Cave Notes

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Cave Notes
Series Title:
Caves and Karst: Research in Speleology
Alternate Title:
Caves and karst: Research in speleology
Cave Research Associates
Cave Research Associates
Tumbling Creek Cave Foundation
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Subjects / Keywords:
Geology ( local )
serial ( sobekcm )


General Note:
Summary of past investigations on typhlotriton spelaeus, the grotto salamander / Ronald A. Brandon -- Water level planes in caves / Arthur L. Lange. Cave Notes(vols. 1-8) and Caves and Karst: Research in Speleology(vols. 9-15) were published by Cave Research Associates from 1959-1973. In 1975, the Tumbling Creek Cave Foundation compiled complete sets of the journals in three volumes. The Foundation sells hardbound copies of the material to support its activities.
Open Access - Permission by Publisher
Original Location:
Windy City Grotto Collection, 1961-2013
Original Version:
Vol. 4, no. 2 (1962)
General Note:
See Extended description for more information.

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University of South Florida Library
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K26-00642 ( USFLDC DOI )
k26.642 ( USFLDC Handle )
13713 ( karstportal - original NodeID )
0008-8625 ( ISSN )

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CAVE NOTES A Review of Cave and Karst Research Volume 4. No.2 Marchi April, 1962 Typhlotriton spelaeus (female with eggs); Wet Cave, Shannon Co., Missouri SUMMARY OF PAST INVESTIGATIONS ON TYPHLOTRITON SPELAE'US. THE GROTTO SALAMANDER by Ronald A. Brandon (Cave Research Association) Department of Zoology, University of Illinois Typhlotriton spelaeus Stejnege r 1892, was described on the basis of an adult and several larvae from Rock House Cave, Berry County, Missouri, as the first cave-adapted salamander discovered in North America. This salamander almost immediately aroused the curiosity of scientists because of its degenerate eyes, and the earliest papers dealing with it were .detailed accounts of eye structure (Eigenmann, 1900. 1909; Eigenmann & Denny, 1898, 1900j and Alt, 1910. Other early anatomical studies were aimed toward determining the taxonomic relationships between Typhlotriton and other caudates (Moore, 1900, and Hilton, 1909). More recently, Hllton (1945, 1953, 9


VOLUME 4, NO. 2 turned out to be hypertrophied jaw muscles, especially evident on large males. These two misinterpretations of material at hand, plus others not mentioned here, place doubt on other conclusions. Smith collected, from one cave, two adult specimens, both of which had large apparently functional eyes, with unfu.s e d eyelids. These two specimens, one alive, were very kindly .s errt to me for examination, and will be reported along with others later, in a more detailed report of my work with this species. P. W. Smith (1948a, b) reported on the food habits of Typhlotriton in a cave near Waynesville, Pulaski County, Missouri, and on a cestode irife a ta-. tion in larvae from the same cave. Wells, et al., (1954) utilized larvae in testing the metamorphosis-inducing propertiesof L-triiodothyronine. Their report made no mention of the condition of the eyes of animals comple ting metamorphosis. Their experiments could easily have been used to test Noble's work, but apparently no attempt was made to do so. Except for numerous articles dealing strictly with distribution. no other report has been publis hed concerning Typhlotriton as far as I am aware. Two general references to the distributlon of thlS species are included in the bibliography (Bishop, 1943; Conant, 1958), References: ALT, A. On the histology of the eye of Typhlotriton spelaeus from Marble Cave. St. Louis Academy of Science, Trans., vol. 19, p.83-96. 1910. BARDEN, R. B. and L. J. KEZER. The eggs of certain plethodontid salamanders obtained by pituitary gland implantation. Copeia, 1944, no. 2, p. 115-118. BISHOP, S. C. Handbook of salamanders. Comstock Publ. Co., Inc. 555p. 1943. BISHOP, S. C. A new plethodont salamander with notes on related species. Copeia, 1944, no. 1, p 1-4. CONANT, r. A field guide to reptiles and ampnibians. Boston. 366~9~ EIGENMANN, C. H. Degeneration in the eyes of the cold-blooded vertebrates of the North American caves. Indiana Acade~ of ~ciences, Froc. 1899, p. 31-46. EIGENMANN, C. H. Cave vertebrates of North America. ~arnegie Institution of Washington, Publ. no. 104, 241p. 1909. Houghton Mifflin Co., EIGENMANN, C. H. and W. A. DENNY. The eyes of Typhlotriton spe I aeus, Indiana Academy of Science, Proc. 1898, p. 252-253. ----EIGENMANN, C. H. and W. A. DENNY. The eyes of the blind vertebrates of North America. Ill. Biological Bulletin, vol. 2, p. 33-41. 1900. HILTON, W. A. The hyo-branchial apparatus of Typhlotriton spelaeus Stejn. Biological BUlletin, vol. 16, p. 167-171. 1909. ------HILTON, W. A. The skeleton of Typhlotriton. Jour. of Entomology and Zoology, vol. 37, p. 79-80. 1945. HILTON, W. A. The choroid plexus of the lateral and third ventricules of tailed amphibia. Jour. of Comparative Neurology, vol. 99, p. 545-559. 1953. HILTON, W. A. Eye muscles of salamanders. Herpetologica, vol. 12, p. 273-276, 1956. 11


CAVE NOTES CAVE NOTES CAVE NOTES is a publication of Cave Research Associates and the Cave Research Association. Subscriptions are available for $1.00 per year (siX issues) or on exchange. Mid-year subscriptions receive the earlier numbers of the volume. Correspondence, contributions, and subscriptions should be addressed to: CAVE RESEARCH ASSOCIATE~ 1911A Berkeley Way, Berkeley 4, California. Editor: Arthur L. Lange, Cave Research Associates Associate Editor: Ronald A. Brandon, Cave Research Association (Department of Zoology, University of Illinois, Urbana, Ill.) Managing Editor: R. deSaussure, Cave Research Associates ~ Copyright 1962, Cave Research Associates. and 1956) published articles containing anatomical information on Typhlotriton. During 1927-30, Noble apparently did much experimental work with larvae and adults (Noble, 1927, 1930; Noble &: Pope, 1928). A brief abstract of his major results (Noble & Pope, 1928) reported that larvae metamorphosing in diffuse and direct light retained welf-deve loped eyes and unfused eyelids as adults, whereas the eyes of larvae metamorphosing in the dark degene r ated, and the eyelids became partially or completely fused, as in nature. Temperature tolerance of larvae was reported to be na ru-owe r than in Eur ycea multiplicata. One sexually mature male was 11 found in nature----wITh open hds and complete retina. II Since no detailed report of Noble's experimente has appeared, I view his results as tentative and in need of verification, even though they have been perpetuated in almost all general and popular works on salamanders. Bishop (1944) described a neotenic population, T. nereus, from. Lawrence County, Arkansas. The taxonomic status of this form LS ati Il questioned and is in need of further investigation. Although the eggs and breeding habits of Typhlotriton have not been reported from nature, Barden and Keae r (1944) descnbed eggs obtained by use of frog pituitary rmpfanta.tion, Hendricks and Kezer (1958) observed a population of Typhlotriton periodically for one year in a cave in Laclede County, Missouri. They found transformed individuals most numerous in the spring and summer, least numerous in the middle of winter. The large number in the cave during summer was attributed to drought conditions at the time. Aside from the two previously mentioned reports, nothing reliable has been published concerning the life history of Typhlotriton. C. C. Smith (1960) reported a study on a population of Typhlotnton in Independence County, Arkansas. This report, although containing much that appears to be good information, is greatly weakened and is amenable to serious question by the inclusion of statements indicating erroneous interpretation of data and observations. In this report, "Ii ght or-gana" on the backs of larvae and "paratoid glands II on the heads of adults were described. The "light organs" were undoubtedly sensory pores reflecting light when filled with water. By actual dissection of specimens obtained from Dr. Smith, the "pa r atodd g lands" 10


CAVE NOTES MOORE, J. P. Postlarval change in the vertebral articulations of other salamanders. Academy of Natural Sciences of Philadelphia, P. 613-622. ----NOBLE, G. K. The plethodontid salamanders; some aspects of their evolution. American Museum Novitates, no. 249, p. 1-26. 1927. Spelerpes and Pz-oc 1900, NOBLE, G. K. What produces species? Natural History, vol. 30, p. 60-70. 1930. NOBLE, G. K. and S. H. POPE. The effect of light on the eyes, pigmentation, and behavior of the cave salamander, Typhlotriton. Anatomical Record, vol. 41, no, 1, p 21. 1928. SMITH, C. C. Notes on the salamanders of Arkansas. No.1. Life history of a neotenic stream-dwelling form. Arkansas Acad. of Science, Proc. 1960, p. 67-74. SMITH, P. W. p. 205-208. Food habits of cave dwelling 1948. amphibians. Herpetologica, vo L, 4, SMITH, P. W. A cestode infestation in Typhlotriton. Herpeto1ogica, vol. 4, p 153. 1948. STEJNEGER, L. Preliminary description of a new genus and species of blind cave salamander from North America. U. S. National Museum, Proc., vol. 15, p. 115117. 1892. WELLS, P. H., WALLEN, T., EDEN, J., and C. TURNER. Salamander metamorphosis induced by L-triiodothyronine. Anatomical ~ecord, vol. 120, no. 3, p. 163. 1954. WATER LEVEL PLANES IN CAVES by Arthur L. Lange, Cave Research Associates The surface of a cave lake, under diverse conditions, can produce mte resting cave structures: among them, planed ceilings, nips or water-level horizons, rims tone terraces, and crusted strands. Since most caves undergo periods during which they are only partly submerged, it is important to r ecognize these structures and the conditions that bring them about, if one is to reconstruct the evolution of the cave after its generating waters have subsided. In this article we analyze the process of uniform solution beneath a water level and its resulting effects on the walls and ceiling of the cave. Theory: The effects of uniform solution in a water-filled chamber were discussed in my introductory paper on cave structures (LANGE, 1959), and are illustrated here in Figure la, b, c as a review. The successive cross-sections (b, c) of the original fracture passage (a) are everywhere equidistant from the initial structure, so that projecting sharp corners remain sharp while inner corners round off. It is the effect produced by graphically rolling a c i r c l.e around the perimeter of the section. Consider next the result of homogeneous solution in a reservoir having a constant water level (Figure Ld e, f). Below the water surface, solution proceeds as in Figure l a b c; while above the water-line, all solutional activity ceases. The evident result is the ceiling planation shown. The development of the corners along the walls of the chamber is illustrated in Figure 2. 12


VOLUME 4, NO.2 1'1 Idl Ibl 1 0 1 1'1 If I Figure 1. Cross-sections of a hypothetical cave passage undergoing uniform solution: (a,b,c) under fUlly submerged conditions; (d,e,f) under partly submerged conditions. (a) and (d) in each row represent the initial fracture passage having a fallen roof block in a floor of insoluble sediment. Wall retreat takes place normal to the original outline unde r wa te r and must terminate at the air/water boundary; thus, the examp.le s of the vertical and overhanging walls are easily understood (a, b). In the case of the po s i ti vesloping wall (c), the segment of rock below angle a retreats normally; within angle a. retreat occurs radially to the infinitesimal hollow cylinder or singularity at 0, according to the method illustrated in Figures 6f and 7d of my introductory paper. Thus, within the angular sector, the wall cross-section assumes the form of a circular arc. Should the water level rise to r ev submerge the structure, a new stage of planation ensues above the first, and the normal effects of uniform solution are superimposed on the submerged corner. This case is illustrated in Figure 2d for the vertical wall example only. I shall refer to the elementary planed ceilings and their terminating wallcorners described above as water-level planes and water-level corners, respectively. These should not be confusecrwITFi:'"""the nips, or water-level horizons. described by Kaye, Corbel, and others (LANGE, 1960), which differ conspicuously in having a lower shelf nnn-parallel to the original wall. Planation, however, does play a role in forrning the visor of the nips, in a manner to be treated in a later report. Water-level mechanisms: One naturally asks, how does the condition of constant water level a r i s e in nature? The simplest mechanism is the spillover. or t rop-cpledn, of a cave-lake system. It is very conceivable to imagine an initially submerged cave becoming tapped (say, by a downcutting outdoor stream) and having its contents rapidly drained to the local base level outside, or to a resistant r-e t ng rock. The spillover may not be apparent, 13


CAVE NOTES --~ -=---Figure 2. Detail of the formation of water-level corners in: (a) the vertical wall, (b) overhanging wall, and (c) positive-sloping wall. The solid line represents the initial wall cross-section; the dashed line, an intermediate stage; and the shaded outline is the final form. In (d), the final stage of (a) has been subjected to a second episode of solution beneath a higher water level. (See also text). Since it could occur in some portion of the cave not connected by air passage. Alternatively, any physical mechanism that can maintain a constancy of water level qualifies as an agent in planation. Such a system requires that inflow balance discharge. Water-level planes could also represent the annual high-water line of a cave lake, when this is traditionally the same or controlled by a spillway. I am thinking here of Empire Cave, Santa Cruz Co , California, where the annual water level is recorded each spring by both flat ceilings and a strand line of floating de bz-i e If a spillover cuts downward in successive steps, the water-level planes forrn an equivalent aeries, like an inverted stairway. If the spillway lowers continuously, no well-defined planes result, but the cave chambers widen with depth, since the lower portions are subjected to solution longer than the upper regions. A little reflection will disclose that a vertical wall is transformed Into an overhanging, inclined, flat wall when the water level drops at a constant rate. Figure 3. Flat ceiling above the lower Magic Pool in Samwel Cave, California. The ceiling is a waterlevel plane now decorated with cave coral. 14


VOLUME 4, NO. 2 .;, ..... Figure 4. Successive cross-sections of a dissolving chamber of constant water volume. The inne~most vertical lines represent the initial walls confining a cave lake at its highest level. As the walls retreat, the water level falls, eventually resulting Ln the shaded profile. In an iso-volumic reservoir, where the water volume does not change while the walls dissolve (this might corne about through stagnation with an ever decreasing ti rne-, rate of solution, or by discharge balancing inflow), the water-line falls as the walls retreat. If the initial reservoir has an initial floor area 4a2. and vertical walls to a height h the water volume is V =4a z h; hence, since the volume remains constant. h=V/4a2., where a is the halfwidth. Figure 4 shows successive cross-sections of this c as'e and the curved walls resulting from continuous wall retreat. At every stage the submerged walls remain vertical. while the exposed wall/ceilings approach horizontality. Water-level planes should p.orrnally correlate from chamber to chamber throughout a cave system, and in some instances their spillways might be found. If the cave rock has been tectonically tilted since being drained, the planes should bear this out; if faulted within the cave, the planes will be shifted accordingly. An interesting exception to the concordance of planes is illustrated in Figure 5, wherein a dome-like cavity has evolved under subrne r g e d conditions. "When the water level in the outer chamber falls to level A, atmospheric pressure at A can still support a water column within the sealedA---=-~-::: pff dome up to level Al (at sea-level this differential is 34 feet maximum). The ceiling thus dissolves upward until this maximum level is reached, at which point a s e mi -vaccuurn determines the water-level plane A I. Thereafter, any drop in outside level; say; to B; produces an equal drop in the interior, to BI, etc. until finally, the s.eparating barrier is exposed to the air at level C_CI, and the remaining suspended water in the dome is released to the common level, forming an integrated cave lake. -=-.----------==...::-_--= (a) (b) (0) (d) Figure 5. Solution dome dissolved out adjacent to cave chamber when cave filled with water. Successive stages of plane generation representing a balanced water column ~ explained in the text. 15


CAVE NOTES Cave examples: Waterlevel planation is best demonstrated in caves of inclined rock. where level ceilings cannot be confused with natural bedding planes. Samwel Cave. California, in almost vertical beds. exhibits water-level planes of which the most prominent corresponds in altitude to a portal 1 ft. in diameter. If the cave could hold water today to the level of this plane water would just spill out the portal to form a spring in the limestone hillside. Othe r plane s occur below the main Figure 6. Water-level plane incised approximately 1 ft. one, possibly r elafing into wall of salt mine (photo courtesy Carey Salt Oo.) to other cliff openings. Some planes are reflected in the water of the Magic Pools. whose dwindling water body is retained by rims tone dams (Figure 3). In the deepest chamber of Samwel Cave, many water lines are registered in overhanging walls. It is hard to conceive of so many successive s pi Hove r s : rather I visualize their formation by a subsiding water body which each year attained a slightly lower maximum. Three explanations come to mind: 1) Rapid entrenching by the adjoining canyon encouraged draining of the cave; 2) Increasing aridity due to climatic change resulted in the lowering of water level (the iso-volurnic process). Pe rhaps two or all three actions participated. Other California caves containing water-level planes are: Cave Man Cave, Tuolumne Co.; Cave of the White Chief, Tulare Co ; Violin Cave, Amador Co ; and Cave of the, Quills. Calaveras Co. Artificial examples: Karl Gripp (1912) produced planar ceilings in laboratory salt blocks. wh11e attempting to model the evolution of the Segeber-ge gypsum caves. He did not, however, distinguish between the ceiling effect and the sloping walls which he attributed to concentration gradients. He referred to the ceilings as "Laugdecken'", or solution ceilings. Figure 6 was taken in the 800 level main haulage way of a salt mine in Winnfield, Louisianna. The mine had been flooded accidentally to the level of the plane for two weeks in 1936. Wall retreat proceeded everywhere beneath this level to a maximum incision depth of one foot. References: LANGE, A. L. Introductory noxes on the changing geometry of cave structures. Cave Studies, no. 11, p. 69-90. 1959. LANGE, A. L. Review of "Shoreline features and Quaternary shoreline changes, Puerto Rico" by C. Kaye. Cave Notes, vo L, 2, no. 5, p 36-39. Sept/Oct. 1960. GRIPP, KARL. Uber den Gipsberg und die in ihm vorhandene H8hle. Hamburg Wissenschaftliche Anstalten, Jahrbuch 30, Beiheft 6, s. 35-51. 1912.-----------16

Summary of past investigations on
typhlotriton spelaeus, the grotto salamander /
Ronald A. Brandon --
Water level planes in caves / Arthur L. Lange.
Cave Notes(vols. 1-8) and
Caves and Karst: Research in Speleology(vols. 9-15)
were published by Cave Research Associates from 1959-1973. In
1975, the Tumbling Creek Cave Foundation compiled complete
sets of the journals in three volumes. The Foundation sells
hardbound copies of the material to support its