The Adaptive Ecology of the Species Groups of the Genus Leptodactylus (Amphibia,

Leptodactylidae)
Author(s): W. Ronald Heyer
Source: Evolution, Vol. 23, No. 3 (Sep., 1969), pp. 421-428
Published by: Society for the Study of Evolution
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 THE ADAPTIVE ECOLOGY OF THE SPECIES GROUPS OF THE
GENUS LEPTODACTYLUS (AMPHIBIA, LEPTODACTYLIDAE)'

W. RONALD HEYER
Division of Amphibiansand Reptiles,
Field Museum of Natural History, Chicago 60605

Received January 29, 1969

Frogs of the Neotropicalgenus Lepto- ica. The relationships among the species
dactylusprovidea clear exampleof one groupsare easiestexplainedvisually(Fig.
way in which a group of amphibians 1).
evolved froman almostaquatic to a ter-
THE ADAPTIVE ECOLOGY
restriallifehistory.The genusis comprised
OF THE SPECIES GROUPS
of approximately 30 species,but the tax-
onomyand relationships of manyof these The principalchange in mode of life
species are confused. My initial interest observablein Leptodactylusis fromripar-
in the genuswas to attemptto determine ian (in the broad sense) to terrestrial
if clusters of related species (species habitats. Fortunately,the species groups
groups) could be recognized, and if so, to presenta seriesof gradesin this process,
determinethe relationshipsamong the allowingtheprobablehistoryof theadapt-
species groups. The followingdiscussion ive shiftto be discerned.
uses as a backgroundthe synthesisof the Simpson(1947) laid down the founda-
species groupsand theirinterrelationshipstions of the adaptive zone concept. The
(Heyer, 1969). adaptive zone hypothesisis a conceptual
means of explainingthe origin of new
SYNOPSIS AND RELATIONSHIPS adaptivekindsof organisms.The useful-
AMONG THE SPECIES GROUPS ness of the concept has been treated
The fivespeciesgroupsand the species recentlyin a symposium(SystematicZool-
in each groupare: 1) Fuscus group-L. ogy,Volume14, Number4, 1965). Simp-
bufonius, fuscus,gracilis,labialis,labrosus, son's (1947) basic featuresof theadaptive
latinasus,mystaceus,mystacinus,poecilo- zone concept, pertinentto the present
chilus, ventrimaculatus;2) Marmoratus discussion,are these: (1) Adaptivezones
group-L. hylaedactylus, marmoratus;3) are subdivisionsof the environment based
Melanonotusgroup-L. dantasi,discodac- on broad adaptive types as evidencedby
taxonomicsegmentsof the biota; (2)
tylus,melanonotus, podicipinus, pustulatus,
Major adaptive zones are separated by
wagneri;4) Ocellatusgroup-L. bolivianus,
discontinuitiesor essentiallyinstableeco-
ckaquensis, ocellatus; 5) Pentadactylus
logical zones; (3) The change fromone
group-L. laticeps, pentadactylus,rho- zone to anotheris usuallyundertakenby
domystax,rhodonotus, rugasus. This is a a smallgroupof organismsat a veryrapid
conservativelist. The only species group
evolutionaryrate; and (4) A radiation
compositionconsidered accurate is the ensues in a newlyenteredadaptive zone.
Melanonotusgroup,as it is theonlygroup
Once thisradiationhas occurred,theforms
to have undergonerecentrevision(Heyer, in the transitional zonesare comparatively
in press). Members of each group are
ill-adapted and rapidly become extinct.
distributed in Centraland/orSouthAmer-
More recently, workers(particularlyBock,
1965; Hecht, 1965; King, 1965; and von
'This paper formedpart of a thesis submitted
in partial fulfillment
of the requirementsfor the Wahlert,1965) have concentrated on the
Ph.D. degree,Universityof SouthernCalifornia. processesinvolvedin the transitionfrom
EVOLUTION 23:421-428. September,1969 421

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422 W. RONALD HEYER

FIG. 1. Relationshipsamong the species groups of the genus Leptodactylus. Dark circles indicate
point locations of the species groups. F = Fuscus group, P = Pentadactylusgroup, Me = Melano-
notus group, 0 = Ocellatus group, Ma = Marmoratus group. Solid concentriccircles indicate total
number of evolutionaryadvancementsfor each of the groups. Dashed lines indicatethe way in which
the groups are related. Of all the groups,the Melanonotus and Ocellatus groups are most closely re-
lated to each other. The Pentadactylusgroup is then relatedto thesetwo groups,and the Fuscus group
is related to the clusterof the threegroups. The Marmoratusgroup is not closelyrelatedto the other
four groups. Taxonomicallyinterpreted,the genus Leptodactylusis composed of two subgenera; the
subgenus Leptodactyluswhich includes the Fuscus, Melanonotus,Ocellatus,and Pentadactylusgroups,
and the subgenusAdenomerawhich includes the Marmoratusspecies group.

one majoradaptivezone to another.These ing considerableamountsof mucus from

workersemphasizethat: (1) Evolutionary theskinand the femalereproductive tract.
changesin transitionalzonesare via micro- Relativelyfew have utilizedsuch mucus
evolutionarymeans,althoughthe rate of to forma foamnest. The adaptationap-
genetic change is rapid; (2) Ecological pears to be mainlyone of behavior,al-
shifts are very important,but are not though concurrentlythere is probably
as easily documentedas morphological selectionfor increasein mucus secretion
changes; and (3) All transitionalforms eitherby an increasein numberof mucous
are fullyadaptedto theparticularenviron- glandsor an increasedproduction of mucus
mentin whichtheyexist,whileat thesame fromexistingmucousglands.
timepossessingcertaincharacteristics(pre- Several selectivefactorsact in concert
adaptations) which allow evolutionary to bringaboutpopulationalmovement into
changeto a new adaptive type given the a differentadaptive zone. The major
proper selective pressuresin a suitable forcesthatwouldoperateon an anuranto
environment. place the eggsin a foamnestout of water
One of the mostunusualfeaturesof the for any portionof developmentinclude:
genus Leptodactylus,and one which is predationon eggs and larvae in a pond-
sharedby certainotherleptodactylid gen- marsh situation,larval competitionin a
era in SouthAmericaand Australia,is the pond-marshsituation,and desiccationof
foam nest. The nest is constructedof a temporary watersource.
glandularsecretionsbeaten into a froth, The main predatorsfeedingin a pond
the consistencyof whichis quite like a utilizedby mostamphibiansforreproduc-
beatenegg white. The eggs are deposited tive and developmental purposesare not
in the nest duringits construction.Many fish,as the ponds used are characteristi-
frogs,probablymost,are capable of exud- cally seasonalor rathertemporary.Water

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 ADAPTIVE ECOLOGY OF LEPTODACTYLUS 423

insects,carnivoroustadpoles,and snakes riparianfrog,feedingand breedingin the

are themajorpredators.A foamnest,even water-land interface. The frogs move
one floatingon top of water,protectsthe about only amongplants characteristic of
nestcontentsfrommanyof thesepredators thewateredgeor marshand breedin water
foras longas thenestis intact. The genus so shallowthat whentheysit on the bot-
Leptodactylusinhabits,and probablyin- tom,theirheads are above water.
habited throughmuch of its evolution, The genusLeptodactylushas been able
regionscharacterized by markedlyseasonal to make the transitionfroma life-history
rainfall. Resistanceof reproductive prod- closelytied to waterto a moreterrestrial
ucts to desiccationis alwaysan important ecology. The Pentadactylusgroup shows
factorin the successof a specieslivingin the firststep towardsa more terrestrial
such an environment, particularlyif the life,if the pre-adaptations
presentin the
rainfallis erraticduringthe breedingsea- Melanonotus-Ocellatus groupsmay be re-
son. While eggs or developinglarvae are garded as primitive.Apparently, L. pen-
withina foam nest, most are protected tadactyluscongregateat bodies of water
fromdesiccationbecause of the insulating only forbreedingpurposes. The frogsare
effectof the nest. Certainof these nests oftenfoundin largeburrowsand breeding
are rathersturdy.In thesummerof 1967, may occurat theseburrowsor in potholes
I preservedsome nestsof L. melanonotus next to water (Breder, 1946) or at the
by placingthe nestson top of a formalin margins of ponds (Bokermann, 1957).
solutionin jars. The jars were violently Breder (1946) observednests of L. pen-
shaken to preservethe eggs,but the nest tadactylusin potholes,eitherin contact
remainedintact. The 3000 mile journey withsmallbodiesofwateror littleremoved
by truckto Los Angeleswas not smooth, frombodies of water. He also observed
thusthe nestswerebouncedconsiderably; that nests were torn apart by torrential
yet,I had to dissectthenestswithteasing rains, and the advanced larvae escaped
needlesin thelaboratoryin orderto count intotherisingwater. The advantagesthat
the eggs. membersof the Pentadactylusgroupshare
One may visualize the followingseries are: (1) theeggsare protectedfrompond
of eventsin a drainagebasin characterized predators,and (2) the eggs are freefrom
by sporadicrainfall.A frogplaces its eggs desiccation.
in a foam nest duringone brief rainy The nextadaptivestep appearingin the
period afterwhichthe pond dries up. A Fuscus groupis placingthe foamnest on
subsequentheavy rain comes which de- land in a burrowfashionedby the male.
stroysthe nest and releasesjust hatched The males call fromtheseburrowslocated
larvae into the newlyrefilledpond. The just under the surfaceof the soil. The
offspring ofsucha frogwouldhave a lower burrowsusuallyhave a singleaccess hole
mortalityrate than thoseof a froglaying to the surface.The femaleis attractedto
its eggs directlyin the water, assuming the burrowwherematingtakes place and
comparableperiodsof egg depositionand the foamnest is formed.Larval develop-
larval development.As it probablywould ment is initiatedin the burrow,but the
take a veryheavyrainto destroythenest, larvaeare eitherliberatedintoan adjacent
the larvae would be assuredof water for pondwhena heavyrainfloodstheburrow,
the durationof theiraquatic development. or perhapsfloodedfromthe burrowwhen
The Melanonotusand Ocellatus groups thelevelof theadjacentpond rises(Dixon
reflecttheprimitive Leptodactyluspattern and Heyer,1968; Fernandezand Fernan-
in that the foamnest is placed on top of dez, 1921; Lutz, 1947).
the water (Fernandez and Fernandez, In additionto the adjustmentto erratic
1921; Lutz, 1947; pers.obs.). Duringthe rainfall,a significantadvantage of the
breeding season,L. melanonotus foamnest was demonstrated
is a typical terrestrial in

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424 W. RONALD HEYER

a series of observationson L. labialis in has withrespectto earlydevelopment are:

westernMexico. On July 11, 1967, Dr. (1) the eggs are not taken by pond or
JamesR. Dixon and I stoppedat a dam groundsurfacefeedingpredators,(2) the
2.4 km E and 9.6 km S of the town of eggs are freerfromdesiccationthan those
Colima on the road fromColima to Teco- of the exposednests of the Melanonotus,
man in the state of Colima. The dam Ocellatus,and Pentadactylusgroups (the
containedno standingwaterand the earth surfaceof a nest will encrustif exposed
was cracked,but the cracks were moist. long enoughand any eggs located in the
The onlyfrogscallingin thedaytimewere encrustedfoam are destroyed),(3) the
several male Leptodactylus.One of the larvaepossiblyavoid muchfoodand space
males capturedat that time was calling competition in thepond,and (4) thelarvae
froma 20 X 30 mm,almostsphericalbur- metamorphose beforemany other species
row. Veryearlyin themorning of July12, usingthesame pond.
after a very heavy thunderstorm, water The Marmoratusgroup has the most
rose behind the dam wall. We returned specializedtype of reproductive behavior
to the dam in theeveningwhileit was still and larval development.Foam nests are
raininglightly,at whichtimetherewas a laid in almostsphericalincubatingcham-
frenzyof breedingactivity. Frogs of six bers like thoseof the Fuscus group. The
species includingL. labialis, were calling larvae hatch and depend entirelyon the
and matingin the pond. I sampled the large yolk stores of the egg for growth
pond on the morningof July 14, and the (Lutz, 1947). The larvae do not have
tadpoles of L. labialis were noticeably toothrowdenticlesin the mouthparts and
advanced over those of Bufo marmoreus lack the spiraclecharacteristic of aquatic
and Hypopachusvariolosus,the otherlar- Leptodactyluslarvae (Heyer and Silver-
vae that had hatched. On July24, meta- stone,in prep.). The larvaemetamorphose
morphosingL. labialis were leaving the within the incubatingchamber without
pond,the firstspeciesthatwe observedto everhavinga free-living aquatic periodof
do so. We found several foam nests of growthand development.The Marmoratus
L. labialis near the dam, some of which groupis entirelyindependentof standing
containedlarvae capable of swimming.L. bodies of waterduringthe life cycle; the
labialishad beenable to matebeforestand- speciesof thisgroupare trulyterrestrial.
ing water was present. The eggs had The shiftfroman aquatic to a terrestrial
matured in the nest and hatched; the life in frogsinvolvesa significant number
larvae had developed functionalmouth- of changes and any intermediateform
parts and enclosed the gills. When the mightbe predictedto be ill-adaptedto its
rainscame and the pond filled,the larvae environment. Nevertheless,a changefrom
were releasedfromtheirnestsand had a a riparianto a terrestrial life history,as
temporaladvantageover the otherspecies evidencedby thegenusLeptodactylus, can
utilizingthe pond. The advancedstage of be explainedby callingon relativelyfew
developmentwhen the pond firstformed genetic changes. Most of the genetic
resultedin an earlymetamorphosis forL. changeswithinLeptodactylusleadingto a
labialis. terrestrial
lifeare behavioral,e.g. whipping
The reproductive behaviorof L. bufo- mucoussecretionsinto foam,or digginga
nius,as reportedby Cei (1949), contrasts burrowforthenest. Associatedwiththese
withthatof L. labialis in certainrespects. modifications of behaviorare largereggs
Cei reportedthat the femalesexcavate a and a reductionin the total numberof
burrowand call males; in L. labialis,the eggsproduced(Fig. 2).
onlyspecimenswe foundin burrowswith- The importanceof the ecologic shift
out nestswerecallingmales. Nevertheless, towardsterrestriality in the genus Lepto-
themainadvantagesthattheFuscus group dactylusis reflectedby certainadaptive

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 ADAPTIVE ECOLOGY OF LEPTODACTYLUS 425

TERRESTRIAL NO FREE LIVING LARVAE

~~~~~~~~~~.~~~~~~~~NEST
/
PLACED IN INCUBATING
CHAMBER. LARVAE NOT
AQUATIC(MARMORATUS
GROUP)

INESTPLACED IN INCUBATING
It HAMBER. AQUATIC LARVAE
........ ~~~~~(FUSCUS
GROUP)

............. NEST PLACED IN CAVITIES

..
................... ADJACENT TO WATER
.................... (PENTADACTYLUS GROUP) 4

. ...........G
S)............*... RO P

AQUATIC EGGSS
LLAIDN INAWATER

FIG. 2. The adaptive shiftdemonstratedby the speciesgroupsof the genusLeptodactylus. The large
terrestrialand aquatic zones are above and below the transitionalriparianzone. Riparian in this con-
text means any freshwater-dryland interface. See text for furtherexplanation.

morphologicalcharacters. Most of the The species do not undergoamplexusin

specialized charactershave been derived an aquatic situation. Thumb spines are
in parallel in the Fuscus, Melanonotus, absent in males of the more terrestrial
Ocellatus,and Pentadactylus groupson the species of the Fuscus and Marmoratus
one hand, and the Marmoratusgroupon groups.
the other. The trendtowarda more ter- Toe fringesin adults are characteristic
restriallife has apparentlyoccurredtwice of membersof the Melanonotusand Ocel-
withinthe genusLeptodactylus. latus groups. Extensive toe ridges are
Male thumbspinesapparentlyfacilitate presentin juvenilesof the Pentadactylus
amplexus. Males of certainspeciesof the group,but are lost duringontogeny.Toe
Melanonotusand Ocellatus groups have fringesor ridgesare absentin bothjuvenile
thumbspines. Matingpairsofthesespecies and adultmembers of theFuscusand Mar-
are quite active,as the male beats up the moratusgroups. Toe fringesgive a larger
foam nest while the pair is in amplexus. surfacearea to thefoot,whichis important
Thumb spineswould seem to help anchor to species associatedwith water,such as
themale to the femalein an aquatic situa- membersof theMelanonotusand Ocellatus
tion. Males of the large-sizedPentadac- groupsare. Toe fringesallow membersof
tylusgroupalso have thumbspines. Pres- the Melanonotusand Ocellatusgroupsto
ence of thumbspines (and chest spines) be effectiveswimmers whenin the water.
wouldappear to,be necessaryin anchoring The fringesadhereto thesides of the toes
a matingpair due to the large adult size. when the frogsare out of water,and do

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426 W. RONALD HEYER

not impede terrestriallocomotionas a Australianleptodactylidgenus Kyarranus

membranous web might. which have terrestriallarvae have large
In the Melanonotus, Ocellatus, and yolkedeggs (diameter3.2 mmat gastrula-
Pentadactylusgroups in which member tion), no tooth denticles,but do have a
speciesdepositthefoamnestin an exposed spiracle (Moore, 1961). Zachaenus par-
situation, the eggs have melanophores vulus,a SouthAmericanleptodactylid with
apparentlyin orderto act as a shield to terrestrial larvae, has a large egg (6 mm
ultravioletlight. Egg melanophoresare includingjelly envelope), tooth denticles
absentin membersof theFuscusand Mar- present(but the total numberis reduced
moratusgroups. in comparisonwith an aquatic, feeding
A trendof decreasingnumbersof eggs tadpole), and no spiracle (Lutz, 1943).
together withan increaseof individualegg Reductionof toothrowdenticlesin a larva
diametercorrelatedwith terrestriality is withlargeyolkstoresis doubtlessa result
evident.Membersof theMelanonotusand of thelarvanothavingto utilizeits mouth-
Ocellatusgroupslay from1000-2000eggs parts for feeding.The loss of a spiracle
of 1.0-1.5 mm diameterper nest. (No may be imposedby the physicalproblems
Pentadactylusgroup nests were available associated with large yolk stores. Large
for analysis.) Members of the Fuscus eggsize appearsto be theimportant deter-
grouplay from50-850 eggsof 1.6-2.5 mm minantin allowingany shifttowardster-
diameterper nest. Membersof the Mar- restrialityin anuran larvae; tooth row
moratusgrouplay from4-25 eggsof 2.1- reductionand loss of the spiraclemay or
3.0 mmdiameterper nest. maynotfollowas a consequenceofincreas-
Larvae of members of the Fuscus, ingtheyolkstores.
Melanonotus,Ocellatus,and Pentadactylus A change apparentlycorrelatedwith a
groupsare all quite generalized,havinga fossoriallifeis a strengthening ofthebones
%/3toothrow formula, and a sinistralspir- of the nasal region.Membersof both the
acle. The terrestrial larvae of the Mar- Fuscus and Marmoratusgroups have a
moratusgrouplack toothrowdenticlesand more rigidbony structureof the anterior
a spiracle. regionof thehead thando membersof the
Large egg size, reductionof larval tooth other species groups. The snout may
rows, and absence of a spiracle do not actuallybe used in the construction of the
necessarilyindicate that a froglarva is incubatingchamber,but thishas not been
terrestrial,or vice versa. Kalophrynus observed.In the Fuscus group,the snout
pleurostigma, a microhylid,has large eggs is strengthened by a calcificationof the
(diameter at mid-gastrulation, including anteriorsphenethmoid region,and in the
envelope,5 mm), has a spiracle (micro- Marmoratus groupthesnoutis strengthened
hylid larvae lack tooth denticles), and by an enlargementof the nasal bones
dependssolelyuponyolkstoresfornourish- (nasal 10.6% of theskullarea in a 26 mm
ment until metamorphosis (Inger, 1956). specimenof L. marmoratus, only 8.6% in
The larvaeare aquatic,however, inhabiting a 32 mmspecimenof L. latinasus,thenext
small waterfilleddepressionsin the forest smallest species of Leptodactylusexam-
(Inger, 1956). Pelophrynebrevipes,a ined).
bufonid,has large yolked eggs (egg size The family Leptodactylidae includes
notavailableforP. brevipes,diameter2.0- membersshowingevery stage in a con-
2.5 mm in P. albotaeniataand P. lighti), tinuumfroman aquatic to a terrestrial life
a reductionof toothrows,no spiracle,and history.Severalof the intermediate forms
probablydoes not feed; yet the larva is have foamnests. The routesto terrestri-
aquatic, inhabitingverysmall water-filled alityin thefamilymayhave beenas shown
depressions, such as brokenbottlesin the withinthe genus Leptodactylus,through
forest (Inger, 1960). Members of the evolutionof a foamnest.

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 ADAPTIVE ECOLOGY OF LEPTODACTYLUS 427

The genus Leptodactylusprovides an forAmphibiasuggeststhatselectionfora

exampleof an almostcompletetransition terrestrialway of life has continuedin-
fromtheaquatic to theterrestrial adaptive tenselythroughout geologichistory.
zones. Its example lends supportto the
adaptive zone concept,particularlywith SUMMARY
respectto four aspects of the transition Five speciesgroupswithinthefroggenus
zone: (1) Each of the individualsteps Leptodactylus are recognized;theadvanced
along the transitionis small; (2) Each Fuscus and Marmoratusgroups,theprimi-
speciesin the transitionzone is adapted to tive Melanonotusand Ocellatus groups,
its environment;(3) Each of the species and the intermediate Pentadactylusgroup.
groupshas certaincharacterswhichserve The speciesgroupsforma seriesofstages
as adaptationsin themselves.These same bridgingthe transitionbetweenthe major
charactersprobablyservedas the base for aquatic and terrestrial habitats. The ex-
furtheradaptation,allowinga more ter- ample of evolutionin Leptodactylusem-
restriallife. The most strikingexample phasizesthat: (1) the adaptivetransition
is theplacementof thefoamnest; and (4) fromwaterto land occursby microevolu-
The most notable featuresof the shift tionarymeans,(2) theformsin thetransi-
pertain to the ecologies of the species tion have certain charactersthat likely
groups. servedas preadaptationsin the ancestral
The Leptodactylusexampleclarifiescer- stocksof the moreterrestrial species,and
tain details of the adaptive zone concept (3) the transition betweenmajor habitats
whichmighthave not been predictedfrom is closelyassociatedwiththe reproductive
the fossilrecordalone. The groupunder- biologyand behaviorof the formsin the
goingtransition betweentwomajoradapt- transitionzone. The Leptodactylusex-
ive zones has usuallybeen thoughtof as ampledemonstrates thatselectionpressures
a small-sizedgroup. Usually,however,it have continuallyoperatedon anurans to
is fossilgenerathatare givenas examples move towards terrestriality, resultingin
for groups going throughthe transition. multipleentriesintotheterrestrial adaptive
The Leptodactylusexample emphasizes zone.
thata smallgroup,belowthegenericlevel,
may pass throughthe transitionbetween ACKNOWLEDGMENTS
two major adaptive zones. More impor- My graduatecommitteeat the Univer-
tantly,however, theLeptodactylus example sity of SouthernCaliforniaofferedmany
indicatesthat the transitionzone may be constructive commentson this sectionas
stable. Simpson'smodel indicatesthat a it appeared in the dissertation-JayM.
transition zone existsfora shortperiodof Savage,JohnS. Garth,JohnL. Mohr,Basil
timeonly. The argumentfora shortlived G. Nafpaktitis, and JohnD. Soule. Robert
transitionzone is that once a transition F. Inger,Field Museum of Natural His-
zone has been passed through,it should tory,criticallycommented on the paper in
neverreopen.MembersofthegenusLepto- its presentform.
dactylushave re-entered a transition zone, Withoutthe help of the followingfield
and are presentlyexploitinga transition companions,my insights of the genus
zone. Goin (1960) indicatedthat there Leptodactyluswould be lessened: James
are numerous amphibianlifehistories inter- R. Dixon,mywife,MiriamHeyer,Roy W.
mediatebetweena completely aquatic and McDiarmid, Norman J. Scott, Jr., and
a completelyterrestriallife cycle. Am- CharlesF. Walker.
phibians,thefirstvertebrate groupto enter I acknowledgethe support from two
the transitionbetweenthe aquatic and the NationalScienceFoundationSummerFel-
adaptivezones,still occupythe lowships for Teaching Assistantsin the
terrestrial
transitionzone. The stabletransition zone summersof 1964 and 1965, an NDEA

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428 W. RONALD HEYER
DissertationTravel Fellowship adminis-
teredby the Universityof SouthernCali-
forniafor travelin the summerof 1967,
and a National ScienceFoundationGrad-
uate Traineeshipfor1966-68.
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