International Journal of 36 (1) 1-21 Bologna (Italy) January 2007

Available online at www.ijs.speleo.it International Journal of Speleology Official Journal of Union Internationale de Spéléologie

Acadian : composition and ecology of fauna of Nova Scotia and southern New Brunswick, Canada. Max Moseley1

Abstract: 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 and disused mines in Nova Scotia and southern New Brunswick are provisionally catalogued and described, based on field collections made over many years. The area was glaciated and the consists of non-troglobites all of which have arrived and colonised the caves during or following final 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 species derived directly from dung communities: another parallel with tropical guano caves. An unanticipated finding is the number of non-indigenous species now utilising local caves. These appear to have colonised unfilled ecological niches, suggesting that post-glacial recolonisation of the subterranean habitat in Nova Scotia has been relatively delayed. Finally the general and regional significance of the subterranean fauna is briefly discussed. Keywords: Cave fauna, Canada, Nova Scotia, New Brunswick, porcupine dung, guano caves, parietal, threshold fauna, introduced taxa, post-glacial, recolonisation.

Received 8 July 2006; Revised 10 October 2006; Accepted 31 October 2006

INTRODUCTION The central focus of biospeleological research and adaptation in the underground environment (e.g. thinking especially in North America has been those Kane & Culver, 1992). highly-adapted specialised troglobiontic Accordingly there is now a clear need for which inhabit deep caves and subterranean waters systematic field surveys and evaluation of the cave in regions to the south of the maximum limits of the and groundwater fauna of neglected regions of the Pleistocene glaciations. Much less attention has been continent, particularly Canada. paid to the subterranean fauna of formerly glaciated The eastern region of Canada comprising northern regions of the continent, or to the many non- Newfoundland, the Maritime Provinces (Nova Scotia, obligate species that occur in cave thresholds there New Brunswick, Prince Edward Island), Quebéc, and elsewhere. and Ontario has many areas of , and natural The limits traditionally imposed by an emphasis on dissolution caves are known from all these provinces troglobionts to the exclusion of other fauna have long except Prince Edward Island. However there have hindered progress in understanding hypogean fauna been almost no specific or regional biospeleological and ecosystems. However this is now changing. Most surveys. The exceptions are a detailed study of the cave ecologists now accept the need to see subterranean fauna and ecology of Frenchman’s Cave, Nova Scotia communities in their entirety encompassing species (Calder & Bleakney, 1965; 1967) and a general survey at many different stages of adaptation (Gibert & of caves and mines in southern Ontario (Peck, 1988). Deharveng, 2002), and evolutionary cave biologists The geographical area dealt with in the present increasingly recognise the value of non-obligate cave- paper, Nova Scotia and the southern part of the dwellers as empirical models of natural selection and adjacent province of New Brunswick (Fig. 1), is the northern end of the Northern Appalachians region. The Maritime Provinces together with areas 1) Research Associate, Nova Scotia Museum of Natural History, of eastern Quebéc are the Canadian part of the 1747 Summer Street, Halifax, Canada B3H 3A6 early French colony of Acadia; hence “Acadian” is a (E-mail: [email protected]) useful term often used in reference to this region.

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007  Max Moseley

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.].

The area is cold-temperate, and somewhat low-lying from present-day patterns of and plant with maximum elevations in Nova Scotia of 532m distribution that some species may have recolonised and 446m in southern New Brunswick. The whole of Nova Scotia from the east by migration from an ice- the present-day land area was subjected to multiple free Wisconsinian Atlantic Coastal Plain Refugium glaciations during the Pleistocene and was ice- (Schmidt, 1986) or more likely from the emergent land covered at the maximum of the last (Wisconsinian) areas which existed during the process of deglaciation glacial advance. The fauna and flora of the region is (King, 1996). The existence of other late-Pleistocene the result of immigration and recolonisation during refugia has also been proposed (e.g. Schmidt, 1986) and following the final retreat of the Wisconsinan but this remains highly speculative. In the historical glaciers, which is believed to have started ~21,000 period European contact resulted in the introduction years BP and to have been complete by ~11,000 years of many exotic species through human migration and BP (King, 1996). Most species must have arrived from sea-borne trade. the south and west. Recolonisation of New Brunswick The area has a number of exposed areas of sul- from these directions was relatively unrestricted by phate (gypsum-anhydrite) and carbonate (limestone major physical barriers but immigration of many and dolostone) bedrock with underground drainage, animals and plants into Nova Scotia is thought to springs, caves and other geomorphological karst fea- have been constrained by the Tantramar Marshes, tures (Moseley, 1976; 1996: McAlpine, 1979). The a narrow marshy isthmus which is the only land southern part of mainland Nova Scotia has no sig- connection with New Brunswick: as a consequence the nificant karst but both gypsum and limestone caves province is zoogeographically an island for many species are generally -distributed throughout the rest of (Fig. 1). The Strait of Canso physically further isolated the region, with gypsum caves and karst predominat- Cape Breton Island until completion of the Canso ing in Nova Scotia and limestone in New Brunswick. Causeway in 1955 (Fig. 1). However there is evidence Dolostone karst is rare and no caves are known.

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 

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 and the many abandoned mines provide ecologically cave-like Fig. 2. Young adult North American Porcupine (Erethizon dorsatum) in habitat. den, dark zone, Cheverie Cave, Nova Scotia. (Photo: F. Vladi) Emerton (1917) reported the Cave Spider Meta ovalis (as M. menardi) in Nova Scotia, although not tracks were reported by Calder & Bleakney (1967) in from caves. Two specimens in the Canadian National Frenchman’s Cave. There are occurrence records of Collection (CNC 3437, 3438) collected in Gays River pallid Brook Trout (Salvelinus fontinalis) in limestone Gold Mine, Nova Scotia, in 1963 appear to be the earliest stream caves (Moseley, 1975; McAlpine, 1979) and underground record. The first published notices Ninespine Stickleback (Pungitius pungitius) and of cave animals in the area are of hibernating bats: Northern Redbelly Dace (Phoxinus eos) in Hayes the Northern Long-Eared Bat (Myotis septentrionalis) Cave (Morris, 1985). Frog (Rana clamitans) tadpoles was reported by Gould (1936) in a cave in Hants were collected in the threshold of Hayes Cave County, Nova Scotia. Bleakney (1965) later found (Morris, 1985). Eastern Pipistrelle (Pipistrellus subflavus) at several The first study of invertebrates was that of the Nova Scotia sites as well as making the first reports terrestrial fauna of Frenchman’s Cave, Nova Scotia of the occurrence of the common Little Brown Bat (Calder & Bleakney, 1965; 1967). Thirty-eight (Myotis lucifugus) in caves and mines in that province. invertebrate taxa were reported from the threshold In New Brunswick, McAlpine (1976) reported P. and deep threshold, the majority associated with subflavusfor the first time (from a cave) and the same decomposing porcupine dung accumulations. Acari author (1979) summarized underground records of were found to be numerically dominant in “poorly this and other bats. There is one sight record of the Big decomposed” dung samples, whilst Collembola Brown Bat (Eptesicus fuscus) in a cave hibernaculum became the most abundant microarthropods later in in Nova Scotia (Scott & Hebda, 2004). Records of bats the ecological succession. at underground sites in Nova Scotia have recently Broader, extensive rather than intensive, inventory been collated by Moseley (in press). of the regional invertebrate cave fauna began in the There has been no systematic effort to document early 1970s with occasional sampling mostly as an other vertebrates, but occasional records and adjunct to other underground work such as cave observations have accumulated. North American mapping, and continued through the 1980s and Porcupine (Erethizon dorsatum) (Fig. 2) and their dung 1990s: a few of the records were published by the have been observed in most caves and a number of former Nova Scotia Speleological Society (Moseley, disused mines in southern New Brunswick and the 1998 and references therein). Earthworms collected mainland of Nova Scotia (Calder & Bleakney, 1965; from several sites in New Brunswick were discussed 1967; McAlpine, 1979; Moseley, 1998). Porcupine did by McAlpine & Reynolds (1977) and a brief summary not reach Cape Breton Island until completion of the of the composition of other New Brunswick cave Canso Causeway. They have since become established fauna was provided by McAlpine (1979). During in the east of the island (Scott & Hebda, 2004) but 1997, a systematic field survey of Nova Scotia cave there are no reported sightings yet in island caves or habitats and invertebrate fauna was performed mines. (Moseley, 1998). Several other vertebrates have been sporadically A few specialised studies have been published recorded underground. McAlpine (1977, 1979) lists based on the collections: Marusik & Koponen (1992); Mink (Mustela vison) scat, Smokey Shrew (Sorex Moseley & Hebda (2001); Moseley et al. (2006); and fumeus), Deer Mouse (Peromyscus maniculatus) and Majka et al. (in press). Christiansen & Bellinger a Beaver (Castor canadensis) den in New Brunswick. (1980, 1998) included records of Collembola in their Beaver have also been seen in a Cape Breton stream comprehensive monographs on the North American cave (Sawatzky, 1986) and Raccoon (Procyon lotor) collembolan fauna.

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007  Max Moseley

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

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 

RESULTS variation. During and early summer as the ambient temperature rises above that underground Fauna recorded relatively cold air is retained inside caves, whilst warm The fauna records are collated and summarised air flows outwards in the autumn, thus skewing the in Tables 1 and 2. Sixteen vertebrate and 170 annual curve (Fig. 3). In certain extreme special cases invertebrate taxa, representing 12 and 99 families this effect results in so-called “ice caves” (see below). respectively, have been identified. An analysis of the Maximum seasonal cave temperatures are recorded in recorded invertebrates by higher taxonomic group is September-October, minimum in January-February. given in Table 3. Eastern Canada experiences a very rigorous winter Excluding accidentals, ectoparasites and those taxa climate. Cave entrances are subject to severe low for which ecological status could not be assessed, temperature conditions throughout the winter and 11 vertebrate and 123 invertebrate cavernicoles there is often a build up of ice and snow. They are (including guanophiles) were recorded in 8 and 72 subjected to repeated freeze-thaw cycles: the average families respectively. Sixty-five of these cavernicolous annual number of cycles (-6°C to +2°C) at Dartmouth, taxa (48%) are categorized as habitual trogloxenes, 44 Nova Scotia has been calculated as 30. As a result (33%) as troglophiles, and 25 (19%) as guanophiles. of this and ongoing dissolution of exposed rock, No troglobites were identified. entrances are often almost filled in by blockfall and Of the cavernicoles 112 (84%) are terrestrial animals, talus derived from within the threshold itself or from 22 (16%) aquatic. Of the 112 terrestrial cavernicoles the cliff face outside (e.g. CB, HC, WIC, and FC). This 38 (34%) are categorized as parietal fauna. physical barrier damps temperature fluctuations 19 (17%) of the terrestrial cavernicoles are almost further inside resulting in fewer, perhaps only one, certainly introduced non-indigenous species (all of them freeze-thaw cycle per annum in the deep threshold. European in origin): the rest are Nearctic, circumpolar The mean annual temperature inside caves in or cosmopolitan. All the non-indigenous species are central Nova Scotia is ~5.7°C. However it is lower at invertebrates. Three further non-indigenous species a few sites. The aspect and physical shape of certain which were collected are considered to occur only as caves cause atypical temperature conditions with ice accidentals in our caves. and snow persisting into mid- or even late summer. In the case of aquatic invertebrate fauna, 20 Snow and ice which accumulate during the winter cavernicolous taxa have been identified in the in WIC for example have been observed to survive collections. Half of these are crustacea: Copepoda until early August. The minimum air temperature (5 species) or Ostracoda (5 species). Most of the rest measured inside the cave on 29 July1997 was 2.5°C, are larvae: Odonata (3 species), Plecoptera and 0.4°C was recorded a few centimeters below the (3 species), and Diptera (1 species). No introduced floor surface on 6 August 1999. Such sites were often aquatic animals were found. 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 Cave environment speleological sense of the term denoting a cave with None of the accessible caves in the region have a permanent ice. constant temperature zone: all are subject throughout The temperature of standing water is usually within to surface environmental influences because of their 0.5°C of the air in the immediate vicinity. There are small size, the presence of a stream, or through drafts several caves (e.g. CB, KC) with active stream sinks due to multiple entrances. Radon gas concentrations that modify the temperature. Temperature may also in the inner areas of one of the largest caves, HC, be affected by the presence of a spring. Some cave (Morris, 1985) suggest almost stagnant air but even streams originate within the cave as eucrenal springs here the temperature changes seasonally, although characterised by cold, clear sediment-free water with the annual range is only ~2°C. This site may be their temperature remaining within two degrees of considered “deep cave” in the sense of Howarth the local annual mean. Examples are a small spring (1988). Annual temperature range in the dark zone in MIC (4.3°C in September 1995) and the F2 cave of a more representative small Maritime Canadian stream (5.1°C in July 1997). cave is ~4°C (Fig. 3): many caves experience greater Cave waters in both limestone and gypsum caves are usually slightly alkaline: pH7.3-7.6. Acidic conditions

8 may occur in stagnant or slowly flowing water where there are accumulations of porcupine dung or plant 7 litter e.g. pH5.8 was measured in seeps with porcupine 6 scat in WIC, pH6.5 in pools with plant flood debris in ºC 5 CB. Due to the high solubility of calcium sulphate (i.e. 2.438g.l-¹ in distilled water @10°C) and the presence 4 of other dissolved solids such as calcium carbonate, 3 gypsiferous waters have high conductivity. Analysis Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan of water samples from ponds in HC (Morris, 1985) Fig. 3. Average monthly air temperature (°C) in a representative showed conductivity readings of 2220±10µS.cm-¹. Nova Scotia cave (dark zone in Frenchman’s Cave). Samples of water from FC were so high in calcium

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007  Max Moseley sulphate that microcrystals of selenite precipitated F2), while the other extreme is represented by a out on cooling in a domestic refrigerator. remarkably simple ecosystem observed in GM where samples of well composted dung from the dark zone yielded only Protaphorura armata (Collembola) and Habitats enchytraeids. Qualitative observational evidence Although characteristically small, caves in the region suggests that moisture content is the most important contain a diversity of terrestrial and aquatic eutrophic variable: both biomass and species diversity decrease (dung), mesotrophic and oligotrophic habitats. in dry material. The abundance of Enchytraeidae North American Porcupine (Fig. 2) dens are almost in particular is affected by the moisture content ubiquitous in caves and mines everywhere except and they are infrequent or absent in drier samples. Cape Breton Island. A den site is typically a small Decompositional sequence is also a major factor. The side passage, cavity or blind pocket within the deep general ecological succession reported by Calder & threshold or dark zone occupied by a single animal. Bleakney (1965) with Acari numerically most abundant Cave dens always have an accumulation of faeces in “poorly decomposed” samples and Collembola either as scattered pellets or more substantial becoming dominant later has been observed at other accumulations (Fig. 4). In areas adjacent to the sites. It was very apparent in GM where mites were porcupine’s access routes sustained inputs of dung abundant in moderately decomposed material (which sometimes result in deposits tens of centimeters deep also contained isotomid Collembola as well as P. covering several square meters of cave floor. These armata and Enchytraeidae) but, as mentioned above, dung piles can occur anywhere from the threshold were absent from samples of well composted dung. through to the cave dark zone. Etiolated seedlings are Acari are the most taxonomically diversified commonly present on fresh dung growing from seeds group in porcupine dung. Most have so far only which have passed through the gut of the porcupine been identified to genus or family. Parasitids (Calder & Bleakney, 1965) (Fig. 4). Decomposing (Parasitus, Eugamasus, Vulgarogasmus) are almost dung fuels a varied community of bacteria, fungi, always abundant, and rhagidids (Rhagidia) are oligochaetes, , and other . usually common. Other mites, some of which may The dung habitat is non-uniform, varying be common to abundant at some sites, include microclimatically and qualitatively with diet, stage of Vegaidae (Vegaia), Zerconidae (Zerconopsis), Ascidae decomposition, and environmental factors including (Arctoseius), Ameroseidae (Epicriopsis), Eviphidae moisture content, ecological zone within the cave, (Alliphis), Macrochelidae (Geolapsis), Pygmephoridae and temperature profile at each site. Fresh dung is (Pygmephorus, Bakerdania), Tetranchidae (Bryobia), acidic (~pH5.1) but in wet areas, such as where there Acaridae (Acarus immobilis), Banksinomidae (Oribella) is seepage water or under roof drips, the acidity is and various unidentified Uropodidae, Erynetidae and neutralized by the buffering effect of cave waters. Histiostmatidae. Also, as observed by Calder & Bleakney (1965), Collembolan populations are less diverse. They are the decompositional sequence is accompanied by almost always dominated by onychiurids (Protaphurura, decreasing acidity, so that well decomposed and/or Tullbergia) and isotomids (Folsomia, Isotoma). wet dung is slightly alkaline: ≤ pH7.3. Neelids (Megalothorax minimus), podurids (Willemia Because of the variability of pH and other empirically scandinavia) and entomobryids (Pseudosinella alba, measurable factors, and because dung piles in the field Tomoceros minor) are found more infrequently. often accumulate over many years, it was not possible The insect fauna of dung is dominated by nematoceran to accurately determine the stage of decomposition of larvae. Larvae of Trichocera maculipennis and individual samples. However visual appearance is a various sciarids are characteristically present and useful approximate guide to the sequence. Fresh scat usually abundant (Figs, 7, 8). At least three different (Fig. 4) consists of scattered, ovoid, greenish-grey pellets types of sciarid larvae are found, none of which have with a mucoid surface. The mucoid material disappears yet been matched with the adults recorded associated rapidly (poorly decomposed). The pellets retain their with dung in the caves. The latter include unidentified shape and physical integrity for some time but turn species of Bradysia, Lycoriella and Scatopsciara. darker brown in colour and the fibrous nature of their Larvae of Limonia cinctipes, Chaoborus, Smittia, and constituents becomes visible (moderately decomposed) Psychoda also occur in some samples, as do those (Fig. 5). In the later stages of decomposition they break of the brachyceran fly Leptocera. Adult Scatopsciara, down physically forming a material rather like dark Chaoborus and Leptocera have been observed well-weathered sawdust in appearance and consistency attracted to fresh scat, presumably ovipositing. A (well decomposed) (Fig. 6). few beetles are also found in this habitat. Larvae of Invertebrate communities in porcupine dung piles Quedius s. spelaeus are often common in moderately are dominated by Acari, Collembola, dipteran larvae decomposed dung, whilst the somewhat rarer adults and enchytraeids (Figs. 7, 8). There are marked are found under stones or running over the cave floor, differences in the species composition and biomass almost always on or near dung (Moseley et al., 2006). of this community from site to site depending on The tiny guanophile Acrotrichis castanea is sometimes microclimatic and qualitative factors. More than 35 abundant, although it may be overlooked because of terrestrial invertebrate taxa are recorded associated its size. Aphodius aleutus and Corticaria pubescens with dung in the Frenchman’s cave system (FC + have been collected from dung: both occur in such

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 

Fig. 4. Entrance of cave den occupied by the porcupine illustrated in Fig. 7. Fauna extracted from a sample of ‘moderately decomposed’ Fig. 2. Numerous fresh droppings are clearly visible. The white areas porcupine dung, Frenchman’s II, Nova Scotia, October 1997. Sciarid are mats of fungal hyphae on older dung, and there are a number (Diptera) larvae are abundant, and there is an adult in the lower right. of etiolated plant seedlings growing from seeds which have passed Parasitid mites and onychiurid Collembola (Protaphorura armata) are through the gut of the porcupine. Several porcupine quills are also scattered throughout the field of view. The large coleopteran larva in visible. (Photo: author). the lower centre is Quedius s. spelaeus, and there are two adult ptilid beetles (Acrotrichis castanea) just above this. (Photo: C. Majka).

Fig. 5. ‘Moderately decomposed’ porcupine dung sample from Frenchman’s II, Nova Scotia (Photo: author).

Fig. 8. Fauna extracted from a second sample of ‘moderately decomposed’ porcupine dung, Frenchman’s II, October 1997. Trichocera maculipennis larvae (Diptera) and an enchytraeid are Fig. 6. ‘Well decomposed’ porcupine dung sample from Frenchman’s prominent. The adult fly near the centre is an unidentified sciarid. II, Nova Scotia (Photo: author). (Photo: C. Majka).

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

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

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 10 Max Moseley with oligotrophic sites e.g. Heteromurus nitidus and shelter and forages outside. There are also two Arrhopalites hirtus. Scoliopteryx libatrix and Triphosa haesitata. Adult As in virtually all cave ecosystems porcupine dung helomyzids (Scoliocentra, Helomyza, Amoebaleria, communities are simplified with two fundamental and Tephrachlamys) are common and are present trophic levels, primary producers and herbivores, year round but their purpose in entering caves absent. They differ from most other cave guano is unknown: their larvae are not found in this ecosystems in that the porcupine is herbivorous habitat. feeding preferentially on the cambium of trees, Several other arthropods (Oniscus asellus especially conifers. The remaining two classic [Isopoda], Polydesmus angustus [Diplopoda]) trophic levels, decomposers and predators, are and gastropods (Arion, Deroceras laeve, Trichia easily recognized. This is usually true of cave guano hispida) are commonly found on rock surfaces in communities but may not be the case in oligotrophic the threshold but rarely much further in. Another cave communities where decreasing resources are gastropod Zonitoides arboreus appears to be associated with blurring of the distinction and even somewhat more cavernicolous in its behaviour than disappearance of obligate predators in favour of the other recorded mollusca, and is sometimes omnivores (DeHarveng & Bedos, 2000). Quedius s. found well inside the dark zone. These species are spelaeus adults and larvae and various predatory also common in dark moist habitats on the surface mites (e.g Alliphis, Geolapsis) are the common and unlike the animals traditionally included in predators in porcupine dung communities. the parietal association are apparently not in caves There is an unusual suite of parietal species for a specific purpose. However, because they are that is ecologically fundamentally different in its characteristically present in this habitat and the nature from the original sense of Jeannel (1926). arthropods are preyed upon by parietal spiders, it It constitutes a distinct faunal component derived seems logical to include them as parietal fauna. directly from the porcupine dung fauna. The larvae The subterranean aquatic fauna is undercollected of several Diptera (Limonia cinctipes, Trichocera and the statistical dominance of terrestrial (80%) maculipennis, Chaoborus, various sciarids, Psychoda, versus aquatic (20%) taxa in the lists certainly and Leptocera) live in porcupine dung, and adult partly reflects this collecting bias. Scatopsciara, Chaoborus and Leptocera have been observed apparently ovipositing in very fresh pellets, Mammals, amphibians and fishes thus, unlike almost all the “traditional” parietal Most of the recorded vertebrates enter caves for insects, these Diptera are probably able to complete definite purposes and thus may be considered to be their life cycle underground. This is reminiscent habitual trogloxenes. of the situation in the humid tropics where there Two fishes, Salvelinus fontinalis and Pungitius is frequently an assemblage of arthropods derived pungitius, are seen frequently enough to suggest from guano communities on cave walls deep inside that in behaviour they are the most cavernicolous caves (Deharveng & Bedos, 2000). vertebrates in the region. Beaver as well as Porcupine A ‘parietal association’ in Jeannel’s (1926) sense use underground sites as dens. McAlpine (1977) can also be readily recognized in Maritime caves. It reports records of beaver in caves from , is very similar to that found in Ontario (Peck, 1988), Missouri and New Brunswick. Two small mammals, northern Europe and other cold temperate regions the Smokey Shrew (Sorex fumeus) and the Deer Mouse of the world and comprises the associations of (Peromyscus) are known from research elsewhere to arthropods living or resting on cave walls and other predate hibernating bats in caves (Banfield, 1974; rock surfaces together with the spiders that prey Trevor-Deutsch, 1973) and are thought to enter local upon them. Parietal predators tend to be specialized caves primarily for this purpose, whilst Raccoon and forms found only in this and similar dark humid Mink enter them to forage. habitats such as cellars, and are usually considered The remaining vertebrates are probably accidentals. to be troglophiles. Most of the parietal association This is certainly the case for the adult frogs seen however consists of habitual trogloxenes that are occasionally, perhaps entering during flooding. They using the habitat temporarily for shelter, summer have however been observed to survive for some time aestivation, hibernation, overwintering, or other where a stream brings in food and offers suitable purposes that are not yet fully understood. Adult habitat (e.g. CB). Phoxinus eos was observed in HC for Diptera predominate both in number of species several years, but this fish has not been seen in any and number of individuals. Overwintering fauna other cave and collected specimens were pallid and includes Nelima elegans (Moseley & Hebda, 2001), undernourished. A single extra-limital winter record female Culex spp. and some species of Bolitophilia, of the migratory bat Lasionycteris noctivagans in PT Rymosia and Tarnania (). Some of probably represented a stranded storm-driven animal the (Mycetophilidae) along with the adult (Hebda, pers. comm.) Limniphilidae occasionally seen in New Brunswick caves are probably there for summer aestivation. Energy inputs: porcupine dung Two spiders (Meta ovalis and Nesticus cellulanus) Porcupine and extensive deposits of their dung specialise in predating other parietal fauna. were first reported in caves in (Graham, Ceuthophilus brevipes uses caves and mines for 1962). Calder & Bleakney (1965, 1967) investigated

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

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 12 Max Moseley

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

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 13 are habitats where species normally living further to the Calder D. R. & Bleakney J. S., 1965 - Microarthropod north or to the south may sometimes be found (Gunn et Ecology of a Porcupine-Inhabited Cave in Nova Scotia. al., 2000). Ecology, 46 (6): 895-899. The cave fauna of Nova Scotia is now probably the Calder D. R. & Bleakney J. S., 1967 - Observations on most extensively and comprehensively sampled and Frenchman’s Cave, Nova Scotia, and Its Fauna. Bull. documented of any geographic region in Canada. Records Nat. Spel. Soc., 29 (1): 23-25. go back to the 1960s and large documented collections Chapman P., 1993 - Caves and Cave Life, New Naturalist, of both terrestrial and aquatic invertebrates have been Harper Collins, London, 219 p. made since then in caves and abandoned underground Christiansen K. & Bellinger P., 1998 - The Collembola mines around the province. of North America North of the Rio Grande: A Taxonomic Invasive and other species in Maritime caves may have Analysis. Grinnell College, Grinnell, Iowa, 1520 p. high potential for investigating and testing biospeleological Christiansen K. & Bellinger P., 1980 - The Collembola evolutionary theories with populations at a very early stage of North America North of the Rio Grande. Grinnell of active colonisation and adaptation to the subterranean College, Grinnell, Iowa, 1322 p. environment. 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.) ACKNOWLEDGEMENTS - Ecosystems of the World: Vol. 30, Subterranean Biota. Any prolonged wide-ranging study is impossible Elsevier, Amsterdam: 603-632. without the help and participation of many individuals, Emerton J. H., 1917 - Spiders Collected in Nova Scotia and it is unfortunately impossible to name them all and New Brunswick by Robt. Matheson in 1912. Proc. here. I would especially like to express my gratitude N. S. Ent. Soc., 1917: 95-96. to the many taxonomists who identified specimens: Gibert J. & DeHarveng L., 2002 - Subterranean there is a list in Moseley (1998). Mention must also ecosystems: a truncated functional . be made of those individuals who accompanied me BioScience, 52: 473–481. on field trips: especially M. Hamilton, C. Murphy, Gnaspini P., 1992 - Bat guano ecosystems. A new the Proctor brothers, Dr. D. Sawatzky, and the late classification and some considerations, with special Dr. P. Schwinghamer. My colleagues amongst the reference to Neotropical data. Mem. Biospéléol., permanent staff and Research Associates of the Nova 19: 135-138. Scotia Museum of Natural History have also helped in Gnaspini P. & Trajano E., 2000 - Guano Communities many ways. Dr. Dale Calder (Royal Ontario Museum) in Tropical Caves. In: Wilkens H., Culver D. C. & commented on the manuscript and also provided useful Humphreys W. F. (Eds.) - Ecosystems of the World: Vol. unpublished data from Frenchman’s Cave. Additional 30, Subterranean Biota. Elsevier, Amsterdam: 251- unpublished records were generously provided by Dr. 268 Don McAlpine (NB Museum) as well as by Calum Ewing Gould V. E., 1936 - Nova Scotia mammal notes. Can. and Andrew Hebda (NS Museum) who also prepared Field-Nat., 50 (6): 103-104. Fig. 1. A mention needs to be made of Harry Bassett Graham R. E., 1962 - Porcupine cave dens in California. and Fred St. Peters who skillfully recovered decades of Cave Notes, Cave Research Associates, USA, notes and raw data from a set of corrupted computer 4 (1): 1-4. diskettes. Finally, thank you to the two anonymous Gunn J., Hardwick P. & Wood P.J., 2000 - The Invertebrate reviewers who critiqued the draft manuscript and made Community of the Peak-Speedwell Cave System, a number of very useful and constructive suggestions. Derbyshire, England - Pressures and Considerations for Fieldwork in 1996/97 was supported in part by a Conservation Management. Aq. Conserv., 10: 353-369. grant from the endowment fund of the Nova Scotia Holsinger J. R. 1980 - Stygobromus canadensis, a new Museum Board of Governors. Fig. 1 was generated subterranean amphipod crustacean (Crangonyctidae) from the NS Museum MIMS database. from Canada, with remarks on refugia. Can. J. Zool., 58(2): 90-97. Howarth F., 1972 - Cavernicoles in Lava Tubes on the REFERENCES Island of Hawaii. Science, 175 (4019): 325-326. Arsenault S. P., Schroeder J., Bérubé D. & Albert R., 1997 Howarth F., 1988 - Environmental Ecology of North - The caves of southeastern New Brunswick (Revised Queensland Caves. In: Pearson, L. (Ed.) - Preprints and Supplemented), Issue 97-7, Open Files, Minerals of papers for the 17th biennial conference, Australian and Energy Division, Dept. of Natural Resources and Speleological Federation Tropicon Conference, Lake Energy, New Brunswick, 33 p. Tinaroo, Far North Queensland, December 27-31, 1988. Banfield A.W., 1974 -The Mammals of Canada. National Jeannel R., 1926 - Faune cavernicole de la France, avec Museums of Canada, Univ. of Toronto Press, 438 p. une étude des conditions d’existence dans le domaine Bleakney J.S., 1965 - First Specimens of Eastern Pipistrelle souterraine. P. Lechevalier ed. Paris, 334 p. from Nova Scotia. J. Mamm., 46: 528-529. Jefferson G. T., 1981 - Diptera in British Caves. In: Bousfield E.L. & Holsinger J. R., 1981 - A second new Beck, B.F. (Ed.) - Proceedings of the 8th International subterranean amphipod crustacean of the genus Congress on Speleology, Bowling Green, , Vol. Stygobromus (Crangonyctidae) from Alberta, Canada. 1: 106-107. Canadian J. Zool., 59: 1827-1830. Juberthie C., 1984 - La colonisation du milieu souterrain:

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theories et methods, relations avec la speciation et Moseley M. & Hebda A., 2001 - Overwintering l’évolution souterraine. Mem. Biospéléol., 11: 65-102. Leiobunum elegans (Opiliones: Phalangiidae) in Caves Kane T. C. & Culver D. C., 1992 - Biological Processes in and Mines in Nova Scotia. Proc. N. S. Inst. Sci., Space and Time: Analysis of Adaptation. In: Camacho, 41 (4): 216-218. A. I. (Ed.) - The Natural History of Biospeleology, Moseley M., Klimaszewski J. & Majka C. J., 2006 Madrid, Monografias del Museo Nacional de Ciencias - Description of the pupa and observations on the Naturales: 423-451. distribution, ecology and life history of Quedius King L.H., 1996 - Late Wisconsin ice retreat from the spelaeus spelaeus Horn (Coleoptera: Staphylinidae) in Scotian Shelf. Bull. Geol. Soc. Am., 108: 1056–1067. Nova Scotia, Canada. Zootaxa, 1226: 61-68. Lee C. E., 2002 - Evolutionary genetics of invasive species. Peck S. B., 1988 - A review of the cave fauna of Canada, Trends Ecol. Evol., 17 (8): 386-391. and the composition and ecology of the invertebrate Majka C. J., Moseley M. & Klimaszewski J., in press fauna of caves and mines in Ontario. Can. J. Zool., - Gennadota canadensis (Casey) (Staphylinidae: 66: 1197-1213. Aleocharinae): new records, a range extension, and Sawatzky K., 1986 - Diving in Diogenes. Can. Caver, bionomic notes. Coleop. Bull. 18 (1): 16-20. Marusik Y. M. & Koponen S., 1992 - A Review of Meta Schmidt R. E., 1986 - Zoogeography of the Northern (Araneae, Tetragnathidae) with Description of Two New Appalachians. In: Hocutt, C.H. & E.O. Wiley (Eds.) Species. J. Arachnol., 20: 137-143. - The Zoogeography of North American Freshwater McAlpine D. F., 1976 - First record of the Eastern Pipistrelle Fishes. John Wiley & Sons, NY: 137-159. in New Brunswick. Can. Field-Nat., 90: 476. Schwert D. P., 1979 - Description and significance of McAlpine D. F., 1977 - Notes on Cave Utilization by a fossil earthworm (Oligochaeta: Lumbricidae) cocoon Beaver. Bull. Nat. Speleo. Soc., 39 (3): 90-91. from postglacial sediments in southern Ontario. Can. McAlpine D. F., 1979 - Preliminary Investigations on the J. Zool., 57 (7): 1402-1405. Solution Caves of New Brunswick. J. New Brunswick Scott F. 1979 - Preliminary Investigations at Hayes Mus., 1979: 99-107. Cave, Hants County, Nova Scotia in 1978. Cur. Rept. McAlpine D. F. & Reynolds J. W., 1977 - Terrestrial 38, N. S. Museum, Halifax, Canada, 14 p. Oligochaeta of Some New Brunswick Caves with Remarks Scott F. & Grantham R., 1985 - The Cave Environment. on Their Ecology. Can. Field-Nat., 91: 360-366. In: Morris, L (Ed.) - The Hayes Cave Site, South Moore K., 1963 - Hayes Cave: a study. Unpub. MS, N. Maitland, Nova Scotia. Cur. Rept. 50, N. S. Museum, S. Museum, Halifax, Canada, 21 p. Halifax, Canada: 101-115. Morris L. (Ed.), 1985 - The Hayes Cave Site, South Scott F. & Hebda A. J., 2004 - Annotated List of the Maitland, Nova Scotia. Cur. Rep. 50, N. S. Museum, Mammals of Nova Scotia. Proc. N. S. Inst. Sci., Halifax, Canada, 128 p. 42 (2): 189-208. Moseley M., 1975 - The Protection of Caves in Nova Shaw P. & Davis M., 2000 - Invertebrates from Caves on Scotia. Nova Scotia Speleo. Soc. Newsletter, 5: 1-23. Vancouver Island In: Darling, L. M. (Ed.) - Proceedings N. S. Museum, Halifax, Canada. of a Conference on the Biology and Management of Moseley M., 1976 - Caves of the Atlantic Region. In: Species and Habitats at Risk, Kamloops, B.C., 15-19 Thompson, P (Ed.) - Cave Exploration in Canada. February, 1999, Vol. One. B.C. Ministry of Environment Canadian Caver, Univ. Alberta, Canada: 8-18. Lands and Parks, Victoria, B.C. and Univ. College of Moseley M., 1996 - The gypsum karst and caves of the the Cariboo, Kamloops, B.C: 121-124. Canadian Maritimes. Cave and Karst Sci., 23 (1): 5-16. Trevor-Deutsch B., 1973 - The role of hibernating Moseley M., 1998 - Invertebrate Fauna of Nova Scotia Caves. bats in the winter diet of Peromyscus spp. (Rodentia: Cur. Rept. 86, N. S. Museum, Halifax, Canada, 37 p. Cricetidae). Unpub. Ph.D. Thesis, Carleton Univ., Moseley M., in press - Records of Bats (Chiroptera) Ottawa, Canada. at Caves and Mines in Nova Scotia. Cur. Rept., N. Wright B., 1979 - Mites, Ticks, Fleas and Lice in the S.museum, Halifax, Canada. Nova Scotia Museum and Acadia University Museum Collections. Proc. N. S. Inst. Sci., 29: 185-196.

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 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: º = Calder & Bleakney, 1965; ¹ = Calder & Bleakney, 1967; ² McAlpine & Reynolds, 1977; ³ = Scott, 1979; ª = McAlpine, 1979; * = Moseley & Hebda, 2001; ** = Moseley et al., 2006; *** = Wright, 1979; ^ = McAlpine, pers. comm.]

CLASS or FAMILY Habitat collected Seasons recorded Distribution ORDER TAXON main Sites th. d.th. d.z. DJF MAM JJA SON & ecology substrate(s) Ciliata Fam. & spp. indet + + + pools, etc. Aq ?

Turbellaria Planaridae CB, PT, GM, Sp. A indet. + pools + + Aq ? KC

Sp. B indet. + pools + + KC Aq ?

Oligochaeta Lumbricidae Dendrodrilus rubidus various + + + + + + Tr, Cp HT (Savigny) organic

Aporrectodea various + + + + + + Tr, Ad HT tuberculata (Eisen) organic

Lumbricus terrestris various CB, GM², + + + + Tr, Ad HT L. organic GP²

Eisenia rosea + + + cave soil + + PT,HO² Tr, Ad HT (Savigny)

Enchytraeidae various Spp. indet + + + + + + + All/most Tr GP organic

Spp. indet. + + + pools, etc. + + + + All/most Aq TP

Naiidae Spp. indet. + pools, etc. + + + + FC Aq TP

Hirudina Glossiphonidae Helobdella papillata + pool + BB Aq, Ne AC (Moore) Cladocera Fam indet. Sp. A indet. + pool + CB Aq AC

Sp. B indet. + pool + CB Aq AC

Ostracoda Candonidae Pseudocandona + + stream, pool + + WB,CC Aq TP albicans (Brady)

Fabaeformiscandona + pool + CC Aq, Cp TP wegelini (Petkovsky) Cypridae Cypria sp. + stream + WB Aq HT

Cavernocypris sp. + stream + WB Aq TP

Cypridopsidae Cypridopsis sp. + stream + WB Aq HT

Copepoda Cyclopoidae Acanthocyclops + pool, sediment + + CB Aq, Cp TP robustus (Sars)

A. brevispinosus + stream + WB Aq, Ne TP (Herrick)

A. venustoides Coker + pool with dung + PT Aq, Ne AC

Eucyclops agilis + pool + CB Aq, Cp TP (Koch)

Diacyclops + stream + + WIC,WB Aq, Cp TP brachycerus (Kiefer)

Paracyclops poppei + pool + CB Aq, Cp AC (Rehberg)

Macrocyclops albidus + sediment + CB Aq, Cp TP (Jurine)

Sp. indet. + sediment + WB Aq ?

Isopoda Oniscidae cave wall, Oniscus asellus (L.) + + + + + + Tr, Ad tTP (P) dung Pauropoda Fam. indet. Sp. indet. no data FC¹ Tr, ?

continued

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 16 Max Moseley

Chilopoda Henicopidae Lamyctes fulvicornis porcupine + + FC Tr, Ad AC Meinert dung

Diplopoda Blaniulidae Proteroiulus fuscus porcupine + + + + FC¹ Tr, Ad HT (Am Stein) dung

Julidae Ophyiulus pilosus porcupine + + + + FC¹ Tr, Ad HT (Newport) dung

Allajulus latestriatus + + + various + + FC¹,F2 Tr, Ad HT (Curtis)

Polydesmidae Polydesmus angustus porcupine + + + + FC Tr, Ad HT (P) Latzel dung Symphyla Symphylella sp. + cave soil + MB Tr TP

Scutigerellida Scutigerella sp. + on pool + KC Tr TP

Collembola Poduridae Hypogastrura + timber + KC Tr, Ne TP pseudarmata

Neanura muscorum + timber + KC Tr, Ad TP (Tempelton)

porcupine Friesea sp. + + GM Tr, ? AC dung

Willemia scandinavia + + + WIC Tr, Cp HT Stach porcupine

Onychiuridae Protaphorura + + + + + HC,WIC,FC Tr, Ne TP pseudarmatus porcupine

porcupine HC,FCº, P. armata (Tullberg) + + + + + + + Tr, Cp TP dung F2,GM

P. cf. boedvarssoni + on pools + + CC Tr, ? ? Pomorski

Tullbergia iowensis porcupine + + + + + + FCº Tr, Ne TP Mills dung

T. roseki Christiansen porcupine + + HC Tr, Ne HT and Bellinger dung

Isotomidae porcupine Folsomia fimitaria (L.) + + + + + + FCº Tr, Ad TP dung

F. stella Christiansen porcupine + + + + + HC,WIC,FC Tr, Ne TP & Tucker dung

dung, pool F. candida (Willem) + + + + + MIC, Tr, Ne TP surface

Isotoma notabilis no data FCº Tr, Ne HT Schäffer

I. caeruleatra Guthrie + plant litter + CB Tr, Ne AC

various Isotoma sp. nova? + + + + + HC,FC Tr TP organic

Entomobryidae Heteromurus nitidus + + + oligotrophic + + + Tr, Ad TP (Templeton)

Pseudosinella alba + + + on pools, dung + HC,FCº Tr, Ne TP (Packard)

P. collina Wray + timber + KC Tr, Ne TP

Entomobrya nivalis + no data FCº Tr, Cp AC (L.)

Tomoceros minor porcupine + + CB Tr, Ad TP (Lubbock) dung

Neelidae Megalothorax porcupine + + + + + + FCº Tr, Ne GP minimus Willem dung

Sminthuridae Arrhopalites nr. + oligotrophic + F2 Tr HT pygmaeus (Wankel)

oligotrophic & A. hirtus Christiansen + + + F2 Tr, Ne TP on pools

Sminthurides + wet plant litter + F2 Tr, Cp AC malmgreni (Tullberg)

Ptenothrix marmorata timber; on + + + + + CB, FCº, KC Tr, Ne HT (Packard) pool 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 17

Ephemeroptera Fam. et sp. indet (nymphs) + stream + HB^ Aq. ?

Odonata Aeshnidae Aeshna umbrosa + pools + HC Aq, Ne HT Walker nymphs

Aeshna sp. indet. + + pools + HC Aq, Ne HT nymphs

Cordulegaster + stream + CB Aq, Ne AC maculata Selys

Macromidae Macromia illinoiensis + + pools + HC Aq, Ne HT Walsh nymphs Plecoptera Capnidae Sp. indet. nymphs + stream + WB Aq AC

Leuctridae Sp. indet. adults + + + cave walls + WB, KC Aq HT

Nemouridae Amphinemura sp. + stream + WB Aq AC nymphs

Taeniopteryx sp. seeps & wet + + + FC, F2 Aq AC nymphs plant litter

Chloroperlidae Haploperla sp. + + + stream + + F2 Aq HT nymphs Perlidae Sp. indet. nymphs + stream + KC Aq HT

Orthoptera Ceuthophilus brevipes + + cave wall + + + TH,PT Tr, Ne, HT(P) (Scudder)

bait C. maculatus (Harris) + + WIC Tr, Ne, AC (molasses)

Coleoptera Dytiscidae Agabus semivittatus + stream CB Aq, Ne AC (Le Conte)

A. larsoni Ferry & + + + stream, pool + HC,DC Aq, Ne HT Nilsson

Dytiscus sp. indet. + stream + CB Aq, ? AC larva

Hydrophilidae Crenitis digesta + cave wall + F2 Tr, Ad AC (LeConte).

Staphylinidae Quedius s. spelaeus porcupine + + + + + + + Tr, Ne TP Horn larvae and dung

Q. mesomelinus + plant litter + KC Tr, Ad TP (Marsham)

Brathinus nitidus various + + FC¹, FH2 Tr, Ad TP LeConte organic

Gennadota bait + + WIC Tr, Ne TP canadensis Casey (molasses)

Atheta sp. indet. + cave wall + PT Tr, Ne ?

Curculionidae Sciaphus asperatus bait + + F2 Tr, Ad AC (Bonsdoff) (molasses)

Dermestidae Dermestus lardarius + cave wall + F2 Tr, Cp AC L.

Scarabaeidae Aphodius leopardus porcupine + + FC¹ Tr, Ne GP Horn dung

Latridae Corticaria serrata porcupine + + MIC Tr, Ad GP (Paykull) dung

Ptilidae Acrotrichis castanea porcupine + + + + + + F2 Tr, Ne GP (Matthews) dung Trichoptera Limnephilidae Sp. indet. adult + cave wall + DC, HB^ Tr, ? HT(P)

Lepidoptera Acrophilidae effrentella + cave wall + PT Tr, Ne AC Clem

Geometridae Triphosa haesitata cave wall Tr, Ne HT(P) affirmaria (Walker)

Xanthotype sospeta + cave wall + FC Tr, Ne AC (Drury)

Noctuidae Scoliopteryx libatrix + + cave wall + + Tr, Cl HT(P) L. GP,GR,HO continued

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 18 Max Moseley

Diptera: Tipulidae Limonia cinctipes Say + cave wall + WB Nematocera adult Tr, Cl TP(P) porcupine - ditto -- larva + + + + WIC, FC, F2 dung Dolichopeza sp. + cave wall + KC Tr, ? HT(P) sensu lato

Tipula sp. larva + veg. litter + F2 Tr, ? AC

Erioptera pilipes + veg. litter + CB Tr, Ne AC (Fabricius) larva

Trichoceridae porcupine Trichocera sp. indet + + WIC Tr, ? TP(P) dung T. maculipennis + + + cave wall + + + most sites Meigen adult Tr, Cp TP(P) porcupine - ditto -- larva + + + + most sites dung Culicidae Culex pipiens L. ♀ + + + cave wall + + CC, MC Tr, Cp HT(P)

C. restuans Theobald cave wall + + Tr, Ne HT(P) ♀

C. territans Walker ♀ cave wall + + Tr, Cp HT(P)

Anopheles sp. ♂ + cave wall + WB Tr HT(P)

Chaoboridae Chaoborus sp. + + various + + F2, WB Tr HT(P)

Chironomidae porcupine Smittia sp. adult, larva + + + FC¹, F2 Tr TP dung

Gen. et spp. indet various + + + FC, F2 Tr ? - adults organic

Gen. et spp. indet + + stream + + F2, WB, KC Aq ? -larvae 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)

HC, CC, PT, Exechia sp. + + + cave wall + + Tr HT(P) MC

Exechiopsis sp. + + cave wall + + CC, PT Tr HT(P)

Tarnania tarnania + + cave wall + PT Tr, Cl HT(P) (Dziedzieki)

Sciaridae porcupine Bradysia sp. adult + + + FC¹ Tr GP dung

porcupine Lycoriella sp. adult + + + CB, FC Tr GP dung

porcupine Scatopsciara sp. adult + + + CB, FC Tr GP dung dung, walls, Unidentified adults + + + + + most sites litter porcupine Larva - type A + + + + + + dung (P) - see porcupine identified adults Larva - type B + + + + HC, WIC, CC dung porcupine Larva - type C + + F2 dung 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 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 19

Diptera: Tabanidae Brachycera Chrysops sp. larva + veg. litter + CB Tr AC

Phoridae Megaselia no data FC¹ Tr, ?? TP meconicera (Specier)

Helomyzidae Scoliocentra fraterna no data FC¹ Tr, Ne HT(P) Loew

Helomyza serrata (L.) + + cave wall + + FC, CC, PT Tr, Cl HT(P)

H. brachypterna + cave wall + PT Tr, Ne HT(P) (Loew) ?

Amoebaleria sp. + + cave wall + + CC, PT Tr HT(P)

Tephrachlamys sp. + cave wall + F2 Tr HT(P)

Leptocera (Limosina) walls and + + + + + FC¹, F2 Tr, GP(P) sp. dung

Siphonaptera Ischnopsyllidae Myodopsylla insignis ex Myotis*** Tr PR (Rothschild) Hymenoptera Fam. indet* no data Tr ?

Psocoptera Liposcelidae Liposcelis + on pool + CC Tr AC bostrychophila Homoptera Aphidae Trama sp. + pitfall + CC Tr AC

Cixidae Gen. et sp. indet. + old fish bones + FH2 Tr AC

Acarina Parasitidae Parasitus sp(p). + dung, timber + + FCº, MC, KC Tr GP

Eugamasus sp. + + dung, timber + FCº, KC Tr TP

Vulgarogasmus sp. nr. porcupine + + Tr GP dung

porcupine Gen. et. spp. indet. + + + + + + most sites Tr GP dung

Vegaidae dung, plant Vegaia sp. + + + FCº, KC Tr TP litter

Zerconidae porcupine Zerconopsis sp. + FCº Tr GP dung

Ascidae porcupine Arctoseius sp. + + + + FCº, KC Tr GP dung

Ameroseidae porcupine Epicriopsis sp. + + + CC Tr GP dung

Eviphididae porcupine Alliphis sp. + + + + FCº Tr GP dung

Macrochelidae porcupine Geolapsis sp. + + FCº Tr GP dung Spinturnicidae Spinturnix sp. ex Myotis*** Tr PR

Macronyssidae Macronyssus crosbyi ex Myotis*** Tr PR (Ewing & Stover)

Uropodidae porcupine Gen. et sp. indet. + + + CB, WIC, FC Tr GP dung

Eupodidae Linopodes motatorius + timber + KC Tr, Cp TP (L.)

Cocceupodes sp. + timber + MC Tr TP

Rhagididae Rhagidia sp(p). + + + dung, timber + + + most sites Tr GP

Ereynetidae porcupine Gen. et sp. indet. + + CC Tr GP dung

Pygmephoridae porcupine Pygmephorus sp. + + FCº Tr GP dung

porcupine Bakerdania sp. + + FC Tr GP dung continued

International Journal of Speleology, 36 (1), 1-21. Bologna (Italy). January 2007 20 Max Moseley

Acarina Tetranychidae Bryobia praetiosa + porcupine dung + FCº Tr AC (Koch)

Gen. et. sp. indet. + + porcupine + HC, WB Tr GP dung, plant

Hydrachnoidea Gen et sp. indet. + stream + KC Aq ?

Glycyphagidae Glycyphagus + timber + MC Tr, Cp TP domesticus (De Geer)

Histiostmatidae Gen et sp. indet. + + porcupine dung + WIC, FC Tr GP

Acaridae Acarus immobilis + porcupine dung + FC Tr AC Griffiths

Banksinomidae Oribella sp. + porcupine dung + CC Tr GP

Araneida Nesticidae Nesticus cellulanus + cave wall + + FC Tr, Ad TP(P) (Clerck)

Tetragnathidae Meta ovalis (Gertsch) + + + cave wall + + + + most sites Tr, Ne TP(P)

Dictynidae Circurina brevis + scree + WIC Tr, Ne HT (Emerton)

Linyphiidae Sisicottus montanus + porcupine dung + FC Tr, Ne AC (Emerton)

Grammonota sp. + porcupine dung + CB Tr, Ne AC

Opiliones Phalangidae Oligolophus tridens (C. scree + WIC, CC* Tr, Ad HT L. Koch)

Sclersomatidae Nelima elegans (Weed) + + cave wall + + + most sites* Tr, Ne HT(P)

Gastropoda Arionidae Arion subfuscus + cave wall + KC Tr, Ad HT(P) (Draparnaud)

A. circumscriptus group + cave wall + + FC, PT, DL, Tr, Ad HT(P) KC

Limacidae Deroceras laeve + + cave wall + F2 Tr, Cl HT(P) (Müller)

Helicidae Trichia hispida (L.) + cave wall + F2 Tr, Ad HT(P)

Zonitidae Zonitoides arboreus + + + cave wall + + FC, F2 Tr, Ne HT(P) (Say)

Endodontidae Discus catskillensis + timber + KC Tr, Ne TP Pilsbry

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 21

CLASS or Habitat collected Seasons recorded Distribution FAMILY TAXON Sites ORDER th. d.th. d.z. substrate DJF MAM JJA SON & ecology limestone Salmonidae Salvelinus fontinalis (Mitchell) + + + + + + + DC, KC Aq,Ne HT streams

Osteichthyes Cyprinidae Phoxinus eos (Cope) + pools HC³ Aq,Ne AC

Gasterosteidae Pungitus pungitus (L.) + + pools + + BB, HC³, FC Aq, Cl HT

Plethodontidae Plethodon cinereus (Green) + stream + MCI Am,Ne AC Amphibia Rana clamitans melanota pools, Ranidae + + + CB, HC³, FC Am,Ne AC (Rafinesque) streams Soricidae Sorex fumeus (Miller) + HOª Tr, Ne HT BB,CB,HC,WIC,FC,F2, Myotis lucifugus (LeConte) + + + + MIC,MC,PT,LCM,OV, Tr, Ne HT LSH,HO, HB, BC,

BB,CB,HC,FC,MIC,CC,LCM, M. septentrionalis (Trouessart) + + + + Tr, Ne HT GRM,OV,LSH,GM,GR,HO, HB,BC, Vespertilionidae CB,HC,FC,MIC,CC, GRM,GM, Pipistrellus subflavus (F. Cuvier) + + + + + Tr, Ne HT GR,KC,UL,TC Mammalia Eptesicus fuscus (Palisot de + + HC (sight record) Tr, Ne HT Beauvois) Lasionycteris + + PT (sight record) Tr, Ne AC noctivagans (Le Conte) Procyonidae Procyon lotor (L.) + + (tracks) CB, HC, FCº Tr, Ne HT Mustelidae Mustela vison Schreber + (scat) KCª Tr, Ne AC Castoridae Castor canadensis Kuhl + DC*, KCª Tr, Ne HT Peromyscus maniculatus Muridae + HC, GMª Tr, Ne HT (Wagner) 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: º = Calder & Bleakney, 1965; ³ = Scott, 1979; ª = McAlpine, 1979; * Sawatzky, 1986. See McAlpine (1979) and Moseley (in press) for sources for records of bats.]

New Brunswick Ontario NS + NB Taxonomic group (McAlpine 1979) (Peck 1988) (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