Hypogean Fauna

The history of zoological investigations at the Batu caves and elsewhere in Southeast Asia from the early 20th century to the present is reviewed. Although there were major surveys and Dark Cave at Batu became the most intensively biologically investigated cave in the region, until the 1970s there was little change in the prevailing view that tropical caves had little of speleobiological importance.  Key advances in speleobiology in the last half century that have changed this perception and drawn attention to tropical cave biology are briefly discussed, and their possible significance for the research potential of the Batu caves assessed.

The Batu caves are the only caves in the Malay Peninsula that are well investigated zoologically, and they are the most thoroughly sampled anywhere in Southeast Asia. However, the records have not been collated to provide a comprehensive overview of the fauna present. This issue is addressed here by presenting an authoritative checklist of all reported zoological taxa, together with ecological annotations and a comprehensive bibliography

Several caves and sinkholes where snow and ice persist well into the summer exist in Nova Scotia and New Brunswick. They were often used in the past as a source of ice or for cold storage, and are known locally as “ice caves” or “ice holes”.  They are not true ice caves in the speleological sense of the term because they do not contain permanent ice, but are similar. Woodville Ice Cave in Hants County, Nova Scotia, described here, is a particularly good example.  It is in an area of unusually well-developed gypsum-anhydrite karst. Invertebrates and bats recorded from such sites are listed, and the possibility of psychrophilic fauna is suggested.

It is proposed that the ecological concept of an ecotone (a zone of steep environmental gradient between adjacent ecological communities or ecosystems) could be applied to caves in their entirety. Arguments are given in support of this proposal, and some of the conceptual, theoretical and empirical implications are highlighted and briefly discussed.

Although Dark Cave (Batu Caves, Selangor) is biologically the most intensively studied cave in the Malay Peninsula, ecological investigations are in their infancy and knowledge of species diversity is incomplete or sometimes inaccurate. A large volume of additional information will be required to informeffective conservation and management plans for this biospeleologically important site. The present paper presents a review of recent investigations of Collembola (springtails), Diplopoda (millipedes), and a general invertebrate survey. Attention is drawn to lessons that can be learnt from these studies.

Caves, as ecological units, are redefined in terms of the ecologically-significant scalar phenomena that, in contrast with such traditional criteria as perpetual darkness, environmental stability and oligotrophy, are the special characteristics of large cavities (macrocaverns) in rocks. When so defined, ‘caves’ are ecologically distinct from the mesocaverns and all other subterranean habitats. Some implications of this perspective for the ecological classification of animals found in subterranean habitats are briefly discussed.

The vertebrate and invertebrate fauna, environment and habitats of caves 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 subterranean fauna 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 includespecies 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 thatpost-glacial recolonisation of the subterranean habitat in Nova Scotia has been relatively delayed. Finally the general and regionalsignificance of the subterranean fauna is briefly discussed.

Millipedes are conspicuous and abundant in the Dark Cave, but there are few published records and thus uncertainty about the species composition of the fauna. There is also doubt about the ecologicalstatus and ecology of the three species reported from the cave (Plusioglyphyiulus grandicollis, Ascetophacus macclurei and Orthomorpha fluminoris). This paper reports some new records and field and laboratory observations that clarify some of these points.

The Isle of Man was heavily glaciated and thus all of its invertebrate fauna is the result of post-Pleistocene recolonization. Accordingly it presents the opportunity to observe an early stage of the active invasion and colonization of hypogean habitats on a maritime temperate island by epigean species. It also has many sea caves some of which have diverse invertebrate communities. This paper reports the results of a preliminary speleobiological survey of the island, which revealed an unexpectedly rich and diverse hypogean fauna.

Several caves and sinkholes where snow and ice persist well into the summer exist in Nova Scotia and New Brunswick. They were often used in the past as a source of ice or for cold storage, and are known locally as “ice caves” or “ice holes”. They are not true ice caves in the speleological sense of the term because they do not contain permanent ice, but are similar. Woodville Ice Cave in Hants County, Nova Scotia, described here, is a particularly good example. It is in an area of unusually well-developed gypsum-anhydrite karst. Invertebrates and bats recorded from such sites are listed, and the possibility of psychrophilic fauna is suggested.

Until the 1970s speleobiologists wrongly believed that troglobionts – highly adapted obligate subterranean animals – were rare or absent in tropical regions. This error probably held back the development of tropical cave biology for several decades. It is argued in this essay that the mistake originated  in the late 1890s during an investigation of Batu caves by the prestigious British Association for the Advancement of Science. Part 1 of this paper describes these events in the context of the development of speleobiology during the 19th century.

Caves, as ecological units, are redefined in terms of the ecologically-significant scalar phenomena that, in contrast with such traditional criteria as perpetual darkness, environmental stability and oligotrophy, are the special characteristics of large cavities (macrocaverns) in rocks. When so defined, ‘caves’ are ecologically distinct from the mesocaverns and all other subterranean habitats. Some implications of this perspective for the ecological classification of animals found in subterranean habitats are briefly discussed.

Early records of aquatic mollusca (Potamopyrgus antipodarum, Ancylus fluviatilis and Dressensia  polymorpha) found in underground water pipes in England are reviewed and discussed. It is possible that if searched for these species will be found in natural caves in Britain.

[FULL TEXT] Boettgerilla pallens is an Eastern European slug that has spread rapidly through North West Europe during the last few decades. It was first found in Britain in 1972 and is now widespread here, though local: mostly in parks, gardens, quarries and similar places.

On 12th September 2000, a specimen was collected in Warton Quarry Mine, Warton, Lancashire (see Northwestern Naturalist 2 (2): 6 (2000)). The animal was crawling on damp bare limestone in the "dark zone" well inside the mine.

It is not unusual to find slugs in mine tunnels and caves: various species have been collected from the threshold regions of such sites. I have for example found the common species Arion fasciatus in cave entrances both in this country and in North America (where it is introduced). It is rare though for slugs to be found well into the dark zone. Whilst it is dangerous to draw conclusions from the find of a single possibly stray individual, it is interesting to see B. pallens in such an unusual habitat and it is perhaps permissible to wonder if this expansive species might prove able to move into and exploit caves and ecologically cavelike habitats such as abandoned mines.

The specimen was identified by Dr. D. Davis, Nova Scotia Museum, Halifax, Canada and has been deposited in the Natural History Collection of that institution (NSMNH 15954).

Although Dark Cave (Batu Caves, Selangor) is biologically the most intensively studied cave in the Malay Peninsula, ecological investigations are in their infancy and knowledge of species diversity is incomplete or sometimes inaccurate. A large volume of additional information will be required to informeffective conservation and management plans for this biospeleologically important site. The present paper presents a review of recent investigations of Collembola (springtails), Diplopoda (millipedes), and a general invertebrate survey. Attention is drawn to lessons that can be learnt from these studies.

The cave-associated invertebrates of Nova Scotia constitute a fauna at a very early stage of post-glacial recolonization. The Coleoptera are characterized by low species diversity. A staphylinid Quedius spelaeus spelaeus, a predator, is the only regularly encountered beetle. Ten other terrestrial species registered from cave environments in the province are collected infrequently. They include three other rove-beetles: Brathinus nitidus, Gennadota canadensis and Atheta annexa. The latter two together with Catops gratiosus (Leiodidae) constitute a small group of cave-associated beetles found in decompositional situations. Quedius s. spelaeus and a small suite of other guanophiles live in accumulations of porcupine dung: Agolinus leopardus (Scarabaeidae), Corticaria serrata (Latrididae), and Acrotrichis castanea (Ptilidae). Two adventive weevils Otiorhynchus ligneus and Barypeithes pellucidus (Curculionidae) collected in shallow cave passages are seasonal transients; Dermestes lardarius (Dermestidae), recorded fromone cave, was probably an accidental (stray). Five of the terrestrial beetles are adventive Palaearctic species. Aquatic beetles arecollected infrequently. Four taxa have been recorded: Agabus larsoni (Dytiscidae) may be habitual in regional caves; another Agabus sp. (probably semivittatus), Dytiscus sp. (Dytiscidae), and Crenitis digesta (Hydrophilidae) are accidentals. The distribution and
ecology of recorded species are discussed, and attention is drawn to the association of beetles found in a Nova Scotia “ice cave”.

The opilionid L eiobunum elegans was found overwintering in caves and mines in mainland Nova Scotia and Cape Breton island. it occurred both as adults and near-adults, usually as single individuals or small clusters but aggregations were formed at one site. It is suggested that aggregation formation is a behavioural adaptation for coping with saturated atmospheres. This is the first published report of overwintering opilionids in such habitats from eastern Canada.

Langness Seacave (Langness peninsula, Isle of Man) contains an invertebrate faunal assemblage dominated by cavernicoles (troglophiles) yet dependent upon marine driftweed as the primary source of food. The biospeleological significance of this unusual site is discussed, and arguments are offered for the possibly subterranean nature of two species Itunella tenuiremis (Copepoda: Canthocamptidae) and Trechus fulvus (Coleoptera: Carabidae)

The woodlouse Trichoniscoides saeroeensis was collected from a mine adit near the shore at Langness, Isle of Man. The record supports the contention that the species is cavernicolous in the British Isles. Field evidence points to it being exceptionally tolerant of water, and this may partly explain its ability to survive in subterranean habitats. These observations add weight to the contention that the species is an example of a coastal invertebrate that is expanding its range by colonising the hypogean, and evolutionary route that might have been followed by some other terrestrial cave invertebrates.

The troglophilic staphylinid beetle Quedius spelaeus spelaeus Horn 1871, has been found in anumber of porcupine dung caves in Nova Scotia where it appears to be the dominant predator onother invertebrates. In culture, late-instar larvae were observed to excavate and remain in cavities excavated in dung, and to pupate in these cavities. The pupa is described for the first time andcompared with other pupae in the genus Quedius Stephens. The apparently disjunct distribution ofthe species in Nova Scotia is discussed and it is suggested that it may have colonized the provincefrom Atlantic glacial refugia.

Pseudoparonella doveri is an unusual troglomorphic collembolan that appears to have no close epigean relatives. It was previously only known from specimens collected in The Dark Cave, Batu Caves, Malaysia almost 70 years ago. The species was rediscovered in 2003 on dripstone/gour pools in the same cave, and a series intended for taxonomic reassessment collected.  A brief qualitative survey of other habitats in The Dark Cave was also carried out to improve our knowledge of the little-known collembolan fauna of this important speleobiological site.

Several caves and sinkholes where snow and ice persist well into the summer exist in Nova Scotia and New Brunswick. In the past they were sometimes used as a source of ice or for cold storage, and are known locally as ‘ice caves’ or ‘ice holes’. Although they are not true ice caves in the speleological sense of the term because they do not contain perennial ice, they are very similar. Woodville Ice Cave in Hants County, Nova Scotia, described here, is a particularly good example. Invertebrates and bats recorded from such sites are briefly discussed and the possibility of finding psychrophilic fauna in them is suggested.Plusieurs grottes et dolines sont présentes là où la neige et la glace ne disparaissent que tard durant l’été en Nouvelle‑Écosse et au Nouveau-Brunswick. Autrefois, elles étaient parfois utilisées comme sources de glace ou aux fins d’entreposage sous froid. À l’échelle locale, elles sont connues sous le nom de glacières ou de puits de neige. Elles ne sont pas de vérit...

The new genus Titanobochica is described for Titanobochica magna sp. nov. from caves of the Algarve karstic massif, in Portugal. The new genus is assigned to the family Bochicidae and its particular characteristics and geographical isolation suggest a relictual condition. A key to the genera of Bochicidae is provided. The composition of the cave-dwelling fauna of the Algarve province is also discussed.

[FULL TEXT] On May 9th, 1999 during a visit to Warton Quarry Mine, Warton Crag, Lancashire (NGR SD 4887.7246 Alt. 88 m) a single moth pupa was found lying on the upper surface of a piece of damp rotting mine timber in the 'deep threshold' of the entrance adit (i.e. the twilight zone beyond the innermost limit of green plants). The specimen was collected and reared: the adult emerged on May 26th and was identified by Steve Garland (Bolton Museum) as the Small Angle Shades Euplexia lucipara (L.) (Noctuiidae). Warton Quarry Mine is an abandoned late Victorian working in limestone and ecologically similar to a natural cave: I am not aware of any previous records of this common moth from any cave or mine.

The caterpillars of E. lucipara feed on ferns, so it is likely that the pupa was from a population feeding on the ferns which grow in the entrance of the adit just a few meters away. According to Heath and Emmet (The Moths and Butterflies of Great Britain and Ireland, Vol. 10, 1983 p. 166) the pupae overwinter in the ground, in a firm cocoon of earth and silk. This specimen was exposed, in a position where, had it not been underground, it would have easily been visible to predators; and there was no cocoon.

The specimens have been deposited in the Lepidoptera collection of Bolton Museum.

The opilionid Leiobunum elegans was found overwintering in caves and mines in mainland Nova Scotia and Cape Breton Island. It occurred both as adults and near-adults, usually as single individuals or small clusters, but aggregations were formed at one site. It is suggested that aggregation formation is a behavioural adaptation for coping with saturated atmospheres. This is the first published report of overwintering opilionids in such habitats from Eastern Canada.

Caves, as ecological units, are redefined in terms of the ecologically-significant scalar phenomena that, in contrast with such traditional criteria as perpetual darkness, environmental stability and oligotrophy, are the special characteristics of large cavities (macrocaverns) in rocks. When so defined, 'caves' are ecologically distinct from the mesocaverns and all other subterranean habitats. Some implications of this perspective for the ecological classification of animals found in subterranean habitats are briefly discussed.

information on museum collections, programs, procedures and research accessible to interested readers. This report contains the preliminary results of an on-going research program of the Museum. It may be cited in publications, but its manuscript status should be clearly noted.