Entomobrya marginata Tullberg, det. Schizothrix calcicola (C. Agardh) D. L. \Vray, USNM; and two others uni­ Komont dentified), three species of flies ( Bradysia sp. Oscillatoria Vaucher sp. det. R. J. Gagne, USNM; M egaselia sp. det. Anabaena Bory sp. W. W. Wirth USNM: Psychoda sp.), and TALUS one beetle were identified. A Psyllipsocidae, Coccochloris Sprengel sp. Psyllipisocus ramburii Selys-Longchamps ROOM Class Chlorophyceae - green algae (det. E. L. Mockford, Ill.) is from a family Mugeotiopsis calospora Palla of known cave-dwellers which feeds on fungi, Chlorococcum humicola (Nageli) pollen and dead insects. Of the larger trog­ Rabenhorst loxenes. only Peromyscus maniculatus (deer mouse) and Eutamias sp. (chipmunks) were Protococcum viridis Agardh seen ( G. Ralston) . Nannochloris Naumann sp. D. Secondary Consumers - eat herbivores Roya anglica G. S. West Sixteen species of protozoans were tenta­ Tentative identification tively identified, but many of these are also Cosmarium corda sp. primary consumers. The pseudoscorpion Chlorella vulgaris Beijerinck M icrocreagris gran dis ( det. R. R. Bridgeman) and four species of spiders belong to the Schmidle Coccomyxa dispar carnivores. Of the insects, there were fleas Palmella miniata Liebl. 0 30 60 ( Dolichophysillidae), bat flies ( Streblidae) , .c:::::====i:======:;::J Class Bacillariophyceae and Culicoides sp. ( det. W. W. Wirth SCALE IN METERS USN:\1). A moth of the family Tineidae, N avicula - 6 species of naviculoid Figure 1. diatoms probably Amydria sp. (det. D. R. Davis, USNM) of uncertain food habit was fairly Coscinodiscus Ehr. sp. Ma~ of Lehman Cave. The entrance section (1) extended from numbtrs 1-6 abundant. Bats were scarce and thought Section 2 (2a, 2b) extends from beyond point 6 to beyond point 1 O and Section Mosses to be mainly insect eaters. 3 {3a, 3b, 3c) extends beyond point 10 to a few yards beyond point 19 Physcomitrium sp. E. Reducers - Large numbers of fungi occur {see Table 1). Campylium chrysophyllum (Bird) J. througho·.It the Cave on moist walls, dead Lange organic matter, dead flies and on stalactites. Pohlia wahlenbergii (Web & Mohr) Bacteria were common in the pools which and animal life may have been transported systems known to exist in Lehman Cave. Andr. dry during winter. The slime bacterium through the Cave. If fish lived in the Cave These are called "microecosystems" because Amblystegium serpens (Hedw.) BSG Dictyostelium sp. occurs on waUs and Ste­ at earlier times, no evidence of their past has they are small in areal extent. Some are monites sp. the slime mold is common on been found and no fish Jive there today, also "terminal" such as the dung ecosystems 3 others unidentified, moss species had wood. Chytrids live on dead algae and poUen. probably because the low energy level of the since no new dung is being added. Once the to be determined on vegetative characters Pack rats were once common in the Caves entire system is inadequate to support fish. dung has no food supply left to support since no sporophytes were present, and cap­ As the Cave became drier, pack rats, bats organisms, the dung microecosystems will sules are rarely formed . before the lights were installed. Deposits of pack rat dung over a foot deep remain as ~nd other organisms spent part of their cease to exist. Others are "pseudo" because hves there. After the lights were introduced they depend on food brought in from the Liverwort - 1 species evidence of their importance in the Caves in the slow colonization by plants and micr~ outside and lack any producers to perpetuate Ferns the past. Beetle galleries and frass are com­ mon under old dung and another type bur­ scopic animals from spores began. The pack food production within the Cave (dung). Cystopteris fragilis (L.) Benth. var. rows into the pellets, but no beetles are known rats left, and the present trogloxenes do not Even the true microecosystems are not fragilis to be active in the dung of the Caves today. concentrate their dung. permanent since two or three weeks with Asplenium sp. the lights left off or herbicide treatment During the early solution stage when would damage the green plants and create B. Producers - Chemotrophs - derive energy flowing water was eroding the Caves, plant Today there are 12 types of microeco- unsightly brown patches. 76 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31, NUMBER 3, }ULY 1969 77 naviculoid diatoms as the producers and 8. Dry dung (moisture under 8%) in dark Oligochaete worms as consumers with bac­ with fungi and bacteria (beetle galleries and teria and fungi as reducers. frass may no longer be active) , terminal and "pseudo." Oligochaete worms Collembola ~ t )Reducers fungi,tbacteria - -.. Navciuloid or Centric diatoms fungi:,,bacteria ...... , nut~ients ..... _,,,nutrilnts t Dry dung, dark I Moist rock,t light t Lost

S. Moist rock in light with algae, some diatoms, no flow dust. Collembola, fungi, and bacteria are usually present with mouse or bat dung (small amounts). Rock may have 9. Moist dung in dark (moisture 12 %+), mosses and liverworts. with larvae and Protozoans feeding on fungi, bacteria - terminal and "pseudo," Collem­ ollembola, flies ? bola may be present. t Pseudo~rpiois, bats larvae, Colle1ola, Protozoans ~educers I Reducers Fungi, bacteria { nutrients,'. Algae, rock mosses, dung""nutrients/ , _ _,< f":.... _.,,,. t 'nutrients Moist rock, light soil) Tno Moist Jung, dark \ ~ 6. Moist rock in light with 0.1 to 1 cm or Lost more soil, with algae, any of the mosses except the rock forms, Collembola, arachnids, flies, Figure 2. fungi, bacteria, mouse and bat dung. 10. Aquatic habitats in the dark with dead "Flow dust," a material composed mostly of diatoms with bits of s~il may. be Arachnids Pseudoscorp1ns, bats flies supporting fungi and chytrids, floating dry and no longer active, brown and moist (dead), or green and a.live. Ol1go­ / •.tsyllipsocids, nr.s on water surfaces, dead organic matter settles chaete worms live within the flow dust and presumably eat diatoms. ~mbol• + to the bottom and provides food for bacteria, Reducers ~Algae, mo ses, dung Reducers Protozoans, chemotrophic iron bacteria are The microecosystems of Lehman Caves are: 3. ''Soda straws," wet, with fungi, bacteria, \ ~ t . I often present (not terminal as long as organic or if in lighted areas there may be algae, n'litrients/ '-...... nutriellts debris is added). 1. Bare rock, slightly moist, in the dark some energy may come from organic or ,_ ---- with fungi, bacteria, living and dead insects, Moist rock 0.1 - 1 cm soil, light inorganic compounds in the water. Water, dark, dead flies, fungi lint, pollen, dust. This is the habitat of Dictyostelium sp. 7. Moist rock in light with soil 1 to many Dead orga'!;;c deb~s ,,.- Dictyostelium, fungi nutrients centimeters deep, with algae, mosses. and ferns. Collembola, arachnids,. flies, fungi, i fungi, Protozoans /-~algae~ fungi, bacteria nutrienrs t bacteria, dung (recent). / b acteria. "' B~re rock-dark; with lint, dust, pollen II t ' II · · \ Bat flies Chytrids fungi, bacteria light / Reduce:S-:- Arachnids \ 2. Moist rock in dark with fungi, Proto­ nutrien/ 11 zoans, bacteria, living on moist dust of dead t / 'nutrients ~ P~doscorpions, bats ~ Prot~zoans insects, pollen, lint, bat and mouse dung. \ t wet "soda straw" \ Collembola, flies, mites Protozoans, (debris feeders) nu~1ents\ . t Chytrids, fungi, U~teria Algae, mosses, fers, protonema, dung Bare rock-dark-moist ""'"dust", bat or 4. Moist rock in light with brown (dead) 11. Aquatic habitats (pH 7.0-8.2) in light mouse dung and green (live) flow dust containing minute Moist rock, 1 + cm soil, light with algae, dead organic matter such as flies, BULLETIN VOLUME 31, NUMBER 3, JULY 1969 78 THE NATIONAL SPELEOLOGICAL SOCIETY 79 I ~ bats, wood, algae, dung providing food for extent and concentration (4x4 cm) to sup­ Protzoans, bacteria, fungi, chytrids. May port them. They are most abundant 30 to have iron bacteria. 0 0 0 00 70 cm from the lights and rarely occur on Water, light steep, wet flowstone. MICROCLIMATES AROUND LIGHTS ~ Reducers The positioning and wattage of individual / Algae - nutrients ! lights strongly influence what lives in the Dead organic matter ~ Protozoans, bacteria Caves. The most severe microclimate occurs around a light which is boxed on four sides fungi resulting in air temperatures at 0.1 m of 0 0 0 0 0 00 0 0 0 0 0 N 22 .5 ° C after 10 minutes of light (an in­ 0 0 N -0 00 00 0 &t') N M N -0 M""" '°' N 12 . Decaying wood (moist) in dark or crease of 11.3 ° C in 10 minutes), with a loss light with a variety of Basidiomycetes includ­ of 89.9% of surface moisture in four hours ing Marasmius sp., slime molds (Stemonites or 0.928g/ hr. (Table 2) . No plant life sp.), flies, Collembola, bacteria, arachnids. occurs at this light because of the sudden Fruiting bodies have Diptera larvae (termi­ temperature and humidity fluctuations and nal). the drying power of the air when the lights are turned on. The relative humidity around Arachnids the boxed light dropped from 98 to 58 % in Collembola, mef'--+Re)ucers 30 minutes with the light on. r- N 0.,... The next most severe habitat occurred at Moo ' nutrie'nts ~~ 0.1 m from the light boxed on 3 sides (Table Bacteria, fungi , Stemonites, BasidiomycetesJ. 2). The temperature range there was only 8.2 ° C, but the drying power at 0.1 m from others J LosT the reflector nearly equalled that of the light Decaying Wood boxed on four sides. The increase in relative humidity occurs when slow air flow and heating allows rapid evaporation. Most transients (mice, chipmunks) are All other light positions studied had lower omitted from the microecosystem diagrams temperature ranges, less evaporation and of since they interact to some degree with most considerable plant life at 0.1 m to 2 m dis­ .r. &t') "'? ,... N ~ 0· tance. The cool temperatures ( 10.2 - l 1.0°C) ~ c-> o 0 organisms present. The differences among microecosystems 1, 2, 3, and 4 are the amount produced at the deep reflectored light were CD > of moisture present progressing from slightly ideal for moss growth, while the warmer 0 c u .o moist to wet. Microecosystems 5, 6, and 7 temperatures ( 15.1 °C ) and fast drip rate at ~ the shallow reflectored light favored algae. c .r. ... differ mainly in the amount of "soil" devel­ 0 opment and the types of plants which each E co 0 -o -o o· """ It) It) The air temperatures around the open .r. ci 0 0 0 ci can support. Microecosystems 8 and 9 differ ..! light (no reflecting surface close by) fluctu­ .,, in their moisture content while 10 and 11 ated only 2.9 °C in 10 minutes with the lights c differ in the presence or absence of light. on. A light over a free water surface caused ::::t CD Collembola occur wherever it is moist and no change in water temperatures after 30 ~ u 0 c- there are fungi, algae, or mosses of sufficient minutes of continuous light. 0 Ill o E .! 't; :- 0 0 .5

lu i

c ... G> en &~ 80 THE NATIONAL SPELEOLOGICAL SOCIETY BULLETIN VOLUME 31 , NUMBER 3, JULY 1969 81 REFERENCES

Mohr, C. E., and T. L. Poulson (Lake of Como), Italy; Memoria 1966 The Life of the Cave: McGraw-Hill V della Rassegna Speleologica Ital­ Book Co., New York, 232 p. iana. Mohr, C. E., and H. N. Sloane 1955 Celebrated American Caves: Rutgers Univ. Press. Parenzan, P. Went, F. W. 1960 Concerning the clayish-slimy forma­ 1968 The mobile laboratories of the Des­ tions termed vermicular: Intnatl. ert Research Institute: BioScience. Symposium on Speleology, Varenna v. 18, pp. 293-297.

Laboratory of Desert Biology Desert Research Institute Reno, Nevada 89507 Manuscript received by the editor. 17 February 1969

82 THE NATION AL SPELEOLOGICAL SOCIETY