Biological Conservation 211 (2017) 45–50

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Biological Conservation

journal homepage: www.elsevier.com/locate/bioc

Conservation relevance of bat caves for biodiversity and ecosystem services

Rodrigo A. Medellin a,⁎, Ruscena Wiederholt b, Laura Lopez-Hoffman c,d
a
Instituto de Ecología, Universidad Nacional Autónoma de México, Ap. Postal 70–275, 04510, Mexico
b
Everglades Foundation, 18001 Old Cutler Road, Suite 625, Palmetto Bay, FL 33157, USA
c
School of Natural Resources & the Environment, The University of Arizona, Tucson, AZ 85721, USA
d
Udall Center for Studies of Public Policy, The University of Arizona, Tucson, AZ 85721, USA

a r t i c l e i n f o a b s t r a c t

Article history: All ecosystems are dotted by salient small natural features that not only characterize them but also significantly
Received 11 August 2016 add to their biodiversity and functions. These small natural features are prominent but easily missed when eco-
Received in revised form 14 January 2017 systems are described. Caves are one key example of this. Cave ecosystems are underrepresented in conservation
Accepted 19 January 2017
planning and implementation around the world and have become mostly overlooked in conservation strategies
Available online 1 February 2017
overall. Caves contain high levels of biodiversity from fungi to invertebrates to vertebrates. This paper empha-
Keywords:
sizes bat caves as providers of ecosystem services to vast areas surrounding them, in the order of hundreds of
Bats thousands of square km just in North America. Their influence extends three-dimensionally via subterraneous
Bat caves water bodies and via the aerial nightly dispersal of the bats that provide a host of services from seed dispersal
Ecosystem services to pollination to pest control. The examples used focus primarily on free-tailed bats in North America, but the
Small natural features same principles apply to any other cave in the world with significant bat colonies. Caves enjoy protection, legal
Cave conservation or actual, in some countries and not in others, and as a result many have suffered damage or been destroyed al-
together. Common threats are vandalism, urbanization, and pollution. Many caves are attractive as ecotourism
destinations and provide unique opportunities to educate the public about unexpected biodiversity values and
ecosystem services. Inventorying caves poses challenges, but efforts are under way to assess caves in need of pro-
tection. Incipient cave protection strategies include legal and educational efforts, and management. Although il-
lustrated with bat caves, given the importance of all caves and their precarious status, it is time to call the
attention of decision makers about the urgent need to launch a worldwide cave conservation initiative.
© 2017 Elsevier Ltd. All rights reserved.

1. Introduction dissolution or aeolic erosion in that type of rock; 3) volcanic caves:

1.1. Defining caves
The definition of a cave has been the subject of significant debate.
Their origin, configuration, scale, contents, and other issues have been
considered, and no single definition has been accepted (Curl, 1964;
Culver and White, 2004). For purposes of this paper, caves are defined
as subterranean cavities in a variety of strata, harboring significant
levels of biodiversity. Most of our discussions deal with bat caves,
given the very strong association between the two, although the impor-
tance of caves is also illustrated through other biodiversity. Because of
this focus, we define bat caves as those that are customarily used by
bats as roosts. Many substrates can contain caves, but for practical pur-
poses of the discussions in this paper we refer to three major substrates
as containing caves (Culver and White, 2004): 1) karst dissolution
caves: underground spaces with an opening to surface, in limestone or
dolomite; 2) sandstone caves: also underground spaces, formed by
⁎ Corresponding author.
E-mail address: medellin@iecologia.unam.mx (R.A. Medellin).
lava tubes, bubbles, or vents. We do not include in this paper sea or gla-
cial caves. We illustrate the value of bat caves focusing on caves harbor-
ing Mexican free-tailed bats (Tadarida brasiliensis mexicana) given the
relatively robust amount of literature on them, but the same is applica-
ble to many other caves across the world, in temperate and tropical
areas containing many species of bats. Although the regional impact of
these caves in other areas of the world has not been modeled, evidently
the principles herein described are clearly applicable to all other caves
with significant bat populations.
1.2. Why are caves important?
We have all seen them, from giant trees to small ponds and streams,
to kopjes, boulders, creeks or caves. But these small natural features,
sites with ecological importance that is disproportionate to its size
(Hunter, 2017) are rarely if ever considered when an ecosystem is de-
scribed. An extreme case of these overlooked but deeply influential fea-
tures are caves. Caves are relatively small to tiny landscape features that
dot huge regions of the world. Few cave maps are available and propor-
tionally, considering the hundreds of thousands of caves in the world,

http://dx.doi.org/10.1016/j.biocon.2017.01.012
0006-3207/© 2017 Elsevier Ltd. All rights reserved.
46 R.A. Medellin et al. / Biological Conservation 211 (2017) 45–50
caves are considered protected areas only in some areas such as Europe,
Puerto Rico, certain caves in Canada, certain states of the U.S.A.,
Australia, the Philippines, and few more (e.g. Fitzsimons and Ashe,
2003; Boulton et al., 2003; Whitten, 2009; Zhalov, 2015). Given that
most conservation assessments and landscape analyses are based on
vegetation cover, forest fragmentation, and other above-ground ecosys-
tem features, subterraneous ecosystems tend to be underrepresented or
ignored in conservation plans and protected areas (Sugai et al., 2015).
Caves are reservoirs of exceptional levels of endemic and threatened
species (Deharveng, 2005), very high levels of genetic uniqueness
(Gibert and Deharveng, 2002), and they harbor unconventional taxa
such blind fish, crustaceans, insects, worms, and many other groups
(Gunn, 2004). In just the contiguous 48 states of the U.S., at least 927
species in 96 families are troglobites (obligate cave dwellers), but only
4% of those species enjoy any level of federal protection (Culver et al.,
2000). Lack of adequate consideration of cave ecosystems in biodiversi-
ty conservation efforts has been pointed out repeatedly (Langer, 2001,
Sugai et al., 2015).
The influence of caves on the surrounding ecosystems is also large.
Given the numerically enormous concentrations they form inside,
once bats emerge they must travel long distances in order to find
food. The movements of bats are associated with their predation pres-
sure on insects, particularly insect pests. A very conservative estimate
of nightly movements of 50 km from the cave roosts of Mexican free-
tailed bats was published by Williams et al. (1973). These bats are
able to easily fly much farther distances, given that they are long-
distance migrants and the fact that they are the fastest flying animals
(horizontally) on earth at 160 km/h (McCracken et al., 2016). If we
adopt 50 km as a benchmark, then the area of influence of bats around
each of their caves can be conservatively estimated at about 7850 km2
(Fig. 1). Applying this to all 29 Mexican free-tailed bat roosts listed in
Table 1, another underestimate given that only a fraction of the bat
caves are known in this region, the total area influenced by these 19
small natural features in the landscape amounts to almost 150,000 km2.
Fig. 1. The 29 roosts (mostly caves) of Mexican free-tailed bats known to date, with their conservative 50-km radius buffer of influence area. Total area covered by these caves adds up to
almost 150,000 km2. The known Winter regions are highlighted in purple (western Mexico), red (central Mexico), blue (Eastern Mexico), and orange (southern Mexico).
Data from Wiederholt et al., 2015.
Although many people associate all bats with caves, only about half
of Mexico's 138 bat species live in caves (Arita, 1993, Medellin et al.,
2008). In other countries the percentage varies; for example, 77% of
the bat species in China live in caves (Luo et al., 2013), 80% of those in
Puerto Rico (Ladle et al., 2012), and 23 out of the 33 bat species in
Bohol island in the Philippines live in caves (Phelps et al., 2016). Other
bats live in large old trees, or in rocky outcrops (Lindenmayer, 2017;
Fitzsimons and Michael, 2017), in tents built by them out of large leaves
in tropical forests (Rodríguez-Herrera et al., 2007), and in a variety of
other natural and anthropogenic roosts (Fenton and Simmons, 2014).
Bats form some of the largest warm-blooded vertebrate concentrations
in the world and their biology is widely diverse (Medellin et al., 2008).
By focusing only on a single, fairly well-known species, the Mexican
free-tailed bat and the ecosystem services they provide, the importance
of caves for ecosystem functioning and for human well-being becomes
obvious. Mexican free-tailed bats suppress pests on important agricul-
tural crops such as cotton and corn in Mexico and the United States
(López-Hoffman et al., 2014) and also provide extensive ecotourism op-
portunities (Bagstad and Wiederholt, 2013). For example, by modeling
the importance of caves to the viability of Mexican free-tailed bat pop-
ulations (Table 1; results discussed in detail in Wiederholt et al.,
2013), we determined that caves in the southern part of the summer
range, those in Texas and northern Mexico, are crucial for maintaining
bat population viability. Several maternity roosts in this area also had
the largest summer population sizes (Wiederholt et al., 2015). In
terms of ecosystem services, the roosts with the highest annual ecosys-
tem service values (N$1 million) were all found in Texas, U.S. (Table 1).
For instance, the value of the pest suppression activities of the bats re-
siding in Bracken Cave, Texas, is $3.42 million per annum, followed by
Frio Cave, Texas ($2.42 million). The total annual ecotourism value of
a subset of cave viewing sites of Mexican free-tailed bats in the south-
western U.S. was conservatively valued at $3.95 million (Bagstad and
Wiederholt, 2013). The site with the highest ecotourism value was
Carlsbad Caverns National Park, with $3,477,072 (Table 1).
 R.A. Medellin et al. / Biological Conservation 211 (2017) 45–50 47

Table 1
Biological and economic importance, risk faced by the roost structure and by the population, and how losing that particular roost would affect the global population of 19 Mexican free-
tailed bat maternity roosts in Mexico and the United States for which sufficient information is known. The scores are standardized (10-based) for equal weighting scenario.
Modified from Wiederholt et al. (2015).

Bat roosts % population loss Scores for Scores for roost Scores for bat Annual pest Scores for pest Annual ecotourism Scores for
from roost removal biological structure risk population risk suppression value suppression value value ecotourism
importance value

Bracken Cave −10.14% 10.000 4.048 10.000 $3,420,256 10.000 $102,320 0.294
Carlsbad Caverns −0.79% 0.650 1.667 4.571 $27,500 0.080 $3,477,072 10.000
Cuatrociénegas de Carranza −2.34% 2.197 2.999 4.897 $0 0.000 $0 0.000
Cueva de Consuelo −2.02% 1.876 2.998 4.897 $0 0.000 $0 0.000
Cueva del Tigre −4.79% 4.652 2.857 6.857 $0 0.000 $0 0.000
Cueva La Boca −5.71% 5.564 1.830 1.656 $0 0.000 $0 0.000
Davis Cave −1.00% 0.859 6.429 8.143 $61,981 0.181 $0 0.000
Devil's Sink Hole −5.17% 5.031 3.003 4.904 $0 0.000 $127,900 0.368
Eagle Creek Cave −0.69% 0.554 2.857 3.143 $0 0.000 $0 0.000
Eckert James River Cave −3.28% 3.140 0.714 3.000 $317,973 0.930 $69,066 0.199
Fern Cave −0.59% 0.452 3.000 4.899 $0 0.000 $0 0.000
Frio Cave −5.68% 5.539 4.643 7.500 $2,421,653 7.080 $142,481 0.410
Maviri −0.23% 0.092 10.000 2.571 $0 0.000 $0 0.000
Merrihew Cave −0.23% 0.092 3.005 4.899 $5605 0.016 $0 0.000
Ney Cave −0.96% 0.818 0.000 0.000 $1,245,623 3.642 $0 0.000
Presa de Amistad −2.33% 2.194 3.000 4.903 $788,847 2.306 $0 0.000
Reed Cave −1.20% 1.063 3.000 4.905 $524,893 1.535 $0 0.000
Stuart Bat Cave −1.20% 1.063 3.003 4.897 $120,438 0.352 $26,808 0.077
Vickery Cave −2.47% 2.331 3.002 4.902 $205,552 0.601 $0 0.000

2. Major threats to cave ecosystems
As with other ecosystems around the world, caves have been
negatively affected by anthropogenic impacts. Examples of bat pop-
ulation declines following an anthropogenic influence abound
(Humphrey, 1978; Tuttle, 1979; Wilson, 1985; McCracken, 1989).
Caves are relatively more vulnerable than other ecosystems
(Elliott, 2000; Hamilton-Smith and Eberhard, 2000), and the most
common threats include urbanization, doline development, soil
erosion, groundwater pollution, and vandalism (Wilson, 1985,
Medellin, 2003, Elliott, 2004, Gunn, 2004).
Given the relatively large body of literature published on their bi-
ology, Mexican free-tailed bats seem to be good indicators of the
conservation status and severity of threats to caves. They can form
some of the largest concentrations of cave life and of bat populations
in the world (Medellin, 2003; Medellin et al., 2004), and their cave
roosts have been severely affected by human encroachment and van-
dalism (McCracken, 2003; Medellin, 2003). Organochlorine pesti-
cides such as DDT may be harmful to many insectivorous bats as
they can acquire high pesticide loads from their diets (McCracken,
1986). Populations are thought to have declined possibly due to
DDT exposure or disturbance from guano mining (Cockrum, 1970;
Bat Conservation International, 1991; Clark, 2001; Betke et al., 2008).
At the same time, caves overall, including bat caves, face serious
threats. For example, excessive visitation of spectacular caves in the
UK is likely causing damages to not only the geological structure of
the caves but to the air and water quality within it (Baker and Genty,
1998). Over-extraction of ground water, saline intrusion, and many
types of pollution in underground ecosystems are creating previously
unrecognized challenges by affecting water quality in caves and other
underground systems (Boulton, 2005).
Bats share many of these threats, but some additional threats af-
fect bats further: White-nose syndrome has wiped out over 6 million
hibernating, cave-dwelling bats since 2006 (Turner et al., 2011).
Other growing threats to bats include bushmeat trade and wind energy
developments (Clements et al., 2006; Baerwald and Barclay, 2009).
3. Conservation activities for cave ecosystems
As with other small natural features, a wide variety of conservation
activities have been applied to caves. Most of these activities have
been undertaken voluntarily by individuals and organizations that
own caves, but in some cases governments are purchasing land contain-
ing caves and developing regulations and incentives for cave conserva-
tion (Hunter et al., 2017). In addition, recent work is paving the road
toward cave prioritization for conservation of biodiversity, from a spe-
cies-area approach (Brunet and Medellin, 2001) to surface-level distur-
bance and cave complexity as good predictors of bat diversity and
abundance (Phelps et al., 2016).
3.1. Educate
Cave ecosystems are still unknown to a large degree and few ed-
ucational efforts have been launched for protection of caves, other
than at specific caves by local experts and conservation profes-
sionals. There are encyclopedias of caves and many other literature
resources (Culver and White, 2004, Gunn, 2004) but mainstreaming
cave protection as a major biodiversity conservation activity will re-
quire considerable focus and energy.
Many authors have pointed out how crucial it is to strengthen and
broaden ongoing efforts to protect bats in caves and to educate the pub-
lic about the benefits they provide (Mickleburgh et al., 2002; Medellin,
2003; Ladle et al., 2012; Sugai et al., 2015). Examples of successful edu-
cational programs do exist; for instance, the Program for the Conserva-
tion of Mexican Bats (PCMM) has a strong environmental education
program that targets elementary school children in Mexican communi-
ties near bat roosts (Medellin, 2003). The educational program involves
school visits several times per year over a three-to-four year time period
(Medellin, 2003). Adults are also targeted through radio shows, travel-
ing exhibits, workshops, capacity building, and posters. In the U.S., Bat
Conservation International (www.batcon.org) has been instrumental
in educating the public in Austin, Texas about bat ecology and in
protecting the now popular Mexican free-tailed bat ecotourism site at
the Congress Avenue Bridge. Given that the Mexican free-tailed bat
has one of the broadest distributions in the Americas, and is abundant
wherever it appears (Wilkins, 1989), the 22-country Latin-American
Network for Bat Conservation (RELCOM) has adopted this species as
an example of what can be synergistically, multinationally accom-
plished in cave conservation by highlighting benefits provided by
caves, specifically bat caves (RELCOM, 2009). Based on these examples,
we recommend that multinational, public educational and outreach
programs be implemented. These programs should be focused not
only on communities surrounding roosts, but also the general public
in areas covered by bat migration, foraging, and scouting. These types
48 R.A. Medellin et al. / Biological Conservation 211 (2017) 45–50
of programs are needed to foster more positive attitudes and behav-
ior toward bats and their conservation, and help the general public
understand why improved roost protection and monitoring is nec-
essary. Although no estimate exists of the number of people visiting
caves around the world, caving both as an “extreme” sport and as a
tourist activity is a very popular activity virtually everywhere there
are caves. Studies from Brazil, Turkey, the United States, all high-
light human visitation is very common and intense, and has signif-
icant negative effects on the conservation of cave ecosystems
(Mann et al., 2002; Paksuz and Ozkan, 2012; Pellegrini and
Ferreira, 2012). A single cave in Brazil is visited by an estimated
45,000 people a year (Lobo and Moretti, 2009), another in Turkey
had over 56,000 visitors in 2011 (Akca et al., 2016). Carlsbad cav-
erns alone, in the state of New Mexico, are visited by about
400,000 people each year (http://www.nationalparkstraveler.
com/2011/02/numbers-carlsbad-caverns-national-park7570), and
Jenolan Caves in Australia has 230,000 visitors annually (Jenolan
Caves Reserve Trust, 2016) and many other caves are promoted as
tourist destinations in that country. We conservatively estimate that
at least a few million people visit caves around the world each year,
thus providing a huge incidental opportunity to educate people and en-
hance cave conservation efforts, if properly conducted.
3.2. Inventory
Research on caves and their conservation needs is beginning to
increase, especially in karstic regions such as the east of the United
States, Brazil, or Southeast Asia. But global efforts to highlight the
relevance of cave ecosystems and the critical situation they face in
many areas are piecemeal and isolated. A search in the Web of Sci-
ence with the keywords “cave” and “conservation” yielded 1988 re-
sults, many of which are focused on marine caves or subaquatic
environments. From that search (Fig. 2), it is evident that published
research for the conservation of caves is on the rise, with the most re-
cent five years showing a 64% growth over the previous five-year pe-
riod, and that the 2010–2015 period represented 40% of the total of
papers published in Web of Science.
Caves are not obvious features in the landscape and are rarely in-
cluded in large-area management plans. Knowledge about the location
and number of caves is only incipient, so inventorying caves is a very
important task that should be promoted everywhere. A few efforts are
in place including via crowdsourcing or groups and societies (http://
www.grottocenter.org/; http://www.cave-research.org/projects.php).
Other efforts are more regional (Harley, 2007; Stafford et al., 2004;
http://www.pcrai.ro/static/engleza_proiect02_04.html). A central re-
pository where cave inventory and science could be stored would help
identify gaps and mainstream success stories.
Fig. 2. Number of papers published and reported in Web of Science with the key words “cave conservation” in five-year increments.
3.3. Protect and influence policy
Few governments around the world have specific policies for the
protection of cave ecosystems. Most efforts focus on specific caves or
the species that inhabit them, particularly endemic or endangered
species such as the four troglobite species included in the U.S. Endan-
gered Species Act. Mexican free-tailed bats have no federal protec-
tion in the U.S. or Mexico, nor is there a North American migratory
bat treaty. However, in May 2015 the Trilateral Committee for Wild-
life and Ecosystem Conservation and Management, composed of fed-
eral wildlife agencies from Canada, Mexico, and the United States,
signed a Letter of Intent acknowledging the need for transborder bat
conservation between the federal governments of Canada, Mexico,
and the United States (U.S. Fish and Wildlife Service, 2015). Subse-
quently several meetings have focused on devising a charter document
(http://www.batconservationalliance.org/about-us/charter). The Latin
American Network for Bat Conservation (RELCOM, 2009), which
currently consists of 22 countries including Mexico (http://www.
relcomlatinoamerica.net/), focuses on four key threats to bats: habitat
loss, destruction and disturbance of roosts, bat-human conflicts and
emerging infectious diseases, and indiscriminate use of toxic substances
(RELCOM, 2009). A similar initiative was launched a few years ago in
North America; the North American Bat Conservation Alliance works
to harmonize and coordinate the work, objectives, and priorities of var-
ious conservation groups. (http://www.batconservationalliance.org/).
A few efforts are spearheading the way to greater ecosystem protec-
tion for caves. Bat Conservation International, Inc., recently underwent a
large and effective private fundraising campaign to purchase the land
around one of the largest caves in the world, Bracken Cave, that was
threatened by the development of 3500 suburban homes. The result is
that now Bracken Cave and its 5000-acre surroundings are protected
for posterity (Moss, 2014). In Australia, a 3-ha purchase of a cave se-
cures one of only three maternity colonies of the eastern horseshoe
bat (Rhinolophus megaphyllus; Fitzsimons and Ashe, 2003)
Interestingly, some countries have created their own laws to protect
cave ecosystems. For example, the United States has a national Federal
Cave Resources Protection Act of 1988 (https://www.law.cornell.edu/
uscode/text/16/chapter-63), several states have cave protection legisla-
tion, Australia has begun highlighting the need to protect caves
(Fitzsimons and Ashe, 2003; Boulton et al., 2003) and Puerto Rico passed
a law in 1985 for the Protection and Conservation of Caves (http://drna.
gobierno.pr/historico/biblioteca/leyes/Ley111.pdf). Zhalov (2015) has
compiled a list of legal federal instruments protecting caves in many
countries in Europe and elsewhere. These laws insert caves in larger
landscape protection schemes and also include the protection of cave
paintings. The document also enlists procedures to protect caves on the
basis of vulnerability, type of underground habitat, protection of the
aquifer, cave development, and policies for cave cleaning.
 R.A. Medellin et al. / Biological Conservation 211 (2017) 45–50
3.4. Manage wisely
Many caves are subjected to human activities; some for the use by
the public as tourist attractions, others for more specialized speleologi-
cal visitation, and others for research or other needs. Although little for-
mal research has been done, there are some indications that limited
public visitation, if conducted with care and closely monitored and con-
trolled, may be compatible with biodiversity conservation objectives.
For example, Faille et al. (2015) describe the effects of public visitation
of the largest cave in France after it had been opened to the public.
The diversity and abundance of all 18 species of troglobitic invertebrates
did not decline after three years of carefully managed visits that were
restricted in duration, intensity, and spatial coverage. Similarly, Galicia
(2004) found that sensible, closely supervised short visits of small
groups of people have no significant detrimental effect on bat or inver-
tebrate diversity or abundance. However, extensive modification with
unsupervised, numerous tourist visits led to a virtual disappearance of
invertebrates, a sharp decline of the bat population and species diversi-
ty, and the disappearance of several species altogether (Galicia, 2004).
Although much is still to be uncovered as to what sustainable manage-
ment of caves implies, it is clear that common sense and short, closely
supervised visits of small groups are compatible with conservation of
some caves.
Some common lessons gleaned from Galicia (2004), Akca et al.
(2016), Paksuz and Ozkan (2012), Pellegrini and Ferreira (2012),
Phelps et al. (2016) and others include: limiting visits to times of the
year when bats and other biological communities are least vulnerable,
for example no visits during mating and maternity seasons; visits by
small groups, accompanied by a keeper or guard; timed visits, limiting
disturbance; avoiding or reducing lighting, noise, trash, and other an-
thropogenic disturbance as much as possible; gating in some cases;
strictly applying regulations by having personnel and infrastructure.
Constant monitoring of biotic and abiotic conditions is very important
as well. For example, studying 56 caves in the Philippines, Phelps et al.
(2016) show that bat diversity and abundance are correlated to sur-
face-level disturbance and cave complexity. This principle would be
helpful to prioritize which caves to protect.
3.5. Restore
Little is known about the restoration of cave environments; howev-
er, restoration itself can be orders of magnitude more expensive than
conservation (Jackson and Hobbs, 2009; Rodrigues et al., 2009). By
working with local inhabitants to win over their support for bat ecosys-
tem services and conservation, some bat populations in caves have re-
covered, in some cases by more than an order of magnitude (Medellin,
2003). Given the difficulty conveyed by geological, hydrological,
bioclimatical, aerial circulation, and other parameters in caves, restora-
tion in them is likely even more difficult and expensive to conduct than
restoration in other small (or large) natural features.
3.6. Create
The creation of cave ecosystems is a difficult task. Given the variables
that determine the way these ecosystems function (rock substrate, hy-
drology, macro and micromorphology, air circulation, geographical lo-
cation, biogeographical province and surrounding ecosystem in which
they are embedded, and more), creating additional cave resources is
not viable logistically, biologically, or financially on a large scale. How-
ever, a few projects have partially created cave ecosystems. One of
them is a project called the Chiroptorium, built by a Texas rancher in
the mid-1990s out of his own personal interest and as an attempt to
promote ecosystem services provided by local bat species, particularly
Mexican free-tailed bats. It took at least five years for the bats to estab-
lish a stable colony that in 2003 began growing, which now has a sizable
maternity colony (Bamberger, 2003). Today the bat colony is composed
49
of two species, including Mexican free-tailed bats, with more than
100,000 bats (Johnson, 2007). Other projects include biologically and
logistically simpler bat houses. For example, the University of Florida
built a large bat house in the early 1990s. Today, after having rotted
away, this bat house has been replaced by two structures. These are re-
putedly the largest purposefully constructed bat houses in the world,
harboring about 300,000 bats of three species, primarily Mexican free-
tailed bats (https://www.flmnh.ufl.edu/index.php/bats/home/).
4. Conclusion
Caves as ecosystems are small landscape features that have huge
effects on ecosystem functioning, and yet are rarely expressly con-
templated in any traditional regional or landscape conservation
plans. Caves dot many parts of the planet and contain extraordinary
levels of biological diversity including endemic species found in only
one or a few caves, and many unique phylogenetic groups. Unfortu-
nately many species that rely on caves are facing extinction risks. In
addition, caves are direct entrances to aquifers and their structure
and geology directly influence local hydrology and provide water re-
sources for human use and ecosystem functioning. Threats to bats
are beginning to be addressed by conservation measures. Recent ac-
tions, such as direct purchase of the land around bat caves produc-
tion and availability of public education resources, legal protection,
and even creation of new cave resources, the conservation commu-
nity seems to be realizing the urgent need to protect caves as integral
ecosystems. An effort to integrate cave conservation efforts that al-
lows for a common platform for sharing information, problems,
tools, solutions, and inventories is of utmost importance to establish
in the next few years.
Acknowledgements
We thank M. Hunter and A. Calhoun for the initiative and invitation
to participate in this publication. J. Fitzsimons, M. Hunter, and two anon-
ymous reviewers greatly improved the previous version of the paper.
Discussions with many presenters during the Small Natural Features
workshop at the 2015 International Congress for Conservation Biology
in Montpellier was very helpful and enriched our final product. We also
thank Tent and Trails, New York City, for equipment support. D. Cafaggi
kindly prepared several versions of Fig. 1.
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