Amphibian Biology and Husbandry

F. Harvey Pough
Abstract
Extant amphibians comprise three lineagessalamanders
(Urodela or Caudata), frogs and toads (Anura), and caecil-
ians (Gymnophiona, Apoda, or Caecilia)which contain
more than 6000 species. Fewer than a dozen species of
amphibians are commonly maintained in laboratory colo-
nies, and the husbandry requirements for the vast majority
of amphibians are poorly known. For these species, a review
of basic characteristics of amphibian biology supplemented
by inferences drawn from the morphological and physi-
ological characteristics of the species in question provides a
basis for decisions about housing and feeding. Amphibians
are ectotherms, and their skin is permeable to water, ions,
and respiratory gases. Most species are secretive and, in
many cases, nocturnal. The essential characteristics of their
environment include appropriate levels of humidity, tem-
perature, and lighting as well as retreat sites. Terrestrial and
arboreal species require moist substrates, water dishes, and
high relative humidity. Because temperature requirements
for most species are poorly known, it is advisable to use a
temperature mosaic that will allow an animal to find an
appropriate temperature within its cage. Photoperiod may
affect physiology and behavior (especially reproduction and
hibernation), and although the importance of ultraviolet
light for calcium metabolism by amphibians is not yet
known, ecological observations suggest that it might be im-
portant for some species of frogs. Some amphibians are
territorial, and some use olfactory cues to mark their terri-
tory and to recognize other individuals of their species. All
amphibians are carnivorous as adults, and the feeding re-
sponse of many species is elicited by the movement of prey.
Diets should include a mixture of prey species, and it may
be advisable to load prey with vitamins and minerals.
Key Words: amphibian; caecilian; care; frog; herpetocul-
ture; husbandry; salamander; toad
Introduction
A
mphibians have a long history of service as research
animals, especially as models for studies of embry-
onic development, regeneration, and physiological
function. Salamanders (especially ambystomatids and sala-
mandrids) and frogs (notably ranids and Xenopus) have con-
tributed much to these studies, and methods for maintaining
them in the laboratory are well established (Nace et al.
1974, ORourke 2002). Reviews of techniques for maintain-
ing, breeding, and rearing these species of amphibians in
captivity appear elsewhere in this ILAR Journal issue
(Browne and Zippel 2007; Browne et al. 2007; ORourke
2007). Discussions of the requirements of some amphibians
that are less commonly kept in captivity include the follow-
ing: Buchanan and Jaeger (1995), Cover et al. (1994), de
Vosjoli (1999), Johnson (1994), Maruska (1994), Pough
(1991), Wake (1994), Zimmermann (1986), and Zimmer-
mann and Zimmermann (1994).
Increased interest in organismal biology and in environ-
mental and evolutionary physiology has brought novel spe-
cies into the laboratory. These species often are selected for
study because they have particular ecological, environmen-
tal, or phylogenetic characteristics. Because investigators
and others have usually collected the animals from the wild,
the individual animals have no history of exposure to cap-
tivity, and as species, they have not experienced selection
for tolerance of captivity.
Currently there are no established husbandry methods
for most species of amphibians, and only scanty information
is available about ecology and behavior for many of them.
Meeting the basic physical needs of a speciesterrestrial or
aquatic conditions, and the appropriate temperature and hu-
midityis obviously important, but other issues may be
equally significant. Stimuli that elicit behavioral responses,
for example, may inadvertently increase stress. The social
behavior of amphibians is substantially more complex and
subtle than one would guess from observations of laboratory
colonies of Xenopus and axolotls (Ambystoma mexicanum
and Ambystoma tigrinum). Social behaviors such as territo-
riality and dominance hierarchies may influence feeding
and reproduction (e.g., Gabor and Jaeger 1995, Jaeger and
Schwarz 1991, Jaeger et al. 1995, Murray and Jenkins 1999,
Simons et al. 1997). Olfactory stimuli deposited on the sub-
strate or carried through the air are elements of the behav-
ioral ecology of some species of salamanders (Jaeger 1986).
Lighting conditions may also be more important than we
F. Harvey Pough, Ph.D., is Professor of Organismal Biology, Department
of Biological Sciences, Rochester Institute of Technology, Rochester, NY.
Address correspondence and reprint requests to Dr. F. Harvey Pough,
Department of Biological Sciences, Rochester Institute of Technology, 85
Lomb Memorial Drive, Rochester, NY 14623, or email [email protected].
Volume 48, Number 3 2007 203
realizephotoperiod as well as the intensity and spectral
quality of light are probably important stimuli for some
amphibians.
Caring for exotic species of amphibians has much in
common with herpetoculture, and herpetocultural resources
can be useful sources of information and advice. Recom-
mended resources include the Herpetocultural Library (de
Vosjoli 2004, 2006; de Vosjoli et al. 2005); web sites (e.g.,
www.pumilio.com); and chat groups (e.g., www.groups
.msn.com/HerpetocultureChatGroup/amphibians.msnw). In
addition, information about the ecology, geographic ranges,
and conservation status of amphibians is available from
Amphibiaweb (www.amphibiaweb.org) and Global Am-
phibian Assessment (www.globalamphibians.org).
Herpetological Review, which the Society for the Study
of Amphibians and Reptiles publishes, includes a section on
Herpetological Husbandry; and the International Zoo Year-
book includes information about the care of amphibians.
Professional staff members at zoos and aquaria may be able
to provide suggestions about the husbandry of particular
species.
When species-specific information about the behavioral,
ecological, and physiological characteristics of a species of
amphibian is not available, inferences based on elements of
the basic biology of amphibians may provide the only
guidelines. Two characteristics of amphibiansectothermal
regulation of body temperature and permeable skinplay
central roles in their biology and provide starting points for
inferences about their requirements in captivity. It is pos-
sible to amplify these general characteristics with informa-
tion about the phylogenetic relationships and by
observations of the morphological characteristics of the spe-
cies in question.
Biological Characteristics and Life Style
of Amphibians
The extant amphibians (Amphibia [Frost 2006] or Lissam-
phibia [Tree of Life Web Project 1995]) appear in the fossil
record during the Mesozoic. A large number of derived
characters (initially identified by Parsons and Williams
[1963] and subsequently restated and modified [Duellman
and Trueb 1985, Frost 2006]) support the monophyly of the
group. The lower Triassic form Triadobatrachus forms a
monophyletic group with frogs and toads (Rage and Rocek
1989). The earliest salamander fossils are from Middle Ju-
rassic sites in England and Kirghizstan, and investigators
have recently reported superbly preserved fossil sala-
manders from the Late Jurassic of China (Evans et al. 1988;
Gao and Shubin 2001, 2003; Gao et al. 1998; Nessov 1988).
The earliest known caecilian, Eocaecilia, is from the Early
Jurassic of Arizona (Jenkins and Walsh 1993). Thus, the
morphological and physiological characteristics of the ma-
jor lineages of amphibians have been established for 150
million years or more. These characteristics are central to
the evolutionary persistence of the lineages and are key
elements of the biology and husbandry of amphibians.
Phylogenetic and Ecological Diversity
A recent revision of the extant amphibians (Frost 2006) has
resulted in a change of the generic names of many species.
The literature of amphibian biology uses the old names, and
literature searches require these names. To minimize con-
fusion, I have used the familiar names with the new names
in parentheses.
The extant amphibians illustrate an approach to terres-
trial vertebrate life that combines ectothermy and a low
metabolic rate with highly efficient secondary production
(Pough 1983). A low metabolic rate reduces the mass-
specific metabolic rate with the result that small body size is
energetically feasible. Most amphibians are small. Twenty
percent of the species of salamanders and 17% of frog spe-
cies have adult body masses of 1 g or less, and 65% of
salamanders and 50% of anurans are smaller than 5 g as
adults. Because they are ectotherms, amphibians do not use
ingested energy to control their body temperature. As a
result, their net secondary production efficiency is high
amphibians convert half or more of the energy they ingest
into their own body mass compared with less than 1% for
birds and mammals. The combination of small body size
and efficient biomass production allows amphibians to play
a unique role in ecosystems by consuming prey items that
are too small for other vertebrates to eat and by repackaging
most of that energy into salamander and frog-size packages
(Pough 1980).
The skin of amphibians lacks scales, is plentifully sup-
plied with mucus and poison glands, and is a site of oxygen
and carbon dioxide exchange for adults as well as for larvae.
Small body size and glandular skin profoundly influence the
ecology, physiology, and behavior of amphibians. Cutane-
ous respiration accounts for a significant proportion of re-
spiratory gas exchange, water evaporates from the skin of
some amphibians as rapidly as from a free-water surface,
and amphibians are sensitive to toxins in the environment
(Alford and Richards 1999; Boutilier et al. 1992; Maerz et
al. 2005; Shoemaker et al. 1992).
The discussion that follows emphasizes features that are
helpful starting points for inferences about the habits, habi-
tats, and husbandry of unfamiliar species of amphibians.
Frogs and Toads
Anurans are the largest group of amphibians. Approxi-
mately 5200 species are in this group (Frost 2006), and they
reside on all of the major continents except Antarctica and
on many oceanic islands (Pough et al. 2004; Zug et al.
2001). Compared with other tetrapods, anurans have short
trunks (only 5-9 presacral vertebrae), broad, flat heads with
extensive reduction of bone, and hind limbs that are sub-
stantially longer than the forelimbs in most species. Al-
though the long hind limbs of anurans are most commonly
associated with jumping, they are also effective for swim-
ming, walking, running, and climbing; and all of these
forms of locomotion are represented among frogs and toads.
204 ILAR Journal
Common names do not match the phylogenetic or eco-
logical diversity of the extant anurans and are not a good
guide to their habits or husbandry. Some animals with frog
in their names are fully aquatic (e.g., the African clawed
frogs, Xenopus), but others are semiaquatic (e.g., many spe-
cies of true frogs, Ranidae) or fully terrestrial (e.g., the
dart-poison frogs, Dendrobatidae). Animals called toads are
often more terrestrial than those called frogs, but exceptions
(e.g., the completely aquatic Surinam toad [Pipa pipa])
make that distinction an unreliable guide for husbandry. An
animal called a tree frog is almost certainly at least partly
arboreal, but many frogs and toads without tree in their
common names have substantial arboreal capacities and are
best maintained in cages that permit them to climb, absorb
water droplets from leaves, and hide in foliage.
Body form is likely to be a more reliable guide to the
habits and needs of an unfamiliar species of anuran than its
name, and some generalizations about form and ecology are
robust (Figure 1). Arboreal species of frogs normally have
long limbs. Familiar examples are found in the hylids (tree
frogs that are most diverse in the New World and occur also
in Europe, parts of Southeast Asia, and Australia) and their
ecological counterparts the rhacophorids (tropical Africa,
parts of Asia, and the Indo-Australian archipelago), as well
as in less familiar groups such as hyperoliids (sub-Saharan
Africa, Madagascar, and the Seychelles) and centrolenids
(Central and South America). Some lineages of arboreal
frogs have expanded toe pads that help them adhere to
smooth surfaces. The bottom surface of a toe pad is formed
by polygonal columnar epithelial cells that are separated by
crevices (Green 1979). Glands secrete a watery fluid that
promotes capillary adhesion between the flat surfaces of the
epithelial cells and the substrate. The adhesive force that is
produced is strong enough to allow small species of frogs to
climb smooth vertical surfaces, including the glass walls of
terraria, and to adhere to the underside of a leaf or a glass
terrarium cover (Green 1981).
Stout bodies and blunt heads are often characteristic of
terrestrial anurans, and species that normally move by walk-
ing or running usually have relatively short legs and fore-
limbs and hind limbs that do not differ greatly in length.
Many species of toads (Bufonidae, a nearly worldwide
group) have stout bodies and blunt heads with relatively
short legs, as do terrestrial frogs (Leptodactylidae, primarily
a Central and South American group, and Microhylidae,
from southern North America to northern South America,
sub-Saharan Africa, Southeast Asia and eastern China, and
islands of the western Pacific Ocean). Anurans that burrow
into the ground generally have stout bodies and short limbs
that place them at the extreme end of the terrestrial walkers,
and some burrowers have pointed snouts and tarsal features
(e.g., the cornified spade of spadefoots) that assist in
burrowing.
Salamanders
The approximately 550 species of salamanders are elongate
animals with long tails. The forelimbs and hind limbs of
terrestrial salamanders are roughly the same length, and
salamanders use alternating movements of the limbs com-
bined with the passage of a sine wave along the body axis
for locomotion. Some fully aquatic species of salamanders
have reduced limbs. Salamanders have a more restricted
geographic distribution than do anurans. They are largely
absent from the Southern Hemisphere, extending southward
only as far as the northern half of South America and the
north coast of Africa (Pough et al. 2004; Zug et al. 2001).
Approximately 70% of extant salamanders are in the family
Plethodontidae. This family is well represented in North,
Central, and South America and has a single species in
southern Europe and another in Korea.
Salamanders exhibit less obvious morphological diver-
sity and habitat specialization than do anurans although
salamanders reside in a variety of habitats, from aquatic to
terrestrial. Plethodontids extend from arboreal through ter-
restrial to aquatic species, with few conspicuous differences
in body form. Most plethodontids (hence, most sala-
manders) dwell in streams, on stream banks, or on the forest
floor, but some tropical plethodontids are arboreal. Arboreal
salamanders have webbed feet that adhere to smooth sur-
faces as the salamanders climb. Salamanders without spe-
cialized feet also climb; Plethodon cinereus climb on plants
in the forest understory to forage on wet nights (Jaeger
1978). Streamside plethodontid salamanders extend from
species that normally live on the edge of a stream or in the
water (e.g., Eurycea and Gyrinophilus), through a series of
Figure 1 Relationship between body form and locomotor mode
among anurans. Reprinted with permission from Pough FH, Janis
CM, Heiser JB. 2005. Vertebrate Life. 7th ed. Saddle River: Pren-
tice Hall.
Volume 48, Number 3 2007 205
increasingly terrestrial species of streamside salamanders
(Desmognathus) that occur in habitats progressively farther
from water, to fully terrestrial species in the genus Pleth-
odon, which never enter water (Hairston 1987).
Salamandrids also span a range from aquatic to terres-
trial. Aquatic salamandridsthe European Triturus (Lisso-
triton) and the North American Notophthalmushave few
features that reveal their aquatic habits except for tails that
are laterally compressed compared with the more rounded
tails of terrestrial forms such as Salamandra in Europe and
Taricha in North America.
Some terrestrial species of salamanders (e.g., Ambys-
toma and Taricha) breed in water. These species develop
aquatic characteristics such as an expanded tail fin during
the breeding season. Fully aquatic species of salamanders
often have paedomophic characteristics that include exter-
nally visible features such as the absence of eyelids and
retention of a functional lateral line system. External gills
are the most conspicuous feature of some aquatic species
such as the mud puppies (Necturus). However, not all
aquatic salamanders have external gills. The largest extant
salamandersthe Chinese and Japanese giant salamanders
(Andrias), which reach adult lengths of 1.5 m or more, and
the North American hellbenders (Cryptobranchus), which
grow to 75 cmare aquatic but lack external gills. An
eel-like body form with extreme reduction in the size of the
limbs is characteristic of two families of aquatic sala-
manders, the sirens (Siren and Pseudobranchus) and the
amphiumids (Amphiuma).
Facultative or obligatory paedomorphosis occurs among
some species of ambystomatids. In some species, entire
populations retain external gills and other larval character-
istics and pass their entire lives in water (Whiteman 1994).
The familiar laboratory axolotls are an example of this phe-
nomenon. In other species, individuals within a population
are paedomorphic. The incidence of paedomorphosis varies
geographically in Abystoma tigrinum (Collins 1981) and
from pond to pond in Ambystoma talpoideum (Semlitsch
1985). Paedomorphosis also occurs in some other families
of salamanders, including salamandrids (Notophthalmus)
and dicamptodontids (Dicamptodon) (Good 1989; Harris
1987).
Caecilians
The Gymnophiona is a lineage of elongate, limbless, and
nearly tail-less amphibians known as caecilians, which in-
cludes both terrestrial burrowing species and aquatic species
(Taylor 1968, Wilkinson and Nussbaum 2007). The ap-
proximately 170 species live in tropical areas of most of the
world, but the group is absent from Madagascar and the
Papuan-Australian region. Their secretive habits make cae-
cilians difficult to study, but new techniques are revealing
important information about the ecology of this group
(Gower and Wilkinson 2005; Gower et al. 2004; Kupfer et
al. 2006a; Measey et al. 2003a, 2003b, 2004; Preswell et al.
2002).
The most important decision about the husbandry of a
caecilian is whether to keep it in an aquarium or a terrarium,
and that question is not always easy to answer. Aquatic and
terrestrial species of caecilians are not always distinguished
by conspicuous morphological differences. The eyes of all
species are greatly reduced and covered by skin, and all
caecilians have a unique chemosensory organthe ten-
taclethat incorporates elements of the eye and the vom-
eronasal organ. Aquatic species are somewhat laterally
flattened, and some of them have colored patterns on their
skin. These characteristics contrast with the cylindrical
body form of terrestrial species and the uniform surface
coloring of most of those species.
Life History
A complex life history that includes an aquatic larval stage
and metamorphosis to the adult form is an ancestral char-
acteristic of amphibians, but enormous interspecific varia-
tion exists even within this pattern. Only fishes have more
diverse life history patterns than amphibians. The diversity
of life history strategies in amphibians complicates gener-
alizations about animal husbandry. Direct development
(omitting a free-living larval stage) and viviparity (retaining
the eggs within the female through metamorphosis) occur in
all groups of extant amphibians.
Salamander larvae are similar to adults in general body
form although they lack some characteristics such as eyelids
and certain jaw bones, which appear only at metamorphosis.
Salamander larvae are carnivorous, as are all adult sala-
manders. The most visible change that accompanies meta-
morphosis of salamanders is loss of the gills and tail fin.
Direct development, which is widespread among plethodon-
tid salamanders, occurs also in Ambystoma and Salaman-
dra. The eggs of direct developing species are permeable
and require high humidity levels to avoid desiccation.
Frog larvae (tadpoles), in contrast to salamander larvae,
are very different from the adults in body form and function.
It is only a mild exaggeration to say that a tadpole is a
feeding machine with just enough locomotor capacity to
move it from one food source to the next. Tadpoles are filter
feeders, and they remove particles of food from the stream
of water that passes inward through the mouth and out past
the gills. Grazers, predatory forms, and egg-eaters use cor-
nified beaks to loosen particles from a large surface, and
midwater feeders sieve particles that are already suspended
in water. Papillae in the buccal cavity direct large suspended
particles to the esophagus, and mucus captures smaller par-
ticles. Strands of mucus and captured particles break loose
and are carried to the esophagus. Algae and bacteria com-
prise the diet of most tadpoles, but some species are mac-
rophagous, consuming zooplankton and small crustaceans.
Carnivorous tadpoles prey on other tadpoles (of their own
species or other species), and egg-eating tadpoles consume
eggs. Oophagous tadpoles live in small bodies of water
known as phytotelmata, such as in the leaf axils of plants.
206 ILAR Journal
Some consume eggs that are incautiously deposited by a
female (of their own species or a different species) in an axil
that already contains a tadpole. Dendrobatid frogs deposit
eggs in terrestrial nests, and one parent remains with the
clutch until the eggs hatch into tadpoles that are transported
to aquatic sites. Female Dendrobates (Oophaga) pumilio
transport tadpoles to phytotelmata and place one tadpole in
each site. The mother returns to her tadpoles and deposits
nutritive eggs during the developmental period (Brust
1993).
Some caecilians deposit large-yolked eggs on land.
These eggs hatch into free-living aquatic larvae that spend
approximately 1 year in the water before they metamor-
phose to become terrestrial adults. Viviparity is widespread
among caecilians; all species in the subfamilies Typhlo-
nectinae and Scolecomorphinae are believed to be vivipa-
rous. The subfamily Caeciliinae includes viviparous
species, species that lay eggs which hatch into aquatic lar-
vae, and species that lay eggs with direct development
(Wake 1977, 1993). Viviparous caecilians produce small-
yolked eggs that hatch into larvae within the mothers ovi-
ducts. When the energy supplied by the yolk has been
consumed, the larvae of viviparous caecilians feed on a
lipid-rich material secreted by the walls of the oviduct or by
the skin (Kupfer et al. 2006b; Wake 1993; Wake and Dickie
1998).
Viviparity appears to be rare among salamanders and
frogs, yet it has evolved repeatedly in both groups (Wake
1993). Salamandra atra gives birth to fully metamorphosed
young, and other species of salamandrids retain the eggs
through hatching and produce aquatic larvae. Viviparity in
the strict sense of retaining eggs in the oviducts through the
entire developmental period is currently known in only two
genera of anuransa Puerto Rican tree frog, Eleutherodac-
tylus jasperi, and a genus of African toads, Nectophrynoi-
des, which includes species that span the entire range from
external fertilization and aquatic tadpoles to internal fertil-
ization and viviparity (Graybeal and Cannatella 1995; Wake
1978). Viviparity is easy to overlook in the field, however,
and it is probably more widespread among anurans than we
currently realize.
Ectothermy and Temperature Regulation
In general, amphibians are regarded as poor thermoregula-
tors compared with other terrestrial vertebrates (Brattstrom
1962, 1963, 1979; Hutchison and Dupr 1992). Some habits
and habitats offer limited opportunities for thermoregula-
tion, at least during the portion of the day when the animals
are active. Nocturnal species, for example, cannot bask in
the sun, and the thermal mosaic of the environment is more
uniform at night than it is during the day. Furthermore,
evaporative cooling counteracts solar heating, and water
loss limits the time most species of amphibians can be in
exposed locations (Tracy 1975).
Amphibians thermoregulation under natural conditions
is probably more common than we appreciate, however,
because much of their thermoregulatory behavior consists
of selecting suitable microenvironments. Newly metamor-
phosed spotted salamanders, Ambystoma maculatum, for
example, take shelter beneath rocks on the banks of their
breeding ponds. The salamanders select sites under rocks
that do not exceed 32C during the day. In laboratory trials
in which rocks were heated, salamanders emerged and
sought cooler shelters when the temperature beneath a rock
reached 32C (Pough and Wilson 1970). Red efts, the ter-
restrial life stage of red-spotted salamanders (Notophthal-
mus viridescens), take shelter beneath pieces of bark in a
forest clearing. They tolerated body temperatures between
26 and 29C although lower temperatures were available
under the same piece of bark; however, an investigation
revealed no salamanders under bark slabs when tempera-
tures exceeded 30C (Pough 1974).
The permeable skin of amphibians both complicates
their thermoregulatory behavior and provides opportunities
that would not be available to animals with lower rates of
cutaneous water loss. Pearson and Bradfords (1976) com-
parison of a toad (Bufo [Chanus] spinulosus) and a lizard
(Liolaemus multiformis) in the high Andes illustrates the
limits that the permeable skin of amphibians places on their
ability to increase their body temperatures by basking in the
sun. Both animals basked in the sun and increased their
body temperatures above air temperature. Although the liz-
ard reached body temperatures between 30 and 35C, the
toads body temperature did not exceed 25C because of the
cooling effect of cutaneous evaporation. Newly metamor-
phosed green toads, Bufo (Anaxyrus) debilis, bask in the sun
on the sides of streams but seek areas of moist substrate to
avoid dehydration (Seymour 1972). Adult bullfrogs, Rana
catesbiana (Lithobates catesbeianus), spend the night in
artificial ponds in California that remain warmer than the
air, and the animals move onto shore during the day (Lil-
lywhite 1970). Frogs that are partly submerged are able to
remain in full sun for prolonged periods because evapora-
tive water loss balances solar heating and allows them to
maintain body temperatures between 25 and 30C whereas
black bulb temperatures exceed 40C. Canyon treefrogs
(Hyla arenicolor) in Arizona spend the day in full sun on
boulders near streams. Apparently cutaneous evaporation
prevents the frogs from overheating, but the investigators
observed that they did become dehydrated. At intervals the
frogs shuttled to the stream to rehydrate and then returned to
a boulder (Brust and Preest 1988). Avoidance of predation
by garter snakes (Thamnophis) is a plausible explanation for
this behavior because the snakes forage along the waters
edge but cannot cross the hot boulders to reach the frogs.
Temperature and Husbandry
Amphibian husbandry requires information about the tem-
peratures at which animals should be maintained, but it is
not possible to provide specific recommendations for am-
Volume 48, Number 3 2007 207
phibians in captivity. Although the temperature relations of
amphibians are flexible, their temperature preferences and
temperature tolerance may change daily or seasonally in
response to the recent thermal history of the animals, to
social interactions, or with age (summarized in Hutchison
and Dupr 1992).
The nocturnal habits of most amphibians and the inter-
action of heat loss (via evaporative cooling) with heat gain
(via radiation and conduction) preclude generalizations
about temperature regulation. Free-ranging animals prob-
ably use microclimate selection to adjust their body tem-
peratures, and they may regulate for only a portion of the
day. Species that are active at night, for example, probably
must choose between being active at whatever body tem-
perature results from the ambient conditions or forgoing
activity for that evening. These species may thermoregulate
during the day, however, by moving within their retreat sites
as some nocturnal lizards do (Mautz and Case 1974).
Aquatic larvae and the adults of fully aquatic species nor-
mally have body temperatures that are very close to water
temperature; however, under natural conditions tadpoles se-
lect different locations during the day in response to tem-
perature changes (Brattstrom 1962).
Temperatures recorded in the field reflect the ranges of
temperatures that amphibians have tolerated and provide a
starting point for husbandry. Based on the temperatures that
Duellman and Trueb (1985) and Hutchison and Dupr
(1992) have tabulated, it is advisable to use the following
guidelines when housing amphibians:
For species that live only in cold aquatic habitats (e.g.,
Ascaphus, Cryptobranchus, Necturus, Gyrinophilus),
initially set the temperature at 10C.
For species from cool temperate and montane tropical
regions, start with temperatures around 15C.
For species from warm temperate and tropical environ-
ments, evaluate the responses of the animals to tempera-
tures between 20 and 25C.
These estimates are intentionally on the cool side because a
temperature that is too high is more dangerous than one that
is too low.
Many species of amphibians are kept by herpetocultur-
ists. Specific information about these species is available on
the web and includes the following suggestions:
www.pumilio.com suggests a temperature of 18C
for dart poison frogs (Dendrobatidae);
www.goecities.com/rafarndell/husbandry.html suggests
temperatures between 26 and 30C for adult Ceratoph-
rys; and
www.peteducation.com recommends 18 to 24C for
Bombina.
Web sites are likely to be particularly useful sources of
information about exotic species, which are not typically
included in publications that focus on the species commonly
used in laboratories.
Observation of the animals color and behavior (activ-
ity, feeding, and production of feces) provides a basis for
adjusting temperature. If animals are dark-colored, inactive,
do not feed, or do not produce feces, it is advisable to
increase the temperature in small increments. Ideally, cap-
tive conditions will replicate the thermal mosaic found in
natural habitats and will allow animals to maintain daily
cycles of temperature and behavior. However, providing
both high humidity and a temperature mosaic simulta-
neously requires care to avoid creating a steamer that risks
overheating the animals it is intended to benefit. One sug-
gestion is to place an under-cage heater at one end of a long
terrarium to establish a temperature gradient that may allow
amphibians to thermoregulate behaviorally. Under-cage
heaters are widely available from herpetocultural suppliers.
Water Uptake and Loss
Evaporation is the major route of water loss for terrestrial
amphibians (Shoemaker et al. 1992). Some species have
very low cutaneous resistances, but others have substan-
tially less permeable skins. Investigators have characterized
anurans according to the following three general categories
of skin resistance to water movement (Buttemer 1990;
Shoemaker et al. 1972, 1989; Wygoda 1984, 1988):
Skin resistance < 1 sec/cm of skin thickness: Many
aquatic and terrestrial species (including species of spa-
defoot toads [Scaphiopus]), which live in deserts)
Skin resistance of 2 to 10 sec/cm: Arboreal species
Skin resistance of 100-300 sec/cm: A few species of tree
frogs, which are relatively waterproof
This pattern of skin resistance appears to include both phy-
logenetic and ecological components. All of the species
with a high level of cutaneous resistance are found among
hylids, rhacophorids, and hyperoliids, but arboreal species
of North American hylids have higher levels of cutaneous
resistances than do nonarboreal species (Wygoda 1984).
Waterproof species have mechanisms that reduce water
movement through the skin. Phyllomedusa sauvagii uses its
limbs to spread a waxy secretion from glands over its body.
Lipids in the skin are probably the basis of the high levels
of cutaneous resistance of Chiromantis, Hyperolius, Litoria,
and Hyla (Barbeau and Lillywhite 2005; Christian et al.
1988; Shoemaker et al. 1992).
The permeable skins of amphibians allow water to move
inward as well as outward, and amphibians rely on cutane-
ous uptake of water rather than on drinking. The pelvic
patch is an area of highly vascularized skin responsible for
most cutaneous water uptake. Water is absorbed by skin in
the pelvic patch from pools or droplets, and also from moist
soil. The relationship between soil water content and water
uptake is complex and depends on the structure of the soil
208 ILAR Journal
and the surface texture of the skin. Rugosity typically char-
acterizes the skin of the pelvic patch of terrestrial amphib-
ians, especially those from relatively dry environments, and
these species can generally absorb water from lower soil
water potentials better than can aquatic species.
The matrix water potential level of soil must be rela-
tively high to permit absorption of water by amphibians. A
frog (Rana [Lithobates] pipiens) that has been dehydrated to
70% of its fully hydrated body mass has a water potential of
-400 kPa (Tracy 1976). Water potentials in the range of -100
to -200 kPa allow water uptake by normally hydrated am-
phibians. (For horticultural potting soil, this water potential
corresponds to soil that can be squeezed into a ball that does
not crumble when the pressure is removed.)
Photoperiod and Light Quality
The light cycles in animal rooms that house amphibians
should accommodate the needs of the resident species, al-
though day and night (photophase and scotophase) can be
shifted for the convenience of the investigator. It is impor-
tant to note that because even nocturnal species need light to
detect prey, the night phase of a photoperiod cycle should
include low-intensity light.
Photoperiod, perhaps perceived by extraocular path-
ways, influences reproductive cycles of amphibians as it
does for other vertebrates. Photoperiod is also involved in
the timing of metamorphosis (Campbell et al. 2001; Pa-
niagua et al. 1990; Wright 2002); however, we do not yet
know about the effects of different spectral distributions of
energy on these functions.
Exposure to ultraviolet-B light (UV-B
1
) is an essential
element of husbandry for some species of lizards (e.g., Ge-
hrmann et al. 1991; Townsend 1979; Townsend and Cole
1985), whereas other species appear to thrive without ultra-
violet light. The role of UV-B in amphibian physiology is
less clear (Gehrmann 1994). Studies of the effect of ultra-
violet light on amphibians have emphasized its deleterious
effects on eggs and embryos (e.g., Blaustein and Belden
2003; Blaustein et al. 1997; Broomhall et al. 2000; but see
also Licht and Grant 1997 for a contrasting opinion); how-
ever, the behavior of some species of frogs exposes them to
direct solar radiation for hours at a time (Brust and Preest
1988; Lillywhite 1970; Seymour 1972). Fluorescent bulbs
with high UV-B output had no apparent ill effects on the
growth of juvenile Wyoming toads (Bufo [ Anaxyus ] bax-
teri) (R.K. Browne, Perth Zoo, South Perth, Western Aus-
tralia, personal communication, 2006), but larval
salamanders (Ambystoma macrodactylum and Ambystoma
gracile) that were exposed to UV-B grew more slowly than
individuals that received only full-spectrum illumination
(Belden and Blaustein 2002). Adults of two species of poi-
son dart frogs (Dendrobatidae) avoided sites with high
UV-B flux in the field and chose low levels of UV-B illu-
mination in cages (Han et al. 2007). This is an area in which
further experiments are needed. Gehrmann (1987) has pro-
vided information about the spectral quality of light from a
variety of lamps.
Chemical Ecology
A variety of noxious or toxic chemicals protect amphibians
and include the following: alkaloids, peptides, biogenic
amines, and bufodienolides (Brodie 1983; Daly 1995; Daly
et al. 1987; Erspamer 1994). Some of these chemicals are
derived from arthropods in the diets of the animals (Clark et
al. 2005; Daly et al. 2000; Darst et al. 2005; Saporito et al.
2003, 2004; Weldon et al. 2006 ). Species with potent
chemical defenses are often warningly colored, but this is
not always the case. Toads (family Bufonidae) contain bu-
fodienolides, which can be toxic to predators, but most
toads are cryptically colored. Other species with noxious or
toxic secretions have cryptic colors and patterns on the dor-
sal surface as well as bright flash colors on the ventral
surface that are revealed by defensive displays. When fire-
bellied toads (Bombina) are attacked, for example, they
bend the trunk upward to display the red or yellow ventral
surface. Similarly, fire-bellied newts (Cynops) and Califor-
nia newts (Taricha) expose brightly colored ventral surfaces
on the trunk and tail when they are threatened. These de-
fensive chemicals can be toxic to other species of amphib-
ians, particularly when individuals are crowded together.
Chemical communication is widespread among pleth-
odontid salamanders and may also occur in other terrestrial
species. Red-backed salamanders (Plethodon cinereus)
identify conspecific individuals by scent and respond dif-
ferently to the scents of neighbors and strangers (Jaeger
1981; Jaeger and Forester 1993; Jaeger and Peterson 2002).
It is important to recognize that moving animals among
cages may be stressful because olfactory communication
may cross species lines and the new cage may contain the
scent of unfamiliar conspecifics. Smith and Pough (1994)
have described intergeneric aggression of plethodontid
salamanders.
Housing and Diet
Amphibians are secretive animals that spend substantial
parts of the day in retreat sites. For this reason, the provision
of hiding places for these animals is a basic element of
successful husbandry. Amphibians are often reluctant to
move into areas where they feel exposed, therefore some
type of cage furnishing is an important component of hous-
ing. Naturalistic terraria with soil, plants, and pools of water
are aesthetically pleasing, but it is difficult to keep such an
enclosure clean and it is not clear whether amphibians ap-
preciate them as much as human observers do.
1
Abbreviation used in this article: UV-B, ultraviolet-B light.
Volume 48, Number 3 2007 209
Anurans and salamanders often thrive in cages with
moist paper as substrate, sections of plastic pipe as hiding
places, and plastic aquarium plants as cover objects. This
utilitarian housing is easier to clean than a naturalistic ter-
rarium, which is an important consideration when large
numbers of individuals are maintained. The use of inert
materials for amphibians is important because of the per-
meability of their skin. For example, the sensitivity of am-
phibians to the acids and bleaches that are used to
manufacture paper make laboratory filter paper a better
choice than paper towels, and distilled, deionized, or spring
water is preferable to chlorinated tap water.
Importance of Live Prey
Aquatic amphibians (e.g., tadpoles, salamander larvae, and
adults of species such as Xenopus and axolotls) accept pre-
pared diets, but terrestrial species often respond only to the
stimuli provided by moving prey items. Most amphibians
feed on insects, although the largest species consume crus-
taceans and small vertebrates. Bullfrogs (Rana [Lithobates]
clamitans) and cane toads (Bufo [Chanus] marinus) are no-
torious for eating other species of anurans, and the horned
toads of South America (Ceratophrys) lure smaller frogs
within reach by wiggling a toe in a way that simulates the
movements of an insect.
Frogs eyes respond preferentially to small, convex,
dark stimuli that move across the field of view. These neural
circuits have been called bug detectors (Lettvin et al. 1959).
Reliance on hard-wired prey detection mechanisms to elicit
a feeding response may explain why anurans and sala-
manders do not respond to motionless prey. Moving an
inanimate food item in front of an amphibian can stimulate
attack; however, the size, shape, and pattern of movement of
the item must fall within the range of stimuli to which bug
detectors respond.
Despite the stereotypy of feeding responses that bug
detector circuitry has implied, at least some amphibians
change their response to prey under different circumstances.
Undisturbed red-backed salamanders (Plethodon cinereus)
selected large fruit flies when they were given a choice
between large and small flies, and this behavior conforms to
the predictions of optimal foraging theory (Jaeger 1990).
Nutritional Quality of Prey
Amphibians consume whole insects. Practical consider-
ations usually limit the choice of food items to several spe-
cies of invertebrates that are readily available commercially,
which include the following: mealworms (Tenebrio moli-
tor), gray crickets (Acheta domesticus), wax worms (Galle-
ria mellonella), cockroaches (usually Blaberus craniiferus),
wingless fruit flies (Drosophila melanogaster), and earth-
worms (Lumbricus terrestris). Insects are generally defi-
cient in vitamins and have low calcium:phosphorus ratios
(Barker et al. 1998). None of these species (with the pos-
sible exception of earthworms) is suitable as the sole com-
ponent of a diet (Bernard and Allen 1997).
It is possible to supply vitamins and minerals by mixing
a bird or reptile multivitamin powder with powdered cal-
cium or a commercial preparation of calcium with vitamin
D
3
and dusting the insects with this mixture just before
offering the food items to the amphibians (Allen and Oft-
edal 1989). Another way to improve the nutritional content
of insects is to feed supplements to thema process called
gut-loading. Ground rodent chow, monkey chow, trout
chow, tropical fish flakes, and high-protein baby cereal have
been used for gut loading In a comparative study, juvenile
Wyoming toads (Bufo [Chanus] baxteri) fed crickets that
had been gut-loaded with an experimental diet that included
spirulina (high levels of vitamin A) and fish oil (high in
omega-3 fatty acids) ate more and grew faster than toads
that ate crickets dusted with powdered vitamins and miner-
als (R. K. Browne, personal communication, 2006).
Because powdered vitamins and minerals adhere to prey
items for only a short time and nutrients in the gut are
transient, it is important to remove uneaten food items
within a few hours. In addition, because hungry insects may
attack amphibians, it is wise to remove insects that are not
eaten immediately as a precaution.
Summary
About a dozen species of amphibians are commonly main-
tained in captivity, and husbandry methods for those species
are well established. Caring for individuals of any of the
remaining 6000 or so species of frogs, salamanders, and
caecilians requires integrating knowledge of the basic bio-
logical characteristics of amphibians with information about
the behavior and ecology of the species in question. Herpe-
toculturists have developed methods for maintaining exotic
species of amphibians, and herpetocultural publications and
web sites may provide useful information. Frequently little
information is available about a species, and in that situa-
tion, it is necessary to draw inferences from morphological
characteristics of the species.
Ectothermy and a permeable skin are basic characteris-
tics of amphibians. In captivity, terrestrial species require a
source of heat for thermoregulation and a closed environ-
ment that maintains a high humidity within the cage. The
combination of heat with a closed container creates a risk of
lethal overheating, therefore cages should provide a thermal
mosaic from which the animal can select conditions that
match its preference.
Many amphibians are nocturnal, spending the day in
hiding places and emerging at night to feed. Terrestrial am-
phibians locate prey by vision, and consequently animal
rooms require a low level of illumination at night. The
significance of ultraviolet light to amphibians is unclear. A
substantial body of literature has documented harmful ef-
fects of ultraviolet radiation on amphibian eggs, and adults
210 ILAR Journal
of some species of tropical forest frogs avoid exposure to
ultraviolet light. However, some species may require expo-
sure to ultraviolet light for vitamin D activation and calcium
metabolism.
Most amphibians feed in response to the movement of
prey items, and live prey is usually necessary. Insects are
the most common food for amphibians in captivity, but
insects do not provide a nutritionally complete diet. Feeding
a variety of insects is advisable, as is supplementing the
insects with vitamins and minerals. Powdered vitamins and
minerals can be dusted onto an insect just before it is fed to
an amphibian, or high concentrations of these substances
can be included in the food of the prey insects. Uneaten prey
should be removed from the cage promptly because the
nutritional supplements are quickly lost, and hungry insects
may attack the amphibian.
Chemicals play a large role in the ecology and behavior
of amphibians, and must be considered in husbandry. Many
species of amphibians are protected by toxic secretions, and
in some situations these substances can be lethal to other
species. For this reason, it is unwise to carry different spe-
cies in the same small container, and it may be unwise to
mix species in cages. Salamanders, at least, have extensive
social behaviors that are mediated by chemical cues from
conspecifics. Some salamanders can recognize other indi-
viduals by scent, for example, and the scent of a strange
salamander elicits behavioral responses different from those
produced by the scent of a familiar individual. Observations
of interspecific aggression suggest that chemical stimuli
might cross species lines as well. It is essential to clean all
of the equipment used to contain and handle amphibians
after each use to remove scent cues.
Because amphibians differ in so many respects from
their human caretakers, successful husbandry requires com-
bining an appreciation of the biology of these animals with
awareness of elements of the physical and biological envi-
ronment that are insignificant or even imperceptible to hu-
mans and sensitivity to subtle changes in the colors,
postures, and behaviors of the amphibians themselves.
Acknowledgments
Larry Buckley, John Cadle, Alan Savitzky, Ellen Smith,
Stephen Smith, and two anonymous reviewers provided
helpful advice and suggestions.
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