International Journal of Speleology

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International Journal of Speleology

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International Journal of Speleology
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Vol. 36, no. 1 (2007)

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International Journal of Speleology 36 (1) 1-21 Bologna (Italy) January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick, Canada. Max Moseley 1 INTRODUCTION The central focus of biospeleological research and thinking especially in North America has been those highly-adapted specialised troglobiontic animals which inhabit deep caves and subterranean waters in regions to the south of the maximum limits of the Pleistocene glaciations. Much less attention has been paid to the subterranean fauna of formerly glaciated northern regions of the continent, or to the many nonobligate species that occur in cave thresholds there and elsewhere. The limits traditionally imposed by an emphasis on troglobionts to the exclusion of other fauna have long hindered progress in understanding hypogean fauna and ecosystems. However this is now changing. Most cave ecologists now accept the need to see subterranean communities in their entirety encompassing species at many different stages of adaptation (Gibert & Deharveng, 2002), and evolutionary cave biologists increasingly recognise the value of non-obligate cavedwellers as empirical models of natural selection and 1) Research Associate, Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Canada B3H 3A6 (E-mail: moleslei@yahoo.ca) Moseley M. 2007. Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick, Canada. International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). ISSN 0392-6672. The vertebrate and invertebrate fauna, environment and habitats of caves and disused mines in Nova Scotia and southern New recession of the Pleistocene glaciers. The statistical composition of the fauna at the higher taxonomic level is similar to that in Ontario, but is less species rich and there are some notable ecological and other differences. Porcupine dung accumulations are an important habitat in the region, constituting a cold-temperate analogue of the diverse guano habitats of southern and tropical caves. Parietal assemblages are, as in other cold temperate regions, an important component of the invertebrate fauna but here include post-glacial recolonisation of the subterranean habitat in Nova Scotia has been relatively delayed. Finally the general and regional Keywords: Cave fauna, Canada, Nova Scotia, New Brunswick, porcupine dung, guano caves, parietal, threshold fauna, introduced taxa, post-glacial, recolonisation. Abstract: Received 8 July 2006; Revised 10 October 2006; Accepted 31 October 2006 adaptation in the underground environment (e.g. Kane & Culver, 1992). Accordingly there is now a clear need for and groundwater fauna of neglected regions of the continent, particularly Canada. The eastern region of Canada comprising Newfoundland, the Maritime Provinces (Nova Scotia, New Brunswick, Prince Edward Island), Quebc, and Ontario has many areas of karst, and natural dissolution caves are known from all these provinces except Prince Edward Island. However there have surveys. The exceptions are a detailed study of the fauna and ecology of Frenchmans Cave, Nova Scotia (Calder & Bleakney, 1965; 1967) and a general survey of caves and mines in southern Ontario (Peck, 1988). The geographical area dealt with in the present paper, Nova Scotia and the southern part of the adjacent province of New Brunswick (Fig. 1), is the northern end of the Northern Appalachians region. The Maritime Provinces together with areas of eastern Quebc are the Canadian part of the early French colony of Acadia; hence Acadian is a useful term often used in reference to this region. Available online at www.ijs.speleo.it International Journal of Speleology International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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2 Max Moseley The area is cold-temperate, and somewhat low-lying with maximum elevations in Nova Scotia of 532m and 446m in southern New Brunswick. The whole of the present-day land area was subjected to multiple glaciations during the Pleistocene and was icecovered at the maximum of the last (Wisconsinian) the result of immigration and recolonisation during glaciers, which is believed to have started ~21,000 years BP and to have been complete by ~11,000 years BP (King, 1996). Most species must have arrived from the south and west. Recolonisation of New Brunswick from these directions was relatively unrestricted by major physical barriers but immigration of many animals and plants into Nova Scotia is thought to have been constrained by the Tantramar Marshes, a narrow marshy isthmus which is the only land connection with New Brunswick: as a consequence the province is zoogeographically an island for many species (Fig. 1). The Strait of Canso physically further isolated Cape Breton Island until completion of the Canso Causeway in 1955 (Fig. 1). However there is evidence from present-day patterns of animal and plant distribution that some species may have recolonised Nova Scotia from the east by migration from an icefree Wisconsinian Atlantic Coastal Plain Refugium (Schmidt, 1986) or more likely from the emergent land areas which existed during the process of deglaciation (King, 1996). The existence of other late-Pleistocene refugia has also been proposed (e.g. Schmidt, 1986) but this remains highly speculative. In the historical period European contact resulted in the introduction of many exotic species through human migration and sea-borne trade. The area has a number of exposed areas of sul phate (gypsum-anhydrite) and carbonate (limestone and dolostone) bedrock with underground drainage, springs, caves and other geomorphological karst fea tures (Moseley, 1976; 1996: McAlpine, 1979). The southern part of mainland Nova Scotia has no sig are generally well-distributed throughout the rest of the region, with gypsum caves and karst predominat ing in Nova Scotia and limestone in New Brunswick. Dolostone karst is rare and no caves are known. Fig. 1. Outline map of Nova Scotia and southern New Brunswick, to show location of study sites. [Provincial and county boundaries are shown by solid lines; the broken line represents the Canada-USA international border. Dots may represent more than one cave or mine. See text for key to alphanumeric codes.]. International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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3 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada Moseley (1996) gives maps showing the surface distri bution of karst-forming rocks. There are scattered notices of caves in various early geological survey reports, newspaper articles and other publications but the past forty years has seen the most intensive exploration and documentation. Approximately 50 dissolution caves are now documented in Nova Scotia and ~20 in southern New Brunswick. All explored caves are small with few exceeding 250m in length. The White Cave system (surveyed length = 515m) in New Brunswick; Hayes Cave (365m) and Point Edward Cave (293m), both in Nova Scotia, are the largest. The existence of more extensive subterranean systems is inferred from surface patterns of sinks and springs (Moseley, 1996). Both Nova Scotia and New Brunswick have long histories of underground hardrock mining and the many abandoned mines provide ecologically cave-like habitat. Emerton (1917) reported the Cave Spider Meta ovalis (as M. menardi ) in Nova Scotia, although not from caves. Two specimens in the Canadian National Collection (CNC 3437, 3438) collected in Gays River Gold Mine, Nova Scotia, in 1963 appear to be the earliest of cave animals in the area are of hibernating bats: the Northern Long-Eared Bat ( Myotis septentrionalis ) was reported by Gould (1936) in a cave in Hants County, Nova Scotia. Bleakney (1965) later found Eastern Pipistrelle ( ) at several of the occurrence of the common Little Brown Bat ( Myotis lucifugus ) in caves and mines in that province. In New Brunswick, McAlpine (1976) reported P. author (1979) summarized underground records of this and other bats. There is one sight record of the Big Brown Bat ( Eptesicus fuscus ) in a cave hibernaculum in Nova Scotia (Scott & Hebda, 2004). Records of bats at underground sites in Nova Scotia have recently been collated by Moseley (in press). There has been no systematic effort to document other vertebrates, but occasional records and observations have accumulated. North American Porcupine ( Erethizon dorsatum ) (Fig. 2) and their dung have been observed in most caves and a number of disused mines in southern New Brunswick and the mainland of Nova Scotia (Calder & Bleakney, 1965; 1967; McAlpine, 1979; Moseley, 1998). Porcupine did not reach Cape Breton Island until completion of the Canso Causeway. They have since become established in the east of the island (Scott & Hebda, 2004) but there are no reported sightings yet in island caves or mines. Several other vertebrates have been sporadically recorded underground. McAlpine (1977, 1979) lists Mink ( Mustela vison ) scat, Smokey Shrew ( Sorex fumeus ), Deer Mouse ( Peromyscus maniculatus ) and a Beaver ( Castor canadensis) den in New Brunswick. Beaver have also been seen in a Cape Breton stream cave (Sawatzky, 1986) and Raccoon ( Procyon lotor ) tracks were reported by Calder & Bleakney (1967) in Frenchmans Cave. There are occurrence records of pallid Brook Trout ( Salvelinus fontinalis ) in limestone stream caves (Moseley, 1975; McAlpine, 1979) and Ninespine Stickleback ( Pungitius pungitius ) and Northern Redbelly Dace ( Phoxinus eos ) in Hayes Cave (Morris, 1985). Frog ( Rana clamitans ) tadpoles were collected in the threshold of Hayes Cave (Morris, 1985). terrestrial fauna of Frenchmans Cave, Nova Scotia (Calder & Bleakney, 1965; 1967). Thirty-eight invertebrate taxa were reported from the threshold and deep threshold, the majority associated with decomposing porcupine dung accumulations. Acari were found to be numerically dominant in poorly decomposed dung samples, whilst Collembola became the most abundant microarthropods later in the ecological succession. Broader, extensive rather than intensive, inventory of the regional invertebrate cave fauna began in the early 1970s with occasional sampling mostly as an adjunct to other underground work such as cave mapping, and continued through the 1980s and 1990s: a few of the records were published by the former Nova Scotia Speleological Society (Moseley, 1998 and references therein). Earthworms collected from several sites in New Brunswick were discussed by McAlpine & Reynolds (1977) and a brief summary of the composition of other New Brunswick cave fauna was provided by McAlpine (1979). During habitats and invertebrate fauna was performed (Moseley, 1998). A few specialised studies have been published based on the collections: Marusik & Koponen (1992); Moseley & Hebda (2001); Moseley et al. (2006); and Majka et al. (in press). Christiansen & Bellinger (1980, 1998) included records of Collembola in their comprehensive monographs on the North American collembolan fauna. Fig. 2. Young adult North American Porcupine ( Erethizon dorsatum ) in den, dark zone, Cheverie Cave, Nova Scotia. (Photo: F. Vladi) International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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4 Except for a preliminary unpublished study of Hayes Cave (Moore, 1963) investigation of the cave environment and ecology started with Calder and Bleakneys (1965, 1967) work, already referred of porcupine dung as a cave habitat in the region. McAlpine (1979) suggested that plant detritus and porcupine dung were the principle energy sources in New Brunswick caves. In Nova Scotia, Hayes Cave and its environmental conditions were investigated by the provincial Museum of Natural History (Scott, 1979; Morris, 1985). The presence of porcupine dung, plant debris, and scattered bat droppings (<250m of passage) as sources of energy was noted. ecological survey of cave habitats and environmental conditions in any substantive geographical region of Canada. It also contributes to information about parietal assemblages, and adds to the very limited knowledge of the ecology of porcupine dung caves. An interim report was issued in manuscript as a museum Curatorial Report (Moseley, 1998) OBJECTIVES AND METHODS This study is intended as an overall assessment of the taxonomic composition and diversity of the invertebrate and vertebrate cave fauna of mainland Nova Scotia, Cape Breton Island and southern New Brunswick, together with a preliminary description of the ecology of the caves and mines. It is based on miscellaneous qualitative collections and ecological a more representative survey in Nova Scotia in 1997. Terrestrial fauna was sampled by hand-collecting, baited pitfall traps, and Tullgren extractions of dung, litter and soil samples: aquatic fauna was taken by dip-nets, giant pipettes, sieves and kick sampling. Field notes were made of habitat, substrate, faunal associations, temperature and other environmental conditions. The stage of decomposition of porcupine dung was estimated using visual appearance, as preserved using standard museum techniques and distributed to appropriate specialist taxonomists in the collections of the Nova Scotia Museum and/ or retained by the relevant taxonomist. Published records, and unpublished records solicited from other workers, are incorporated into the text and tables. There are faunal records (i.e. at least one taxon determined to genus or species) from a total of 26 natural caves and 11 abandoned mines and Brunswick. Study sites were distributed throughout the geographical area of investigation in order to obtain representative taxonomic coverage, although and some collections were made in all seasons. The approximate locations of study sites are shown in Fig. 1, and a list follows: the-Bats (CB), Lake Charlotte Gold Mine (LCM), Tunnels, York Redoubt (YRT), Gays River Gold Mine (GRM) [Halifax Co.]; Hayes Cave (HC), Woodville Ice Cave (WIC), Frenchmans Cave (FC), Frenchmans II (F2), Weir Brook Cave (WB), Minasville Ice Cave (MIC), Millers Creek Cave (MC), Cheverie Cave (CC), The Honeycombs (TH), Peddlars Tunnel (PT), Centre Rawdon Gold Mine (CRM), Walton Barite Mine (WBM) Brook Cave (MB), New Laing, adit # 1 (NL1), New Laing, adit # 3 (NL3) [Pictou Co.]; Diogenes Cave (DC); Mabou Cave (MCI) [Inverness Co.]; Fairy Hole II (FH2) Lear Shaft (LSH) [Colchester Co.]. (GM), Glebe Pot (GP), Kitts Cave (KC) [Kings Co.]; Greenhead Cave (GR), Howes Cave (HO), Harbells Cave (HB) [St. John Co.]; Hillsborough Bat Cave (BC), Co.]. Descriptions and/or maps of the more important caves may be found in Moseley (1976, 1996) and Arsenault et al. (1997). For convenience collection sites are referred to hereinafter by their alphanumeric codes, as given above. Taxa were categorized using the widely recognized Schiner-Racovitza categories of troglobite troglophile and habitual trogloxene for cavernicolous forms and accidental for strays and animals brought into the status of taxa found underground is empirically about the status of those in this study are tentative. Taxa were assessed based on such evidence as the frequency of records from subterranean collecting sites and occurrence records of juveniles or larvae. Repeated long-term observations of a species from the same cave or mine and multiple collections from different sites are particularly valuable (Moseley, 1998). The known bionomics of a species elsewhere is also useful. All animals that are regularly found underground including threshold dwellers and guanophiles (see Discussion) are herein regarded as cavernicolous. Sampling was not comprehensive enough to satisfactorily map geographical distribution or accurately ascertain seasonal occurrence of most cave-occurring taxa within the study area. In this report threshold means that zone within support vascular green plants, deep threshold means the low light area beyond the inner limit of the threshold and dark zone that part of the cave permanently in absolute darkness. Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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5 RESULTS Fauna recorded The fauna records are collated and summarised in Tables 1 and 2. Sixteen vertebrate and 170 invertebrate taxa, representing 12 and 99 families recorded invertebrates by higher taxonomic group is given in Table 3. Excluding accidentals, ectoparasites and those taxa for which ecological status could not be assessed, 11 vertebrate and 123 invertebrate cavernicoles (including guanophiles) were recorded in 8 and 72 taxa (48%) are categorized as habitual trogloxenes, 44 (33%) as troglophiles, and 25 (19%) as guanophiles. Of the cavernicoles 112 (84%) are terrestrial animals, 22 (16%) aquatic. Of the 112 terrestrial cavernicoles 38 (34%) are categorized as parietal fauna. 19 (17%) of the terrestrial cavernicoles are almost certainly introduced non-indigenous species (all of them European in origin): the rest are Nearctic, circumpolar or cosmopolitan. All the non-indigenous species are invertebrates. Three further non-indigenous species which were collected are considered to occur only as accidentals in our caves. In the case of aquatic invertebrate fauna, 20 collections. Half of these are crustacea: Copepoda (5 species) or Ostracoda (5 species). Most of the rest are insect larvae: Odonata (3 species), Plecoptera (3 species), and Diptera (1 species). No introduced aquatic animals were found. Cave environment None of the accessible caves in the region have a constant temperature zone: all are subject throughout small size, the presence of a stream, or through drafts due to multiple entrances. Radon gas concentrations in the inner areas of one of the largest caves, HC, (Morris, 1985) suggest almost stagnant air but even here the temperature changes seasonally, although the annual range is only ~2C. This site may be considered deep cave in the sense of Howarth (1988). Annual temperature range in the dark zone of a more representative small Maritime Canadian cave is ~4C (Fig. 3): many caves experience greater variation. During spring and early summer as the ambient temperature rises above that underground relatively cold air is retained inside caves, whilst warm annual curve (Fig. 3). In certain extreme special cases this effect results in so-called ice caves (see below). Maximum seasonal cave temperatures are recorded in September-October, minimum in January-February. Eastern Canada experiences a very rigorous winter climate. Cave entrances are subject to severe low temperature conditions throughout the winter and there is often a build up of ice and snow. They are subjected to repeated freeze-thaw cycles: the average annual number of cycles (-6C to +2C) at Dartmouth, Nova Scotia has been calculated as 30. As a result of this and ongoing dissolution of exposed rock, talus derived from within the threshold itself or from the cliff face outside (e.g. CB, HC, WIC, and FC). This further inside resulting in fewer, perhaps only one, freeze-thaw cycle per annum in the deep threshold. The mean annual temperature inside caves in central Nova Scotia is ~5.7C. However it is lower at a few sites. The aspect and physical shape of certain caves cause atypical temperature conditions with ice and snow persisting into midor even late summer. Snow and ice which accumulate during the winter in WIC for example have been observed to survive until early August. The minimum air temperature measured inside the cave on 29 July1997 was 2.5C, and 0.4C was recorded a few centimeters below the used historically for cold storage or as a source of ice and are known locally as ice caves. However, none of the known local sites are true ice caves in the speleological sense of the term denoting a cave with permanent ice. The temperature of standing water is usually within 0.5C of the air in the immediate vicinity. There are several caves (e.g. CB, KC) with active stream sinks that modify the temperature. Temperature may also be affected by the presence of a spring. Some cave streams originate within the cave as eucrenal springs characterised by cold, clear sediment-free water with their temperature remaining within two degrees of the local annual mean. Examples are a small spring in MIC (4.3C in September 1995) and the F2 cave stream (5.1C in July 1997). Cave waters in both limestone and gypsum caves are usually slightly alkaline: pH7.3-7.6. Acidic conditions there are accumulations of porcupine dung or plant litter e.g. pH5.8 was measured in seeps with porcupine CB. Due to the high solubility of calcium sulphate (i.e. 2.438g.l in distilled water @10C) and the presence of other dissolved solids such as calcium carbonate, gypsiferous waters have high conductivity. Analysis of water samples from ponds in HC (Morris, 1985) showed conductivity readings of 2220S.cm Samples of water from FC were so high in calcium 3 4 5 6 7 8 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan C Fig. 3. Average monthly air temperature (C) in a representative Nova Scotia cave (dark zone in Frenchmans Cave). International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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6 sulphate that microcrystals of selenite precipitated out on cooling in a domestic refrigerator. Habitats Although characteristically small, caves in the region contain a diversity of terrestrial and aquatic eutrophic (dung), mesotrophic and oligotrophic habitats. North American Porcupine (Fig. 2) dens are almost ubiquitous in caves and mines everywhere except Cape Breton Island. A den site is typically a small side passage, cavity or blind pocket within the deep threshold or dark zone occupied by a single animal. Cave dens always have an accumulation of faeces either as scattered pellets or more substantial accumulations (Fig. 4). In areas adjacent to the porcupines access routes sustained inputs of dung sometimes result in deposits tens of centimeters deep dung piles can occur anywhere from the threshold through to the cave dark zone. Etiolated seedlings are commonly present on fresh dung growing from seeds which have passed through the gut of the porcupine (Calder & Bleakney, 1965) (Fig. 4). Decomposing dung fuels a varied community of bacteria, fungi, oligochaetes, insects, and other arthropods. The dung habitat is non-uniform, varying microclimatically and qualitatively with diet, stage of decomposition, and environmental factors including moisture content, ecological zone within the cave, acidic (~pH5.1) but in wet areas, such as where there is seepage water or under roof drips, the acidity is neutralized by the buffering effect of cave waters. Also, as observed by Calder & Bleakney (1965), the decompositional sequence is accompanied by decreasing acidity, so that well decomposed and/or Because of the variability of pH and other empirically often accumulate over many years, it was not possible to accurately determine the stage of decomposition of individual samples. However visual appearance is a useful approximate guide to the sequence. Fresh scat (Fig. 4) consists of scattered, ovoid, greenish-grey pellets with a mucoid surface. The mucoid material disappears rapidly ( poorly decomposed ). The pellets retain their shape and physical integrity for some time but turn constituents becomes visible ( moderately decomposed ) (Fig. 5). In the later stages of decomposition they break down physically forming a material rather like dark well-weathered sawdust in appearance and consistency ( well decomposed ) (Fig. 6). Invertebrate communities in porcupine dung piles are dominated by Acari, Collembola, dipteran larvae and enchytraeids (Figs. 7, 8). There are marked differences in the species composition and biomass of this community from site to site depending on microclimatic and qualitative factors. More than 35 terrestrial invertebrate taxa are recorded associated with dung in the Frenchmans cave system (FC + F2), while the other extreme is represented by a remarkably simple ecosystem observed in GM where samples of well composted dung from the dark zone yielded only Protaphorura armata (Collembola) and enchytraeids. Qualitative observational evidence suggests that moisture content is the most important variable: both biomass and species diversity decrease in dry material. The abundance of Enchytraeidae in particular is affected by the moisture content and they are infrequent or absent in drier samples. Decompositional sequence is also a major factor. The general ecological succession reported by Calder & Bleakney (1965) with Acari numerically most abundant in poorly decomposed samples and Collembola becoming dominant later has been observed at other sites. It was very apparent in GM where mites were abundant in moderately decomposed material (which also contained isotomid Collembola as well as P. armata and Enchytraeidae) but, as mentioned above, were absent from samples of well composted dung. group in porcupine dung. Most have so far only ( Parasitus, Eugamasus, Vulgarogasmus ) are almost always abundant, and rhagidids ( Rhagidia ) are usually common. Other mites, some of which may be common to abundant at some sites, include Vegaia ), Zerconidae ( Zerconopsis ), Ascidae ( Arctoseius ), Ameroseidae ( Epicriopsis ), Eviphidae ( Alliphis ), Macrochelidae ( Geolapsis ), Pygmephoridae ( Pygmephorus Bakerdania ), Tetranchidae ( Bryobia ), Acaridae ( Acarus immobilis ), Banksinomidae ( Oribella ) Histiostmatidae. Collembolan populations are less diverse. They are almost always dominated by onychiurids ( Protaphurura Tullbergia ) and isotomids ( Folsomia, Isotoma ). Neelids ( Megalothorax minimus ), podurids ( Willemia scandinavia ) and entomobryids ( Pseudosinella alba, Tomoceros minor ) are found more infrequently. The insect fauna of dung is dominated by nematoceran Trichocera maculipennis and various sciarids are characteristically present and usually abundant (Figs, 7, 8). At least three different types of sciarid larvae are found, none of which have yet been matched with the adults recorded associated species of Bradysia, Lycoriella and Scatopsciara. Larvae of Limonia cinctipes Chaoborus Smittia and Psychoda also occur in some samples, as do those Leptocera Adult Scatopsciara, Chaoborus and Leptocera have been observed attracted to fresh scat, presumably ovipositing. A few beetles are also found in this habitat. Larvae of Quedius s. spelaeus are often common in moderately decomposed dung, whilst the somewhat rarer adults almost always on or near dung (Moseley et al., 2006). The tiny guanophile Acrotrichis castanea is sometimes abundant, although it may be overlooked because of its size. Aphodius aleutus and Corticaria pubescens have been collected from dung: both occur in such Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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7 Fig. 4. Entrance of cave den occupied by the porcupine illustrated in Fig. 2. Numerous fresh droppings are clearly visible. The white areas are mats of fungal hyphae on older dung, and there are a number of etiolated plant seedlings growing from seeds which have passed through the gut of the porcupine. Several porcupine quills are also visible. (Photo: author). Fig. 5. Moderately decomposed porcupine dung sample from Frenchmans II, Nova Scotia (Photo: author). Fig. 6. Well decomposed porcupine dung sample from Frenchmans II, Nova Scotia (Photo: author). Fig. 7. Fauna extracted from a sample of moderately decomposed porcupine dung, Frenchmans II, Nova Scotia, October 1997. Sciarid (Diptera) larvae are abundant, and there is an adult in the lower right. Parasitid mites and onychiurid Collembola ( Protaphorura armata ) are the lower centre is Quedius s. spelaeus, and there are two adult ptilid beetles (Acrotrichis castanea) just above this. (Photo: C. Majka). Fig. 8. Fauna extracted from a second sample of moderately decomposed porcupine dung, Frenchmans II, October 1997. Trichocera maculipennis larvae (Diptera) and an enchytraeid are (Photo: C. Majka). International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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8 habitats on the surface. Other terrestrial invertebrates recorded associated with dung include Oniscus asellus (Isopoda), Lamyctes fulvicornis (Chilopoda), Proteroiulus fuscus Ophyiulus pilosus Polydesmus angustus (Diplopoda), and two linyphiid spiders Sisicottus montanus and Grammonota. Two earthworms ( Dendrodrilus rubidus and Aporrectodea tuberculata ) have also been collected, but as porcupine dung is almost unpalatable to earthworms it is at best a marginal habitat (McAlpine & Reynolds, 1977). Most local caves are at shallow depth and in consequence detritus and plant debris seep in from the surface through crevices. Leaf and other plant litter also often accumulate particularly in cave and mine thresholds. Damp, often rotting, support timbers are found in disused mines. Beavers denning in caves (e.g. KC) store woods such as willow and alder as food. In stream sink caves such as CB, freshets and plant debris and organic sediment. The varied invertebrate community of vegetation litter and detritus includes Dendrodrilus rubidus Aporrectodea tuberculata and Eisenia rosea (Oligochaeta); Hypogastrura pseudarmata, Neanura muscorum, Willemia scandinavica, Protaphorura pseudarmatus, Isotoma caeruleatra, Isotoma sp. nova? Pseudosinella collina, Sminthurides malmgreni, and Ptenothrix marmorata (Collembola); Quedius mesomelinus, Brathinus nitidus, and Gennadota canadensis (Coleoptera); various dipteran larvae; Parasitus, Eugamasus, Vegaia, Linopodes motatorius, Cocceupodes, Rhagidia, and Glycyphagus domesticus (Acari); and Discus catskillensis (Gastropoda). Flood debris is often rich in many otherwise unexpected aquatic stages of insects and other accidentals. Most are dipteran larvae e.g. Tipula, Erioptera pilipes, and Chrysops other arthropods and gastropods occur regularly on cave walls and ceilings. Several arthropods ( Oniscus asellus [Isopoda], Polydesmus angustus [Diplopoda]) and gastropods ( Arion Deroceras laeve, and Trichia hispida) are commonly found within or near the threshold but almost never further inside. With the exception of these this assemblage tends to be richest both in number of species and in number of individuals in the deep threshold, but it extends into the dark zone. The species composition changes seasonally. Diptera predominate: there is an especially species rich fauna of mycetophilids ( Boletina, Bolitophilia, Rhymosia, Exechia, and Exechiopsis ) and helomyzids ( Scoliocentra, Helomyza, Amoebaleria, and Tephrachlamys ). By far the most numerically frequent Diptera in these caves are Trichocera maculipennis and various sciarids: adults of these Culex females are abundant at many sites in winter. Other common Limonia cinctipes, Chaoborus, Psychoda and Leptocera Dolichopeza, Anopheles and Peromyia As already mentioned the larvae of L. cinctipes, T. maculipennis, Chaoborus, Psychoda, and Leptocera live in porcupine dung. Other insects and arachnids found on cave walls include Ceuthophilus brevipes (Orthoptera); Scoliopteryx libatrix and Triphosa haesitata (Lepidoptera); Nelima elegans (Opiliones); Meta ovalis and Nesticus cellulanus (Aranea). One gastropod, Zonitoides arboreus is found further inside than other gastropods. Oligotrophic habitats are uncommon in caves in this region. They are more usual in limestone caves than in gypsum but can occur in the latter (e.g. F2) where they typically comprise areas of pebbles, gravel and/or cold spring. Most terrestrial taxa recorded from such sites are Collembola: Heteromurus nitidus, Arrhopalites hirtus, and Arrhopalites nr. pygmaeus Allajulus latestriatus have also been collected. The pool surface association comprises various Collembola ( Protaphora cf. boedvarssoni Folsomia candida, Isotoma sp. nova? Heteromurus nitidus, Pseudosinella alba and Arrahopalites hirtus ), occasional Symphyla ( Scutigerella ), and a number of Liposcelis ) was collected at one site. Aquatic habitats comprise standing water, ranging (e.g. HC); running water, ranging from tiny seeps and rivulets (e.g. WIC) to large streams (e.g. KC), and interstitial water. Cave streams may originate from the surface, or from an underground spring. In most grained chocolate-brown sediment representing the insoluble residue of gypsum dissolution. These aquatic habitats support a diverse fauna dominated by copepods ( Acanthocyclops spp., Eucyclops agilis Diacyclops crassicaudis Paracyclops poppei and Macrocyclops albidus ) ostracods ( Pseudocandona albicans Cypria Cavernocypris Cypridopsis and Fabaeformiscandona wegelini ), microdrile oligochaetes and aquatic insect larvae: particularly Odonata, Plecoptera and chironomid capnids, Amphinemura (Nemouridae), Taeniopteryx (Taeniopterygidae), Haploperla (Chloroperlidae) and from the threshold although Haploperla can live ( Aeshna, Macromia ) are found frequently enough in cave thresholds to be considered habitual trogloxenes in that habitat. Planarians may also be present in pools in cave dark zones, and a water-beetle Agabus larsoni has been collected from dark zone pools and streams in both gypsum and limestone caves and present in cave pools near the entrance, sometimes straying well into the dark zone, but most of these are essentially part of the threshold fauna or accidentals. The leech Helobdella papillata and another waterbeetle A. semivittatus are examples of the latter. Haploperla sp. nymphs (Plecoptera) and Simulium Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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9 sp. larvae (Diptera) occur in the oligotrophic eucrenal streams arising from underground cold springs. looking Brook Trout ( Salvelinus fontinalis ) are sometimes seen in clear well-oxygenated streams well inside the dark zone in limestone caves. The Ninespine Stickleback ( Pungitius pungitius ) is at home in the thresholds of gypsum caves in large pools that have a connection with outside waters. It does not however stray far into the dark zone. A population of Northern Redbelly Dace ( Phoxinus eos ), was observed in HC for other cave. Several different food inputs support animal communities in aquatic habitats. Dead insects are an important energy source. Insect corpses, especially Diptera, at times accumulate in large numbers on and in cave pools. A sample collected from a pool in F2 in October yielded Trichocera maculipennis, Leptocera, numerous sciarids and several Quedius s. spelaeus larvae and adults. These insects originate from the dung fauna and thus aquatic ecosystems are indirectly supported by porcupine dung. Dung may also be present as scattered droppings, or, sometimes (e.g. PT and GM) in substantial accumulations. Associated fauna includes planarians, aquatic microdriles, copepods ( Acanthocyclops venustoides ) and dipteran larvae. Bat droppings never form substantive accumulations: Scott & Grantham (1985) observed cyclopoid copepods associated with droppings in ponds in HC and Moseley (unpublished) made the same observation in MC. sinking streams is also an important source of food in some caves. This input tends to be seasonal, with most material being brought in by spates during the spring snowmelt. The aquatic invertebrate fauna found associated with such material is more diverse than where this food supply is not available, but it the many accidentals carried in along with the plant debris. Beaver living quarters with stored willow and alder were found well inside the dark zone of KC (McAlpine, 1977). The site had been abandoned when it was examined in 2005. Sites that meet appropriate microclimatic and morphological conditions and are relatively species of insectivorous vespertilionid bats as winter hibernacula. Three gregarious non-migratory species ( Myotis lucifugus, M. septentrionalis and Pipistrellus ) commonly use caves and mines as winter hibernacula, entering in late September/early October and leaving in early summer. Another non-migratory bat Eptesicus fuscus preferentially hibernates in buildings but may occasionally use underground sites (Scott & Hebda, 2004). Hibernating are solitary and only the two Myotis spp. form hibernating colonies. The known colonies are not large: the population (>95% Myotis spp.) in the largest known hibernaculum is estimated to be <10,000: other sites rarely contain more than a few hundred animals. Small numbers of male bats also sometimes roost in underground sites in mid-summer, whilst in latesummer (September and early October) bats roost in caves and mines by day and continue feeding at night in preparation for hibernation. Bat droppings that accumulate in late summer have been seen in several form. Acari and Collembola are often present on bat droppings (Scott & Grantham, 1985). reported from caves and mines are listed in Table 2. Wright (1979) reported two species of ectoparasitic Acari and one siphonapteran collected exMyotis from non-cave sites. DISCUSSION Composition of the invertebrate fauna The invertebrate fauna is in many ways similar in composition to that of Ontario and northern Europe but there are also some important differences. Taxonomically it is dominated by insects and Collembola, followed by arachnids. Mollusca form from the two previous eastern Canadian regional surveys (Table 3). Pooled results from all three studies indicate that in eastern Canadian caves, insects and Collembola typically constitute ~60% of all invertebrate species, arachnids ~20%, molluscs ~5% and all other invertebrates ~15%. In the taxonomic groups Diptera, Aranea, Opiliones and Mollusca, as well as in the overall total of all taxa, the number of taxa recorded in the present southern Ontario caves (Peck, 1988) (Table 3). This the present survey area, particularly Nova Scotia which has a somewhat impoverished provincial fauna due at least in part to the Tantramar Marshes zoogeographical barrier. More intensive collecting will The four taxonomic groups referred to are dominated by species from the parietal assemblage (see below) which constitutes a major faunal association in caves and mines in both Ontario and the Maritimes. In the Acari and the Collembola the opposite is the case: the local cave fauna is more species rich than that of Ontario. Both these taxonomic groups are wellrepresented in communities living in porcupine dung, a habitat not reported in southern Ontario caves by Peck (1988). invertebrates found associated with different organic terrestrial troglophiles than is porcupine dung. Rotting timbers in abandoned mines and elsewhere provide a habitat for a few earthworms, Acari and Collembola, but the fauna found in them does not appear to be very rich in this geographical area. Some invertebrate species appear to be almost exclusively associated International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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10 with oligotrophic sites e.g. Heteromurus nitidus and Arrhopalites hirtus. As in virtually all cave ecosystems porcupine dung trophic levels, primary producers and herbivores, absent. They differ from most other cave guano ecosystems in that the porcupine is herbivorous feeding preferentially on the cambium of trees, especially conifers. The remaining two classic trophic levels, decomposers and predators, are easily recognized. This is usually true of cave guano communities but may not be the case in oligotrophic cave communities where decreasing resources are associated with blurring of the distinction and even disappearance of obligate predators in favour of omnivores (DeHarveng & Bedos, 2000). Quedius s. spelaeus adults and larvae and various predatory mites (e.g Alliphis Geolapsis ) are the common predators in porcupine dung communities. There is an unusual suite of parietal species that is ecologically fundamentally different in its nature from the original sense of Jeannel (1926). It constitutes a distinct faunal component derived directly from the porcupine dung fauna. The larvae of several Diptera ( Limonia cinctipes, Trichocera maculipennis, Chaoborus, various sciarids, Psychoda, and Leptocera ) live in porcupine dung, and adult Scatopsciara, Chaoborus and Leptocera have been observed apparently ovipositing in very fresh pellets, thus, unlike almost all the traditional parietal insects, these Diptera are probably able to complete their life cycle underground. This is reminiscent of the situation in the humid tropics where there is frequently an assemblage of arthropods derived from guano communities on cave walls deep inside caves (Deharveng & Bedos, 2000). A parietal association in Jeannels (1926) sense can also be readily recognized in Maritime caves. It is very similar to that found in Ontario (Peck, 1988), northern Europe and other cold temperate regions of the world and comprises the associations of arthropods living or resting on cave walls and other rock surfaces together with the spiders that prey upon them. Parietal predators tend to be specialized forms found only in this and similar dark humid habitats such as cellars, and are usually considered to be troglophiles. Most of the parietal association however consists of habitual trogloxenes that are using the habitat temporarily for shelter, summer aestivation, hibernation, overwintering, or other purposes that are not yet fully understood. Adult Diptera predominate both in number of species and number of individuals. Overwintering fauna includes Nelima elegans (Moseley & Hebda, 2001), female Culex spp. and some species of Bolitophilia, Rymosia and Tarnania (Mycetophilidae). Some of the Exechiini (Mycetophilidae) along with the adult Limniphilidae occasionally seen in New Brunswick caves are probably there for summer aestivation. Two spiders ( Meta ovalis and Nesticus cellulanus ) specialise in predating other parietal fauna. Ceuthophilus brevipes uses caves and mines for shelter and forages outside. There are also two moths Scoliopteryx libatrix and Triphosa haesitata Adult helomyzids ( Scoliocentra, Helomyza, Amoebaleria, and Tephrachlamys ) are common and are present year round but their purpose in entering caves is unknown: their larvae are not found in this habitat. Several other arthropods ( Oniscus asellus [Isopoda], Polydesmus angustus [Diplopoda]) and gastropods ( Arion Deroceras laeve, Trichia hispida) are commonly found on rock surfaces in the threshold but rarely much further in. Another gastropod Zonitoides arboreus appears to be somewhat more cavernicolous in its behaviour than the other recorded mollusca, and is sometimes found well inside the dark zone. These species are also common in dark moist habitats on the surface and unlike the animals traditionally included in the parietal association are apparently not in caves characteristically present in this habitat and the arthropods are preyed upon by parietal spiders, it seems logical to include them as parietal fauna. The subterranean aquatic fauna is undercollected and the statistical dominance of terrestrial (80%) versus aquatic (20%) taxa in the lists certainly Most of the recorded vertebrates enter caves for habitual trogloxenes. Salvelinus fontinalis and Pungitius pungitius are seen frequently enough to suggest that in behaviour they are the most cavernicolous vertebrates in the region. Beaver as well as Porcupine use underground sites as dens. McAlpine (1977) reports records of beaver in caves from Alabama, Missouri and New Brunswick. Two small mammals, the Smokey Shrew ( Sorex fumeus ) and the Deer Mouse ( Peromyscus ) are known from research elsewhere to Trevor-Deutsch, 1973) and are thought to enter local caves primarily for this purpose, whilst Raccoon and Mink enter them to forage. The remaining vertebrates are probably accidentals. This is certainly the case for the adult frogs seen have however been observed to survive for some time where a stream brings in food and offers suitable habitat (e.g. CB). Phoxinus eos was observed in HC for other cave and collected specimens were pallid and undernourished. A single extra-limital winter record of the migratory bat Lasionycteris noctivagans in PT probably represented a stranded storm-driven animal (Hebda, pers. comm.) Energy inputs: porcupine dung Porcupine and extensive deposits of their dung 1962). Calder & Bleakney (1965, 1967) investigated Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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11 the ecology of a porcupine-inhabited cave in Nova Scotia. Despite the wide distribution of the porcupine porcupine dung habitat in caves has been overlooked or neglected. Graham (1962) stated that it was poor in invertebrate life. In a recent review of cave guano communities Gnaspini & Trajano (2000) do not mention porcupine dung. Substantial and sustained inputs of porcupine dung are the principle energy source supporting subterranean communities in New Brunswick and mainland Nova Scotia. These communities often display considerable species diversity. This is well illustrated by the St. Croix group of caves (FC + F2 + WB) in Nova Scotia where some 89 taxa in 64 families the humid tropics where inputs of externally derived organic energy sources such as bat and cricket faeces support some of the most diverse cave ecosystems known (Gunn et al., 2000). There are further ecological and faunal similarities with tropical guano caves. Although species-rich in comparison to other cave habitats, guano communities are in general simpler than those above ground (Gnaspini & Trajano, 2000). Calder & Bleakney (1965) demonstrated that the microarthopod fauna of FC is less diverse than that of nearby epigean habitats. Acari, as noted by Gnaspini & Trajano (2000) in Brazil by Collembola. Mites are almost never dominant organisms in caves except in guano. The presence of an unusual parietal association derived directly from dung communities has already been referred to. Accordingly this fauna can be seen as a rare coldtemperate North American analogue of tropical guano cave faunas. It must be emphasized that porcupine dung does not only directly support a guanobious fauna, but indirectly contributes substantially to other aquatic and terrestrial communities through the export of this fauna to other areas of the cave. The habitat thus has a more important status regionally, species diversity than hitherto recognized. Historically, the guano habitat has not been seen as a true cave habitat and guanophiles have been categorized as false cave-dwellers (see e.g. Gnaspini & Trajano, 2000 and references therein). This view can be challenged as an extreme extension of the a priori assumption that the cave environment is per se oligotrophic (itself an extension of the troglocentric dismissal of the guano fauna in tropical caves seems to be at least in part an unconscious derivation from a belief, now known to be false, that there were no or very few troglobites in the tropics. Troglobites are in fact not restricted to oligotrophic habitats. In Hawaii a rich fauna of specialized troglobites is found in the food-rich habitat provided (Howarth, 1972). Some animals which are dependent on guano are traditionally accepted as troglophiles. This is almost certainly the case for instance with Trichocera maculipennis distinctly uncommon in surface habitats (Jefferson, 1981). It also seems somewhat illogical to exclude organisms that complete their life cycle underground as not cavernicolous whilst accepting those which for example merely use caves as temporary seasonal shelter. Finally, in order to reach and colonise guano piles organisms must be able to orient and survive in the wider cave environment (Gnaspini, 1992). Thus, as proposed by Gnaspini (1992), guano should be treated simply as a substrate within a cave. All animals regularly found in and utilizing caves and cavernicolous. Those guanophiles regularly occurring in subterranean habitats should be treated as one ecological category of cave fauna. Other energy inputs main energy source in Cape Breton sites, where the constitutes a special habitat because it is seasonally pulsed. Rotting timbers are available at some sites, especially abandoned mines. Insectivorous bat droppings are only of incidental importance as a food source, being localized and never forming substantial guano deposits. Where bat colonies occur they are small and droppings only accumulate in hibernacula for a few weeks during the autumn. Seasonality Circannual rhythms in cavernicoles have not been observed or investigated in the Canadian Maritimes. However the cave communities are subject to several seasonal environmental changes and cues. These include annual temperature cycles, pulsed inputs of organic matter into stream caves during the spring thaw and bat droppings in late summer. The composition of parietal assemblages is seasonal. Origin of the subterranean fauna The present survey indicates that the subterranean fauna of the region consists of communities of nonobligate species which have all arrived in the area and colonised hypogean habitats at various different times and via several routes during the past ~11,000 years, or perhaps in some cases earlier during deglaciation (~21,000-11,000 BP). No convincing example of a preglacial survivor has been found. The apparently disjunct distribution of Q. s. spelaeus suggests that this troglophilic beetle may have arrived from the emergent land areas that existed on the present-day Atlantic continental shelf during deglaciation (Moseley et al., 2006) but evidence based The subterranean aquatic fauna needs more sampling as it is possible that one or more aquatic troglobites International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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12 (stygobites) could have survived and reinvaded Nova Scotia groundwaters from Atlantic refugia. An Cavernocypris (Ostracoda) may hint at this possibility. Stygobitic amphipods of the genus Stygobromus and on offshore islands as far north as southeastern Alaska (Shaw & Davies, 2000) and in glaciated regions 1981). of recent invasive non-indigenous taxa present in terrestrial cave communities. Of the cavernicolous terrestrial invertebrates 17% are probably European in origin (Table 1). Notable examples are the European threshold troglophile Nesticus cellulanus which is now established here, being found in cellars and similar dark damp places in the region, and the springtail records of N. cellulanus in North America (Ewing, North American locality for (Christiansen & Bellinger, 1980). It is also worth noting that specimens of the circumpolar Protaphorura armata (Collembola) from local caves are morphologically very similar to European examples from anthropogenic (agricultural) habitats (Pomorski pers. comm. 2006) and thus may represent another non-indigenous population. Some care must be taken in accepting a taxon as nonindigenous. Some North American species previously later been found to be distinct, and there are examples in the Acadian cave fauna. The spider Meta ovalis was distinguished from the European M. menardi only recently (Marusik & Koponen 1992) and the widespread North American collembolan Folsomia stella was long confused with the morphologically very similar Old World (Christiansen & Bellinger, 1980). In other cases species previously thought to be introduced have later been shown to be native e.g. the earthworm Dendrodrilus rubidus (Schwert, 1979). Nevertheless, taken as a whole, the list of probable introductions is convincing. The eastern Canadian seaboard has been subject to times, and has long been known as the probable point of introduction of many exotic invertebrates as a consequence of human migration and trade. Many of these are now widespread in surface habitats. Apparently a subset of pre-adapted species that have arrived and successfully established in the Maritimes has subsequently been able to enter and survive in subterranean habitats here. Many of the invasive species were collected in FC and/or F2 (Table 1), but it is uncertain whether this is related to the fact that these caves are located in one of the earliest areas of Acadian French settlement in Nova Scotia or is a result of collecting bias. Although recent work with epigean invasive species elsewhere suggests that many have colonized environments that are radically different from their sources (Lee, 2002) no evidence of this has been found in our non-indigenous cave-inhabiting fauna: all the introduced species listed in Table 1 are also reported either as common in caves (as troglophiles or habitual trogloxenes) or as guanophiles in their region of origin. There can be little doubt that rather than competing with established fauna, some or most of these exotic species are exploiting empty biotopes in the subterranean environment. In the case of the Isopoda and Diplopoda there are no native species in the lists, all are introductions. Most of the cave-collected earthworms and mollusca are non-indigenous taxa, and almost half the terrestrial beetles are European in origin. The widespread Nearctic threshold spider Nesticus pallidus was not found: it is replaced in this habitat by the closely related European N. cellulanus. niches must be at least in part due to the taxonomic impoverishment of the Nova Scotian fauna resulting from zoogeographical isolation of the province. Several authors (e.g. Chapman, 1993) have pointed out that cave faunas in formerly glaciated regions are in an early dynamic phase of recolonisation and adaptation. It appears that recolonisation of hypogean habitats in Nova Scotia has been relatively even more delayed and thus that subterranean communities are in general at an earlier stage of this process than those in other otherwise similar geographical areas. In Maritime Canada we also have the unusual situation where there are eutrophic dung communities in this early phase: guano caves in the humid tropics are ancient systems never subjected to glaciation. It is believed that the porcupine is a relatively late postglacial arrival in Nova Scotia (Calder & Bleakney, 1965). Colonisation, adaptation and speciation in cave faunas can be rapid processes that may be taking place on a human timescale. Lava tubes on the geologically very young island of Hawaii, which is less than 700,000 years old, already harbour a rich fauna of highly-adapted troglobites consisting of representatives of native groups in the process of adaptive radiation (Howarth, 1972). Before 1970 only a few local caves were well-known to naturalists and cave ecosystems in the region were assumed to be isolated, localised and restricted. Maritime Canadian cave fauna was not recognised as ecologically many more caves together with the important insight from elsewhere that most so-called cave fauna is not restricted to caves (an anthropocentric concept) as such but is widely distributed throughout mesocavernous voids 1984 and references therein) means that we must now recognize subterranean ecosystems as provincially and regionally notable. The porcupine dung habitat is exceptional and thus particularly important. fascinating, and cave ecosystems are natural systems man. Also, due to the protection that they offer from large oscillations in climate, caves and subterranean waters Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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13 are habitats where species normally living further to the north or to the south may sometimes be found (Gunn et al., 2000). The cave fauna of Nova Scotia is now probably the most extensively and comprehensively sampled and documented of any geographic region in Canada. Records go back to the 1960s and large documented collections of both terrestrial and aquatic invertebrates have been made since then in caves and abandoned underground mines around the province. Invasive and other species in Maritime caves may have high potential for investigating and testing biospeleological evolutionary theories with populations at a very early stage of active colonisation and adaptation to the subterranean environment. ACKNOWLEDGEMENTS Any prolonged wide-ranging study is impossible without the help and participation of many individuals, and it is unfortunately impossible to name them all here. I would especially like to express my gratitude there is a list in Moseley (1998). Mention must also be made of those individuals who accompanied me the Proctor brothers, Dr. D. Sawatzky, and the late Dr. P. Schwinghamer. My colleagues amongst the permanent staff and Research Associates of the Nova Scotia Museum of Natural History have also helped in many ways. Dr. Dale Calder (Royal Ontario Museum) commented on the manuscript and also provided useful unpublished data from Frenchmans Cave. Additional unpublished records were generously provided by Dr. Don McAlpine (NB Museum) as well as by Calum Ewing and Andrew Hebda (NS Museum) who also prepared Fig. 1. A mention needs to be made of Harry Bassett and Fred St. Peters who skillfully recovered decades of notes and raw data from a set of corrupted computer diskettes Finally, thank you to the two anonymous reviewers who critiqued the draft manuscript and made a number of very useful and constructive suggestions. Fieldwork in 1996/97 was supported in part by a grant from the endowment fund of the Nova Scotia Museum Board of Governors. Fig. 1 was generated from the NS Museum MIMS database. REFERENCES Arsenault S. P., Schroeder J., Brub D. & Albert R., 1997 The caves of southeastern New Brunswick (Revised and Supplemented) Issue 97-7, Open Files, Minerals and Energy Division, Dept. of Natural Resources and Energy, New Brunswick, 33 p. The Mammals of Canada National Bleakney J.S., 1965 First Specimens of Eastern Pipistrelle from Nova Scotia J. Mamm., 46 : 528-529. A second new subterranean amphipod crustacean of the genus Stygobromus (Crangonyctidae) from Alberta, Canada. Canadian J. Zool., 59 : 1827-1830. Calder D. R. & Bleakney J. S., 1965 Microarthropod Ecology of a Porcupine-Inhabited Cave in Nova Scotia. Ecology, 46 (6) : 895-899. Calder D. R. & Bleakney J. S., 1967 Observations on Frenchmans Cave, Nova Scotia, and Its Fauna. Bull. Nat. Spel. Soc., 29 (1) : 23-25. Chapman P., 1993 Caves and Cave Life, New Naturalist, Harper Collins, London, 219 p. Christiansen K. & Bellinger P., 1998 The Collembola of North America North of the Rio Grande: A Taxonomic Analysis. Grinnell College, Grinnell, Iowa, 1520 p. Christiansen K. & Bellinger P., 1980 The Collembola of North America North of the Rio Grande. Grinnell College, Grinnell, Iowa, 1322 p. DeHarveng L. & Bedos A., 2000 The Cave Fauna of Southeast Asia. Origins, Evolution and Ecology. In: Wilkens H., Culver D. C. & Humphreys W. F. (Eds.) Ecosystems of the World: Vol. 30, Subterranean Biota Elsevier, Amsterdam: 603-632. Emerton J. H., 1917 Spiders Collected in Nova Scotia and New Brunswick by Robt. Matheson in 1912. Proc. N. S. Ent. Soc., 1917: 95-96. Gibert J. & DeHarveng L., 2002 Subterranean ecosystems: a truncated functional biodiversity. BioScience, 52 : 473. Gnaspini P., 1992 Bat guano ecosystems. A new reference to Neotropical data. Mem. Biosplol., 19 : 135-138. Gnaspini P. & Trajano E., 2000 Guano Communities in Tropical Caves. In: Wilkens H., Culver D. C. & Humphreys W. F. (Eds.) Ecosystems of the World : Vol. 30, Subterranean Biota. Elsevier, Amsterdam: 251268 Nova Scotia mammal notes Can. Field-Nat., 50 (6) : 103-104. Graham R. E., 1962 Porcupine cave dens in California. 4 (1) : 1-4. Gunn J., Hardwick P. & Wood P.J., 2000 The Invertebrate Community of the Peak-Speedwell Cave System, Derbyshire, England Pressures and Considerations for Conservation Management. Aq. Conserv., 10 : 353-369. Holsinger J. R. 1980 Stygobromus canadensis, a new subterranean amphipod crustacean (Crangonyctidae) from Canada, with remarks on Wisconsin refugia. Can. J. Zool., 58(2) : 90-97. Howarth F., 1972 Cavernicoles in Lava Tubes on the Island of Hawaii Science, 175 (4019): 325-326. Howarth F., 1988 Environmental Ecology of North Queensland Caves. In: Pearson, L. (Ed.) Preprints of papers for the 17th biennial conference, Australian Speleological Federation Tropicon Conference, Lake Tinaroo, Far North Queensland, December 27-31, 1988. Jeannel R., 1926 Faune cavernicole de la France, avec une tude des conditions dexistence dans le domaine souterraine P. Lechevalier ed. Paris, 334 p. Jefferson G. T., 1981 Diptera in British Caves In: Beck, B.F. (Ed.) Proceedings of the 8th International Congress on Speleology, Bowling Green, Kentucky, 1 : 106-107. Juberthie C., 1984 La colonisation du milieu souterrain: International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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14 theories et methods, relations avec la speciation et lvolution souterraine. Mem. Biosplol., 11 : 65-102. Kane T. C. & Culver D. C., 1992 Biological Processes in Space and Time: Analysis of Adaptation. In: Camacho, A. I. (Ed.) The Natural History of Biospeleology Naturales: 423-451. King L.H., 1996 Late Wisconsin ice retreat from the Scotian Shelf Bull. Geol. Soc. Am., 108 : 1056. Lee C. E., 2002 Evolutionary genetics of invasive species Trends Ecol. Evol., 17 (8) : 386-391. Majka C. J., Moseley M. & Klimaszewski J., in press Gennadota canadensis (Casey) (Staphylinidae: Aleocharinae): new records, a range extension, and bionomic notes. Coleop. Bull. Marusik Y. M. & Koponen S., 1992 A Review of Meta (Araneae, Tetragnathidae) with Description of Two New Species. J. Arachnol., 20 : 137-143. McAlpine D. F., 1976 First record of the Eastern Pipistrelle in New Brunswick Can. Field-Nat., 90 : 476. McAlpine D. F., 1977 Notes on Cave Utilization by Beaver Bull. Nat. Speleo. Soc., 39 (3) : 90-91. McAlpine D. F., 1979 Preliminary Investigations on the Solution Caves of New Brunswick. J. New Brunswick Mus., 1979: 99-107. McAlpine D. F. & Reynolds J. W., 1977 Terrestrial Oligochaeta of Some New Brunswick Caves with Remarks on Their Ecology. Can. Field-Nat., 91 : 360-366. Moore K., 1963 Hayes Cave: a study S. Museum, Halifax, Canada, 21 p. Morris L. (Ed.), 1985 The Hayes Cave Site, South Maitland, Nova Scotia Cur. Rep. 50 N. S. Museum, Halifax, Canada, 128 p. Moseley M., 1975 The Protection of Caves in Nova Scotia Nova Scotia Speleo. Soc. Newsletter, 5 : 1-23. N. S. Museum, Halifax, Canada. Moseley M., 1976 Caves of the Atlantic Region. In: Thompson, P (Ed.) Cave Exploration in Canada Moseley M., 1996 The gypsum karst and caves of the Canadian Maritimes Cave and Karst Sci., 23 (1) : 5-16. Moseley M., 1998 Invertebrate Fauna of Nova Scotia Caves Cur. Rept. 86 N. S. Museum, Halifax, Canada, 37 p. Moseley M., in press Records of Bats (Chiroptera) at Caves and Mines in Nova Scotia Cur. Rept., N. S.museum, Halifax, Canada. Moseley M. & Hebda A., 2001 Overwintering Leiobunum elegans (Opiliones: Phalangiidae) in Caves and Mines in Nova Scotia Proc. N. S. Inst. Sci., 41 (4) : 216-218. Moseley M., Klimaszewski J. & Majka C. J., 2006 Description of the pupa and observations on the distribution, ecology and life history of Quedius spelaeus spelaeus Horn (Coleoptera: Staphylinidae) in Nova Scotia, Canada. Zootaxa, 1226 : 61-68. Peck S. B., 1988 A review of the cave fauna of Canada, and the composition and ecology of the invertebrate fauna of caves and mines in Ontario Can. J. Zool., 66 : 1197-1213. Sawatzky K., 1986 Diving in Diogenes. Can. Caver, 18 (1) : 16-20. Schmidt R. E., 1986 Zoogeography of the Northern Appalachians. In: Hocutt, C.H. & E.O. Wiley (Eds.) The Zoogeography of North American Freshwater Fishes John Wiley & Sons, NY: 137-159. Schwert D. P., 1979 a fossil earthworm (Oligochaeta: Lumbricidae) cocoon from postglacial sediments in southern Ontario Can. J. Zool., 57 (7) : 1402-1405. Scott F. 1979 Preliminary Investigations at Hayes Cave, Hants County, Nova Scotia in 1978. Cur. Rept. 38 N. S. Museum, Halifax, Canada, 14 p. Scott F. & Grantham R., 1985 The Cave Environment In: Morris, L (Ed.) The Hayes Cave Site, South Maitland, Nova Scotia Cur. Rept. 50 N. S. Museum, Halifax, Canada: 101-115. Scott F. & Hebda A. J., 2004 Annotated List of the Mammals of Nova Scotia. Proc. N. S. Inst. Sci., 42 (2) : 189-208. Shaw P. & Davis M., 2000 Invertebrates from Caves on Vancouver Island In: Darling, L. M. (Ed.) Proceedings of a Conference on the Biology and Management of Species and Habitats at Risk, Kamloops, B.C., 15-19 February, 1999, Vol. One B.C. Ministry of Environment the Cariboo, Kamloops, B.C: 121-124. Trevor-Deutsch B., 1973 The role of hibernating bats in the winter diet of Peromyscus spp. (Rodentia: Cricetidae). Ottawa, Canada. Wright B., 1979 Mites, Ticks, Fleas and Lice in the Nova Scotia Museum and Acadia University Museum Collections. Proc. N. S. Inst. Sci., 29 : 185-196. Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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15 Table 1. Summary of records of invertebrates from caves and mines in Nova Scotia and southern New Brunswick. [Abbreviations: th. = threshold, d.th. = deep threshold, d.z. = dark zone, Aq.= aquatic, Tr. = terrestrial, Am. = amphibian, PR = bat ectoparasite, Ne = Nearctic, Cp = cosmopolitan, Cl = circumpolar, Ad = non-indigenous Palaearctic introduction), AC = accidental (stray), HT = habitual trogloxene, TP = troglophile, GP = guanophile, (P) = parietal fauna. For key to alphanumeric site codes, see text]. [References: = Calder & Bleakney, 1965; = Calder & Bleakney, 1967; McAlpine & Reynolds, 1977; = Scott, 1979; = McAlpine, 1979; = Moseley & Hebda, 2001; ** = Moseley et al., 2006; *** = Wright, 1979; ^ = McAlpine, pers. comm.] continued CLASS or ORDER FAMILY TAXON Habitat collected Seasons recorded Sites Distribution th. d.th. d.z. main substrate(s) DJF MAM JJA SON & ecology Ciliata Fam. & spp. indet + + + pools, etc. Aq ? Turbellaria Planaridae Sp. A indet. + pools + + CB, PT, GM, KC Aq ? Sp. B indet. + pools + + KC Aq ? Oligochaeta Lumbricidae Dendrodrilus rubidus (Savigny) + + + various organic + + + Tr, Cp HT Aporrectodea tuberculata (Eisen) + + + various organic + + + Tr, Ad HT Lumbricus terrestris L. + + various organic + + CB, GM, GP Tr, Ad HT Eisenia rosea (Savigny) + + + cave soil + + PT,HO Tr, Ad HT Enchytraeidae Spp. indet + + + various organic + + + + All/most Tr GP Spp. indet. + + + pools, etc. + + + + All/most Aq TP Naiidae Spp. indet. + pools, etc. + + + + FC Aq TP Hirudina Glossiphonidae Helobdella papillata (Moore) + pool + BB Aq, Ne AC Cladocera Fam indet. Sp. A indet. + pool + CB Aq AC Sp. B indet. + pool + CB Aq AC Ostracoda Candonidae Pseudocandona albicans (Brady) + + stream, pool + + WB,CC Aq TP Fabaeformiscandona wegelini (Petkovsky) + pool + CC Aq, Cp TP Cypridae Cypria sp. + stream + WB Aq HT Cavernocypris sp. + stream + WB Aq TP Cypridopsidae Cypridopsis sp. + stream + WB Aq HT Copepoda Cyclopoidae Acanthocyclops robustus (Sars) + pool, sediment + + CB Aq, Cp TP A. brevispinosus (Herrick) + stream + WB Aq, Ne TP A. venustoides Coker + pool with dung + PT Aq, Ne AC Eucyclops agilis (Koch) + pool + CB Aq, Cp TP Diacyclops brachycerus (Kiefer) + stream + + WIC,WB Aq, Cp TP Paracyclops poppei (Rehberg) + pool + CB Aq, Cp AC Macrocyclops albidus (Jurine) + sediment + CB Aq, Cp TP Sp. indet. + sediment + WB Aq ? Isopoda Oniscidae Oniscus asellus (L.) + + + cave wall, dung + + + Tr, Ad tTP (P) Pauropoda Fam. indet. Sp. indet. no data FC Tr, ? International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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16 Chilopoda Henicopidae Lamyctes fulvicornis Meinert + porcupine dung + FC Tr, Ad AC Diplopoda Blaniulidae Proteroiulus fuscus (Am Stein) + + porcupine dung + + FC Tr, Ad HT Julidae Ophyiulus pilosus (Newport) + + porcupine dung + + FC Tr, Ad HT Allajulus latestriatus (Curtis) + + + various + + FC,F2 Tr, Ad HT Polydesmidae Polydesmus angustus Latzel + + porcupine dung + + FC Tr, Ad HT (P) Symphyla Symphylella sp. + cave soil + MB Tr TP Scutigerellida Scutigerella sp. + on pool + KC Tr TP Collembola Poduridae Hypogastrura pseudarmata + timber + KC Tr, Ne TP Neanura muscorum (Tempelton) + timber + KC Tr, Ad TP Friesea sp. + porcupine dung + GM Tr, ? AC Willemia scandinavia Stach + + + WIC Tr, Cp HT porcupine Onychiuridae Protaphorura pseudarmatus + + + + + HC,WIC,FC Tr, Ne TP porcupine P. armata (Tullberg) + + + porcupine dung + + + + HC,FC, F2,GM Tr, Cp TP P. cf. boedvarssoni Pomorski + on pools + + CC Tr, ? ? Tullbergia iowensis Mills + + porcupine dung + + + + FC Tr, Ne TP T. roseki Christiansen and Bellinger + porcupine dung + HC Tr, Ne HT Isotomidae (L.) + + porcupine dung + + + + FC Tr, Ad TP F. stella Christiansen & Tucker + + + porcupine dung + + HC,WIC,FC Tr, Ne TP F. candida (Willem) + + dung, pool surface + + + MIC, wells Tr, Ne TP Isotoma notabilis Schffer no data FC Tr, Ne HT I. caeruleatra Guthrie + plant litter + CB Tr, Ne AC Isotoma sp. nova? + + various organic + + + HC,FC Tr TP Entomobryidae Heteromurus nitidus (Templeton) + + + oligotrophic + + + Tr, Ad TP Pseudosinella alba (Packard) + + + on pools, dung + HC,FC Tr, Ne TP P. collina Wray + timber + KC Tr, Ne TP Entomobrya nivalis (L.) + no data FC Tr, Cp AC Tomoceros minor (Lubbock) + porcupine dung + CB Tr, Ad TP Neelidae Megalothorax minimus Willem + + porcupine dung + + + + FC Tr, Ne GP Sminthuridae Arrhopalites nr. pygmaeus (Wankel) + oligotrophic + F2 Tr HT A. hirtus Christiansen + + oligotrophic & on pools + F2 Tr, Ne TP Sminthurides malmgreni (Tullberg) + wet plant litter + F2 Tr, Cp AC Ptenothrix marmorata (Packard) + + + timber; on pool + + CB, FC, KC Tr, Ne HT continued Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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17 Ephemeroptera Fam. et sp. indet (nymphs) + stream + HB^ Aq. ? Odonata Aeshnidae Aeshna umbrosa Walker nymphs + pools + HC Aq, Ne HT Aeshna sp. indet. nymphs + + pools + HC Aq, Ne HT Cordulegaster maculata Selys + stream + CB Aq, Ne AC Macromidae Macromia illinoiensis Walsh nymphs + + pools + HC Aq, Ne HT Plecoptera Capnidae Sp. indet. nymphs + stream + WB Aq AC Leuctridae Sp. indet. adults + + + cave walls + WB, KC Aq HT Nemouridae Amphinemura sp. nymphs + stream + WB Aq AC Taeniopteryx sp. nymphs + + seeps & wet plant litter + FC, F2 Aq AC Chloroperlidae Haploperla sp. nymphs + + + stream + + F2 Aq HT Perlidae Sp. indet. nymphs + stream + KC Aq HT Orthoptera Ceuthophilus brevipes (Scudder) + + cave wall + + + TH,PT Tr, Ne, HT(P) C. maculatus (Harris) + bait (molasses) + WIC Tr, Ne, AC Coleoptera Dytiscidae Agabus semivittatus (Le Conte) + stream CB Aq, Ne AC A. larson i Ferry & Nilsson + + + stream, pool + HC,DC Aq, Ne HT Dytiscus sp. indet. larva + stream + CB Aq, ? AC Hydrophilidae Crenitis digesta (LeConte). + cave wall + F2 Tr, Ad AC Staphylinidae Quedius s. spelaeus Horn larvae and + + + porcupine dung + + + + Tr, Ne TP Q. mesomelinus (Marsham) + plant litter + KC Tr, Ad TP Brathinus nitidus LeConte + various organic + FC, FH2 Tr, Ad TP Gennadota canadensis Casey + bait (molasses) + WIC Tr, Ne TP Atheta sp. indet. + cave wall + PT Tr, Ne ? Curculionidae Sciaphus asperatus (Bonsdoff) + bait (molasses) + F2 Tr, Ad AC Dermestidae Dermestus lardarius L. + cave wall + F2 Tr, Cp AC Scarabaeidae Aphodius leopardus Horn + porcupine dung + FC Tr, Ne GP Latridae Corticaria serrata (Paykull) + porcupine dung + MIC Tr, Ad GP Ptilidae Acrotrichis castanea (Matthews) + + porcupine dung + + + + F2 Tr, Ne GP Trichoptera Limnephilidae Sp. indet. adult + cave wall + DC, HB^ Tr, ? HT(P) Lepidoptera Acrophilidae Amydria effrentella Clem + cave wall + PT Tr, Ne AC Geometridae Triphosa haesitata (Walker) cave wall Tr, Ne HT(P) Xanthotype sospeta (Drury) + cave wall + FC Tr, Ne AC Noctuidae Scoliopteryx libatrix L. + + cave wall + + Tr, Cl HT(P) GP,GR,HO continued International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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18 Diptera: Nematocera Tipulidae Limonia cinctipes Say adult + cave wall + WB Tr, Cl TP(P) ditto -larva + + porcupine dung + + WIC, FC, F2 Dolichopeza sp. sensu lato + cave wall + KC Tr, ? HT(P) Tipula sp. larva + veg. litter + F2 Tr, ? AC Erioptera pilipes (Fabricius) larva + veg. litter + CB Tr, Ne AC Trichoceridae Trichocera sp. indet + porcupine dung + WIC Tr, ? TP(P) T. maculipennis Meigen adult + + + cave wall + + + most sites Tr, Cp TP(P) ditto -larva + + porcupine dung + + most sites Culicidae Culex pipiens + + + cave wall + + CC, MC Tr, Cp HT(P) C. restuans Theobald cave wall + + Tr, Ne HT(P) C. territans cave wall + + Tr, Cp HT(P) Anopheles + cave wall + WB Tr HT(P) Chaoboridae Chaoborus sp. + + various + + F2, WB Tr HT(P) Chironomidae Smittia sp. adult, larva + + porcupine dung + FC, F2 Tr TP Gen. et spp. indet adults + + various organic + FC, F2 Tr ? Gen. et spp. indet -larvae + + stream + + F2, WB, KC Aq ? Mycetophilidae Boletina sp. + + cave wall FC Tr HT(P) Bolitophilia sp. + + + cave wall + FC, KC Tr HT(P) Rymosia sp. + + + cave wall + GM Tr HT(P) Exechia sp. + + + cave wall + + HC, CC, PT, MC Tr HT(P) Exechiopsis sp. + + cave wall + + CC, PT Tr HT(P) Tarnania tarnania (Dziedzieki) + + cave wall + PT Tr, Cl HT(P) Sciaridae Bradysia sp. adult + + porcupine dung + FC Tr GP Lycoriella sp. adult + + porcupine dung + CB, FC Tr GP Scatopsciara sp. adult + + porcupine dung + CB, FC Tr GP + + + dung, walls, litter + + most sites (P) see Larva type A + + + porcupine dung + + + Larva type B + + porcupine dung + + HC, WIC, CC Larva type C + porcupine dung + F2 Cecidomyidae Peromyia sp. + cave wall + KC Tr HT(P) Gen. et sp. indet. + cave wall + YRT Tr ? Simulidae Simulium sp. larva + stream gravel + F2 Aq HT Psychodidae Psychoda sp. adult + + dung, wall + + F2 Tr HT(P) -ditto --larva + veg. litter + F2 Max Moseley continued International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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19 Diptera: Brachycera Tabanidae Chrysops sp. larva + veg. litter + CB Tr AC Phoridae Megaselia meconicera (Specier) no data FC Tr, ?? TP Helomyzidae Scoliocentra fraterna Loew no data FC Tr, Ne HT(P) Helomyza serrata (L.) + + cave wall + + FC, CC, PT Tr, Cl HT(P) H. brachypterna (Loew) ? + cave wall + PT Tr, Ne HT(P) Amoebaleria sp. + + cave wall + + CC, PT Tr HT(P) Tephrachlamys sp. + cave wall + F2 Tr HT(P) Leptocera (Limosina) sp. + + + walls and dung + + FC, F2 Tr, GP(P) Siphonaptera Ischnopsyllidae Myodopsylla insignis (Rothschild) ex Myotis*** Tr PR Hymenoptera Fam. indet* no data Tr ? Psocoptera Liposcelidae Liposcelis bostrychophila + on pool + CC Tr AC Homoptera Aphidae Trama sp. + pitfall + CC Tr AC Cixidae Gen. et sp. indet. + + FH2 Tr AC Acarina Parasitidae Parasitus sp(p). + dung, timber + + FC, MC, KC Tr GP Eugamasus sp. + + dung, timber + FC, KC Tr TP Vulgarogasmus sp. nr. + porcupine dung + Tr GP Gen. et. spp. indet. + + + porcupine dung + + + most sites Tr GP Vegaidae Vegaia sp. + + dung, plant litter + FC, KC Tr TP Zerconidae Zerconopsis sp. + porcupine dung FC Tr GP Ascidae Arctoseius sp. + + porcupine dung + + FC, KC Tr GP Ameroseidae Epicriopsis sp. + porcupine dung + + CC Tr GP Eviphididae Alliphis sp. + + porcupine dung + + FC Tr GP Macrochelidae Geolapsis sp. + porcupine dung + FC Tr GP Spinturnicidae Spinturnix sp. ex Myotis*** Tr PR Macronyssidae Macronyssus crosbyi (Ewing & Stover) ex Myotis*** Tr PR Uropodidae Gen. et sp. indet. + porcupine dung + + CB, WIC, FC Tr GP Eupodidae Linopodes motatorius (L.) + timber + KC Tr, Cp TP Cocceupodes sp. + timber + MC Tr TP Rhagididae Rhagidia sp(p). + + + dung, timber + + + most sites Tr GP Ereynetidae Gen. et sp. indet. + porcupine dung + CC Tr GP Pygmephoridae Pygmephorus sp. + porcupine dung + FC Tr GP Bakerdania sp. + porcupine dung + FC Tr GP continued International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada

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20 Acarina Tetranychidae Bryobia praetiosa (Koch) + porcupine dung + FC Tr AC Gen. et. sp. indet. + + porcupine dung, plant + HC, WB Tr GP Hydrachnoidea Gen et sp. indet. + stream + KC Aq ? Glycyphagidae Glycyphagus domesticus (De Geer) + timber + MC Tr, Cp TP Histiostmatidae Gen et sp. indet. + + porcupine dung + WIC, FC Tr GP Acaridae Acarus immobilis + porcupine dung + FC Tr AC Banksinomidae Oribella sp. + porcupine dung + CC Tr GP Araneida Nesticidae Nesticus cellulanus (Clerck) + cave wall + + FC Tr, Ad TP(P) Tetragnathidae Meta ovalis (Gertsch) + + + cave wall + + + + most sites Tr, Ne TP(P) Dictynidae Circurina brevis (Emerton) + scree + WIC Tr, Ne HT Linyphiidae Sisicottus montanus (Emerton) + porcupine dung + FC Tr, Ne AC Grammonota sp. + porcupine dung + CB Tr, Ne AC Opiliones Phalangidae Oligolophus tridens (C. L. Koch) scree + WIC, CC* Tr, Ad HT Sclersomatidae Nelima elegans (Weed) + + cave wall + + + most sites* Tr, Ne HT(P) Gastropoda Arionidae Arion subfuscus (Draparnaud) + cave wall + KC Tr, Ad HT(P) A. circumscriptus group + cave wall + + FC, PT, DL, KC Tr, Ad HT(P) Limacidae Deroceras laeve (Mller) + + cave wall + F2 Tr, Cl HT(P) Helicidae Trichia hispida (L.) + cave wall + F2 Tr, Ad HT(P) Zonitidae Zonitoides arboreus (Say) + + + cave wall + + FC, F2 Tr, Ne HT(P) Endodontidae Discus catskillensis Pilsbry + timber + KC Tr, Ne TP Max Moseley International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007

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21 CLASS or ORDER FAMILY TAXON Habitat collected Seasons recorded Sites Distribution & ecology th. d.th. d.z. substrate DJF MAM JJA SON Osteichthyes Salmonidae Salvelinus fontinalis (Mitchell) + + + limestone streams + + + + DC, KC Aq,Ne HT Cyprinidae Phoxinus eos (Cope) + pools HC Aq,Ne AC Gasterosteidae Pungitus pungitus (L.) + + pools + + BB, HC, FC Aq, Cl HT Amphibia Plethodontidae Plethodon cinereus (Green) + stream + MCI Am,Ne AC Ranidae Rana clamitans melanota + + + pools, streams CB, HC, FC Am,Ne AC Mammalia Soricidae Sorex fumeus (Miller) + HO Tr, Ne HT Vespertilionidae Myotis lucifugus (LeConte) + + + + BB,CB,HC,WIC,FC,F2, MIC,MC,PT,LCM,OV, LSH,HO, HB, BC, Tr, Ne HT M. septentrionalis (Trouessart) + + + + BB,CB,HC,FC,MIC,CC,LCM, Tr, Ne HT GRM,OV,LSH,GM,GR,HO, HB,BC, (F. Cuvier) + + + + + CB,HC,FC,MIC,CC, GRM,GM, Tr, Ne HT GR,KC,UL,TC Eptesicus fuscus (Palisot de Beauvois) + + HC (sight record) Tr, Ne HT Lasionycteris noctivagans (Le Conte) + + PT (sight record) Tr, Ne AC Procyonidae Procyon lotor (L.) + + (tracks) CB, HC, FC Tr, Ne HT Mustelidae Mustela vison Schreber + (scat) KC Tr, Ne AC Castoridae Castor canadensis Kuhl + DC*, KC Tr, Ne HT Muridae Peromyscus maniculatus (Wagner) + HC, GM Tr, Ne HT Erethizontidae Erethizon dorsatum (L.) + + + + + + + most: except Cape Breton Tr, Ne HT Table 2: Summary of records of vertebrates from caves and mines in Nova Scotia and southern New Brunswick. [Abbreviations: th. = threshold, d.th. = deep threshold, d.z. = dark zone, Aq.= aquatic, Tr. = terrestrial, Am. = amphibian, Ne = Nearctic, Cl = circumpolar, AC = accidental (stray), HT = habitual trogloxene, TP = troglophile, GP = guanophile, (P) = parietal fauna. For key to alphanumeric site codes, see text]. [References: = Calder & Bleakney, 1965; = Scott, 1979; = McAlpine, 1979; Sawatzky, 1986. See McAlpine (1979) and Moseley (in press) for sources for records of bats.] Taxonomic group New Brunswick (McAlpine 1979) Ontario (Peck 1988) NS + NB (this study) Collembola* 7 (2%) 25 (15%) Coleoptera 15 (5%) 14 (8%) Diptera 149 (50%) 33 (19%) Other insects 33 (11%) 23 (14%) TOTAL INSECTA 54% 204 (68%) 95 (56%) Acari 8 (3%) 27 (16%) Aranea 31 (10%) 5 (3%) Opiliones 7 (2%) 2 (1%) TOTAL ARACHNIDA 22% 46 (15%) 34 (20%) MOLLUSCA 4% 21 (7%) 6 (4%) OTHER INVERTEBRATA 20% 30 (10%) 35 (21%) TOTAL INVERTEBRATES 100% 301 (100%) 170** (100%) Table 3: Invertebrate faunal composition of caves in eastern Canada: statistical breakdown by number of species and percentages. [* Collembola is included in the Insecta in order to permit comparison with McAlpine (1979). ** Accidentals are included to permit comparison of all three studies.] International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 Acadian biospeleology: composition and ecology of cave fauna of Nova Scotia and southern New Brunswick,Canada



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Available online at www.ijs.speleo.it International Journal of Speleology International Journal of Speleology 36 (1) 23-30 Bologna (Italy) January 2007 Sails: a new gypsum speleothem from Naica, Chihuahua, Mexico Tullio Bernabei 1 Paolo Forti 2 Roberto Villasuso 3 INTRODUCTION th Bernabei T., Forti P. 2007. Sails: a new gypsum speleothem from Naica, Chihuahua, Mexico. International Journal of Speleology, 36 (1), 23-30 Bologna (Italy). ISSN 0392-6672. The caves of Naica (Chihuahua, Mexico) are perhaps the most famous mine caves of the world due to the presence of gigantic gypsum crystals. Nevertheless, very little research has been carried out on this karst area until now. A multidisciplinary investigation aspects of these caves. This paper describes a completely new type of gypsum speleothem: the sails, observed only inside the Cueva de las Velas, one of the caves of the Naica system. This speleothem consists of extremely thin, elongated skeleton crystals that have grown epitaxially only on the tips of the gypsum crystals pointing upward. The genesis of sails is strictly related to the environmental conditions set speleothems will disappear entirely and therefore this study is fundamental to preserve at least the memory of them. Keywords: gypsum speleothems, speleogenesis, mine caves, Naica, Mexico Abstract: Received 15 September 2006; Revised 6 November 2006; Accepted 7 November 2006 velas GEOLOGICAL SETTING Available online at www.ijs.speleo.it International Journal of Speleology 1 2 3

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Fig. 1. Sketch of the mine in which the main natural cavities are related to the original groundwater level. Fig. 3. A sail growing epitaxially over a pre-existing large gypsum crystal with its apex pointing upward (photo by Tullio Bernabei, Archivio La Venta & S/F). Tullio Bernabei Paolo Forti Roberto Villasuso International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007

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Fig. 2. Cueva de los Cristales: the largest gypsum crystals of Naica (photo by Giovanni Badino, Archivio La Venta & S/F). Fig. 4. Plan and vertical sections of the Cueva de las Velas: the grey area is where the sails have developed. International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007 Sails: a new gypsum speleothem from Naica, Chihuahua, Mexico

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Fig. 5. A portion of the cave wall in which a thick deposit of metallic oxides-hydroxides are exposed (photo by Tullio Bernabei, Archivio La Venta & S/F). Fig. 6 A sail developing over a re-dissolved gypsum crystal with the characteristic stalk shape (photo by Tullio Bernabei, Archivio La Venta & S/F). Fig. 7. Main genetic steps for sails: Athe growth of macrocrystals continued as long as the crystals were completely submerged; Bwhen the CSupersaturation at the crystal apex induced the deposition of gypsum and a sail developed, with its characteristic shape being controlled by strong upward air; Dthe process ended when the distance between the crystal apex and the water surface exceeded the capillary fringe. Tullio Bernabei Paolo Forti Roberto Villasuso International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007

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important of them are the Gibraltar, about 1 m 3 springing in the mine galleries has a EXPERIMENTAL OBSERVATIONS Fig. 8. The evolution of a sail over a partially re-dissolved gypsum crystal: Ainitial stage: condensation over the macrocrystals induced capillary uplifting and evaporation from the crystal was the consequence of capillary uplifting of unsaturated water. Sails: a new gypsum speleothem from Naica, Chihuahua, Mexico International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007

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DISCUSSION Tullio Bernabei Paolo Forti Roberto Villasuso International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007

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2+ 4 = Finally it is reasonable to suppose that a rather FINAL REMARKS ACKNOWLEDGEMENTS REFERENCES Condensation as a microclimate process: Measurements, numerical simulation and prediction in Glowworm Cave, New Zealand 23(5) Sails: a new gypsum speleothem from Naica, Chihuahua, Mexico International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007

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Le grotte di miniera tra economia mineraria ed economia turistica 17 Les grottes a cristaux de gypse de Naica 32 The problem of condensation in karst studies. 60(1) Compositionally distinct, saline hydrothermal solutions, Naica Mine, Chihuahua, Mexico 74 The Pulp gigantic geode (Almeria, Spain): geology, metal pollution, microclimatology and conservation 50 Microclimate processes characterization of the giant geode of Pulp (Almeria, Spain): technical criteria for conservation. International 26 miniera. serie II, 17 Le grotte di Naica e i loro giganteschi cristalli di gesso Mineralogy of Cueva de las Velas (Naica, Chihuahua, Mexico) The selenite caves of Naica, Mexico 12 Cave minerals of the World 31(3) A geological evaluation of the Naica deposit, Chihuahua, Mexico. Internal Report of New Cave of the Crystals at Naica, Chihuahua, Mexico Hightemperature, carbonate-hosted Pb-Zn-Ag (Cu) deposits of northern Mexico 83 The largest crystals. 66 76(5) Ore genesis in the Naica District, Chihuahua, Mexico 54 18 International Journal of Speleology, 36 (1), 23-30. Bologna (Italy). January 2007 Tullio Bernabei Paolo Forti Roberto Villasuso



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International Journal of Speleology 36 (1) 31-38 Bologna (Italy) January 2007 Greg Hodgins 1 George A. Brook 2 and Eugene Marais 3 INTRODUCTION Caves are natural traps and as such are important repositories for both animal and human remains. Hominid remains dating to >4 Ma have been recovered from breccias in South African caves such as Sterkfontein, Swartkrans, and Kromdraai (e.g. Brain, 1981). Uncemented remains of animals are commonly found in the caves of arid lands. For example, bones of a Giant Ground Sloth ( Glossotherium harlani ), that may have weighed 900 kg, were recovered from a chamber ca. 64 m underground in Grand Canyon Caverns, in western Arizona. The Giant Sloth became extinct between 20 and 11 ka. In the same cave a presumed to be about 100 years old at that time ( www. desertusa.co m ). encountered in arid land caves due to localized dry microclimates that prevent decay. Commonly, the animal falls into the cave or enters via steep or slippery access to scavenging animals, thus preserving the carcass. In arid areas, animals trapped in caves can Keywords: Abstract: Received 20 June 2006; Revised 17 October 2006; Accepted 16 November 2006 due to the exceptionally dry conditions. For example, Tasmanian tiger ( Thylacinus cynocephalus ) was discovered in Thylacine Hole in the Nullarbor Plain of Western Australia (Lowry & Lowry, 1967; Lowry & Merrliees, 1969). The mummy was radiocarbon dated to 4600-4700 14 C yr BP. in caves in Namibia, including a porcupine ( Hystrix africanus ) in Arnhem Cave (Marais, unpublished), and hyrax ( Procavia capensis ) and klipspringer ( Oreotragus oreotragus ) in Gauab Cave (Marais et al., 1996). In Sesfontein (Martini et al., 1991; Martini & Marais, ever been dated. baboon, subspecies Papio cynocephalus ursinus was discovered on a ledge in Ludwig Cave in Namibia (Figs. 1 and 2A) by Mr. J. Loots, the owner of the cattle ranch on which the cave is situated (pers. comm. during the wet season, explaining their advanced state of decomposition. The microclimate of the ledge, rather than decomposition of the female baboon. As the mummy had three upper and three lower molars, Available online at www.ijs.speleo.it International Journal of Speleology 1 ghodgins@physics.arizona.edu) 2 3Entomology Centre, National Museum of Namibia, P.O. Box

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32 and as the M 3 upper molars do not appear in female baboons until around 7 years of age (Smith et al., 1994), the animal had reached adulthood before a male juvenile, was discovered on a ledge near the adult female; it was not there in 1998. As none of has ever been dated, we did not know if the adult female baboon mummy was ancient or recent. This paper outlines our attempt to learn more about this LUDWIG CAVE Ludwig Cave is close to the border of the Etosha National Park, 40 km from Otavi, and 120 km from Grootfontein (Fig. 1). Average rainfall at Grootfontein coming in the six austral summer months November to April. The mean annual temperature is 20.2C (1968-1989), with mean monthly values ranging from local residents were searching for a burglar named Ludwig who according to local legend lived in the developed in slightly metamorphosed blue, Maieberg Formation limestone of the Tsumeb Subgroup. The entrance to the cave is a vertical solution pit 9 x 4 m and 9 m deep located near the top of a limestone ridge. Mamba Crawl, a low, wide passage, leads into the cave from the bottom of the entrance shaft but rapidly decreases in height to become impassable before connecting to the rest of the cave (Fig. 1). Access to the main system is through a larger smooth by generations of baboons roosting in a small chamber behind it. A tunnel-like passage from this chamber continues for about 30 m and then slopes into a passage running east to west. To the east this which contained three decomposing baboon corpses curving north to join a larger northeast to southwest oriented passage. At the western end of the eastwide and 20 cm high, leads to Mamba Crawl. The female baboon mummy was found in the northeasthumidity were recorded at various sites throughout the cave. Temperature averaged 24.2 C (n = 10, = = 7.3) at mummy. Greg Hodgins, George A. Brook and Eugene Marais International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007

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33 International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007 tail root.

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34 Greg Hodgins, George A. Brook and Eugene Marais International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007 RADIOCARBON DATING To determine the age of the baboon mummy, bone surface of the metatarsal was cleaned by abrasion and bone powder milled from the dense cortical tissue (Fig. 2B). Collagen was extracted from the bone powder using methods described by Bronk Ramsey et mass, indicated that the bone was reasonably well a sealed, evacuated ampoule in the presence of CuO. The resulting carbon dioxide was then cryogenically ratio mass spectrometer. The 13 C was found to be fractionation correction calculation. The remaining carbon dioxide was catalytically al. (1984). Graphite 14 13 C ratios were measured at the Center for Applied Isotope Studies, University of Georgia. The sample ratios were compared to the ratio measured from Oxalic Acid I (National Bureau of Standards, Standard Reference Material #4990). The both statistical and experimental errors (Table 2). By convention, the radiocarbon content of the Since radioactive decay reduces the 14 C content of organic remains over time, the age of organic materials The 14 C content of the baboon bone collagen was found a different location and the measurements repeated. measurement. These super modern values indicate the presence of 14 C from atomic bomb testing, and Radiocarbon measurements were then carried out on tendon and skin samples in an attempt to more precisely identify when the animal lived. A sample of tendon was initially given the standard extraction of collagen from bone. The recovery was unacceptably low suggesting that the tendon collagen was in a fragmentary state. A second tendon sample hour at room temperature. The swelled tendon was then washed in deionized water until the pH of the wash solution increased to 4.0. The tissue was then lyophilized, combusted, and graphitized for AMS measurement in the manner described for bone collagen. A sample of skin was also prepared using the same acid-only treatment. This minimal treatment would remove bulk contaminants, carbonate salts, and a fraction of the weakly bound contaminants. Omitting the high pH treatment seemed reasonable soluble, soil-derived humic acids seemed unlikely. AMS of graphitized tendon and skin samples carbon (Table 2). Comparison of the skin value with Southern Hemisphere atmospheric 14 C values shown in Figure 3 indicates that atmospheric levels passed through these values in September 1963 and between occurred after this, and a discussion of the lag time is presented below. DISCUSSION Above ground nuclear testing raised atmospheric 14 natural levels. The incorporation of bomb carbon Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Year Rain (mm)* 122 1 0 1 3 16 81 Temp ( C) 23.6 22.6 21.7 16.6 13.7 13.6 16.8 21.2 23.8 20.2 Material Laboratory ID Fractionation-corrected pMC 13 C ( PDB) Bone collagen UGAMS-0022 Bone collagen UGAMS-0022B Tendon UGAMS-0112 Skin UGAMS-132

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indicates that the animal lived during this period (Fig. 3). This brief pulse of 14 C into the atmosphere has been referred to as the bomb-spike and it has been used in 14 C dating for authentication studies and forensic science. Forensic studies have been aimed at establishing either the date of death, or the age at time of death, of 14 C from the atmosphere into tissues. The factors that affect this are the nature of the food chains incorporating 14 C into diet, the metabolic routing of dietary carbon between energy production and tissue carbon within different tissues. Sex, genetics, behavior, disease and other factors are likely to modulate tissue 14 C content. Broecker et al . (1964), and Stenhouse & Baxter (1977) all studied the uptake of bomb 14 C in various human tissues. The estimates attempted to account for the omnivorous human diet, and included assessments of the lag time between food growth and consumption for various categories of foodstuffs. In their estimates, dietary inputs from fruits, vegetables, and dairy products were most tightly linked to atmospheric 14 C levels whereas 14 C levels lagging well behind the contemporary atmosphere. They all estimated about a one-year lag for 14 C levels in soft 14 C content of soft tissues provides the most reliable data for establishing the date of death. This approach is well illustrated by Wild et al (2000). They measured the bomb14 C content of human hair, bone lipid, and bone collagen and from individuals of known age and date of death. They determined that the 14 C content of hair (a continuously synthesized, non-turning over tissue) lagged approximately one year behind the atmospheric levels, whereas lipid 14 C content lagged six years and bone collagen 14 C between twenty and thirty years behind atmospheric levels. Clearly collagen is a less than ideal material for bomb-spike determinations of date of death. However its longer turnover time suggests it might be useful for establishing age at time of death, although a detailed Studies have shown that collagen residence time in human bone is variable. Much of this variability has development, stable at about 20 years in adult life (Stenhouse & Baxter 1977) but possibly extending to greater than 30 years in elderly individuals (Stenhouse & Baxter 1977, Wild et al 2000). Jull et al presented data showing residence time could vary within adults by a factor of two. Although the reason for this was unknown, differences in nutritional history were suspected. This variability confounded attempts to determine age at time of death based upon collagen 14 C content. International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007

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36 of the year of death for the baboon as 1977. In spite of the fact that our understanding of the rate of 14 C into the baboons tissue is limited, encompassing by Wild et al seems reasonable. Our measurements were on baboon skin rather than hair, and 14 C in skin may have a longer residence time. However, we feel this potential increase in residence time is offset by the fact that the baboon is frugivorous (fruit-eating). Thus, its dietary 14 C intake is more tightly linked to atmospheric 14 C levels than the average human diet. We also carried the study further and attempted to estimate the year of birth, and hence age at time of death. Observation indicated that the baboon was of adult size. Female baboons reach full size at about baboons reach full size at 7-8 years and life expectancy is 22 years (Susan Alberts, pers. comm. 2001). If the of bomb 14 C during early development would result in bone collagen 14 C levels being higher than tendon and skin. The fact that the bone 14 C levels are lower than soft tissue points toward the baboon developing to adulthood before or during the early part of the bomb spike, and slowly incorporating bomb 14 C through steady-state turnover. This reasoning places the birth bomb spike curve, some 6 years prior to 1963 (6 years to adulthood) when atmospheric 14 C was the same as that of the baboons bone collagen at the time of death in 1977. A rough age for the baboon at death is therefore 20 years, which suggests that the residence time of baboon bone collagen is similar to that of humans. Modeling collagen radiocarbon uptake using estimates of collagen turnover rates generated a more sophisticated estimate. Although collagen turnover in that allometric scaling from human to baboon might be used to approximate a residence time for baboon bone collagen. If steady-state collagen turnover is linked to basal metabolic rate then a scaling factor, such as Kleibers Law, which correlates body mass with basal metabolic rate, or Hofmans correlation between longevity, brain mass and basal metabolic rate, might be used to estimate collagen turnover time (see discussions in Austad & Fischer 1992). Most formulae imply that turnover rates in baboons should Jull et al 14 C uptake into human bones using box modeling. We took a similar approach. Our model is based on three important assumptions. First, bone collagen 14 C levels are determined directly from food, and that herbivore food 14 C levels lag one year behind atmospheric levels (see Broecker et al , 1964; and Stenhouse & Baxter, 1977). Second, during skeletal growth (the linear increase in body mass to adulthood) and newly formed bone 14 C levels track food 14 C levels. Third, after adult size is attained, bone collagen turns over at turnover rate is based upon turnover rates in human collagen determined by Libby et al (1964), Stenhouse & Baxter (1977), and Wild et al. (2000). The 14 C content of the replacement bone is assumed to follow atmospheric 14 C levels with a one-year lag as explained above. Jull et als model also incorporates a slowfrom our model. In modeling, the accumulation of 14 C in baboon skeletal collagen was calculated from the atmospheric 14 C values plotted in Figure 3 using the scenarios were examined by assuming different birth years, modeling the accumulation of 14 C in bone over time, and selecting the birth year that most closely predicted bone collagen 14 predicts the observed bone 14 C level at the time of death (1977). From this an age at time of death was calculated to be 19 years. When we visited the cave in 2002 there was a second the mummy of the adult female. There were pupal cases of of the new mummy, which was not there when we visited the cave in 1998. As the animal had been in rapid in this semiarid cave despite relatively high cave humidity. into Ludwig Cave. One possibility is that unresponsive females, immature males, or low status males, may have tried to escape the hassling of higher status males or females by running deeper into the cave (there is a lot of noise and general mayhem before baboons settle down at night). Another possibility is that leopards, their main predator, attempted to prey on the troop at night. If attacked by leopards during the day baboons will usually defend themselves and have been known to kill their attackers. However, at night they do not defend themselves but rather run off in all directionssome possibly deeper into the cave. Leopards are mainly nocturnal hunters and relatively common in the area around Ludwig Cave so this explanation is extremely plausible. the ca. 6 m slope. We believe that under wet conditions to climb out. They may have been attracted to the passage and a small amount of light during the day, cm wide) to pass. Being lost in the dark, the animals probably ended up sitting on ledges eventually dying of dehydration or famine. Greg Hodgins, George A. Brook and Eugene Marais International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007

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37 collapse entrances to lava tube caves and their bones are numerous in some caves, possibly carried there by leopards (Glover et al., 1964). Bones of baboons, many of them extinct, are common in breccias of caves such as Sterkfontein, Swartkrans, and Kromdraai in South Botswana (Callum Ross, pers. comm. 2000). For example, Brain (1981) reports 181 baboons among 331 animals represented in Sterkfontein Member 4, Australopithecus africanus In Swartkrans Member 1, 89 of 312 animals are baboons, 3 Homo Australopithecus robustus while in Kromdraai B, 32 of 86 animals are baboons and six are Australopithecus robustus One possibility for the preponderance of baboons in these cave breccias is that these South African caves, like Ludwig Cave in Namibia, could be entered easily many caves in the dolomites of Southern Africa, including several with Australopithecus and baboon remains, have or used to have vertical entrances with steep to overhanging walls. This very common cave morphology results from solutional development at or near the groundwater table, followed by roof collapse and opening of the cave to the surface (e.g. it is possible that a large number of baboons were chased into these caves where they became trapped and died from dehydration and starvation. Some of these animals were probably injured by the fall into the vertical cave entrance; perhaps others could not climb out because of the steepness or overhanging nature of the entrance walls. CONCLUSIONS corpses in Ludwig Cave, Namibia suggest that baboons ran into the cave and became trapped because of harassment by other baboons or because a leopard attempted to prey on the troop at night. We believe that this may have occurred during the wet season and that once the animals slid down the result of dehydration or famine. As the male juvenile semiarid climate despite relatively high humidity in the cave. pMC values of greater than 100 for bone, tendon and skin from the adult female mummy suggest death in late 1977 according to the atomic bomb spike calibration curve of Manning et al. (1990) and Manning & Melhuish (1994). The animal was likely around 19 years old at the time of death, based upon the relative 14 C content of bone collagen and soft tissue collagen. The young age of the baboon mummy caves may also be of relatively recent age. Our studies indicate that where there are no other indications of an animals age, pMC measurements of skin plus that of bone collagen, together with knowledge of the animals growth characteristics, can provide an estimate of age at the time of death. The bomb-spike age calibration Ludwig Cave can be used to date the remains of any years. ACKNOWLEDGEMENTS This research was supported by NSF grant BCSMuhle and Mr Jan Loots for permission to visit the REFERENCES Growth rates in a maternal effects Behavioral Ecology and Sociobiology, 57 Austad, S.N. & Fischer K.E., 1992 Primate longevity: its place in the mammalian scheme American Journal of Primatology, 28 Deposits Transvaal Museum Memoirs No. 11. Brain C.K., 1981 The Hunters or the Hunted? An Introduction to African Cave Taphonomy. University of Chicago Press, Chicago. Transvaal karst: some considerations of development and morphology The South African Geographical Journal, 47 Bomb carbon-14 in human beings Science, 130 Bronk Ramsey, C., Pettitt P.B., Hedges R.E.M., Hodgins Radiocarbon dates from the Oxford AMS system: Archaeometry Datelist 30. Archaeometry, 42(2) 1964 The lava caves of Mount Suswa, Kenya Studies in Speleology, 1 mb-derived 14 th Annual Meeting, J.F., 1964 Replacement rates for human tissue from atmospheric radiocarbon Science, 146 Discovery of a Thylacine (Tasmanian tiger) carcass in a cave near Eucia, Western Australia Helictite, 5 Age of the desiccated carcass of a Thylacine (Marsupiala Dasyuriodea) from Helictite, 7 Wallace G., Brenninkmeijer C.A.M. & McGill R.C., 1990 The use of radiocarbon measurements in atmospheric studies Radiocarbon, 32 Manning M.R. & Melhuish W.H., 1994 Atmospheric 14 International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007

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38 Bulletin of the South African Spelaeological Association, 36 : 58-78. Splognse et implication dans le dveloppement des karsts en climat arid. Karstologia, 28 (2) : 13-18. Spelaeological Association, 31 : 25-41. global historical climatology network temperature data base. Bulletin of the American Meteorological Society, 78 : 2837-2849. Smith H.B., Crummett T.L. & Brandt K.L., 1994 Ages of eruption of primate teeth: A compendium for aging individuals and comparing life histories Yearbook of Physical Anthropology, 37 Stenhouse M.J. & Baxter M.S., 1977 Bomb 14 biological tracer Nature, 267 1984 Performance of catalytically condensed carbon for use in accelerator mass spectrometry Nuclear Instruments and Methods in Physics Research, B5 289-293. Wild E.M., Arlamovsky K.A., Golser R., Kutschera W., W., 2000 14 to forensic medicine Nuclear Instruments and Methods in Physics Research B172 Greg Hodgins, George A. Brook and Eugene Marais International Journal of Speleology, 36 (1), 31-38. Bologna (Italy). January 2007



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International Journal of Speleology 36 (1) 39-50 Bologna (Italy) January 2007 Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon) Fadi H. Nader 1 INTRODUCTION A considerable amount of papers have documented concentrically laminated grains termed as cavepisoids, or pearls found in low energy rimstone pools and high energy splash pools in caves (e.g. Baker & Frostick, 1947; Gradzinski & Radomski, 1967; Donahue, 1969; Jones & Kahle, 1986; Jones & MacDonald, 1989; Hill & Forti, 1997 and references herein; Gradzinski, 2001). Only a very few contributions, strictly descriptive, have reported similar cave speleothems from the Middle-East (e.g. Abdul-Nour, 1991; Choppy, 1991; Karkabi, 1991). This and geochemical studies on cave pearls found in Lebanon collected from the Kanaan cave (central Lebanon). New data about the formation of this type of speleothems in a typical Mediterranean setting is provided. Also, the various controlling factors on the genesis and growth of cave pearls with a special focus on the inherent role of diagenesis are discussed. Nader F.H. 2007. Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon). International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). ISSN 0392-6672. The Kanaan cave is situated at the coastal zone, north of Beirut City (capital of Lebanon). The cave is located within the upper part of the Jurassic Kesrouane Formation (Liassic to Oxfordian) which consists mainly of micritic limestone. Twenty seven cave pearls were subjected to petrographic (conventional and scanning electron microscopy) and geochemical analyses (major/trace elements and stable isotopes). The cave pearls were found in an agitated splash-pool with low mud content. They are believed to have formed 18 O V-PDB values 13 C V-PDB 18 O V-PDB ranging between 13 C V-PDB between -12.3 and -12.1. A genesis/diagenesis model for these speleothems is proposed involving invading the inner micrite cortical laminae and the nuclei (cross-cutting the pre-existing mud-envelopes), and the slight depletion in 18 18 O V-SMOW of the water (-4.2) matches with data on meteoric water signature for the central eastern Mediterranean region. Keywords: 18 O, Lebanon. Abstract: Received 5 May 2006; Revised 14 October 2006; Accepted 14 November 2006 GEOLOGICAL BACKGROUND Lebanon covers 10452 km 2 of surface area and stretches along the central eastern coast of the Mediterranean Sea (Fig. 1A). Geomorphologically, it consists of two mountain chains (Mount-Lebanon and Anti-Lebanon) separated by a high inland plain (the Bekaa; Fig. 1A). The western chain (MountLebanon) borders the Mediterranean Sea, displaying in Lebanon is located in the northern part of this mountain chain; i.e. the Qornet es Saouda, 3083 m above sea level (asl). Precipitation (rain and snow) falls in abundance on the Lebanese mountain chains, especially on Mount-Lebanon. The precipitation rate varies between 700 and 1200 mm/year with increasing elevations (i.e. higher altitudes) across Mount-Lebanon, and about 80% of the yearly precipitation falls from November through February (Edgell, 1997). Carbonate rocks (limestone and dolostone) dominate the known Lebanese rock succession. The oldest exposed rocks are of Liassic age (part of the Early-Middle Jurassic Kesrouane Formation; Dubertret, 1975). The Jurassic strata constitute Available online at www.ijs.speleo.it International Journal of Speleology 1 Box: 11-0236/26 Beirut, Lebanon E-mail: fadi.nader@aub.edu.lb Paper presented during the 14th International Congress of Speleology, Kalamos (Greece) 21-28 August 2005. 21-28 AUGUST 2005 Anniversary 40 th

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40 the cores of the Mount-Lebanon and Anti-Lebanon. The Cretaceous strata especially the CenomanianTuronian Sannine and Maameltain Formations most of the country surface-area. The stratigraphy of Lebanon has been investigated by a number of authors; e.g. Dubertret (1955), Saint-Marc (1974 and 1980), Walley (1997), Beydoun (1988), and Nader (2000). The Late Jurassic is characterized by a period emergence, the Jurassic rock-mass was deeply Subsequently, volcanic deposits and continental depressions accentuating the palaeotopography (Renouard, 1955; Nader et al., 2003). During the Cretaceous times, the Early Jurassic strata were buried to a depth reaching 3 km (Nader et al., occurred during the Oligocene (Dubertret, 1975; Nader & Swennen, 2004). Since the Miocene, the morphology of Mount-Lebanon was not very different than that of today (Dubertret, 1975; Walley, 2001). diagenesis processes, is expected to have affected the exposed Mount Lebanon at least since that time (i.e. Miocene). KANAAN CAVE SETTINGS The Kanaan cave is located in the Kassarat area to the east of the coastal town of Antelias, a few kilometers north of Beirut City (capital of Lebanon; Fig. 1B). The cave is located within the deeply Formation (Fig. 1B). The deepest known sinkholes (Houet Fouar ed Dara, -622 m and Houet Qattine Azar, -450 m) as well as the longest cave in Lebanon (Magharet Jeita, 9 km) are found within this rock unit. The Kanaan cave was discovered in 1997, when its entrance became exposed during rock-quarrying. The entrance is situated at around 100 m above sealevel, almost at the foot of a steep limestone cliff with a height exceeding 150 m (Fig. 2A). The monotonous blue-grey (fossiliferous) micritic limestone lithology characterizing the Kesrouane Formation (Dubertret, 1975; Walley, 1997) is, nevertheless, disrupted with minor clayey/marly horizons in some places evidence supporting paleosols and subaerial exposure were also suggested (see Nader & Swennen, 2004). A marly horizon, rich in brachiopods, including relatively thin seams of coal, was observed underlying the Kanaan cave entrance and its underground network. This horizon provides a local, impermeable substratum for role with respect to the cave speleogenesis. The Kanaan cave, which amounts to some 120 m International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader

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41 of underground passages and chambers, is divided into three parts (Fig. 2B): (i) the Entrance, (ii) the Mud Gallery, and (iii) the Calcite Gallery. The cave entrance overlies a rock-collapse slope (quarrying debris) and a room with a high chimney, hosting a community of bats. A low passage trending northward leads to another room with another chimney (ca. the Northern Chamber, Fig. 2B). The Mud Gallery is named after the considerable amount of mud existing in this part of the cave forming small hills of slippery mud. Here, relatively thick rock slabs (with thickness exceeding 1 m) collapsed from the cave-ceiling. These massive is covered with mud. The southernmost part of the Kanaan cave, the Calcite Gallery, is loaded with a wide variety of speleothems. It consists of a big hall with a high ceiling (more than 20 m) and a lateral development with reverse shape (see Fig. 2B). The to the mud found in the Mud Gallery). A wide diversity of speleothems is observed (stalagmites, stalactites, among others). The investigated cave pearls were collected from the middle part of this hall (Fig. 2B; small depressions hosting cave pearls (Fig. 3C, D). To the opposite side, a relatively large, but shallow, water pool exists (Fig. 2B). MATERIAL AND METHODS Twenty-seven cave pearls were collected from the Kanaan cave. Petrographic observations included conventional microscopy and scanning electron microscopy. Ca, Mg, Na, Sr, Fe, Mn, Zn, and K concentrations in powdered samples, micro-drilled from the various concentric growth layers within cave spectrometry (AAS). Powdered samples (1g of each) were leached in 40 ml (1 molar) HCl and left on hot plates until evaporation the strength of the acid was chosen to be as low as possible in order to leach the minimum possible quantity of the noncarbonate fraction. The residues were dissolved for a diluted before AAS analysis. Analytical precision was (B) cave survey showing the three distinct sections and the location of the cave pearls (from Nader, 1998). International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon)

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42 and the detection limit was 1 ppm. Water, sampled from the cave pools, was also subjected to chemical analyses (including Ca, Mg, Na, Sr, Fe, Mn, Zn, and K concentrations). Various diagenetic phases were microsampled for measurement of their carbon and oxygen stable isotopic composition. Stable isotope analyses (Institute of Geology and Mineralogy). The carbonate powders were subjected to reaction with phosphoric acid (density >1.9; Wachter & Hayes, 1985) at 75C in an online carbonate preparation line (Carbo-Kiel single sample acid bath) connected to a Finnigan Mat 252 mass-spectrometer. All values are reported in per mil () relative to Vienna Pee Dee Belemnite (V-PDB). Reproducibility based on replicate analysis of laboratory standards is better than 0.02 for 13 C and 0.03 for 18 O. KANAAN CAVE PEARLS The approach followed by Jones & MacDonald (1989) to characterize cave pearls according to their size and shape, external appearance, nucleus, and cortical laminae is applied here as well. The observed and collected Kanaan pearls range in size from about 1 to 3 cm (with rare exceptions exceeding 5 cm; Figs. 3 & 4). The shape of these cave pearls varies between and ellipsoidal forms. Some of them are attached to observed cave pisoids show an external smooth, wellpolished, lustrous morphology (Fig. 3C) advocating 1947; Donahue, 1969; Hill & Forti, 1997). Less commonly, some cave pearls show rough, irregular crenulated surfaces made up of tiny trigonal calcite crystals (Fig. 4). The majority of the cut and polished cave pearls show distinct, relatively spherical nuclei (Fig. 5). Only a few samples revealed elongated ellipsoidal nuclei irrespectively of the external shape of the cave pearls. The nuclei are relatively small with respect to the cave pearls (less than 10 mm; Fig. 5). Yet, a relatively thick, yellowish aureole usually surrounds the nucleus. The investigated cave pearls revealed several concentric growth layers around the nuclei. These cortical laminae consist of a repetition of grayish translucent and massive milky white laminae (Fig. 5). The number of these growth zones (and their thickness) is not related to the size of the cave pearls (Fig. 5). PETROGRAPHY Five thin-sections were prepared out of the twentyseven sampled cave pearls. The thin-sections were pearls consist only of calcite. They were examined with conventional microscopic techniques. The thin-sections show two types of cave pearl in shape. It consists of terra-rossa with some pyrite (opaque minerals shining under induced light) and probable organic matter as well as oxides/ hydroxides (yellowish-brown stained material; Fig. 6A). In the second group, the nucleus is made up of clear micritic, anhedral calcite crystallites including clear calcite spars (Fig. 6B, C). Relatively thin, dark cortical laminae envelop the nucleus displaying micritic calcite and a considerable amount of terrarossa. In both cases, the nucleus is surrounded by an aureole of darkish (impurity-rich) micritic calcite. It is worth mentioning that the second group of cave pearls clearly shows under polarized view a divergent outward, pseudo-uniaxial cross extinction (Fig. 6D), resulting from the convergent fabric of elongated columnar or palisade length-fast calcite crystals (cf. Kendall & Tucker, 1973; Folk & Assereto, 1976). The cortical laminae which are close to the nucleus (inner laminae) consist of micritic calcite crystals (often trigonal in shape; Fig. 7A, B). Here the intercrystalline porosity is occluded with impurities (probably terra-rossa, clay and/or organic matter). The outer cortical laminae consist of elongated, radial, sparry calcite crystals (size exceeding 1 mm; Fig. 7C, D) similar to length-fast (palisade) calcite crystals (cf. Folk & Assereto, 1976), with the original intercrystalline porosity occluded by other generations of calcite spars. Similar crystal fabrics were called feather-like calcite crystals by Gradzinski & Radomski (1967) and dendrite calcite crystals by Jones & MacDonald (1989). Alternatively, the growth competition may have resulted in similar textures where some crystals did not grow fast as the others, and remained of smaller sizes due to lack of space (Fig. 7C, D). Fig. 8A shows the transition from inner micritic calcite cortical laminae to outer sparry calcite laminae. A faint syntaxial growth is revealed, as the same extinction is displayed. The impurity-rich thin layer (mud-cover) separating these cortical laminae show a discontinuous pattern, where extensions of the sparry calcite crystals prevail (Fig. 8B). This pattern i.e. the local removal of impurity-rich thin laminae suggests that recrystallization during and/or after the sparry calcite formation must have occurred. The external morphology of the cave pearls was mainly investigated through scanning-electron microscopy (Fig. 9A, B). The calcite crystals of the outermost growth layer display a general trigonal morphology (crystal size: 150 to 500 m). Further investigation show that the apexes of these crystals are rhombohedra faces that are combined to the trigonal prism faces (Fig. 9A, B). This crystalline framework results in a considerable amount of intercrystalline porosity (Fig. 9A). Further investigations of the calcite crystals revealed that they consist of several generations of smaller, embedded trigonal crystals welded together (Fig. 9B). This could be viewed as an aggradational crystal growth, keeping the same crystalline morphology and leading to larger crystals. Note that the sharp extremities of the trigonal crystals are often broken, probably due to corrosion induced by movement of the pearl. International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader

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43 pearls at a closer zoom (C, D). Note that some pearls are well polished (C) others are more or less crenulated (D). Fig. 4. The twenty-seven cave pearls collected from the Kanaan cave. The sizes of the cave pearls range between 1 and 3 cm. International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon)

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44 are added for reference. inner micrite laminae to the outer spar laminae. International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader

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45 are of smaller sizes and seem embedded within the calcite spars.are added for reference. Fig. 8. Photomicrographs showing the transition from inner micrite laminae Fig. 9. SEM Photomicrographs showing crystal microfabrics of the crystals with rhombohedral apexes and a considerable amount International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon)

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46 GEOCHEMISTRY Table 1 shows the major and trace element composition of the only water sample (A) collected from the rimstone pool located opposite to the location of the pearls (cf. Fig. 2), the inner and outer parts of the cave pearls and a typical spelean calcite from a nearby coastal cave (B). Table 2 shows the 18 O and 13 C results (in V-PDB) of the cortical laminae from four different cave pearls. The stable isotopic values of a recent spelean calcite (less than 30 years old; precipitated in a tunnel dug in 1971) are also added as a reference for the actual isotopic signature of speleothems in central coastal Lebanon. Temperature measurements inside the cave were conducted during several visits. The air temperature in the Calcite Gallery (cf. Fig. 2B) is constantly about 20C, while the water temperature measured in the rimstone pool opposite to the location of the pearls, cf. Fig. 2B is about 18C with a pH of 7.65. The cave water has a calcium concentration ranging between 54.6 and 57.1 wt.%, hence it is over-saturated with Ca. The same water possesses about 10 ppm of Na, about 1 ppm of Sr. The measured Fe, Mn, Zn and K concentrations in the pool-water were all below the detection limit. The inner and outer layers of the cave pearls display almost the same Ca concentrations (ca. about 36.4 wt.%). The inner layers show a relatively high Na content (54 ppm) with respect to that of the outer layers (37 ppm). The Sr, Fe, Mn, Zn, and K contents are almost the same for inner and outer layers, and they are higher than those of the cave water (see Table 1A, B). The insoluble residue (IR%, after dissolution in 1 molar HCl) is slightly higher for the outer layers (1.2 wt.%) with respect to that of the inner layers (1.7 wt.%). When compared to similar trigonal spelean calcite from a nearby coastal cave, Ca and Fe contents in the cave pearls are found higher, while Na and Sr concentrations are much lower, respectively. The higher Na and Sr values could be related to sea-water incursions into that coastal cave. Figure 10 is a crossplot featuring the 18 O versus 13 C values for the cortical lamina from different cave pearls collected from the Kanaan cave (Table 2). The data show clearly a decreasing trend for both 18 O and 13 C from inner to outer cortical layers. The average oxygen isotopic values of the inner layers are in the order of -5.0 V-PDB, and the corresponding 13 C from Table 2). International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader

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47 values are around -11.8 V-PDB. The outer layers exhibit 18 O values ranging between -5.3 and -5.2 V-PDB, and 13 C between -12.3 and -12.1 V-PDB. from the tunnel dug in 1971 in Jurassic limestones in order to reach the underground network of a nearby Jerita cave, reveals an average 18 O value in the order of -6.1 V-PDB and 13 C value around 9.1 V-PDB. While this oxygen isotope signature is lighter than those recorded in the Kanaan cave pearls, the carbon isotopic value is less depleted. DISCUSSION According to Baker & Frostick (1947) and Donahue (1969) the Kanaan spherical to subspherical concretions, which have smooth (polished) and lustruous shining surface, may be termed cave pearls and are believed to have formed in agitated water. Donahue (1969) related the genesis of ooid and pisoid grains to the prevailing energy (agitated versus nonagitated conditions). The agitated grains were, thus, characterized by distinct concentric laminations, a pseudo-uniaxial cross (under polarized view), nucleus, smooth polished surfaces, low insoluble residue. All of these characteristics match well with the investigated cave-pearls. Some of the Kanaan cave pearls contain nuclei surrounded with yellow-brownish aureoles (Fig. 5). These are impurity-rich micritic calcite cortical laminae directly enveloping the nucleus. Such pearls usually include a nucleus with preserved terra-rossa. They do not display uni-axial cross under polarized light. The impurity-rich nucleus and lamina in some pearls could suggest the presence of organic matter as well as clay and oxides/hydroxides. However, their occurrence is relatively limited when compared to the overall volumes of the investigated pearls. Jones & MacDonald (1989) have discussed some similar micrite laminae in spearls from a cave in the British West Indies. These authors suggested seven alternatives for the origin of the micritic calcite; out of which several scenarios could also work for the Kanaan cave pearls e.g. inorganic precipitation from cave water, formation of destructive coatings, and/or formation of constructive envelopes. The role of bacteria in the genesis of the Kanaan cave is refuted based on: (1) the cave pearls were found in splash-pools where water 18 O composition of the water from which the cave pearls precipitated for temperatures between 18 and 20C. The 18 O V-SMOW V-SMOW ). International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Petrographic and geochemical study on cave pearls from Kanaan Cave (Lebanon)

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48 motion is considerable and agitation prevails, and (2) (Jones & MacDonald, 1989; Gradzinski, 2001). Yet, effectively destroyed pre-existing bacterial patterns. The clear micritic calcite crystallites, devoid of impurities, and the presence of sparry calcite crystals with extinction pattern similar/uniaxial to the lengthfast crystals in the surrounding cortical laminae (see Figs. 6 and 7) advocate for recrystallization phases that are believed to have occurred after the formation of the nucleus. This also suggests that the uniaxial cross pattern could be associated with some degrees of recrystallization of the pearls, especially when length-fast calcite crystals are observed in the nucleus and the different cortical lamina (Fig. 7A, B). The spar calcite laminae consist of elongate fastcrystals (Fig. 7C, D). This fabric may be explained by growth competition, whereby some crystals grew faster than the others in a radial, outward direction (e.g. Fig. 6D). Alternatively, the spars were formed and then the inter-/ intracrystalline porosity was occluded by another phase of calcite cement. The feather-like arrangement (Gradzinski & Radomski, 1967) has been observed where the nucleus is made up of calcite micrite and spars rather than terra-rossa. Here, the spars seem to invade the inner cortical laminae and the nucleus (Fig. 6). In addition, the mud envelopes (impurity-rich thin layers thought to have covered the pearl during its growth) are incorporated within the length-fast sparry calcite crystals (Fig. 8). Such envelopes appear like ghost inclusions within the calcite and may invoke recrystallization. The sparry calcite represents rapid growth from supersaturated solutions, in response to certain temperature conditions (Jones & Kahle, 1986; Jones & MacDonald, 1989). The water in the Kanaan caves rimstone is highly saturated with Ca (about 56 wt.%) and its temperature is about 18C and the pH around 7.65. The outermost l ayers of the investigated cave pearls consist of crystals having rhombohedra apexes combined to trigonal prism faces, where a considerable amount of intercrystalline porosity remains preserved. Similar crystalline fabrics to the ones discussed by Jones & MacDonald (1989) are observed in the Kanaan cave pearls. The prismatic trigonal crystals reveal through SEM investigation that they consist of crystallites displaying similar trigonal morphology (Fig. 9B). This aggradational to form coarser crystals, invokes the role of recrystallization as well. Trigonal crystals are believed International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader

PAGE 11

49 to form exclusively in meteoric vadose environments salinity and through CO 2 degassing (Binkley et al., 1980; Given & Wilkinson, 1985). The actual water in the Kanaan cave has a relatively high Ca/Mg ratio, suggesting that its low salinity and especially CO 2 degassing are the controlling factors for the precipitation of the trigonal crystals. The Jurassic Kesrouane limestones are characterized by Na concentrations ranging between 103 and 214 ppm (Nader et al., 2004), higher than the Na contents measured in the cave pearls. These rocks have Sr contents between 110 and 193 ppm (Nader et al., 2004). Na and Sr concentrations measured in the investigated cave pearls could have originate from the dissolution of the host-rock (i.e. the Kesrouane limestones) and/or clays and impurities. Iron and manganese contents are believed to be related to the vadose and near-surface environments (cf. Lohmann, 1988). The oxygen and carbon isotope values from the various layers from four cave pearls showed that a decrease in both 18 O and 13 C generally prevails from inner to outer layers. The 18 O depletion could be related to precipitation from water that has lower 18 O values, higher temperature or recrystallization (Lohmann, 1988). Degassing of CO 2 may also result in a depletion in 13 C values (Given & Wilkinson, 1985), this is further supported by the trigonal crystalline pattern observed at the outer part of the pearls (see above). The approach of Woronick & Land (1985) was used in order to estimate the 18 O SMOW of 11). The temperature was set between 18 and 20C and the 18 O PDB of the calcite was measured between -5.33 and -4.94 This resulted in 18 O SMOW around -4.2 Note that the 18 O of the meteoric water in the eastern Mediterranean region ranges between -6 and -4 V-SMOW (Emery & Robinson, 1993). Cave pearls Genesis/Diagenesis Model In general, the nuclei of the investigated pearls show a yellowish aureole with either terra-rossa (preserved at the center) or micrite. The original terra-rossa fabric within the nucleus and the cortical laminae seems to have been destroyed by calcite crystal formation and recrystallization. The micrite calcite cortical laminae which envelope the nuclei, possess relatively higher 18 O and 13 genesis during spelean evaporative conditions, i.e. CO 2 degassing (Gradzinski & Radomski, 1967). The overlying spar calcite laminae resulted from fast precipitation in water highly saturated with Ca and during further degassing of CO 2 the relative depletion in 18 O and 13 C with respect to the inner part of the pearl. Petrographic investigation suggests the predominance of recrystallization in some pearls (especially where the nucleus is devoid of terra-rossa). This is somehow supported by the corresponding lower 18 O values. Figure 12 shows two cut-faces of two distinct cave pearls and a sketch presenting a tentative model explaining the formation of the cave pearls and their related recrystallization. After the precipitation of the nucleus and impurityrich, inner micrite cortical laminae, the surrounding spar laminae (length-fast calcite) were formed in water highly saturated with Ca with enhanced CO 2 degassing. Subsequently, trigonal prismatic calcite crystals were precipitated on the outermostlayers. Such process has also triggered selective recrystallization of the inner layers. With each new concentric layer added some inner layers would have severed some degrees of (re)crystallization; e.g. destruction of mud envelopes, occlusion of porosity and crystal aggradations CONCLUSIONS Based on petrographic and geochemical data of some cave pearls from the Kanaan cave (Jurassic, central Lebanon), the following points can be concluded:1. The environment of cave pearl formation consists of an agitated splash-pool, with low mud content.2. The formation of the cave pearls is believed to over-saturated with calcium here biogenic related precipitation is dismissed. The internal nucleus and micritic laminae ( 18 O V-PDB : -5.0 ; 13 C V-PDB :-11.8 ) include impurity-rich calcite crystal framework; while the surrounding length-fast calcite spar laminae ( 18 O V-PDB : -5.3 to -5.2 ; 13 C V-PDB : -12.3 to -12.1 ) have precipitated from water with low salinity, highly saturated with Ca and during enhanced CO 2 degassing. 3. A model for the formation of the Kanaan cave pearls is proposed involving recrystallization which has selectively affected the inner layers of the cave pearls during the growth of the successive outermost layers. 4. The calculated 18 O V-SMOW of the water (-4.2) matches with data on meteoric water signature for the central eastern Mediterranean region. ACKNOWLEDGEMENTS The author is grateful to Joanna Doummar (Dept. of without which this contribution would have never of Erlangen Germany) is thanked for performing the for the technical assistance. This work is partially acknowledged for providing data related to the Kanaan cave (central Lebanon). Critical reviews and valuable comments made by Dr. Calaforra, an anonymous reviewer and the editor (Dr. De Waele) have improved the presentation and content of this contribution, and are greatly appreciated. International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Petrographic and geochemical study on some pearls from Kanaan Cave (Lebanon)

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50 REFERENCES Abdul-Nour H., 1991 Perles cubiques Liban Souterrain (Bulletin du Groupe dEtudes et de Recherches Souterraines du Liban), 3 : 64-65. Baker G. & Frostick A.C., 1947 Pisolites and ooliths from some Australian caves and mines Journal of Sedimentary Petrology, 17 : 39-67. Beydoun Z.R., 1988 The Middle-East: Regional geology and petroleum resources p. Binkley K.L., Wilkinson B.H. & Owen R.M., 1980 Vadose beachrock cementation along a southeastern Michigan marl lake Journal of Sedimentary Petrology, 50 : 953-961. Choppy J., 1991 Perles des Cavernes Cubiques et Polyedriques Al-OuatOuate (Revue Libanaise de 6 : 43-47. Donahue J., 1969 Genesis of oolite and pisolite grains Journal of Sedimentary Petrology 39 : 1399-1411. Dubertret L., 1955 Carte gologique du Liban au 1/200000 avec notice explicative Ministre des Travaux Publiques, Beirut, 74 p. Dubertret L., 1975 Introduction la carte gologique au 1/50000 du Liban Orient, 23 : 345-403. Edgell H.S., 1997 Karst and hydrogeology of Lebanon Carbonates and Evaporites, 12 (2) : 220-235. Emery D. & Robinson A., 1993 Inorganic Geochemistry: Applications to Petroleum Geology Folk R.L. & Assereto R., 1976 Comparative fabrics of length-slow and length-fast calcite and calcitized aragonite in a Holocene speleothem, Carlsbad Cavers, New Mexico Journal of Sedimentary Research, 46 (3) : 486-496. Given R.K. & Wilkinson B.H., 1985 Kinetic controls of morphology, composition, and mineralogy of abiotic sedimentary carbonates Journal of Sedimentary Petrology, 55 : 108-119. Gradzinski M., 2001 Role of bacteria in the growth of cave pearls In: Proceedings of the 13 th International Congress of Speleology Brasilia 2001 Gradzinski R. & Radomski A., 1967 Pisolites from Cuban caves Rocz. Polsk, Tow. Geol., 37 : 243-265. Hill C. & Forti P., 1997 Cave Minerals of the World 463 p. Jones B. & Kahle C.F., 1986 Dendritic calcite crystals environment Journal of Sedimentary Petrology, 56 : 217-227. Jones B. & MacDonald R.W., 1989 Micro-organisms and crystal fabrics in cave pisoliths from Grand Cayman, British West Indies Journal of Sedimentary Petrology, 59 : 387-396. Kendall A.C. & Tucker M.E., 1973 a replacement after acicular carbonate. Sedimentology, 20 : 365-389. Karkabi S., 1991 La Perle de Caverne Hexagonale. Al-OuatOuate Karstologie), 6 : 48-53. Lohmann K.C., 1988 Geochemical patterns of meteoric diagenetic systems and their application to studies of paleokarst. In: James N.P. & Choquette P.W. (Eds.) Paleokarst, Springer-Verlag, New York: 58-80. Nader F.H., 1998 Mgharet Kanaan: the temple of speleology. Al-OuatOuate (Revue Libanaise de 11 : 54-59. Nader F.H., 2000 Petrographic and Geochemical Characterization of the Jurassic-Cretaceous Carbonate Sequence of the Nahr Ibrahim Region, Lebanon. M.Sc. Nader, F.H. & Swennen R., 2004 Petroleum prospects of Lebanon: Some remarks from sedimentological and diagenetic studies of Jurassic carbonates. Marine and Petroleum Geology, 21 : 427-441. Nader F.H., Swennen R. & Ottenburgs R., 2003 Karstmeteoric dedolomitisation in Jurassic carbonates, Lebanon. Geologica Belgica, 6 : 3-23. Nader F.H., Swennen R. & Ellam R., 2004 Stratabound dolomite versus volcanism-associated dolomite: an example from Jurassic platform carbonates in Lebanon. Sedimentology, 51 : 339-360. Renouard G., 1955 Oil Prospects of Lebanon. American Association of Petroleum Geologists Bulletin, 39 (11): 2125-2169. Saint-Marc P., 1974 Etude stratigraphique et micropalontologique de lAlbien, du Cnomanian et du Turonien. Tome XIII, CNRS, Paris/Beirut, 298 p. Saint-Marc P., 1980 Le passage Jurassique-Crtac et le Crtac infrieur de la rgion de Ghazir (Liban central). 7 : 237-245. Wachter E. & Hayes J.M., 1985 Exchange of oxygen isotopes in carbon-dioxide phosphoric acid systems. Chemical Geology, 52 : 365-374. Walley C.D., 1997 The Lithostratigraphy of Lebanon: A Review. 10 : 81-108. Walley C.D., 2001 The Lebanon passive margin and the evolution of the Levantine Neothethys. In: Ziegler P.A., Cavazza W., Robertson A.H.F. & CrasquinSoleau S. (Eds.) Peri-Tethys Memoir 6: Peri-Tethyan 86 : 407-439. Woronick R.E. & Land L.S., 1985 Late burial diagenesis, Lower Cretaceous Pearsall and Lower Glen Rose formations, South Texas. In: Schneidermann N. & Harris P.M. (Eds.) Carbonate Cements, Special Publications of the Society of Economical Paleontology Mineralogists, Tulsa, 36 : 265-275. International Journal of Speleology, 36 (1), 39-50. Bologna (Italy). January 2007 Fadi H. Nader


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