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Amblyopsis rosaeOzark Cavefish

By Mark Levi Johnston

Geographic Range

With the only recorded populations in the world found in seven adjacent counties in the midwestern United States, Amblyopsis rosae is truly a locally endemic species. The only three states that contain counties with the cavern environment necessary to sustain their delicate niche are in Arkansas (Benton County), Missouri (Barry, Greene, Jasper, Lawrence, Newton, Spring, and Stone Counties), and Oklahoma (Delaware and Ottawa Counties). Due to decades of both natural and human-related pressures, historical populations of A. rosae have experienced a reduction in their range, which may still be occurring today. The common name, Ozark cavefish, is derived from the Ozark Highlands, a large plateau spanning three states. In states where they are found, populations have been verified in at least 21 different cave localities. (Noltie and Wicks, 2001; Romero, 1998)

Habitat

Ozark cavefish are extremely specialized to their cave ecosystem, requiring a pristine water source that remains at a temperature between 12.8 and 15.6°C. Because of this, they typically occupy caves whose water source comes from swelling groundwater, as opposed to surface-running streams. Access to this groundwater is possible due to the geologic characteristics of the plateau under which they live. Lying under a layer of exposed rock and soil, various types of eroded limestone including Burlington, Pierson, and Keokuk, make up the walls of individual cave habitats. Beneath this layer of limestone lies a bed of Chattanooga shale, which the groundwater settles on, allowing the groundwater to swell into the caves. In addition to the natural purification of the groundwater, the physical isolation of the cavern environment keeps out competing organisms and/or pathogens that are more prevalent in surface stream environments.

Because of lack of sunlight, streams in these caves are relatively barren, with mainly rocky, small-pebbled bottoms. Species richness is low, limited to a few species of salamanders, invertebrates, and copepods. (Culver and Pipan, 2009; Graening and Brown, 2003; Noltie and Wicks, 2001; Willis and Brown, 1985)

  • Aquatic Biomes
  • rivers and streams
  • Other Habitat Features
  • caves

Physical Description

With an average length of approximately 38 mm and mass of approximately 2.5 g, Amblyopsis rosae has evolved into a species that lacks pigmentation. Their de-pigmented epithelial layer is absent of melanin, thus they have a translucent appearance. As a result, the visceral organs can often be seen through the fish, consequently giving it a pinkish tint. Ozark cavefish lack a pelvic fin as well as functional eyes. This lack of sight is a result of the environment, which has rendered sight useless in the dark depths of caverns. To compensate for the absence of vision, A. rosae has a nervous system that is highly sensitive to smell and sound. (Noltie and Wicks, 2001; Poulson, 1963; Romero and Conner, 2007; Romero, 1998)

  • Range length
    24.0 to 42.5 mm
    0.94 to 1.67 in
  • Average length
    38.0 mm
    1.50 in
  • Average basal metabolic rate
    0.011 cm3.O2/g/hr

Development

The Ozark cavefish life cycle begins as an egg that is held within the gill chambers of the female. For four to five months after hatching, the young remain within the confines of the gill chambers. By doing this, young fry have the opportunity to grow and, upon leaving the gill chamber, are less likely to be victims of cannibalism. Maturation through their intermediate stages is very slow; both sexes grow an average of 0.6 mm per month. Regardless of age, individuals that are smaller in size tend to grow at faster rates compared to those of larger size. (Noltie and Wicks, 2001; Poulson and White, 1969; Poulson, 1963; Poulson, 2001)

Reproduction

Ozark cavefish are polygynandrous species, mating at random with multiple partners in an attempt to increase population fitness, although not all females may have ova ready for fertilization. (Noltie and Wicks, 2001; Poulson, 1963; Romero, 2001)

Ozark cavefish are considered sexually mature at age 4. Regardless of their sexual maturity, about 20% of the mature female population develop ova each year. It is believed that this reproductive limitation results from limited food sources in the cave. This birth limitation also helps to control food availability; lower population sizes do not demand as much food, which is limited in the cave environment. Eggs produced during each breeding period tend to be large in size and low in number when compared to similar species. (Poulson, 1963; Romero, 1998; Romero, 2001)

  • Breeding interval
    Breeding in individuals is non-annual and sporadic, with 20% of mature females becoming fertile each year.
  • Breeding season
    Ozark cavefish may breed at any time throughout the year.
  • Range number of offspring
    20 to 25
  • Average time to hatching
    4-5 months
  • Average time to independence
    Immediate upon hatching. minutes
  • Range age at sexual or reproductive maturity (female)
    4+ (low) years
  • Range age at sexual or reproductive maturity (male)
    4+ (low) years

Though juvenile Ozark cavefish are independent, parental investment before and after hatching is high. Once the eggs fertilized, they are transferred to the female's gill chamber, where they remain for four to five months until hatching. During that time, the female is responsible for protecting the eggs from harm. This, in turn, temporarily reduces her fitness as she is unable to be as mobile as she would be without her offspring. Males provide no parental investment outside of fertilizing the female ovum. (Poulson, 1963; Romero, 1998; Romero, 2001)

  • Parental Investment
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

Ozark cavefish grow at a slow rate, approximately 0.6 millimeters every month, and have a long lifespan. Because they are only viable in the wild and conservation measures restrict the capture of this species, a measure of average lifespan in captivity is not available. However, the average lifespan of wild populations is approximately 10 years. (Poulson, 1963)

  • Average lifespan
    Status: wild
    10 years

Behavior

Ozark cavefish have an extremely low metabolism. They spend the majority of their time hovering motionless in stream waters. Ozark cavefish exhibit aggressive behavior when provoked. However, they are relatively sedentary, conserving energy in their resource poor environment. (Graening, et al., 2009; Noltie and Wicks, 2001; Poulson, 1963; Romero, 1998)

Home Range

Home range size in Ozark cavefish is not described.

Communication and Perception

Because populations of Ozark cavefish are small and individuals are relatively solitary, communication with individuals outside of mating is limited. However, perception is vital for this species, as they have no vision. Ozark cavefish rely on concentrated bundles of nerves in their peripheral nervous system known as neuromasts. These neuromasts, located on the caudal fin, are able to detect motion in the surrounding water in the form of ripples or waves. They also have a lateral line across the side of their body which is far more sensitive than that of other fish species. Through a function known as hydrodynamic imaging, Ozark cavefish are able use this sensitive lateral line to create a picture of their surrounding environment. (Noltie and Wicks, 2001; Romero, 1998; Romero, 2001)

Food Habits

Ozark cavefish are carnivorous. The diet is relatively specialized, as prey choice is limited to the few organisms that are found in their cave environments. Ozark cavefish consume crayfish species including cave crayfish and spot handed crayfish; eggs from darksided salamander and cave salamanders; stygobitic arthropod species such as Stygobromus onondagaensis and Stygobromus ozarkensis; as well as a species of styobitic isopod. Additionally, they are cannibalistic. A portion of the diet, during periods of hatching, may consist of newly hatched young and/or developing juvenile cavefish. (Graening and Brown, 2003)

  • Primary Diet
  • carnivore
    • piscivore
    • eats non-insect arthropods
    • eats other marine invertebrates
  • Animal Foods
  • amphibians
  • fish
  • aquatic crustaceans

Predation

Ozark cavefish do not have any natural predators. Most deaths are due to dying from old age or from cannibalism in younger stages. Humans harm Ozark cavefish populations as a result of disruption from cave spelunking or collection of specimens for research. (Brown and Johnson, 2001; Graening and Brown, 2003; Poulson, 1963; Poulson, 2001)

Ecosystem Roles

Cave streams inhabited by Ozark cavefish are nutrient-poor because of their isolation from natural nutrient inputs, such as decomposing materials that make up the base of the food chain, including rotting leaves, soil, and dead organisms. The primary input of decomposing organic matter in these cave streams is guano from gray bat (Myotis grisescens) roosts. (Cambray, 2000; Graening and Brown, 2003; Poulson, 2001)

Mutualist Species

Economic Importance for Humans: Positive

Ozark cavefish are often targeted for academic research because of their unique ecology, evolution, and their endangered status. This research has the potential to generate positive economic roles in the form of research grant money. (Graening, et al., 2009; Stephens and Horton, 2007)

  • Positive Impacts
  • research and education

Economic Importance for Humans: Negative

Ozark cavefish do not negatively effect humans in general, although protection efforts can disrupt certain cave sports. A handful caves in Missouri, Oklahoma, and Arkansas have been quarantined in an attempt to help preserve the species. (Brown and Johnson, 2001; Graening, et al., 2009; Stephens and Horton, 2007)

Conservation Status

Both the United States Federal List and the International Union for Conservation of Nature's Red List indicate that Amblyopsis rosae is a threatened species. The primary threat is pollution of cave stream habitats. Habitat disturbance is also a major threat; both through over-collection of A. rosae specimens and caving and spelunking. Also, declining groundwater levels are negatively affecting populations.

To combat these effects on the Ozark cavefish, the states in which they are found have taken measures to ensure their conservation, such as blocking off the entrances to caverns. Additional monitoring techniques are also being practiced, such as water quality and population monitoring. (Brown and Johnson, 2001; Cambray, 2000; Graening, et al., 2009; Romero, 1998; Stephens and Horton, 2007)

Other Comments

Historically, Ozark cavefish have acted as biological indicators. Early settlers of the Ozark plateau would search for Ozark cavefish in the buckets of water they drew from their wells. Because they are extremely sensitive to water quality, only surviving in pristine waters, settlers would know that the drawn well water was clean and safe enough to consume if they found an Ozark cavefish. This important role gave A. rosae the title of "well keepers" or "spring keepers." (Graening, et al., 2009)

Contributors

Mark Levi Johnston (author), Radford University, Karen Powers (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

external fertilization

fertilization takes place outside the female's body

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

freshwater

mainly lives in water that is not salty.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

indeterminate growth

Animals with indeterminate growth continue to grow throughout their lives.

motile

having the capacity to move from one place to another.

natatorial

specialized for swimming

native range

the area in which the animal is naturally found, the region in which it is endemic.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

piscivore

an animal that mainly eats fish

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

sedentary

remains in the same area

semelparous

offspring are all produced in a single group (litter, clutch, etc.), after which the parent usually dies. Semelparous organisms often only live through a single season/year (or other periodic change in conditions) but may live for many seasons. In both cases reproduction occurs as a single investment of energy in offspring, with no future chance for investment in reproduction.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

threatened

The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).

vibrations

movements of a hard surface that are produced by animals as signals to others

year-round breeding

breeding takes place throughout the year

References

Adams, G., J. Johnson. 2001. Metabolic rate and natural history of Ozark cavefish, Amblyopsis rosae, in Logan Cave, Arkansas. Environmental Biology of Fishes, 62: 97-105. Accessed February 01, 2010 at http://www.springerlink.com/content/v042148477468033/fulltext.pdf.

Brown, J., J. Johnson. 2001. Population biology and growth of Ozark cavefish in Logan Cave National Wildlife Refuge, Arkansas. Environmental Biology of Fishes, 62: 161-169. Accessed February 02, 2010 at http://www.springerlink.com/content/x5m6671346gx5556/fulltext.pdf.

Cambray, J. 2000. Threatened fishes of the world series, an update. Environmental Biology of Fishes, 59: 353-357. Accessed February 02, 2010 at http://www.springerlink.com/content/p754k4g056700v82/fulltext.pdf.

Culver, D., T. Pipan. 2009. The Biology of Caves and Other Subterranean Habitats. New York, New York: Oxford University Press, Inc.. Accessed February 01, 2010 at http://books.google.com/books?hl=en&lr=&id=2Bg5BG_z12AC&oi=fnd&pg=PR16&dq=ozark+cavefish&ots=jw_Wixi7s_&sig=tviPoMvpPpjviTsRrl91uMUK3WE#v=onepage&q=ozark%20cavefish&f=false.

Graening, G., A. Brown. 2003. Ecosystem dynamics and pollution effects in an Ozark cave stream. Journal of the American Water Resources Association, 39: 1497-1507. Accessed February 02, 2010 at http://www3.interscience.wiley.com/cgi-bin/fulltext/118849605/PDFSTART.

Graening, G., D. Fenolio, M. Niemiller, A. Brown, J. Beard. 2009. The 30-year recovery effort for the Ozark cavefish (Amblyopsis rosae): Analysis of current distribution, population trends, and conservation status of this threatened species. Environmental Biology of Fishes, 87: 55-88. Accessed February 01, 2010 at http://www.springerlink.com/content/p747g1726u426332/fulltext.pdf.

Noltie, D., C. Wicks. 2001. How hydrogeology has shaped the ecology of Missouri’s Ozark cavefish, Amblyopsis rosae, and southern cavefish, Typhlichthys subterraneus: insights on the sightless from understanding the underground. Environmental Biology of Fishes, 62: 171-194. Accessed February 01, 2010 at http://www.springerlink.com/content/j77x470895nq20w1/fulltext.pdf.

Poulson, T. 1963. Cave Adaptation in Amblyopsid Fishes. American Midland Naturalist, 70: 257-290. Accessed February 02, 2010 at http://www.jstor.org/stable/2423056.

Poulson, T. 2001. Morphological and physiological correlates of evolutionary reduction of metabolic rate among amblyopsid cave fishes. Environmental Biology of Fishes, 62: 239-249. Accessed February 01, 2010 at http://www.springerlink.com/content/x1r8j14w000j6n18/fulltext.pdf.

Poulson, T., W. White. 1969. The cave environment: limestone caves provide unique natural laboratories for studying biological and geological processes. Science, 165: 971-981. Accessed April 23, 2010 at http://www.jstor.org/stable/pdfplus/1727057.pdf?cookieSet=1.

Romero, A. 2001. An introduction to the special volume on the biology of hypogean fishes. Environmental Biology of Fishes, 62: 7-12. Accessed February 02, 2010 at http://www.springerlink.com/content/q0t36823p121671n/fulltext.pdf.

Romero, A. 1998. Threatened fishes of the world: Amblyopsis rosae. Environmental Biology of Fishes, 52: 434. Accessed February 01, 2010 at http://www.springerlink.com/content/nq67032tk018301p/fulltext.pdf.

Romero, A., L. Bennis. 1998. Threatened fishes of the world: Amblyopsis spelaea. Environmental Biology of Fishes, 51: 420. Accessed February 01, 2010 at http://www.springerlink.com/content/g674066416330017/fulltext.pdf.

Romero, A., M. Conner. 2007. "Status Report for the Southern Cavefish, Typhlichthys subterraneus in Arkansas" (On-line pdf). Arkansas State University. Accessed February 04, 2010 at http://www.clt.astate.edu/aromero/493.ReportStatusSouthernCavefish.pdf.

Stephens, B., R. Horton. 2007. Ozark cavefish (Amblyopsis rosae) conservation in Missouri: a proactive approach to recovery. 2007 National Cave and Karst Management Symposium: 62-67. Accessed February 05, 2010 at http://www.utexas.edu/tmm/sponsored_sites/biospeleology/nckms2007/Papers/stephens.pdf.

Willis, L., A. Brown. 1985. Distribution and Habitat Requirements of the Ozark Cavefish, Amblyopsis rosae. American Midland Naturalist, 114: 311-317. Accessed February 01, 2010 at http://www.jstor.org/stable/2425606?seq=1.

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