Transcript Lecture 9 Lissamphibia diversity

Lissamphibia
~4,900 species
(more than number of extant mammal
species)
>4,300 species frogs (Anura)
>415 species salamanders (Urodela)
>165 species caecilians (Gymnophiona)
Lissamphibia
Gymnophiona
(caecilians)
Typhlonectes
Typhlonectes
CAECILIANS
The Unknown Amphibian
Who are they?
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Order: Gymnophiona
Family: 6 families
Genus: 34
Species: 165
Habitat: loose soil,
leaf litter in tropical
forests or rivers and
streams
• Length: up to 2.4m;
shortest 3.5 in.
• Up to 2.2 lbs
• Life span: up to 13 yrs
• Offspring: 30 – 60
eggs; 2 - 25 young
born
Range
Central Africa, Southeast Asia, Southern Mexico to Argentina
Description
• Caecilians are tropical amphibians that
look like large worms or slick snakes. They
have no arms or legs, and sometimes it’s
hard to tell which end is the head and
which is the tail! Their shiny skin is ringed
with skin folds called annuli, and they
usually come in shades of gray, brown,
black, orange, or yellow. Some species
have tiny, fishlike scales (dermal scales)
within the rings.
Gymnophiona (caecilians) (see Pough CH3)
- elongate, distinctly annulated
-primary etc.
- terrestrial, burrowing, or aquatic (derived)
- tails reduced or absent
- eyes reduced (covered by skin or even bone)
- limbs and girdles absent in all extant taxa
(still present in fossil ancestor)
evidence of tentacles
Fig. 3-11
Pough et al. 2001
Fossil Gymnophiona ancestor
(Eocaecilia)
Annuli and Smooth
Dermal Scale Present in Some
Caecilians
Gymnophiona (caecilians)
dermal scales in some spp.
- pouches in dermis below annular groove
- scales like this not found in other Lissamphibia
Fig. 3-9
Pough et al. 2001
Caecilian Head
Gymnophiona (caecilians)
- Skulls unique
- Well ossified (but openings for eyes, nares, tentacles)
stegokrotaphic = skull completely roofed
zygokrotaphic = skull weak in temporal region
vs.
gymokrotaphic = temporal region open (anurans + urodelans)
stegokrotaphic =
skull completely roofed
zygokrotaphic =
skull weak in
temporal region
stegokrotaphic
Fig. 3-10
Pough et al. 2001
Lots of bones fused
Caecilian skulls vs. Anuran skull
Z
S
S
S
G
S
Fig. 13-12
Duellman and Trueb, 1986
S
Fig. 13-15
Duellman and Trueb, 1986
Gymnophiona (caecilians)
- many bones of skull fused
- Why have well ossified skull?
- unique dual jaw closing mechanism
Fossorial
Adult caecilian can
bury itself in a matter
of minutes. Fossorial
animals spend most of
their time
underground.
Aquatic Species
Gymnophiona (caecilians)
- left lung often reduced or absent
- aquatic eggs and larvae, or terrestrial
oviposition with female parental care
- some species viviparous
- internal fertilization (phallodeum)
Male Typhlonectes
(cloaca)
Typhlonectes
(mating)
Oviparous Species
Some female caecilians lay eggs in damp
holes near water. When the larvae hatch
they have gills and a short, finned tail to
help them swim in the water, feeding on
plankton. Through a series of changes the
gills are replaced by a single lung, the skin
becomes thicker, the annuli develop, and
sensory tentacles appear. At this point the
newly developed adult returns to the land
and goes underground.
Caecilian Embryo
Embryo removed from
egg. Yolk is visible as
well as sensory
structures along head
and side of body.
These include
mechano- and
electrical receptors
of the lateral line.
Eggs are laid in
burrows near streams.
hatch, larval stage lives
in streams until
metamorphosis a year
later.
Icthyophis kohtaoensis
Viviparous Species
• Some species give birth to live young that
are fully developed inside the mother before
they are born.
Gymnophiona (caecilians)
In viviparous species,
- young nourished by secretions
from oviduct
- mother’s skin may produce edible
secretions after birth
Fig. 17-13
Stebbins and Cohen, 1995
Fig. 5-5
Duellman and Trueb, 1986
Larval Caecilian
Aquatic stage with gills that are transient.
Senses
• A hard, thick, pointy skull helps these
amphibians dig in soft dirt. Because of their
underground lifestyle, caecilians have little need
to see or hear. Therefore, their eyes are very
tiny in some species, or hidden under the skin or
skull in other species, making just tiny gray
bumps for eyes. Some species in South America
spend their lives in water instead of on land.
Caecilians don’t have ear openings, so it is
doubtful they can hear sounds.
Tentacles
• Tentacles, located between the nostrils and
the eyes, are used to locate prey and detect
surroundings.
Gymnophiona (caecilians)
TENTACLE
- chemoreception
- between eyes and nostrils
- protrusible in some taxa
- involves structures normally associated
with eye and Jacobsen’s organ
Diet
• Inside a caecilian’s mouth are dozens of
needle-sharp teeth. The teeth are used to
grab worms, termites, beetle pupae,
mollusks, small snakes, frogs, lizards, and
even other caecilians. All food is
swallowed whole.
Eating tubifex worms
Skin Glands
• Caecilians have toxic glands in their skin
that sometimes protect them from being
eaten by other animals.
Mucus
glands
Toxic glands
Families of Caecilians
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Rhinatrematidae South America
Ichthyophiidae Southeast Asia
Uraeotyphlidae Southern India
Scolecomorphidae Subsahara Africa
Caeciliaidae South and Central America,
Subsahara Africa, Southern Asia
• Typhlonectidae South America
Gymnophiona
Pough et al. 2001
Lissamphibia
Gymnophiona
(caecilians)
6 families, 34 genera, 165 spp.
Fig.3-12
Pough et al. 2001
Gymnophiona (caecilians)
Rhinatrematidae
(2 genera, 9 species; northern South America)
- ancestral (‘primitive’) lineage
- true tail
- mouth at tip of snout
- tentacle next to eye
- zygokrotaphic skull (weaker)
- many scales
- not well-known
- likely not fossorial
- relatively small
- oviparous (egg-laying), with aquatic larvae
Rhinatrematidae
• Small, terrestrial caecilian with aquatic larvae
@1998 Marvalee
H. Wake
Gymnophiona (caecilians)
Uraeotyphlidae
1 genus, 5 species; Southern India
Stegokrotaphic skulls
oviparous (with aquatic larvae)
small (300 mm)
true tail
Tentacle underneath the nostril
Uraeotyphlidae
Gymnophiona (caecilians)
Ichthyophiidae
2 Genera, 38 species+; SE Asia, India, Islands of
the Indo-Pacific, and the Philippines
Terrestrial
Ovivparous (up to 100 eggs)
True Tail
Tentacle closer to eye than nostril
500 mm
Stegokrotaphic skull
Dermal Scales
Ichthyophiidae
• Large terrestrial caecilian with aquatic
larvae
Gymnophiona (caecilians)
Scolecomorphidae
(2 genera, 5 species; parts of central Africa)
- unusual lineage
- vestigial eyes attached to tentacles
- orbits absent
- zygokrotaphic skull lacking elements
- calcified spines on male phallodeum
- no scales
- oviparous or viviparous
Scolecomorphidae
Gymnophiona (caecilians)
“Caeciliidae”
(21 genera, 90 species; central and S.
America, parts of Africa and Asia)
- no tail
- mouth recessed under snout
- tentacular location varies
- scales variable
- mostly burrowers
- wide range of sizes
- some brightly colored
- life history variable
Caeciliaidae
• Very varied with small
(10cm) to large
(1.5m) organisms.
• Terrestrial, aquatic,
oviparous, viviparous.
• Most are gray – black
but some are more
colorful.
Gymnophiona (caecilians)
Typhlonectidae
(5 genera, 13 species; South America)
- often aquatic
- tracheal lungs
- narial plugs
- secondarily zygokrotaphic skulls
- includes largest lungless tetrapod
- nocturnal
- viviparous with aquatic larvae
Typhlonectes
Sometimes sold
in fish stores.
Aquatic.
Female with
young.
Typhlonectidae
• Aquatic caecilian,
viviparous
• Larval gills
Ambystoma californiense
Lissamphibia
Urodela
(salamanders)
10 families, 61 genera, 415 spp.
Ambystoma tigrinum
Ambystoma californiense
Fig. 13.5
Stebbins and Cohen, 1995
Unken reflex
Urodela (salamanders) (see Pough CH3)
- Urodela (extant)
- Caudata (extant plus fossil relatives)
- North America and temperate Eurasia
- Plethodontids have radiated in tropics
- extensive salamander fossil record
(unlike Gymnophiona)
Urodela (salamanders)
- typically 4-limbed, short bodied, long tail
some with limbs reduced or lost
- costal grooves
- most species terrestrial,
but need water for reproduction
(some fully aquatic or terrestrial)
- some arboreal
Urodela (salamanders)
- unique jaw musculature
- skulls reduced (bones absent)
- skull bone ossification sequence
- late appearance of maxillae in skull
Urodela (salamanders)
- lack middle ear cavities and tympana (drum)
- various opercular components missing
Pough et al. 2001
Urodela (salamanders)
- mucous and poison glands
(pheromones)
- aggregations of mucous glands =
courtship glands (e.g., mental
glands)
Fig. 17.3
Stebbins and Cohen, 1995
mental glands
Urodela (salamanders)
- some external fertilization
- mostly internal fertilization
(via spermatophore)
female
male
Duellman and Trueb, 1986
Urodela (salamanders)
- large genomes (and large cell nuclei)
- usually have larval stage
larvae with true teeth, gill slits, external gills
- larvae rather similar to adults
(compared to anurans)
palate is remodelled during metamorphosis
- many with direct development
Salamander Larvae
Pond
Stream
Terrestrial
Stebbins and Cohen, 1995
Urodela (salamanders)
- Paedomorphosis common
all adults retain larval
characteristics
OR
some retain larval characteristics
(facultative)
OR
none retain larval characteristics
- makes phylogenetic reconstruction
difficult
Urodela
Duellman and Trueb, 1986
(salamanders)
- Unique
defensive
adaptations
Urodela (salamanders)
- Unique defensive adaptations
Fig. 10-7
Duellman and Trueb, 1986
Lissamphibia
Urodela
(salamanders)
10 families, 61
genera, 415 spp.
Fig. 3-1
Pough et al. 2001
Urodela (salamanders)
- Unique defensive adaptations
Fig. 10-7
Duellman and Trueb, 1986
Echinotriton chinhaiensis
Chinhai Salamander
Salamandridae
© Max Sparreboom
Urodela
Pough et al. 2001
Sirenidae
Hynobiidae
Salamandridae
Urodela
Pough et al. 2001
Urodela
Plethodontidae
Proteidae
Pough et al. 2001
Urodela (salamanders)
Sirenidae
(2 genera, 4 species; SE U.S., NE Mexico)
- ancestral lineage
(sister to all other
salamanders)
- long, slender, eel like
- lack pelvic girdle and hindlimbs
- keratinized beak
- fully aquatic
- prey on invertebrates (insects,
crayfish, worms)
- Paedomorphic (no eyelids, external
gills, non-pedicillate teeth,
reduction in number of
digits on forelimbs)
- mucous cocoon and aestivation
- external fertilization?
- oviparous
The dwarf-siren
Pseudobranchus striatus
(photo J. White)
Urodela (salamanders)
Cryptobranchidae
(2 genera, 3 species;
Japan, China, E N. America)
- incomplete metamorphosis
adults lack eyelids,
retain one pair gill slits
- flattened bodies and heads
- aquatic
- asymmetrical suction
feeding
- cutaneous respiration
- largest salamanders (up to
1.8 m)
- cold mountain streams
- external fertilization
- males make nests and
guard eggs
(Den Master)
© Dr. Eric J. Routm
The Hellbender
Cryptobranchus alleganie
(photo M.A. Donnelly)
Urodela (salamanders)
Hynobiidae
(7 genera, 36 species; disjunct Asia)
- complete metamorphosis
eyelids, lack gill slits as adults
- lungs may be reduced or absent
Batrachuperus mustersi
Photo by Max Sparreboom
- some fully aquatic
- some with terrestrial adults and migration to breed
- relatively small (100-250 mm)
- some breed while wetlands still ice and snow covered
- males have territories
- males may compete (mating ball)
- external fertilization
Urodela (salamanders)
Amphiumidae
(1 genus, 3 species; S, SE U.S.)
- elongate, paedomorphic
lack eyelids, lack
external gills (but have slits)
- pedicellate teeth
- retain both pairs of limbs (and
girdles),
but small limbs
- aquatic
- eat invertebrates and
vertebrates
- ocassionally move overland or
aestivate
- up to 1.1 m long
- internal fertilization
- oviparous on land (alligator nest
mounds?)
female parental care
Two-toed amphiuma
Amphiuma means
© John White
Urodela (salamanders)
Plethodontidae
(28 genera, 266 species; disjunct N, Central, S America, ~ Italy)
- diverse and specious
(only group to radiate in tropics)
- lungless
(adaptation to mountain streams)
- nasolabial groove (chemoreception)
- subterranean, aquatic, terrestrial, arboreal
- variable body forms
(arboreal often with webbed feet and prehensile tails)
- variable life history strategies
- include smallest and
~largest terrestrial Urodela
Batrachoseps stebbinsi
Tehachapi Slender Salamander
© Tim Manolis
Plethodontidae
Figs 7.2 and 6.1
Stebbins and Cohen, 1995
Fig.3-6
Pough et al. 2001
Urodela (salamanders)
Rhyacotritonidae
(1 genus, 4 species; Pacific Northwest U.S.)
- inhabit old-growth forest
(indicator species)
- very old lineage
Rhyacotriton olympicus
Olympic Torrent Salamander
© William Flaxington
Urodela (salamanders)
Proteidae
(2 genera, 6 species; NE U.S., S Central Europe)
- diploid chromosome count of 38
- paedomorphic
large gills and caudal fins
- lack maxillary (~upper jaw) bone
- mostly aquatic
- Proteus live in caves
(slender;
whitish skin;
reduced eyes)
Proteus anguinus
Urodela (salamanders)
Salamandridae
(15 genera, 55 species; N America, Europe,
Asia, N Africa)
- includes ‘newts’
- may have extremely rugose skin
- life history variable
- some species with efts (terrestrial
immature)
Taricha granulosa
Rough-skinned Newt
© Harry Greene
Unken reflex
Newts
Taricha spp.
Fig. 13.5
Stebbins and Cohen, 1995
Salamandra salamandra
Fire Salamander
© Arie van der Meijden
- defensive displays
- aposematic coloration
(bright warning)
- tetrodotoxins (neuro)
Urodela (salamanders)
Ambystomatidae
(1 genus, 32 species; N America)
- several hybrids and unisexual forms
variable ploidy and number of parental species
- hybrids generally of ancient origin
- Ambystoma mexicanum often used in research
- variable metamorphosis (none, some, all)
- robust and ‘medium sized’
Ambystoma californiense
Urodela (salamanders)
Dicamptodontidae
(1 genus, 4 species; ~NW U.S.)
- similar to Ambystomatidae
- relatively large
- inhabit coniferous forest habitats
cold streams and
cold mountain lakes
- species very difficult to differentiate
morphologically
Dicamptodon tenebrosus
Pacific Giant Salamander
© Henk Wallays
Rana pipiens
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, 4,300 spp.
Fig. 3-20
Pough et al. 2001
caecilian
salamander
frog
Zug et al. 2001
Anura (frogs) (see Pough CH3)
- Salientia (clade that includes extinct
Triadobatrachus)
extra
vertebrae
Pough et al. 2001
tail
Anura (frogs)
- most extant families first appear in the Cretaceous
- radiation in the early Cenozoic
Ascaphus truei
Anura (frogs) (see Pough CH3)
GENERAL
- tail absent
- metamorphosis
larvae and adults very different
- specialized skin...
- jumpers (saltation)
- 88% of Lissamphibia are Anura
- found ~worldwide except Antarctica, poles,
and very dry deserts
Anura (frogs)
Jumping
Most anurans jump
Likely ancestral state was terrestrial protofrog that
moved into water in riparian system to avoid
danger.
‘Jumping’ is fast way
to get to safety.
Derived jumper
Stebbins and Cohen, 1995
Anura (frogs)
Jumping
(lose mass, add rigidity)
- inflexible vertebral column
- caudal vertebrae fused to form urostyle
- loss and reduction of skull bones
- loss of teeth in lower jaw (except one species)
- anterior ribs reduced or lost
- radius-ulna and tibia-fibula fused
- elongate ankle bones (astragulus-calcaneum)
transfers energy to trunk during push-off
- limbs folded under body (greater propulsion)
Anura (frogs)
Jumping etc.
- skull reduced
- 9 or fewer vertebrae
- fused bones
limbs, urostyle
- no tail
- elongate hindlimbs
ankles, feet
Fig. 3-16
Pough et al. 2001
Anura (frogs)
Pectoral Girdles
1. arciferal
- widespread
- ancestral
2. firmisternal
- derived
- systematics
- intermediate conditions
- reversals
Anura (frogs)
Reproduction (typically external)
Amplexus:
1. inguinal
2. axillary ------>
3. cephalic
Eggs deposited:
- in water
- on land
- suspended in air
Stebbins and Cohen, 1995
Development:
- complete metamorphosis
- direct development
- viviparity
Anura (frogs)
Reproduction
Parental Care:
not typical, but...
Darwin frog, vocal sac
Stebbins and Cohen, 1995
Stebbins and Cohen, 1995
Anura (frogs)
Reproduction
Metamorphosis:
Tadpole (Larvae)
- herbivorous filter feeders
- some carnivorous
- or cannibalistic
- some don’t feed at all
Adult
- mostly carnivorous
- two frugivorous species
- tongue attachment
Modify gut, body plan,
mouth, jaws, limbs etc.
Stebbins and Cohen, 1995
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, > 4,300 spp.
Fig. 3-20
Pough et al. 2001
Anura (frogs)
Larvae
Surface film
Internal Gills
Stebbins and Cohen, 1995
Suctorial
scraper
Carnivore
Use to ID
Stebbins and Cohen, 1995
Anura (frogs)
Larvae
General types based on morphology and diet
Keratinous mouth parts
1
2
3
spiracles
4
spiracle
Pough et al., 2001
Anura (frogs)
Sound
Most important anuran communication tool
- Most anurans vocalize
- Usually male
- Inflate throat sacs
- species specific
calls
- mate attraction
- territoriality
- alarm calls
- more later...
Anura (frogs)
Skin
Specialized for multiple functions
- hydration, defense, locomotion, thermoregulation,
reproduction, respiration etc.
2
1
glands
- mucus (2 types)
- poison = granular
pigmentation
Fig 2.1
Stebbins and Cohen, 1995
Anura (frogs)
Skin
Specialized glands
A. mucus
-1 moist coating
prevent dehydration
-2 sexual dimorphism
nuptial pads
Dendrobates
tinctorious
Nuptial
pads
Pough et al., 2001
B. poison = granular
-defensive secretions
(sticky, antibiotics, toxic, etc.)
Anura (frogs)
Skin
Pigmentation (Chromatophores)
Xanthophores
Iridophores
Melanophores
-reflectance
-infrared
-aposematic
Stebbins and Cohen, 1995
Pigmentation under hormonal control
Anura (frogs)
Skin
Respiration
- blood vessels
- epidermal thickness
- versus hydration
gas transfer requires ‘solution’
Stebbins and Cohen, 1995
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, 4,300 spp.
Fig. 3-20
Pough et al. 2001
Anura (frogs)
Ascaphidae
(1 species; ~NW U.S., into Canada)
- monotypic Family and Genus
- ‘tailed frog’
- tail is extension of cloaca
- internal fertilization (1 of ~3 spp.)
- do not call
- no tympana
- highly turbulent aquatic habitat
Ascaphus truei
Anura (frogs)
Leiopelmatidae
(1 genus, 3 species; New Zealand)
- only amphibians native to NZ
- ribs embedded in ventral musculature
- lack ear components and vocal sac
- direct development
- male parental care in 2/3 species
Leiopelma hochstetteri
Leiopelma hamiltoni
Anura (frogs)
Bombinatoridae
(2 genera, 8 species; Europe, E Asia)
- Bombina
toxic skin secretions
aposematic coloration (~orange + yellow)
unken reflex
Bombina orientalis
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, 4,300 spp.
Fig. 3-20
Pough et al. 2001
Anura (frogs)
Pelobatidae
(3 genera, 11 species; N America, Eurasia)
- include ‘spade foots’
- well-developed keratinous, spade-like
metatarsal
tubercle on hind feet
- fossorial
- often with enlarged parotoid glands on dorsum
- desert adapted
aestivate
explosive breeders
egg to metamorph in ~8 days
Anura (frogs)
Pelobatidae (con’t)
Scaphiopus couchii
spade
Anura (frogs)
Rhinophrynidae
(1 species; Texas to Costa Rica)
- monotypic family and genus
- ultra fossorial
spade tubercles
pointed head with cornified skin
small eyes
no tympanum
reinforced skull
- feed on ants and termites
underground
- unique tongue
Rhinophrynus dorsalis
Anura (frogs)
Pipidae
(5 genera, ~30 species; Panama, S America, sub-Saharan Africa)
- no tongue
- keratinous ‘claws’
- aquatic
- modified ears and calling
- Xenopus laevis
research
polyploids
- Pipa spp.
eggs in dorsal
skin of female
X. tropicalis
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, 4,300 spp.
Fig. 3-20
Pough et al. 2001
Anura (frogs)
Bufonidae
(33 genera, ~400 species; widespread, not in Australopapuan region)
- Bidder’s organ
rudimentary ovary on male testes
(paedomorphic)
- no teeth
- parotoid glands
toxins
- Bufo marinus
pest
Anura (frogs)
“Leptodactylidae“
(49 genera, > 900 species; Americas, West Indies)
- not monophyletic
- variable
- Eleutherodactylus
> 500 species!!
most species rich
genus of vertebrates
some with direct dvpt.
2 spp. with internal fert.
Ceratophrys
cornuta
Eleutherodactylus
auriculatus
Anura (frogs)
Myobatrachidae
(21 genera, 120 species; Australia, New Guinea, Tasmania)
- variable
- one species with
male inguinal pouches
for brood rearing
- two species with
gastric brooding
in female
Rheobatrachus
(extinct?)
Rheobatrachus vitellinus
Anura (frogs)
Rhinodermatidae
(2 species; Argentina and Chile)
- fleshy proboscis at
tip of snout
- males carry larvae
to water, or brood
in vocal pouch
Rhinoderma darwinii
Anura (frogs)
Hylidae
(38 genera, 760 species; Americas, W indies, Australopapuan region)
(Hyla in Americas, Eurasia, Japan, N Africa)
- arboreal generally
- well-developed toe discs
- claw-shaped terminal phalanges
- many spp. with females that
transport young dorsally
- Phyllomedusa
toxin for native rituals
lipid glands in skin
wiping behavior
Phyllomedusa sauvagi
Anura (frogs)
Centrolenidae
(3 genera, > 130 species; Mexico to S America)
- medial process on 3rd metacarpal
- toe discs
- terminal phalanges T-shaped
- eggs attended by males
- transparent venter =
‘glass frogs’
Centrolenella
fleischmanni
Centrolenella oyampiensis
Lissamphibia
Anura
(frogs)
~28 families, 310
genera, 4,300 spp.
Fig. 3-20
Pough et al. 2001
Anura (frogs)
Dendrobatidae
(6 genera, 185 species; Cent. America, N South America)
- pair of dermal scutes on dorsal surface of fingers
- generally diurnal and terrestrial
- cephalic amplexus (if amplex)
- parental care
move larvae around
- Dendrobates
may feed tadpoles eggs
- Phyllobates terribilis
lipophilic alkaloids
dietary origin
Dendrobates
tinctorious
Anura (frogs)
Arthroleptidae
(7 genera, 75 species; sub-Saharan Africa)
- includes Trichobatrachus robustus (hairy frog)
male sits on clutch and aids aeration?
Stebbins and Cohen, 1995
Anura (frogs)
“Ranidae”
(46 genera, > 700 species; most everywhere, except many islands)
- likely not monophyletic
- variable
- some unique Rana esculenta hybrids
both sexes represented
alternate generations
sexual <---> asexual
Rana pipiens
Anura (frogs)
Hyperoliidae
(19 genera, 230 species; sub-Saharan Africa, Seychelles, Madagascar)
- many arboreal
- toe discs
- ~ brightly colored
- Afrixalus
eggs on leaf
then ‘taco’
Afrixalus osorioi congicus
Anura (frogs)
Rhacophoridae
(15 genera, 315 species; Africa, Madagascar, Asia)
- mostly arboreal
- enlarged toe discs
- some with foam nests
- some communal nests
Buergeria japanicus
- Mantella with skin alkaloids
and aposematic coloration