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Home , Organotroph, Troglofauna

Reprintedfrom: ECOSYSTEMSOF THE WORLD 30

SUBTERRANEANECOSYSTEMS

Editedby Prof. Dr. Horst Wilkens Zoologisches Institut und Zoologisches Museum der UniuersitcitHamburg, M artin-Luther- Kin g-P latz 3, 20146 Hamburg, Germany

Prof. David C. Culver The American UniuersiQ, 4400 MassachusettsAuenue, Washington,DC 200I 6-8002, USA

Dr. William F. Humphreys WesternAustralian Mus eum, Francis Street, Perth, WesternAustralia 6000, Australia

2000 ELSEVIER Amsterdam- Lausanne- New York- Oxford - Shannon- Singapore- Tokyo Chapter0

BACKGROUNDAND GLOSSARY

W.F.HUMPHREYS

INTRODUCTION within ,and the issuewill not be pursued here at length. As with other specialized subjects, a plethora of Trogiobiteswere long consideredto be phylogenetic terminology and definitions has come to surround, or distributional relicts driven into by climatic sometimesto obscure,the discipline of biospeleology change,especiaily in the Pleistocenerefugium model. (speleobiology,stygobiology, . .), terms sometimesso Recently, colonization has been viewed as an subtly different - even synonymous as to confound active process - the invasion of a vacant . even researcherswithin the discipline (for a fuller Notwithstanding,the undergroundrealm does harbour treatmentsee, for instance,Camacho, 1992). many distributionaland phyletic relicts. The terminology coverssix main themes: Anchialine waters (inland groundwaterwith current ( I ) How a speciescame to be underground. marine connections)harbour a distinct (Sket, 1996) (2) The degree of dependenceof a specieson the and, in some areas! a predictable (Yager and undergroundrealm. Humphreys,1996). (3) Whether a speciesshows to the under- Stygofaunawith marine ancestryare consideredto ground realm. colonize the shore line and to becomeisolated inland (4) Whether a species inhabits subterraneanvoids by marine regressionfollowing a period of manne filled r,vithwater or with air, and which part(s) of transgression(Stock, 1977; Boutin and Coineau,1990; each it inhabits. Notenboom,l99l ; Holsinger,1994). (5) The delimitation of the various hypogean sub- systems. The degreeof dependenceof a specieson the (6) The amountand origin of the energysource for the underground realm ecosyslem. As a backgroundto this volume,the main themesre- have many pathways of interaction with curring in biospeleologyare introduced below, followed subterraneanspaces (Gibert et al., 1994); some are by a Glossary. only found underground,while others move in and The subjectof cavesis well introducedby Gillieson out ol the hypogean realm. More than twenty spe- cial nomenclature schemes have proposed (1991), while the literature pertaining to hy- been to describethese relationships,based on morphological, drology, encountered especially in the context of behavioural,and ecological adaptationsof to ,is accessiblein Fordand Williams (1989). undergroundlife and on their presenceor absencein caves(; see review in Camacho,1992). How a speciescame to be underground ln the Schiner-Racovitd system (Schiner, 1854; Racovitza,1907), cave-inhabiting animals are grouped This issueis coveredfrom dilferent stancesin several by their degreeofdependence on the caveenvironment: contributions to this volume (e.g. Chapter 21 by troglobitesare specieswhich do not existoutside caves; Holsinger,Chapter 22 by Humphreys,and Chapter3l troglophiles may live and reproduce underground as by Deharvengand Bedos) as the topic is all-pervasrve well as in the epigean domain; whlle n'ogloxenesdo 4 W.F.HUMPHREYS not normally feed underground, but may enter caves which togetherproduce the convergence,characteristic actively (regular trogloxene) or passively (acciden- of cave-adaptedanimals, that is termed troglomorphy. tal trogloxene).Additional criteria have subsequently These adaptations include morphological, ecologi- entered these purely ecological definitions, so that cal, physiologicaland behaviouralcharacteristics (Ta- they have become a variable mix of biological and ble0.1). morphologicalcriteria (Howarth, 1973:p. 142). The lack of detailed ecological information for Whether a speciesshows adaptations to the most cave-inhabitingspecies leads to ambiguity, and underground realm this is to some extent overcome by the concept of troglomorphy (Christiansen,1962), which permits Organisms have many pathways of interaction with classificationof the cave inhabitantsusing a iess am- the subterraneanrealm. While some are only found biguous morphologicaldefinition (troglomorphic/non- underground,others move in and out of the hypogean troglomorphic species)rather than the ecologicalone environmentas indicatedin the ecologicalclassification (troglobitic/troglophilic/trogloxenicspecies); but this of speciesby the terms troglo(stygo)bitic,troglophilic term is itself being infiltrated by behavioural and and trogloxenic. physiologicalattributes (Tabie 0.1). It is important to note that functionally these categoriesare more diffuse than is suggestedby their Table 0.1 ecological context. For example,while stygo(troglo)- Characteristicsof cave organismsrelative to surface organlsms xenesmay occur undergroundonly accidentally,they troglomorphies I may influencehappenings in the hypogeanrealm, for example,functioning either as predatorsor prey (e.g. Morphological Gibert et al. 1994: p. l1), as well as constituting Specializationof sensoryorgans (chemo-, hygro- thermo- and a subset of the allochthonousinputs from epigean baro-receptors) sources. Elongationo1' appendages In consideringtroglomorphic correlates (e.g. reduced Reductionof eyes,pigments and rvings eyes,pigment, wings etc.) it is necessaryto distinguish Cuticle thinning (in terrestrralarthropods) whether the traits are different from close epigean (in Foot modification Collernbolaand planthoppers) ancestors(apomorphic) or simply characteristicsof the Scalereduction (in fish) group or lineage(.plesiomorphic). E co I ogic al and l:ehoo io tu'uI Slowing Whether a speciesinhabits subterranean voids Starvationresistance filled with water or with air, and which part(s) of each it inhabits Relaxctionand degencratronof circadranrhyhrns

I n*,^.o.4 fe,",-.lit., While the Schiner Racovitrdsystem is often used for lncreasedegg volume both terrestrialand aquaticspecies! Gibert et al. (1994: Increasedlife span p. 13) have proposed stygo- (.sensulato) as the most Behauioural ecologicallydescriptive prefix for groundwaterfauna. Decreasedaggregation (in Collembola) Hence, troglofauna becomes restricted to terrestrial Reducedreaction to alarm substances(in fish) systems (see review in Camacho, 1992), while the Increasedsensitivity to vibratlon aquatic fauna is referred to generally as stvgofauna (Ginet Reducedintraspecific aggre:sion and Decou, 1977; Botosaneanu,1986; Gibert et al., 1994).There are complex parallel classifications I From Culver et al. (1995), after Chdstiansen(1992). in use for aquatic subterraneanorganisms (e.g. Boto- saneanu,1986: endpaper; Williams, 1984;Stanford and Many species found living underground display Ward,1993). certain characteristictraits that are thought to be Gibert et al. (1994) followed the main Schiner adaptiveto undergroundlife. These include both the Racovitrdsystem, replacing the prefix "troglo-" by reduction or loss of characters(regressive evolution) "stygo-" (hence,stygoxene, stygophile, stygobite). ln and the enhancementof others(constructive evolution), porous aquifers,stygophiles are subdividedinto three BACKGROUNDAND GLOSSARY 5 categories:occasional hl,porheos, which comprises in some tropical areas(Cape Range, Australia), highly essentiallythe larvae, especially of aquatic , troglomorphic (and stygomorphic)species are found, of most of the benthos;amphibites, whose life cycle under certain conditions,to be leeding in sunlight at requires the use of both surface and groundwater the entrance to caves. However, even within caves, systems;and permanent hyporheos,which comprises the boundariesbetween zones are dynamic, shifting, a diverse assemblageof organismspresent in all life inter alia, accordingto the degreeof biogenic carbon stagesin the groundwateror in the benthic dioxide production(dependent on periodicity of water an aerial epigeanstage is not requiredto completethe and energy inputs), the weather, the season and life cycle. major climatic events. The extent of the zones and Stygobitesare subdividedinto two classesofobliga- their vagility varies between caves according to the tory hypogeanforms: ubiqtdtousstygobites are widely relative sizes of entrancesand volume, and the size, presentin both karstic and alluvial groundwatersand shapeand arrangementof the chambers(see Racovita, are sometimes found very close to the surface. ln Chapter28, this volume). contrast,phreatobite species are stygobites that are restrictedto the deepgroundwater substrata of alluvial aquifers(Gibert et al., 1994). The amount and origin of the energy source for the Delimitation of the various hypogean sub-systems In the absenceof light subterraneanecosystems have The hypogeanrealm used to be defined in terms of been consideredto be dependenton organic matter cavesand the degreeof darkness.The realizationthat fixed by and reaching the hypogean most caves had no openings to the surface (Curl, realms by various direct and indirect pathways. Routes 1966), and that troglobitic animals may inhabit the of entry are as diverseas logs being washedinto open immediatesubsurface spaces even in non-karsticareas caves,dissolved organic matter percolating downwards (Ueno, 1977; Jtberthie et a1., 1980; Juberthie and through a porous medium into cavesotherwise closed Delay, i981), meansthat the limits to the hypogean to the surface.and animals such as bats and crickets realm aredifficult to define.At the surface,epigean and moving between the cave and the exterior. edaphicenvironments may gradeimperceptibly into the Diverse subterraneanecosystems dependent upon hypogeanrealm, while at the lower end of the hypogean the chemoautotrophicbacteria that rely on sulphides realm the metazoansprogressively fade away,leaving a have recently been demonstrated(see Chapter 17 by microbiologicalworld that may penetratedeep into the Sarbu,Chapter 18 by Pohlmanet al., and Chapter 19 Earth'scrust (Gold, 1992). by Sarbu et al.). Sulphidesmay be of magmatic or This complexity is apparentalso at a finer scale.For petroleum origin (see Chapter 12 by Poulson and example, the hyporheic zone is an between Lavoie) and thesecaves are thus analogousto the deep- surface water and groundwater, but this ecotone is seaenvironments of hydrothermalvents (Childressand difficult to define. It has been variously interpreted (Arp in both physical and biological terms: physically by Fisher, 1992) and cold seeps and Fisher, 1995; the depth to which the river water penetrates into Scott and Fisher, 1995),respectively. the hyporheic zone, and biologically by the depth to Undergroundenvironments are typically considered which amphibitespenetrate into it. In reality, it is a to be sparselyprovided with food energy to have region of great complexity, the origin of the water a low energy density and it is hypothesizedthat being from either groundwater (upwelling zone) or this is a major selective pressure involved in the surfacewater (downwellingzone). It is best considered loss of characters,such as eyes, in troglobites.This as a dynamic ecotone or transition zone (see Gibert low energy density may be evenly distributed and et al., 1994). so difficult to harvest, or it may occur as sparsely In large cavesthe highly troglomorphic specresare distributedbundles ofhigh energyand thus be difficult generallyconsidered to be restrictedto the deep cave to find. A few systemshave profuse and predictable environmentswhere the temperatureis stableand the suppliesof energy usually transportedinto the system air is saturated with water vapour, and frequently by animals (bats,birds and crickets) or flowing water containselevated levels ofcarbon dioxide.Nonetheless. (some chemoautotrophicsystems). o W.F.HUMPHREYS

TERMINOLOGY in the United States;locally in Francethe term for pothole. As indicated above, biospeleologyhas developed a Biofilm. Accumulationof bacteriaattached to solid sur- range of special terms to cover its subject matter. In facesby meansof appendagesand exopolymers,in addition, the terminology of speleologyis frequently which condition cross-feedingis favouredand they encounteredin biospeleologicalliterature. This vocabu- are protectedfrom dislodgment, and ad- lary is coveredwell in Lowe and Waltham (1995), and versephysico-chemical conditions (Gounot, 1994). the equivalentterms in English, German"French and Blue hole. (1) Non-turbulent resurgence(Jamaica); Hungarianare listed by Attila (1996). (2) drowned sinkholeson the BahamasBanks from Specializedterms are listed and explained in the which coolerwater rises during high tide (Bahamas); Glossary below, together with a range of terms tn (3) generally,shaft-like depressionsdeeply flooded related disciplines which may be unfamiliar to the by seawater, typically in low-lying coral islands. reader.Additional terms can be found in Lincoln et al. Boundstone. Carbonaterock whose components (r were e82). bound together during deposition and remain in position of growth. such as many reef rocks. Calcarenite. Wackstones,q.u. (Gary, McAfee and GLOSSARY Woll 1972). Calcrete. Carbonatedeposits that form in the soil Aesthetasc. Sensory or ollactory bristles on the or in the vicinity of the groundwater table as a antennaor antennulesof many arthropods. result of the evaporationof soilwateror groundwater Allochthonous. Originating outside the habitat under respectively.Groundwater calcretes form in arid discussion. climates(annual rainfall lessthan 200 mm) with high Allopatric. Used of speciesor populationsoccupying potential evaporation(more than 3000mm per year: a geographicalarea different from that of another Mann and Horwitz, 1979). speciesor population. Cave (sensu luto). Underground void, generally of Amensalism. An interspecificinteraction in which one a size sufficient for people to enter. However,the speciespopulation is inhibited, typically by toxin word means different things to different people. producedby the other,which is unaffected. Caverstreat as cavesnatural holes large enoughto Amphibite. Particular type of stygophile species enter - sometimesthe term pothole is applied to whose life cycle requires the use of both surface vertical developments- or which extendto darkness. water and groundwatersystems; also amphibionts. Archaeologistsoften use the term loosely to include Anchialine (or anchihaline). Anchialine habitats rock-shelters.In geomorphologicalterms, a cave consist of bodies of haline waters, usually with a is an opening of diameter or width greater than restrictedexposure to open air, alwayswith more or 5-15 mm - that is, a void of suficient dimensrons less extensivesubterranean connections to the sea. to allow turbulent rather than laminar flow of water and showingnoticeable marine as well as terrestrial (Ford and Williams, 1989). These differencesare influences(Stock et al., 1986).They typicallyoccur well reconciledby Howarth(1983: p. 370) "ln both in volcanic or limestonebedrock (Iliffe, Chapter3, soluble and volcanic rocks. therefore,a complex of volume). this interconnectedvoids of varying sizes anastomoses Apomorphic. Pertainingto a derived characterwithin throughout the rock in a great labyrinthine sys- the taxa being considered,in contrastto the ancestral tem. Within this system there is a continuum of (plesiomorphic)state. various sized voids from the microscopic to the Aquiclude. Body of relatively impermeablerock or largest caverns.The existenceof these voids, their sedimentacting as a boundaryto an aquifer. size, depth, and extent, dependson the geological Autapomorphy. The possessionof a unique derived history of the area. From a biological perspective, character by a species or monophyletic taxon; this continuum can be divided into three srze c/. Synapomorphy q.u. classes:microcavernous (<0.1 cm), mesocavernous Aven. Hole in a caveroof, which may be a blind pocket (0. I -20 cm) and macrocavernous( >20 cm)". or a tributary inlet shaft, and known as dome-pits Cave lint. The accumulationon cave formations of BACKGROUND AND GLOSSARY

(skin, fibres, hair, etc.) shed by human Deep (zone) cave environment. Region of cave with visitors to caves;mainly a problem in busy tourist constant temperatureand near-saturatedhumidity; caves. the most troglomorphic speciesare often restricted Cavernicole.Pertaining to an inhabitantofcaves sensu to this region. lato, without implying any particulardependence on, Distrophic. Pertainingto cavesmostly supplied with or associationwith the caveenvironment. vegetalmatter during floods. . Floodedcollapse depressions on flat limestone Doline. Simple closedcircular depressionin karst area plain - originally from Yucat5n,Mexico. with subterraneandrainage and commonly funnel- Chaetotaxic. Pertainingto the arrangementor pattern shaped, varying from a few to many metres in of bristles. dimensions. I Chemoautotrophs. Organismsderiving nourishment DOM. Dissolvedorganic matter, typically <1mg1- in from chemicalreactions of inorganic substances,as pristine groundwater. in sulphur and iron . Dripstone. - ,q.0,. Chemoheterotrophs. Organisms that obtain energy Dysaerobic. Environments in which the oxygen from oxidation/reductionreactions and use organic concentrationis between0. I and l.0mll-1 . electrondonors; cf. chemoorganotrophs. Edaphobite. Deep soil inhabitant. Chemolithoautotrophs. Organismsthat obtain energy Edaphon. The biota living within the soil or substrate from oxidation/reductionreactions and use inorganic cf Psephon,Edaphobite, q.r.r. electrondonors. Embryonic genesis (embryogenesis). Embryonic . : Chemoautotrophs. developmentfrom an ovum. Cleitric. The fissure system in the rock matrix in Endobenthos.Organisms living within the substratum, contradistinctionto the cave(speleic) system. as, for example,burrowing species. Cockpit karst. Synonym of Cone karst, referring to Endogenouszone. The zone immediatelybeneath the the depressions. Earth'ssurface. Compurtintent souterrain superJiciel. : Superficial Endopsammic. Pertaining to inhabitants of water- undergroundcompartment, 4.u. saturatedsand. Cone karst. Karst type of tropical regions character- Entrance zone of cave. Area of cave where outside ized by an extentof steep,conical hills and adjacent influencesdominate both the physicaland biological star-shapeddepressions, or cockpits having steep, environment. convex sidesand slightly concavefloors. Epibenthic. Living on the surface of the sea bed or CPOM. Coarseparticulate organic matter [for exam- lake floor. ple, dead leavesand twigs; c/ FPOM, q.r,r]. Epigean. The surfaceenvironment as opposedto the Craton. Part of the Earth'scrust little deformed for a subsurface(subterranean, hypogean) environment. prolongedperiod (shield region). Epikarst. Upper part of the percolationzone. Crenobionts.Inhabitantsofspringsandspringbrooks. Epiphreatic. Inhabiting the superficialpart of the Crevicular. Pertainingto the finer cracks,crevices and Phreaticzone, q.u; the vadose phreaticboundary. spaceswithin rock matrix. Euryece. lnhabiting a wide range of habitats. Cryptokarst. Karst secondarily covered by sedi- Eurytopic. Having a wide geographicaldistribution. ments' Eutrophic. Pertainingto a large amount of available Cyclothemic. In Jurassiclimestone of central ltaly, organic matter, high energy density or nutrient pertaining to a cyclic sedimentation controlled enrichment;in cavesespecially that providedby bat by climatic events (Cycle of Milankovitch: Mi- or flood debris. lankovitch, 1920). Exaptation. for one function serving for Dark zone. Region of cave in perpetual total anotherfunction. darkness. Exokarst. All featuresthat may be found on the surface Decoration. ,q.u. of a karst landscape,ranging in size from ttny karren . Any that feeds by degrading to massivepoljes. organlcmatter. Exploitative . An interaction between 8 W.F.HUMPHREYS

organismsin which one benefits at the expenseof Haematophagy.Feeding on blood. the other. Halocline. More or less abrupt changein salinity with tr'lowstone.Deposit formed from trickles or films of depth in water column. water on walls and floors of caves, normally of Heterochronv. Change in timing or rate of devel- calcite. opmental events,relative to the same event in the Fluviokarst. A karst landscapewhere the dominant ancestor. landforms are valleys cut by surface rivers - the Hom[o]eotic mutation. Mutation from one state to valleys become dry as karst developmentimproves aqotherof a structurethat exist in a seriesof states, undergrounddrainage. such as the mutation of an wine into a Fossil cave. Cave no longer containing a stream or haltere. active speleothems. Homoplasy. Structuralresemblance due to parallelism Foul-air zone. Still-air zone of a cave? usually or convergent evolution rather than to common in a depressionor pot, with high humidity and ancestry. elevated carbon dioxide levels, which may result Hypogean. The subsurfaceor subterraneanenviron- from biogenic or physical effects (see,for instance, ment as opposedto the surface (epigean)environ- Howarthand Stone,1990). ment. FPOM. Fine particulateorganic matter resulting from Hyporheos [hyporheic zone]. Interstitialspaces within abrasionand mechanicaldisruption by biologicaland the sedimentsol a stream bed; a transition zone physical agents;c/ CPOM. between surface water and groundwater (Orghidan, Gelifraction. Mechanical disintegration of rock or 1959). soil causedby the freezingof containedwater (also Impounded karst. Karst system in which the under- congelifraction). ground drainage is dammed, typically by another Ghyben-Herzberg principle. Expressionof the rela- lithology, sometimesresulting in dammedsprings. tionshipwhich occursin near-coastalporous aquifers Interstitial. Living in the spacesbetween particles, wherefresh water overlies sea water; the depthbelow especiallyin alluvia. sea level of the fresh water/saltwater interface is Iridophore. Coiour cells containing guanine in, for -40 times the height of the water table above sea example, fishes, reptiles, and butterflies; level;thus a minor drawdownof the groundwatercan iridocytes. lead to a major intrusionof seawater into an aquifer Isopiestic. Regions of an aquiler where the pressure by upward coning of the salt water in responseto head of ground water is the same. a thinner freshwater layer. Hence, Ghyben-Herzberg Karren. Synonymof Lapiaz. systemor lens. (SeeChow, 1964.). Karst. Soluble-rocklandscape; terrain with distinctive Grainstones.Mud-free carbonatesedimentary rock. hydrology and landforms arising from a combination Grieta. Crevice,fissure, crack or cleft in Spanish. ofhigh rock solubility and well-developedsecondary Guanobite. Speciesthat, when in caves,inhabit guano porosity.The distinctivelandforms above and below deposits(from bats,swiftlets, etc.), and whoseentire ground that are the hallmark of karst result from life cycle takesplace in this substrate(see Gnaspini the solutionof rock (mainly by carbonicacid) along and Trajano,Chapter 13, this volume). pathwaysprovided by the structure.The unusualfea- Guanophages.Species that feed directly on guano,or turesof the Kras (Karst in the period of the Austro- on microorganismsand fungi that grow on it (see Hungarian empire) region on the Italo Slovenran Gnaspini and Trajano,Chapter 13, this volume). border becameknown as "karst phenomena".Such Guanophile. Speciesthat may inhabit and reproduce areas are characterizedby sinking streams,caves, both in guanopiles and in othersubstrates in the cave encloseddepressions, fluted rock outcropsand large environment(see Gnaspini and Trajano,Chapter 13, springs(Ford and Wil1iams,1989). this volume). Karstification. Production of karstic featureson the Guanoxene. A speciesfeeding and/or reproducingin landscape. guanodeposits but dependingon other substratesin Karst window. A sectionof groundwaterexposed by the caveto completeits life cycle (seeGnaspini and roof coliapseor dissolution,especially in karstlands, Trajano,Chapter 13, this volume). as" for example,an anchialinepool. BACKGROUNDAND GLOSSARY q

Kipuka. Isolatedprotrusion of bedrock (in Hawaii) in profound implications for the aquatic fauna, hence lava flow, such as the summit of a hill; synonym: the MessinianSalinity Crisis. steptoe. Microvoid. Seemesocaverns. Lapiaz. In karst terrain, a general term for solutional Milieu souterrain superficiel. : Superficial under- furrows in a limestonesurface, with depthsbetween ground compartment. a few millimetres and more than a metre, usually Mixohaline. Waterwith a salinity between5 and 30o/oo separatedby knifelike ridges; synonym of karren. under the Venice System (equivalentto 5000 and (Pyr6n6esdialect, alsolapi6s.). 30000mg1 I total dissolvedsolids). Lava tubes. Tubularcaves within lava flows, subjectto Mixture corrosion. The ability of two calcite-saturated mechanicalcollapse of the roof (vulcanokarst). water massesof different carbon dioxide contentto Lavicole. Pertainingto an inhabitantof lava beds. dissolveadditional calcite when mixed. lake, pond. Lentic. Standingwater as in swamp, MSS. - Superficialunderground compartment. Limnicoid. Inhabiting lakes; also, limnicole. Nappes phreatiques, Pertaining to sheet flow in deep Limno- (Prefix). Restrictedto fresh water. groundwater. to fresh Limnostygobionts. Stygobionts restricted Nectarivory. Feedingon nectar. waters. Neogeoaeolian. Pertaining to an ecosystemwithin Lotic, Running\\ater. as in streamor river. a newly formed landscape(lava flows, sand cays) Lucifuge. lntolerant of light. supported by windborne allochthonous material Macrocavernous. Underground voids >20 cm, espe- (Howarth, 1979). passagesin karst and cially cavesand underground Neoteny. Paedomorphosis(q.u.) produced by re- volcanic substrates. tardation of somatic development,so that sexual Mafic. Said of igneousrock composedof one or more maturity is attainedin an organismretaining juvenile ferromagnesianminerals. characters:cf, Progenesis. insects Manna. Honeydew secretedby some scale Neuromast. Sensoryreceptor, part of the acoustico- (Coccidae). lateralis system of aquatic chordates, to detect MAST (or M.A.S.T.). The mean annual surface r ibration and movementin water. temperatureof a region the expectedtemperature Neutral mutation. A geneticmutation that proffersno of the deep cave environment in the absenceof advantageor disadvantageto the organism. significant geothermal heat or internal climatic Nomocoenosis.Pasture . effects. Normoxic. Pertainingto habitatswith the normal level Meiobenthic. Small benthic (q.u.) organisms that of oxygen found in the atmosphereor dissolvedin pass through a 500 pm sieve but are retainedby a 40 ptm sieve; cf. metofauna, q.u. water in equilibrium with it. Meiofauna. Assemblageof animalsthat passthrough Nycthemeral. Occurring ai night. a 500 pm sieve but are retainedby a 40 ptm sieve, Oligotrophic. Pertainingto low amount of available often interstitial;prefix meio-, hencemeiobenthic. organic matter, low energy density. Mesocaverns. Underground voids in the size range Outgroup. A distinct but closely related speciesor 0.1-20cm, especially in karst and volcanic sub- lineage used as an anchor in phylogenetic and strates. cladistic analyses. Mesopsammon.Psammon q.u. Packstones.Sedimentary carbonate rock whosegranu- Mesovoid shallow stratum. See superficial under- lar materialis self-supportingyet containscalcareous ground compartment. mud. Mesotrophic. Pertainingto an intermediateamount of Paedogenesis. Paedomorphosis(q.u) produced by availableorganic matter, medium energydensity. precocious sexual maturity in an organism that is Messinian. Part of Miocene epoch-6.5-5.5 million still at a morphologicallyjuvenile slage;cf . Neoteny, years ago. The Mediterraneanarea was landlocked Progenesis4.0,. and almost completely dried out at this time, Paedomorphosis. Retention of sub-adult ancestral leaving a series of hypersaline lakes which had traits in the descendantadult. 10 W.F.HUMPHREYS

Palaeokarst. A rock or area that has been karstified epigenotype.Subsequent subterranean dispersal ex- and subsequentlycovered by sediments. pandsthe geographicrange as adaptationimproves. Paludicole. Pertainingto an inhabitantof marshes. Plesiomorphic. Pertaining to an ancestralcharacter Parafluvial. Pertainingto groundwateradjacent to the within the taxa being considered,in contrastto the river channel. derived (apomorphic)state. Parapatric. Pertaining to populations whose geo- Poecilotrophic. Pertaining to variable amounts of graphic rangesare contiguousbut not overlapping, organic matter, as in iong caveswith sectionsdif- so that gene flow betweenthem is possible. fering in energy density,ranging from oligotrophic to eutrophic. Peripatric. Peripheralto the main distribution,as used Polje. Large closed depression- with flat floor in 'speciationin peripatricpopulations'. a that makesa sharp break with surroundingslopes Philopatric. In the context of bats, loyalty to caves draining underground,often containing a stream and (maternityor hibernal)or otherroosts (Tuttle,1916). liable to flood, becominga lake. It is applied to bats which return to the samecaves Pre-adaptation. Possessionby an organism of the on a daily and./oron a seasonalbasis; thus, the bat necessaryproperties to permit a shift into a new coloniestend to be stable. niche or habitat.A structureis preadaptedif it can Photoautotroph. Organisms that obtain metabolic assumea new function before it becomesmodified - energy from light by a photochemicalprocess; itself-. ,q.u'. Progenesis. Early cessationof developmentalevents . Organismthat utilizes radiant en- in the descendant;produces paedomorphic traits ergy and organic electrondonors (synonym:photo- when expressedin the adult phenotype;early sexual ). maturationwill produceglobal progenesis,but early Photo-negativity.The avoidanceof light. cessationin local growth fields can also produce Phototroph. : photoautotroph,q. u. progenesis(McNamara, 1990). Phreatic. That part of a cave systemlying below the Provincialization. Progressiveformation of morpho- local groundwatertable; below the unsaturatedor logically or genetically distrnct subpopulationsin vadosezone, q.u. a region having previously a more homogeneous population. Phreatic loop. Water-filledcave tube rejoining itself. Psammolittoral. Pertaininsto sandsalons lake Phreatobiological net. Net designedto sample deep and sea shores. groundwater,usually through bores and wells. Psammon. Organismsliving between sand grains of Phreatobite. Stygobitesthat are restrictedto the deep freshwateror marine shores;Mesopsammon. groundwatersubstrata of alluvial aquifer (phreatic Psephon. Interstitial water dwellers (equivalent water) (Gibert et al., 1994). to Edaphon;unusual). Phreatomorphic. Pertaining to the convergentmor- Pseudokarst. Karst-like landforms producedby pro- phological,behavioural and physiologicalfeatures of cesses other than solution or corrosion-induced phreatobites Stygomorphy, Troglomorphy, q.u. subsidenceand collapse; e.g. ice caves (Ford and Phreatos. Assemblageof organismsinhabiting deep Williams,1989). groundwater(phreatic zone). Pycnocline. More or less abrupt change in water : Phytophilic. Plant-eating phytophagous,herbivo- density with depth, typically at the halocline or rous. thermoclinein the water column. Pleiotropic. Pertainingto a gene that has more than Regressiveevolution. The loss of morphologicaland one phenotypiceflect. behaviouralcharacters that accomoaniesisolation in Pleistocene refugium model. Traditional model for caves. isolationofspecies in cavesincluding both dispersal- Refuge (refugium, refugia). A region in which certain ist and vicariant elements.Ancestral populations are typesor suitesof organismsare ableto persistduring isolatedin cavesby climatic changewithout active a period in which most of their original geographic invasion.The geneticbottleneck resulting from small range becomes uninhabitablebecause of climatic founderpopulation is followed by reorganizationof change(Morton et al., 1995). T2 W.F.HUMPHREYS

Syntopically. Pertainingto populationsor speciesthat Trogloxene. Species that do not normally feed occupy the same microhabitat, are observablein underground,but may enter caves actively (regu- close proximity, and could thus interbreed. lar trogloxene)or passively(accidental trogloxene) Tabular karst. Broad exoanse of more or less (Racovitza,1907). horizontalkarst. Tropical winter effect. The drying of the air in Thalasso-. (Prefix) Pertainingto marine or brackish tropical cavesin winter when the cave temperature waters,hence thalassostygobionts, thalassopsammon and the partial pressureof water vapour inside the and thalassopsephon. cave exceedthose outsidethe cave,resulting in the (Howarth, Thalassoid. Having marine affinities. loss of water from the cave air 1980). Thermocline. More or less abrupt changein tempera- Trou technique. Sampling method for fauna sediments percolated ture with depth in air or water column. involving filtering the water that has into a hole dug in stream bank; synonym: Karaman Thigmotactism. The tendencyof many small organ- Chappuismethod. isms to seekmaximum surfacecontact. as in corners Twilight zone of cave.Region of caveextending from and crevices. the limit of green vascularplants to total darkness; Tower karst. Residualkarst hills with near-verticalto physical climate not stable. overhanginglower slopes,often separatedby areas Upper hypogeanzone. Region immediatelyabove the of alluvial deposits. bedrock where the high porosity provides suitable Transition zone of cave.A dynan,iczone in total dark- hypogeanhabitat for troglobites(U6no, 1987);: Su- ness where the microclimate is still conspicuously perficial underground compartment, q.r,,. affectedby short-termmeteorological events at the Vadosezone. The zone of rock abovethe water table, surface. mostly free-draining; also called the unsaturated Troglo-. (Prefix) For subterraneanterrestrial and zone, and lying above the saturatedor phreatic aquatic systems,now more commonly restrictedto zone)q.u. terrestrial systems,being replaced by "stygo-" for Vauclusian. A spring rising lrom a vertical or very aquaticsystems. steepflooded bedrock cave passage directly draining types Troglobiont. inhabiting the various ol the phreatic zone and en,ergingin dayiight; named subterraneanspaces; sometimes restricted to air- after the type example,La Fontainede Vauclusein filled voids; and hence terrestrial as opposed to southernFrance. aquaticspecies. Vicariance. Speciationas a result of range disruption. Troglobiontic stygobionts. Obligate inhabitants of typicallythe resultof somenon-biological process; undergroundwaters found in caves. also termed passiveallopatric speciation. Troglobite. Specieswhich do not exist outside caves Vicariant event. The division of the biota or taxon (Schiner, 1854); they may, ho"vever,occur in the through the development of a natural barrier. superficialunderground compartment or in the upper Wackstones. Mud-supported carbonate sedimentary hypogeanzone. rocks containing >10% grains (particles>20ptm in : Troglobitization. Troglogenesis. diameter.l.e.g. calcarenite. Troglogenesis.The morphological,behavioural, physi- ological and other changesfound in species/lineages as they populate/adaptto cave or hypogeanlile. REFERENCES Troglomorphic. Pertaining to morphological, be- havioural and physiologicalcharacters that are con- Arp, A.J. and Frsher,C.R., 1995. Introduction to the symposiurn: Life ,4n. Zool.. vergent in subterraneanpopulations (Christiansen, with sul{ide. 35: 81-82. Attila, K.. 1996. The Cauers'Liuing Dictiorun,. Hungarran t962). SpeleologicalSocicty, Budapest. Troglophile. Species able to live and reproduce Botosaneanu, L. (Editor), 1986. Stygofuuna Mundi: A Fuunistic, undergroundas well as in the epigeandomain. Distribtrtional, and Ecologicul Swtie,v.r rf the Wrld Fuuna inhuhiting Suhten'anean Waters(intluding the nturine interstitial). Troglos. Assemblageof organismsinhabiting under- 'terrestrial' E.J.Brill. Leiden, 740 pp + endpapers. ground voids, often restricted to the Boutin. C. and Coineau.N., 1990. "RegressionModel", "Modele orsanismsin air-filled voids. Biphase" d'6volution et origine des micro-organismesstygobies BACKGROUND AND GLOSSARY 13

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