Advertisement Calls And Relationships Of Chilean Frogs Eupsophus contulmoensis And E-Insularis (Amphibia, Anura, Leptodactylidae)

7 Pages·1994·2.2 MB·English

by  J R Formas

PROC. BIOL. SOC. WASH.
107(2), 1994, pp. 391-397
ADVERTISEMENT CALLS AND RELATIONSHIPS OF
CHILEAN FROGS EUPSOPHUS CONTULMOENSIS
AND INSULARIS
E.
(AMPHIBIA: ANURA: LEPTODACTYLIDAE)
J. Ramon Formas and Lila Brieva
ylfo?rac/.—Advertisement calls oiE. contulmoensis and E. insularis are de-
scribed.Thecallsofbothspeciesconsistofasingleshortnote.Calldata,together
with chromosomal information, indicate that E. contulmoensis, and E. insu-
laris, E. roseus, E. calcaratus and E. migueli comprise a related group distinct
from E. vertebralis and E. emiliopugini.
Calls of frogs can be useful in revealing Methods and Materials
evolutionary relationships at the species
level. Ingeneral,closelyrelatedspecieshave Field recordings were made at 19 cm/sec
some acoustic characteristics in common on an Uher 4000 Report-IC portable tape-
(Blair 1958, Barrio 1965, Mecham 1971, recorderandanUher 517 microphone. Call
Martin 1972,Schiotz 1973, Kuramoto 1974, durations,numberofnotespercall,andnote
Ryan 1988, Straughan & Heyer 1971). In duration were analyzed with a storage os-
cilloscope Tektronix 5113. Audiospectro-
order to establish evolutionary trends re-
grams were made with a Kay Elemetric au-
latedtothediversificationofthegenusEup-
diospectrograph model 675, using 85-8000
sophus, weanalyzedtheadvertisementcalls
Hzfrequencyscaleandnarrow(45 Hz)band
ofE. contulmoensis and E. insularis. Both
calls are compared with those other mem- filter. Callrepetitionratesweremeasuredin
the field by counting consecutive calls over
bers ofthe genus that were previously de-
scribed by Formas & Vera (1980), Formas one minute. Water temperature, location
(1985, 1989), and Penna & Veloso (1990). and behavior ofthe individuals used in the
analysisweretakenatthetimeofrecording.
Eupsophus is the most speciose taxon of
Specimens and tapes were deposited in the
the rather reduced anuran fauna of the
collection ofAmphibians ofthe Institute of
Nothofagus forests of southern Chile and
Zoology, Universidad Austral de Chile
Argentina. Seven species have been de-
(IZUA), Valdivia.
scribed: E. roseus, E. calcaratus, E. migueli,
E. contulmoensis, E. insularis, E. vertebralis
Eupsophus contulmoensis Ortiz,
and E. emiliopugini. Ibarra-Vidal, & Formas, 1989
E. contulmoensishasbeencollectedinthe
type locality of Contulmo (37°52'S, Theadvertisementcallof£". contulmoen-
73°12'W), Nahuelbuta Range (Ortiz et al. siswasrecordedatthetypelocality(Natural
1989), andE. insularisisendemictoMocha Monument of Contulmo, Malleco Prov-
W
Island (38°22'S, 73°56'S; 38 km of the ince). Description is based on 25 calls from
coast ofArauco Province) (Formas & Vera seven specimens recorded at 1 1.8°C (water
1982). temperature)duringNovember 1990. Males
392 PROCEEDINGSOFTHE BIOLOGICALSOCIETY OFWASHINGTON
Table L—Call characteristic(mean and range)ofEupsophusspecies (S, shortcall; L, longcall)
Call Notes Calllength Dominant
Species type percall sec Repetitionrate frequency(Hz)
E. contulmoensis 1 0.18(0.15-0.20) 23.3(15-34) (1100-2000)
E. insularis 1 0.16(0.14-0.18) 7.8(4-12) (1500-2100)
E. roseus t s 1 0.20(0.19-0.21) 64.0(60-72) (1600-2900)
** s 1 0.10(0.07-0.16) 25.1 (11.1-60) (1250-1350)
** L 32.1(8-47) 2.73(0.65-40.0) 10.2(93-112) (1220-1470)
E. migueli ft S 1 0.24(0.20-0.35) 6.0(3-8) (1500-2500)
** s 1 0.20(0.16-0.26) 4.2(2.4-6.6) (1170-1820)
tt L 24.0(19-33) 3.40(3.30-4.40) 6.0(5-8) (900-1500)
** L 12.3(4-23) 1.07(0.30-2.16) 5.4(2.4-8.4) (1210-2000)
E. calcamtus tt 1 0.19(0.15-0.21) 19.0(16-25) (1100-2700)
E. vertebralis t 5.0(4-6) 0.60(0.40-0.80) 4.0(2-10) (1100-2500)
** 5.6 (3.8) 0.64(0.40-0.88) 27.8(18.6-36.6) (700-1110)
—
E. emiliopugini * 2 0.50 (729-1320)
tFrom Formas & Vera (1980); ttFormas (1985); *Formas (1989); **Penna& Veloso(1990).
wereobservedandcollectedwhiletheywere E. roseus (Dumeril & Bibron),
calling from cavities in the ground at the E. migueli Formas, E. calcaratus (Giinther),
border of a stream close to the forest. In- E. vertebralis Grandison,
dividualswere organizedinto duetsandtri- and E. emiliopugini Formas
os spaced apart by 40-180 cm. Generally
Callcharacteristicsoftheabovecitedspe-
males called in alternation. The advertise- cies were described previously (Formas &
ment call ofE. contulmoensis is composed Vera 1980; Formas 1985, 1989; Penna &
loafstosnl0y.1o5n-e0.n1o8tseec(Foingd.s.laT,hde;dToambilnean1)ttfhraet- Veloso 1990) and are depicted in Table 1.
quency is spread between 1100-2000 Hz
and the repetition rate is 15-34 calls/min- Discussion
ute. All calls showed modulation and de- It is noteworthy that all Eupsophus spe-
fined harmonics are present at about 500 cies call during the spring-time from cavi-
Hz intervals. ties along margins of small streams in the
forest (Formas & Vera 1980; Formas 1985,
Eupsophus insularis (Philippi) 1989; Penna & Veloso 1990). Among the
Description is based on 18 calls recorded anuran fauna ofthe temperate Nothofagus
from three individuals at Isla Mocha, dur- forest of South America, males of Eupso-
ing December 1989. Calls were recorded at phus species are unique in calling from
a water temperature of 12°C. Males were flooded underground cavities near streams.
observedandcapturedwhile theywerecall- Based only on temporal parameters (call
ing from cavities in the ground close to a length and notes per call) ofadvertisement
stream at the border ofthe forest. The in- calls, Formas (1985) established two clus-
dividualscall isolatedand neitherduets nor ters ofspecies within the genus Eupsophus.
trios were detected. Calls consist ofa single The first group contains E. roseus, E. mi-
note (Fig. lb, e; Table 1) lasting 0.14-0.18 gueli and e. calcaratus, whose calls consist
seconds. The call is modulated and defined ofonlyonenotelasting0.10to0.20 seconds
harmonics are present at about 500 Hz in- (Table 1). Calls off. contulmoensis and E.
tervals. Maximum energy is spread over a insularis (a single note lasting 0.14 to 0.18
frequency range of 1500-2100 Hz. seconds) provide good evidence for group-
—
VOLUME 107, NUMBER 2 393
kH:
8
7
6
5
10'
;-#«^
4
^
3 -
2 -
1 -
"T
1
sec.
sec.
1
Fig. 1. Audiospectrograms ofthe advertisement call ofEupsophus contulmoensis (a), E. insulahs (b), and
E. roseus(c), band filter45 Hz. Oscillogramsofthe advertisementcalloff. contulmoensis(d)and E. insularis
(e).
ingboth ofthese species within this species Both species groups (i.e., roseus and ver-
group. The second group includes E. ver- tebralis) are also supported when spectral
tebralis, whose call contains 4-6 notes last- parameters of the calls are considered.
ing 0.09 seconds. The advertisement call of Membersofthe vertebralisgroup showlow-
E. emiliopuginishowsthatthisspeciesemits er mean values of dominant frequencies
two notes lasting 0.20 seconds. Based on (1024-1350 Hz) than do members of the
this information, we consider that E. emi- roseusgroup (1550-1900 Hz) (Table 1). On
liopugini should be grouped with E. verte- the other hand, members ofthe vertebralis
bralis, rather than the roseus group. group exhibit marked pulsed calls (11-34
394 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OFWASHINGTON
3.7 -r ularphysicalfeaturesofthecallsitesofthese
frogs. When the mean dominant frequen-
cies oiEupsophus species are compared, E.
emiliopugini and E. vertebralis show lower
r =0.996
u values (1024-1350 Hz) than those present-
c y= 6.33-1.90 X ed by E. contulmoensis, E. insularis, E. ro-
a3> seus, E. calcaratus and E. migueli (1550-
cr
a> 1900 Hz) (Table 1). The relationship be-
tween the mean dominant frequency ofthe
Dc call varies inversely with the logarithm of
C animal size (r = -0.996) (Fig. 2). The ad-
oE vertisement calls of£". emiliopugini and E.
TJ vertebralis have been described as conspic-
o> uously pulsed (Formas 1989). The former
O
_J species emits 25.45 (17-34) pulses pernote
andthelatter 15.90 (1 1-23) pulsespernote.
In contrast, calls of E. roseus, E. migueli,
E. contulmoensis, E. insularis and E. cal-
caratus are poor in pulsation; however the
1.4 1.5 1.6 1.7 1.8 "B" call (orlongcall) of£". migueliis mark-
edly pulsed; 6-9 pulses per note (Formas
Log snout-vent length
1985).
Fig. 2. Relation between mean snout-vent length Eupsophus species exhibit considerable
and mean dominant frequency ofthe calls ofEupso- variation in calls, especially in the number
phus species. Snout-vent length measurements corre- ofnotes and their durations. Extremes are
spond to samples ofmales from localities where re- the single note (short call) ofE. roseus, E.
cordings were made. Abbreviations (sample size in migueli, E. calcaratus,E. contulmoensis, and
parentheses): E.r., Eupsophus roseus (12), E.m., Eup-
sophusmigueli(7), E.ca., E. calcaratus(6), E.i.,E. in- E. insularis (0.10-0.20 seconds), and the
sularis(5), E.co., E. contulmoensis(7), E.v., E. verte- complex trilled calls (long call) with more
bralis(14), E.e.,E. emiliopugini(17). than 40 notes ofE. roseus and E. migueli
(3.3-4.4 seconds). This latter vocalization
has been interpreted as a territorial call
pulsespernote), whilethe roseusgroup spe- (Formas 1985, Penna & Veloso 1990). Be-
cies show calls poor in pulsations. tween these extremes are the advertisement
Asindicatedearlier,differentauthorshave calls ofE. emiliopugini(2 notes)andE. ver-
suggested that calls of frogs are useful in tebralis (4-6 notes lasting 0.40 to 0.80 sec-
revealing systematic and evolutionary re- onds). Ifthe length ofeach individual note
lationships. Temporal and spectral param- is considered, the individual notes of the
eters of the advertisement calls of Eupso- call ofE. emiliopuginilasts0.20(0.19-0.23)
phus species may also reveal information seconds in duration, a value similar to the
concerningtheevolutionaryhistoryofthese single notes ofthe advertisement calls ofE.
species. Spectral components of the call roseus, E. calcaratus, E. migueli, E. con-
(dominant frequencies) show wide overlap lolmoensis. andE. insularis{X=0.18; range
within Eupsophus (Table 1). The frequency 0.14-0.26 seconds). On the other hand, in-
range (700-2900 Hz) at which these species dividual notes ofthe advertisement call of
call is also shared by other frogs ofthe tem- E. vertebralis have a duration {X = 0.089;
perate Nothofagus forest (Penna & Veloso range0.062-0.187)similartotheindividual
=
1990), and thus cannot be related to partic- notes of the long call of E. migueli (X
VOLUME 107, NUMBER 2 395
0.083; range 0.051-0.092). In spite ofthis E. emiliopugini) are also allopatric. In spite
similarity, both calls are different in the ofthe fact that species ofeachgroup {roseus
number ofpulses per note {X= 15.90 in E. and vertebralis) are allopatric, it is possible
vertebralis; X = 7.5 in £". migueli), number to find members ofthe two different species
ofnotes percall (4-6 inE. vertebralis; more groups in sympatry. For example, at the lo-
than 40 notes in E. migueli), and duration cality ofMehuin, E. vertebralis and E. mi-
ofthe call {X = 0.6; range 0.4-0.8 seconds gueliwere observed while calling from cav-
in E. vertebralis and X= 3.4; range 3.3-4.4 ities separated by two or three meters
A
seconds in E. migueli). (Brieva, personal observation). similar
Although members of the roseus group case of sympatry between members ofthe
exhibit some similar patterns in advertise- two different species group was observed at
ment calls, differences can be found when Cuestade Soto(7 kmbyroadfromValdivia
intervals among harmonics are examined. city) between E. roseus and E. vertebralis.
Calls ofE. contulmoensis and E. insularis In three localities from southern Chile (La
have harmonics at about 500 Hz intervals, Picada, 42°07'S, 73°49'W; Rio Rollizo,
whileE. calcaratus, E. migueliandE. roseus 4r27'S, 72°27'W; and Puntra, 42°07'S,
showharmonicsatabout 1000Hzintervals. 73°49'W) E. emiliopuginiand E. calcaratus
A comparison among the calls of£. roseus, were observed calling from cavities sepa-
E. contulmoensis and E. insularis is shown rated by 25-50 cm (Formas 1989). Finally,
inFig. 1. Inthecaseofthevertebralisspecies E. vertebralisandE. contulmoensiswereob-
group, both members {E. vertebralis and E. servedwhile callingfrom cavities separated
emiliopugini) have harmonics at about 250 bytwo to fourmeters atthe localityofCon-
Hz intervals (Formas 1989). tulmo (Natural Monument of Contulmo)
Recognizingtwo speciesgroupswithinthe (Formas, personal observation).
genus Eupsophus based on some temporal The four pairs ofEupsophus species re-
(call length and notes per call) and spectral cordedtooccurin sympatry{E. migueliand
(mean dominant frequencies and pulses) E. vertebralis; E. roseus and E. vertebralis;
characteristics ofadvertisementcalls agrees E. calcaratus and E. emiliopugini; E. ver-
with the same proposal based on chromo- tebralisandE. contulmoensis) are members
somal data (Formas 1980). Members ofthe of the two different species groups. These
vertebralisgroup {E. vertebralisandE. emi- pairs of sympatric species exhibit a strong
liopugini) share a 28-chromosome karyo- divergenceamongtheiradvertisementcalls.
type (Formas 1991), while members ofthe Passmore (1981) found that sympatric spe-
roseusgrouphavea 30-chromosomekaryo- cies of the African frog genus Ptychadena
type (Formas 1980, Iturra & Veloso 1981, showremarkabledifferencesintemporaland
1989). Cuevas (personal communication) spectral components oftheir calls. In order
foundthatE. insularisandE. contulmoensis to explain the originofthisphenomenon he
have a 30-chromosome karyotype, as oc- used the notion of the Specific Mate Rec-
curs in E. roseus, E. migueli, and E. cal- ognition System (SMRS) (Paterson 1978,
caratus. 1982). According to this view, interspecific
MembersoftheEupsophusroseusspecies differences in signals do not agree with the
group{E. roseus, E. calcaratus, E. insularis, traditional hypothesis that call divergence
E. contulmoensis, and E. migueli) are al- among closely related species develops due
lopatric species; however, E. roseus and E. to selection for increasing reproductive iso-
migueli are sympatric but not syntopic at lation upon re-encounter ofpreviously iso-
the locality of Mehuin (39°26'S, 73°10'W) lated populations (Dobzhansky 1970).
(Iturra & Veloso 1981). Members ofthe E. Passmore's (1981) explanation for Ptycha-
vertebralis species group {E. vertebralis and dena species is that the "pattern of vocal
396 PROCEEDINGS OFTHE BIOLOGICALSOCIETY OFWASHINGTON
divergenceprobablyresultedfromindepen- the roseus group have the same chromo-
dent evolutionary histories, and concom- somal formula (2N = 30), although differ-
itant with the development oftheir respec- ences have been found among the karyo-
tive acoustical recognition mechanism, types of some species. For example, the
rather than from selective pressure for in- karyotypesof£. roseusandE. migueli(Itur-
creasing reproductive isolation" (Penna & ra & Veloso 1989) exhibit marked similar-
Veloso 1990). ities, but sex chromosomes differ. In these
In light ofthe SMRS concept we propose two species, the Y chromosome is a small
a hypothesis addressing the divergence of metacentric one without paracentromeric
vocal patterns and the evolutionary history heterochromatin, and corresponds to pair
of the genus Eupsophus. As shown above No. 14. In E. roseus, the sex chromosomes
(Table 1)thevertebralisgroup{E. vertebralis havethesamemetacentricmorphology,but
and E. emiliopugini) exhibits two different in E. migueli the X chromosome is telo-
advertisement calls. Both members ofthis centric. This example ofchromosomal di-
species group occur allopatrically (Formas vergence between two members ofthe ro-
1989)andtheirkaryotypesshowdifferences seus group (which share the same temporal
inrelationtochromosomepairNo. 14(telo- and spectral characteristic ofthe advertise-
centric in E. vertebralis and metacentric in ment calls) suggests that chromosomal dif-
E. emiliopugini; Formas 1991). From an ferentiation has played a more important
evolutionary point ofview, the genus Eup- role inthe diversification ofsome members
sophushasalonghistory, with earlygeneric ofthe roseus species group than has the ad-
differentiation documented by fossil rec- vertisement call. Finally, based on chro-
ordsdatingbacktotheOligocene(Schaeffer mosome data, we suggest that vocal differ-
1949, Baez & Gasparini 1979). Conditions encesnotedforEupsophusspeciesthatoccur
for further acoustical and chromosomal di- in sympatry results from the presumed in-
versification within these species existed dependentevolutionaryhistoriesofthetaxa
duringPleistoceneglaciations,duringwhich involved.
reduced areas free ofice provided scattered
refugia for the survival of flora and fauna
Acknowledgments
(Vuilleumier 1968, Heusser 1974, Ash-
worth et al. 1991). Since the present geo- This study was supported by Fondo Na-
graphicalrangesof£. vertebralisandE. em- cional de Investigacion Cientifica y Tec-
iliopugini (especially the southermost part) nologica (Proyecto 0050-89) and Direccion
are in an area that was affected by glacia- de Investigacion y Desarrollo, Universidad
tions,wehypothesizethatice-freeareaswere Austral de Chile (Proyecto RS 89-06). We
the appropriate scenarios where acoustical aregrateful toCorporacionNacional Fores-
andchromosomaldifferentiationtookplace. tal (CONAF) for its hospitality and field
Frogsofthe roseusgroup{E. roseus, E. mig- facilities at Parque Nacional Contulmo and
ueli, E. calcaratus, E. contulmoensis, andE. Isla Mocha. The audiospectrograms were
insularis) are allopatric species sharing a done by Alfredo Brain, Institute de Acus-
similar one note advertisement call; how- tica, Universidad Austral de Chile. Mario
ever,E. contulmoensisandE. insularishave Pennaassisteduswiththeosilloscopicanal-
harmonics at about 500 Hz intervals while yses. We wish to thank Carlos Jara for his
E. roseus, E. migueliandE. calcaratushave useful comments on the manuscript. Raul
harmonics at about 1000 Hz intervals. Vo- Arriagada aided with collections and tape
cal differences among these allopatric spe- recordings in the field. Our thanks to Ron-
cies suggest that these species represent a ald Heyerandtwoanonymousreviewersfor
recent stage of speciation. All members of their suggestions.
VOLUME 107, NUMBER 2 397
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