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Mammoth Cave: a Hotspot of Subterranean Biodiversity in the United States

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Mammoth Cave: a Hotspot of Subterranean Biodiversity in the United States diversity Article Mammoth Cave: A Hotspot of Subterranean Biodiversity in the United States Matthew L. Niemiller 1,*, Kurt Helf 2 and Rickard S. Toomey 3 1 Department of Biological Sciences, The University of Alabama in Huntsville, 301 Sparkman Dr NW, Huntsville, AL 35899, USA 2 Cumberland Piedmont Network, National Park Service, Mammoth Cave National Park, 61 Maintenance Rd., Mammoth Cave, KY 42259, USA; [email protected] 3 Division of Science and Resources Management, Mammoth Cave National Park, P.O. Box 7, Mammoth Cave, KY 42259, USA; [email protected] * Correspondence: [email protected] or [email protected] Abstract: The Mammoth Cave System in the Interior Low Plateau karst region in central Kentucky, USA is a global hotspot of cave-limited biodiversity, particularly terrestrial species. We searched the literature, museum accessions, and database records to compile an updated list of troglobiotic and stygobiotic species for the Mammoth Cave System and compare our list with previously published checklists. Our list of cave-limited fauna totals 49 species, with 32 troglobionts and 17 stygobionts. Seven species are endemic to the Mammoth Cave System and other small caves in Mammoth Cave National Park. The Mammoth Cave System is the type locality for 33 cave-limited species. The exceptional diversity at Mammoth Cave is likely related to several factors, such as the high dispersal potential of cave fauna associated with expansive karst exposures, high surface productivity, and a long history of exploration and study. Nearly 80% of the cave-limited fauna is of conservation concern, many of which are at an elevated risk of extinction because of small ranges, few occurrences, Citation: Niemiller, M.L.; Helf, K.; and several potential threats. Toomey, R.S. Mammoth Cave: A Hotspot of Subterranean Biodiversity Keywords: checklist; karst; species richness; stygobiont; troglobiont in the United States. Diversity 2021, 13, 373. https://doi.org/10.3390/ d13080373 Academic Editors: Tanja Pipan, David 1. Introduction C. Culver and Louis Deharveng The Mammoth Cave System in central Kentucky, USA is the most extensive cave sys- tem in the world with over 663 km (412 miles) of mapped passaged, including 27 entrances Received: 10 July 2021 and 10 significant caves that have been connected since explorations began in the late 1700s: Accepted: 3 August 2021 Colossal, Crystal (=Floyd Collins’ Crystal), Donkey, Hoover, Mammoth, Morrison, Proctor, Published: 12 August 2021 Roppel, Salts, and Unknown caves. Colossal, Crystal, Salts, and Unknown caves comprise the 206 km (128 mile) Flint Ridge Cave system (Figure1). Mammoth Cave National Park Publisher’s Note: MDPI stays neutral was created in 1941 and includes two-thirds of the Mammoth Cave System [1]. The Mam- with regard to jurisdictional claims in moth Cave System was recognized as a UNESCO World Heritage Site in 1981 because of its published maps and institutional affil- uniqueness as the world’s longest cave system as well as its extensive geological, mineral, iations. and biological resources. The region was recognized as the core of an UNESCO Biosphere Reserve—Mammoth Cave Biosphere Region—in 1990. The Mammoth Cave System is developed in three major limestone layers at the northwestern extent of the Pennyroyal Plateau, an expansive flat karst plain within the Copyright: © 2021 by the authors. Interior Low Plateau physiographic province. The limestone layers include, from youngest Licensee MDPI, Basel, Switzerland. to oldest, the Girkin Formation (40 m thick), Ste. Genevieve Limestone (35 m thick), and This article is an open access article St. Louis Limestone (53–60 m thick) [2–5]. The Girkin Formation is capped by resistant distributed under the terms and sandstone and shale of the Big Clifty Formation that form the Mammoth Cave, Flint, Joppa, conditions of the Creative Commons and Toohey Ridges. Most of the cave system is developed in the Ste. Genevieve Limestone Attribution (CC BY) license (https:// and the upper 40 m of St. Louis Limestone [5]. The limestone strata gently slope from creativecommons.org/licenses/by/ the southeast to the northwest. The Pennyroyal Plateau is exposed at the surface to the 4.0/). Diversity 2021, 13, 373. https://doi.org/10.3390/d13080373 https://www.mdpi.com/journal/diversity DiversityDiversity2021 2021, 13, 13, 373, x FOR PEER REVIEW 2 of 21 2 of 21 fromsoutheast, the southeast while insolubleto the northwest. strata ofThe the Pennyroyal Chester Upland, Plateau includingis exposed theat the Big surface Clifty to Formation, theform southeast, a rugged while hilly insoluble terrain thatstrata overlies of the Chester the cave Upland, system including to the northwest. the Big Clifty The For- Green River, mation, form a rugged hilly terrain that overlies the cave system to the northwest. The a tributary of the Ohio River, has cut into the Pennyroyal Plateau about 60 m such that Green River, a tributary of the Ohio River, has cut into the Pennyroyal Plateau about 60 mmost such of that the most Mammoth of the Mammoth Cave watershed Cave watershed now occurs now underground occurs underground [6]. The [6]. karst The watershed karstof Mammoth watershed Caveof Mammoth includes Cave seven includes groundwater seven groundwater basins (Pike basins Spring, (Pike Great Spring, Onyx, Echo GreatRiver, Onyx, Double Echo Sink, River, River Double Styx, Sink, Floating River Styx, Mill Floating Hollow, Mill and Hollow, Turnhole and Bend); Turnhole in addition, 2 Bend)flood; in overflow addition, occurs flood overflow into an eighth occurs basininto an (Sand eighth Cave). basin (Sand These Cave). basins These encompass basins 317 km encompassand ultimately 317 km drain2 and ultimately at springs drain at base at springs level into at base the level Green into River the Green [7,8]. River [7,8]. FigureFigure 1. 1.MapMap showing showing the location the location of Mammoth of Mammoth Cave National Cave National Park (MCNP) Park (MCNP)and the extent and theof the extent of the MammothMammoth Cave Cave System System in and in and adjacent adjacent to MCNP. to MCNP. The major The major segments segments of the Mammoth of the Mammoth Cave Sys- Cave System temare are shown shown as as line line plots plots in in various various colors.colors. The different different segments segments explored explored from from different different en- entrances trances (27 total). Line plot data from Cave Research Foundation. MCNP also contains over 500 smaller(27 total). caves Line developed plot data in various from Cave karstified Research limestones Foundation. that are MCNPnot attached also containsto the Mammoth over 500 Cave smaller caves System.developed These in are various grouped karstified on the map, limestones but include that the are St. not Louis, attached Ste. Genevieve, to the Mammoth Haney, Glen Cave Dean System. These andare Girkin grouped Formations. on the map,These butsmaller include caves the contain St. Louis, a variety Ste. of Genevieve,habitats from Haney, epikarst Glen to base Dean-level and Girkin streams.Formations. These smaller caves contain a variety of habitats from epikarst to base-level streams. TheThe Ma Mammothmmoth Cave Cave System System is characterized is characterized by a complex by a complex network network of vadose of and vadose and phreaticphreatic passages passages with with at at least least five five primary primary horizontal horizontal levels levels of ofpassages passages (four (four fossil fossil stream stream levels and the modern base level) representing distinct stages of development in levels and the modern base level) representing distinct stages of development in association association with past periods of water table stability and intervening periods of downcut- with past periods of water table stability and intervening periods of downcutting of ting of the Green River valley through the resistant caprock into the soluble limestone layersthe Green below River[1,6]. The valley evolution through of the the Mammoth resistant Cave caprock system into is thelinked soluble to the limestoneincision layers hibelowstory of [1 ,the6]. TheGreen evolution River, drainage of the Mammothreorganizations, Cave and system significant is linked climatic to the changes incision history fromof the the Green Pliocene River, through drainage the Pleistocene, reorganizations, with the oldest and significantupper-level climaticpassages changesdating to from the 3.2Pliocene Mya and through the lower the levels Pleistocene, developing with over the the oldest past 2 upper-levelMya [9]. passages dating to 3.2 Mya andMammoth the lower Cave levels has developing long been a over focal theregion past of 2 study Mya for [9]. North American subterra- nean biodiversityMammoth and Cave for hasadvancing long been our foundational a focal region knowledge of study of the for ecology North Americanand evo- subter- lutionranean of biodiversitycave fauna. Studies and for of the advancing biodiversity our in foundational the Mammoth knowledge Cave System of have the ecologyan and extensiveevolution history of cave dating fauna. back Studies to the 1820s of the (see biodiversity [10]) when Constantin in the Mammoth S. Rafinesque Cave first System have visitedan extensive Mammoth history Cavedating [11]. Darwin back to [12] the even 1820s mentions (see [10 ])cave when life Constantin from the Mammoth S. Rafinesque first visited Mammoth Cave [11]. Darwin [12] even mentions cave life from the Mammoth Cave region in On the Origin of Species by Means of Natural Selection. Much of our early knowl- edge of the North American cave fauna was derived from visits and studies by biologists to Mammoth Cave in the 1800s, such as DeKay [13], Wyman [14–19], Tellkampf [20–22], Agas- siz [23–25], Von Motschulsky [26,27], Call [28], and Packard [29–36] (reviewed in [10] and [37]). Diversity 2021, 13, 373 3 of 21 Additional significant early publications on the fauna and ecology of Mammoth Cave in- clude Putnam [38], Eigenmann [39–42], Bolivar and Jeannel [43], Bailey [44], Buchanan [45], Park [46], Dearolf [47], Hubricht [48–53], Jeannel and Henrot [54], and Barr [55–60].
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