BONA ABIGAIL B.

HILARIO
2008-81298
ENRM 230

REVIEW PAPER
Functions of Caves in a Karst Landscape: Patch or Corridor?

ABSTRACT - Understanding the karst landscape is important in the conservation and management
strategies of caves. Whatever changes and disturbances in the landscape can affect the dynamics of
the caves. This review paper attempts to answer the functions of caves in a landscape perspective,
whether they are a patch or a corridor. Studying through available materials, the author concludes
that caves are habitat patches because they have a well-defined boundary and inhabit diverse types
of species. However, caves are corridors for two cases. Trogloxenes and troglophiles can move in
and out of the caves if caves are simple, short and with an exit passage. For long and complex with
twilight and deep zones, only troglobites or stygobites with their adaptation of bioluminescence or
enlarged sensory systems are able for subterranean dispersal. The author recommends that caves
should be studied in a landscape perspective with an emphasis on the patch-corridor-matrix model
because of its effect on the energy flow and movement of individuals.

INTRODUCTION

Karst landscapes comprise approximately 15% of the earth’s surface (Culver & Pipan, 2009). Caves are
found throughout the world, but only a small portion of them have been explored and documented.
They are often described as one of the harshest and unforgiving environments on earth because of dark
habitats with very little food. Caves are often part of a karst landscape formed from the dissolution of
soluble rocks such as limestone, dolomite, and gypsum.

Each cave is unique according to form and geological and geographical location (Gamble, 1981).
Although cave ecosystems and habitats are poorly sampled when compared with surface ecosystems
(Niemiller & Zigler, 2013), studies have shown that caves comprise unique biodiversity and a
taxonomically diverse fauna has been documented from these habitats. For example, more than 1,138
cave-restricted species and subspecies from 112 families and 239 genera have been described in the
United States alone. Because of the fragile nature of the cave ecosystem, and the fact that cave regions

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tend to be isolated from one another, caves harbor a number of endangered species, such as the tooth
cave spider, liphistius trapdoor spider, and the gray bat.

The different animals that consider caves as home are adapted to the natural extremes of the caves, a
life in perpetual darkness and generally limited food resources. They have developed traits such as loss
and reduction of eyes and pigmentation, elongation of appendages, increased longevity, and
enhancement of nonvisual sensory modalities (Niemiller & Zigler, 2013).

These cave faunas are categorized as trogloxenes, troglophiles and troglobites or stygobites generally
referred to as cavernicoles by speleologists. Trogloxenes are animals that live in caves during part of
their life cycles, but come out to the surface to forage or migrate. The most notable trogloxenes are
bats and cave crickets. Troglophiles (cave-loving) are animals that may complete their entire life
cycle within caves but can survive equally well in surface habitats. These include some spiders,
crickets, beetles, and salamanders. Troglobites (terrestrial) and stygobites (aquatic) are species that
spend their entire lives in the dark zone of caves. They mostly display albinism and are blind but their
smell and hearing are acutely developed. A remarkable example is the olm, an aquatic salamander.

Karst cave systems have three different terrestrial zones used as habitat by cavernicoles. The
“entrance zone” which is 0-10m from the entrance of the caves is located at the littoral opening and
is the most influenced by surface conditions. The “twilight zone” extends from the entrance to mid -
depth with a decreasing light and connectivity to the exterior. The “deep zone” is isolated from the
surface with no light, low temperature, and 100% humidity.

This review paper will attempt to discuss the author’s hypotheses on the functions of caves from a
landscape ecological perspective. A landscape is defined as a heterogeneous area composed of
elements or spatial components which include a patch, a corridor, and a matrix. Patch is a relatively
discrete areas or periods of relatively homogeneous environmental conditions, with boundaries
distinguishable or relevant to the organism or ecological phenomenon under consideration. While a
corridor is defined as an elongated patch that connects other patches together thereby increasing the
movement of organisms within a landscape. This is important especially for mobile organisms because
they need greater space for their physiological activities such as feeding, dispersal or migration.

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Hypothesis 1: Cave ecosystems function as a patch because they support a variety of species and
connected with other nearby ecosystems. They are either a source or a sink patch for species.

Hypothesis 2: Caves can be used as corridors by trogloxenes and troglophiles if caves have long
tunnels/passages with entrance and exit zones.

Karst landscapes diagram

Published: 08 October 2012
Copyright: Bruce W. Hayward. From Karst in Stone, Bruce W.Haywood & Jill A.Kenny. Geoscience Society of NZ Guidebook
No. 15.w

DISCUSSION

Biologists have traditionally-regarded caves as environments whose boundaries are well defined
and within which the resident flora and fauna can be assessed, their trophic relationships defined
and the flows of mass and energy measured (Gillieson, 1996).

The different terrestrial zones of caves determine the composition and distribution of species. The
outside environment with its terrestrial community or water flowing in and out of the caves

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influences the ecology of the caves, its species composition and abundance. The entrance zone can
serve as an ecotone between the cave and the surface ecosystem. Vegetation like ferns abundantly
grows near cave entrances which influences the cave biota.

The entire epigean (above the ground) environment and cave entrance show the higher habitat diversity
compared to the other regions in the cave. In a study conducted by Prous et al (2015) about arthropod
species composition and spatial distribution inside a Brazilian limestone cave, after 30 m away from the
entrance, habitat diversity showed a decline, reaching the lowest values starting from 40 meters. They
further concluded that cave entrances are important for the cave ecosystem as a whole, considering
that a considerable amount of trophic resources are transferred from the epigean to the hypogean
environment through this area.

Other faunas which are considered opportunistic are found at the entrance zone and use the caves
temporarily. These include the deer, bears, wolves and small mammals which use the caves for birth
and shelter and shield from predators. Bird species of dippers, thrushes, and swallows utilize the
caves for feeding and nesting. Caves with aquatic habitats support fishes, and amphibians. Many
species of algae, mosses, and cyanobacteria also inhabit caves.

According to Gillieson (1996), nutrients are generally limited in cave systems and thus utilize external
sources of energy and nutrients. These are through the foraging efforts of trogloxenes (e.g. crickets,
bats, etc.) and troglophiles (e.g. beetles, spiders, etc.), as well as because of floods, air transport and
other environmental processes. Trogloxenes and troglophiles provide a link between surface and
subterranean ecosystems (Rector, 2009).

To establish the function of caves as a corridor in the landscape, it is important to study its connectivity
and opportunities for dispersal. Niemiller & Zigler (2013) who studied patterns of cave biodiversity and
endemism presumed that areas of high cave density have higher connectivity between caves because of
the greater opportunities for dispersal. If caves have poor connectivity with other habitats, species
diversity and abundance are likely to be low. Dispersal may decrease extinction rates and differences in
species composition among localities or regions so corridors are important. A cave is either a strip
corridor because it contains a patch interior environment in which interior species may migrate or live or
a network because of its intersection and passages.

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However, the actual geometry of caves varies widely from a few meters in length to the 652-km-long
(405 mi) Mammoth Cave in Kentucky, USA, the longest cave ever recorded. With the twilight and
deep zones, caves as a corridor for movement and dispersal of cavernicoles into other ecosystem is
limited to certain circumstances.

Among the cavernicoles, troglobionts and stygobionts display the highest endemism in caves studied.
Two obvious explanations are drawn out by Christman, et al (2005). First is because of the reduced
opportunities for dispersal afforded cave animals. Second is not from the lack of dispersal, but from
subterranean dispersal followed by speciation. After the initial invasion of a cave, the isolated cave
species may occasionally move to nearby caves or habitats.

CONCLUSION

Caves as patches
For the first hypothesis, the author concludes that caves are habitat patches because caves are
discrete areas with relatively homogeneous environmental conditions and with boundaries relevant to
the organism it inhabits. They are ecosystems in a karst landscape and are home to diverse species
inhabiting the entrance zone, the twilight zone and the deep zone. However, there is no sufficient
data available to be able to establish if caves are a source patch or a sink patch.

Caves as corridors
For the second hypothesis, the author has derived two conclusions. First is that if caves are simple,
short in length and with an exit passage, they can be used as corridors by trogloxenes and
troglophiles for dispersal in and out the cave. Absence of light in complex and long caves can hinder
their movement. On the other hand, if caves are very long and complex with twilight and deep
zones, they can be used as corridors by troglobites for subterranean dispersal. Even in the absence
of light, troglobites with their adaptation of bioluminescence or enlarged sensory systems can move
from one ecosystem to another ecosystem inside the cave. For example, an amphibian troglobite
can move from an epigean ecosystem to a subterranean river inside the cave.

Caves are important ecosystems that provide a home to rare and endemic species which in turn provide
significant ecosystem services. Bats, for example, provide insect control, seed dispersal, and pollination.

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Its unique biodiversity makes them of special interest in conservation biology. However today, caves are
explored more for tourism and industrial purposes such as mining and quarrying. Just like all other
ecosystems, humans can alter the caves directly or indirectly, disturbing and threatening the
trogloxenes, troglophiles and the troglobites. Because of the disturbance, some species especially
the rare troglobites may be forced to come out of hiding and are unlikely to survive outside the
caves. Serious damage not only affects the cave biology but also its geology as stalactites and
stalagmites are cut to give way to visitors. Human disturbance at caves is a persistent problem
internationally and has been documented as a major cause of the decline of cave-dependent bats
(Barbour & Davis, 1969)

Understanding the karst landscapes is important in the conservation and management strategies of
caves. Changes and impacts of anthropogenic nature, such as deforestation, residential
construction, material deposits, dams, shrines, tourist infrastructure and even scientific research
(e.g., paleontological and archaeological excavations) in the landscape can affect the dynamics of
the caves (Prous, et al 2015). Therefore, it is recommended that caves should be studied in a
landscape perspective with an emphasis on the patch-corridor-matrix model because of its effect on
the energy flow and movement of individuals. Because of the subterranean connections, protection
of cave biodiversity should be on a landscape level.

LITERATURE CITED

Barbour, R.W. & Davis, W.H. (1969). Bats of America. University Press of Kentucky, Lexington.

Christman, M.C., Culver, D.C., Madden, M.K., & White, D. (2005). Patterns of endemism of the eastern
North American cave fauna. Journal of Biogeography (J. Biogeogr.), 32, 1441–1452.

Culver, D.C. & Pipan, T. (2009). The Biology of Caves and Other Subterranean Habitats. Oxford
University Press.

Forman, R.T.T & Godron, M. (1981). Patches and Structural Components for a Landscape Ecology.
BioScience, 31(10), 733-740.

Gamble, F. M. (1981). Disturbance of underground wilderness in karst caves. International Journal of

Environmental Studies, 18(1), 33-39.

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Gillieson, D. (1996). Caves: Processes, Development and Management. Blackwell Publishers Ltd.

Niemiller, M.L. & P Zigler, K.S. (2013). Patterns of Cave Biodiversity and Endemism in the Appalachians
and Interior Plateau of Tennessee, USA. PLoS One, 8(5).

Prous, X. & Ferreira, R.L. (2015). The entrance as a complex ecotone in a Neotropical cave. International
Journal of Speleology, 44 (2), 177-189.

Rector, M.A. (2009). Foraging in the Cave Environment: The Ecology of the Cave Spider (Meta ovalis).
Evolution, Ecology, and Organismal Biology Graduate Program. The Ohio State University.

ADDITIONAL REFERENCES

Alaska Wildlife Action Plan 2015. Retrieved November 12, 2016, from
https://www.adfg.alaska.gov/static/species/wildlife_action_plan/appendix5_karst_cave_habitats.pdf

Models of Landscape Structure. Retrieved November 12, 2016, from

http://www.umass.edu/landeco/teaching/landscape_ecology/schedule/chapter7 models.pdf

Romero, A. (2009). Cave Biology: Life in Darkness. Cambridge University Press.