Bijdragen tot de Dierkunde, 64 (2) 75-85 (1994)

SPB Academie Publishing bv, The Hague

Advertisement calls of Bolivian species of (Amphibia, Anura,

Hylidae)

Ignacio de la Riva Rafael Márquez & Jaime Bosch

Departamento de Biología Evolutiva, Museo Nacional de Ciencias Naturales, 28006 Madrid, Spain

Keywords: Anura, , Scinax, , advertisement calls, ecology, phylogeny

Abstract Myers, 1991; Duellman& Salas, 1991). Among the

South American countries, Bolivia contains some

of least known its fauna has The advertisement calls of eight Bolivian species of Scinax are the areas, and anuran

described information the behaviour of each including on calling been the subject of several recent discoveries (De la

A species. characteristic audiospectrogram and oscillogram are Riva, 1990a, 1990b, 1992a, 1992b, 1993; Reynolds presented for each species, as well as numerical information & Foster, 1992). about the spectral and temporal features of the calls. Two

In this study we contribute to the of phenograms based on the characteristics of the mating calls are knowledge constructed, the first one using a traditional multivariate tech- Bolivian herpetofauna by presenting the charac- nique (UPGMA) and the second one using audiospectrogram teristics of the advertisement calls of eight species in correlation,a new technique that allows holistic comparisons of the hylid genus Scinax, previously known as Ololy- single vocalizations. gon (see Duellman & Wiens, 1992). The advertise-

ment calls of two of these species are described for

the first time. Resumen The genus Scinax was known for many years

as Ololygon Fitzinger, 1843. Ololygon was resur- Se describen las llamadas de apareamiento de ocho especies de rected from the synonymy of Hyla by Fouquette & Scinax de Bolivia, aportando información sobre su comporta- based characters. Un Delahoussage (1977), on sperm miento fonador. audiospectrogramay un oscilograma son

cada información numérica the of this feature presentados para especie juntocon Although validity was questioned sobre las características temporales espectrales de las vocaliza- y by some authors, some characteristic morphologi- ciones. Se presentan dos fenogramas basados en las característi- cal features shared all the are by species in this cas de las llamadas, uno de ellos usando metodologíamultivari- genus. Recently the correct nomenclature for the ante tradicional (UPGMA), y el segundo usando una novedosa

genus was considered to be Scinax (see Duellman metodologíabasada en la correlación de los archivos de imagen de los audioespectrogramas. & Wiens, 1992). Among other morphological fea-

tures, this genus differs from Hyla mainly by lack-

ing a membrane between toes I and II or having

Introduction only a vestigial fringe along the inner margin of the

second toe. Additionally, they have truncate disks,

The Neotropics containby far the most diverse anu- which are wider than long, on the hands (Duellman ran fauna in the world, and a great number of spe- & Wiens, 1992). cies is described being each year. Central and South The genus Scinax is one of the most abundant

America the of numberof and of in are subject a exploratory conspicuous groups the Neotropics. studies that provide an ever more complex view of Frost (1985) reports 54 species in the genus (as the richness in anuran diversity (see Donnelly & Ololygon) and several more have been described

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since then. De la Riva (1990) lists eight species In the present paper, the calls of eight species of

in additional Scinax from with present Bolivia, although two species Bolivia are compared previously have been described since then (De la Riva, 1990b, published descriptions from other areas of South

America, to to inconsistencies in tax- 1993). The genus has been divided into several point possible

considered in the future of taxonomie research. phenetic groups. The eight species onomy, or areas

least three different used to present study belong to at (possibly four) Finally, two methodologies are

of affinities vocalisa- different species groups. The first group is the S. generate phenetic trees among

tions which allow to discuss some of rubra groupthat includes S. rubra, as well as proba- relationships bly S. chiquitana, S. fuscovaria, and S. nasica. The the species within Scinax. The first method is a second is the S. rostrata that would multivariate and the second method is group group, technique,

based the correlationof include S. garbei and S. nebulosa. The third group an innovative technique on

The is the S. staufferi group which includes S. parkeri audiospectrograms. relationships resulting and possibly S. nasica. The fourth potential group from the study of the advertisement calls are com- is the S. x-signata group to which S. nasica has been pared with the currently accepted species groups. assigned according to some authors. Of these four groups only the S. rostrata group appears to have Material and methods solid phylogenetic consistency (see Duellman &

Wiens, 1992, for further detailson the of first author from Recordings were obtained in Bolivia by the the group). These four groups include all eight spe- 1987 to 1990. Unless otherwise specified, calls were recorded in cies considered in the present study except for S. Puerto Almacén, northwestern part of Santa Cruz Department

which has been in castroviejoi, not assigned to any (15°46'S 62°15'W), an Amazonian locality wet subtropical

(Tosi et al., 1975). Recording equipment included either group (De la Riva, 1993). Ml and a Sony WM D6C or a Sanyo 120 tape recorder a Senn- Since the inception of sound analysis techniques, heiser Me 80 directional microphone. A representative audio- anuran mating calls have been considered to be im- spectrogram and oscillogram were presented for a 2.5 seconds

determinantsdue to theirrole portant taxonomie as recording segment for each species. A longer recording (20-60 pre-zygotic species isolating mechanisms (Blair, seconds) was analysed to generatenumerical information on the spectral and temporal characteristics of the sounds. For each 1955, 1958, 1959; Jameson, 1955; Johnson, 1959). species, we selected for analysis the calls of a single representa- The richness of the anuran tropical fauna allows for tive individual which emitted vocalisations which appeared to be studies of the vocalisations, either comparative "normal" to the human ear. The criteria for the choices were from ecological (Hödl, 1977; Drewry & Rand, also based on the certainty of the identification of the recorded

individual and in the quality of the recording. 1983; Zimmerman, 1983) or from taxonomiestand-

Recordingswere processed with a Macintosh-based digitalsig- points (Barrio, 1965, 1966). nal analysis system. Digitalisation and editing were completedat In general terms, it is accepted that the mating a sampling frequency of 44.1 kHz and 16 bit resolution with calls of anurans do not show homologies above the Sound Tools hardware and software. Signalyze software was

used to obtain numerical information and to generate audio- genus level, but some similarities among calls that

spectrograms and oscillograms. Frequency information was at and may have phylogenetic significance generic obtained through fast Fourier transform (FFT, width 1024 infrageneric level may be expected. The validity of points). The terminologyused for the description ofthe adver- vocalisations in studies phylogenetic is, however, tisement calls follows Heyer et al. (1990). limited A total of 12 different call characteristics were compared in a and depends on the group of anurans con-

phenetic analysis. The variables considered were: note duration, sidered. According to Schiotz (1973:313) "the voice pulse rate (pulses per second), dominant frequency, fundamen- the certain which is follows taxonomy to a extent, mini- tal frequency, other frequency with substantial energy, quite natural as the voice is a product of the mor- mum number of notes per call, maximum number of notes per

of the rate phology species." However, convergences call, number of pulses per note, call repetition (calls per

minute), note repetition rate (notes minute within the call), are often found between species of different fami- per number of different types of notes, and ratio pulse dura- lies and acoustic partitioning in allopatric anuran transformed in tion/interpulse interval. Variables were linearly communities seems to have similar characteristics the original matrix in order to standardise the scales of the

(Duellman & Pyles, 1983). different variables. The standardised matrix was used to calcu-

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late the "average taxonomie distance" between each species pair The mating call always consisted of two similar

The unrooted (after Sneath & Sokal, 1973). resulting phenetic notes emitted together in rapid succession (Fig. la). tree was generated through the application of an unweighted Both were short (26 and 28 milliseconds), with a pair-group-method arithmetic average (UPGMA) on the previ- high number of pulses, low fundamentalfrequency ously calculated matrix (NTSYS statistical package, Rohlf,

and a dominant 1992). The methodology and the parameters used were similar (891 Hz), relatively high frequency to those used for the comparison of advertisement calls of 18 (2400 and 2850 Hz). For the quantitative compari- species of Hyla from Bolivia by Márquez et al. (1993). sons withother calls the second note in the sequence In addition to the UPGMA analysis, a second method for was considered as the primary note because of its quantitatively comparing vocalisations was used, based intensity. De la Riva (1993) described the on the comparison of the audiospectrograms of single notes of greater the advertisement call. This method, described in Clark et al. calls of this species from the same locality and from

(1987) allows for a comparison of a full representation of a the same recordings. sound's structure in time and frequency rather than ofindividu- al acoustical features. The software used for the correlation Scinax chiquitana (De la Riva, 1990a) process was available in "Canary" (Cornell Lab. of Ornitholo-

This medium-sized SVL: males gy, Ithaca, New York). To the best of our knowledge, it is the species (maximum firsttime that this method is appliedto the quantitativecompari- 32.0 females35.3 to the mm, mm) probably belongs son of the vocalisations of different taxa. When the advertise- S. rubra group (De la Riva, 1990b) and occurs in the ment call was composed of more than onenote, the primarynote rain of northern Bolivia and southern was used for comparisons (we defined the primary note as the

the & At Puerto Almacén it note which was never omitted from the call or loudest note). (Duellman Salas, 1991).

Unless otherwise specified, collected individuals were deposit- was an explosive breeder, forming large aggrega- ed in the Centro de Estudios Tropicales, Sevilla, Spain, and/or tions of males during the rainy season. However, in the Museo de Historia Natural "Noel Kempff Mercado", mating calls were only heard sporadically. Santa Cruz de la Sierra, Bolivia.

Males usually called while perched on aquatic

plants or bushes emerging from temporal or

Results and discussion in their near vicinity. The advertisement call was

composed of a single type of note of intermediate Description of the advertisement calls duration (243 ms), pulsed, and repeated at regular

intervals (Fig. lb). The mean dominant frequency The audiospectrograms and oscillograms of a 2.5

was of 2172 Hz. No previous records of the vocali- second section of the calls of the different species

sations of this species were available. are shown in Figs. 1 and 2. A summary of the nu- merical information from the sound analyses is Scinax fuscovaria (Lutz, 1925) shown in Table I. The phenetic trees produced by This males large species (maximum SVL: 50.2 mm, the UPGMA analyses and by audiospectrogram females 53.2 mm) probably belongs in the S. rubra correlation are shown in Figs. 3 and 4. group (Fouquette & Delahoussaye, 1977; Frost,

Scinax castroviejoi (De la Riva, 1993) 1985). It occurs in open and forest through-

This recently described large species of Scinax out the Chaco, Cerrado, and the subtropical forests

(maximum snout-vent length (SVL): males 41.9 of , Argentina, Paraguay, and Bolivia. At mm, single female known 48.7 mm) occurs in tem- Puerto Almacén S. fuscovaria was an explosive

choruses in perate valleys of the eastern slopes of the Andes breeder forming large temporal ponds from Bolivia to northern Argentina. Calls were following heavy rains. recorded in a small lagoon near Bermejo (Dept. of Males called from bushes or small trees, usually

Santa Cruz, Bolivia, 18°97'S 63°38'W). Males perching directly on trunks or thick branches. The migrated to the shore at dusk, and called perched call was short and consisted of one note with low on small bushes or rushes. Scinax castroviejoi is frequency (860 Hz) and a low pulse rate (50 pulses probably a prolonged breeder because it did not per second) (Fig. lc). The call was emitted in fast rain the visited during two days when the was succession at regular intervals. De la Riva (1993) and males were active and calling. provided basic parameters of the calls of S. fus-

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bottom and of 2.5 second of characteristic call. Fig. 1. From to top: audiospectrogram oscillogram a recording a mating Top: expanded oscillogram showing the temporal details of the notes; a, Scinax castroviejoi; b, Scinax chiquitana, c, Scinax fuscovaria; d, Scinax garbei.

covaria from Puerto Almacén which are coincident 3715 Hz and a fundamental frequency of 1695 Hz with our description. (Fig. Id). The calls were separated by long intervals

of variable duration, averaging 1300 ms.

Scinax garbei (Miranda-Ribeiro, 1926) Duellman (1970a, 1972, 1978) described the

This call of Scinax from Santa is a medium-sized species (maximum SVL: mating garbei Cecilia, males 39.8 mm, females 42.1 mm) of the S. rostrata (in 1970a and 1972, Duellman considered group (Duellman, 1972). It is widespread in the the species to be in the genus Garbeana; in 1978,

described the upper Amazon Basin from to Bolivia. Duellman referred to it as Hyla). He

Apparently, there are marked differencesin size be- call as being a single, moderately long pulsed note, tween populations (Heyer, 1977). At Puerto Alma- repeated at a rate of 14 notes per minute. The note cén S. garbei was a prolonged breeder during the repetition rate of our recordings is nearly twice that rainy season. figure, whereas the duration of the note reported

Males called for the Ecuadorian 160-260 is perched head-down on trunks or frogs (range ms)

the shore shorter than the found leaves near of temporal or semi-perma- markedly duration by us nent ponds. The call consisted of one long note (950 (range 295—1469 ms). The pulse rates do not show ms), highly pulsed, with a dominant frequency of much agreement either; calls from Ecuador had

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Fig. 2. From bottom to top: audiospectrogram and oscillogram of a 2.5 second recording of a characteristic matingcall. Top: expanded

oscillogram showing the temporal details of the notes; a, Scinax nasica; b, Scinax nebulosa, c, Scinax parkeri; d, Scinax rubra.

pulse rates ranging from 195 to 240 pulses/second, of the variables are not even within the ranges of the

but Bolivian recordings showed pulse rates ranging previously published data for recordings obtained

site from 53 to 57 pulses/second. The only parameter in the same (Duellman, 1978). The discrepan-

that shows slightly less dramatic differences be- cies include the following parameters: note repeti-

tween the calls described, is the dominantfrequen- tion rate ranging from 54.5 to 72.3 notes/minutein

cy: 3250 Hz according to Duellman (1978) and 1983 and reported to be 14 notes/minutein 1978;

from 3453 in the call from 370 650 in 1983 ranging Hz to 3958 Hz present duration, ranging to ms

study. In spite of the differences in pulse rate and and from 160-260 ms in 1978; pulse rate, ranging

duration, the audiospectrograms provided (Fig. 1 from 80 to 100 pulses/second in 1983 and from 195

in Duellman, 1970 and Fig. 3 in Duellman, 1972) to 240 pulses/second in 1978; dominantfrequency,

show some structural similarities with our Fig. Id, ranging from 1564 to 1926 Hz in 1983 and estimated emphasising two wide, flat, frequency bands, the to be 3250 Hz in 1978. In any event, our data on the

upper one raising in frequency at the beginning of mating calls from Bolivian S. garbei do not coincide the call. with most of the numerical parameters reported for

Duellman & Pyles (1983) provided numerical the mating calls of Ecuadorian frogs of this species, data on the calls of S. garbei (then Ololygon) from either as reported by Duellman(1970a, 1972, 1978)

Cecilia several described later Duellman Santa (Ecuador). Curiously enough, or as by & Pyles (1983).

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a Further research on this widespread and highly The note had a mean duration of 160 ms, pulse

and harmonics variable species could provide information about rate of 58 pulses/second emphasised the actual taxonomie status of different popula- at 720 and 1050 Hz. Whereas the pulse rate reported

the tions. is similar to the values found by us (mean 55.6),

durations (our mean 240 ms), and particularly the

Scinax nasica (Cope, 1862) emphasised frequencies (2374-2867 Hz in the

Bolivian far from the values This is a medium-sized species (maximum SVL: recordings), are re-

37.0 mm) of uncertain phylogenetic relationships ported by Duellman (1972). However, the audio-

been considered member of the S. 1972: which has a spectrogram provided (Duellman, 195, fig. 1)

duration and with rubra, x-signata, and staufferi groups (Duellman & shows a call over 200 ms in

from 2500 Hz 5000 Wiens, 1992; De la Riva, 1993). It occurs through- emphasised frequencies to Hz,

structure similar to out the same range as S. fuscovaria. Its mating sys- showing a general very our tem is explosive and breeding occurs in temporary Fig. 2b. ponds. Hödl (1977) described the calls of H. egleri from

of He Recordings were obtained at Santa Cruz de la the area Manaus (central Amazon, Brazil).

Sierra(14°48'S 63° 10' W) where males called after described a call composed of two types of notes:

in middle of a note and a much shorter note. heavy rains from short grasses, the a primary secondary

notes pond, or on the ground near the shore. The mating We did not detect the existence of secondary

of short in from Bolivia. Furthermore, call consisted of one pulsed note, duration, our recordings with low dominant and fundamental frequencies neither the temporal nor the spectral values re-

call in Hödl with our of (970 Hz) and it was emitted in long groups ported by (1977) agree recordings

in- Bolivian rapid succession (337 calls per minute) at regular frogs.

numerical tervals (Fig. 2a). De la Riva (1993) described the Duellman & Pyles (1983) provided

calls of S. nebulosa vocalisations of this species recorded at the same data on (then Ololygon egleri)

from Belém de Pará The data time and locality. His description is based on the (Brazil). descriptive

the call S. nebulosa do with the same recordings as ours. for of not agree

previously published description by the senior

Scinax nebulosa (Spix, 1824) author (Duellman, 1972). According to that de-

has This is a relatively small species (maximum SVL: scription S. nebulosa only one note per call,

with note rate of 62.3 notes/ males29.5 mm, females 36.0 mm) of the S. rostrata an average repetition

inhabits in durationof 130 group (Duellman, 1972). It open areas minute, an average ms, an average the lower Amazon Basin from the Guiana region to pulse rate of 60 pulses/second, and dominant fre-

Mato Grosso (Brazil) and Bolivia. quencies ranging from 3840 to 4147 Hz. While the

close those found Recordings were obtained at Puerto Almacén values for pulse rate are to by us,

the other values do show similarities with where the species had a prolonged breeding system, not our lasting during the whole rainy season and probably results.

of calls of S. beyond it. Males called perched on riverine plants In summary, the descriptions the

to rather conflictive. However, on river banks or near semi-permanent ponds. Calls nebulosa appear be

of the characteristics were emitted by males spaced at long distance from the uniformity morphological

consisted of of this its la each other. The advertisement call one species throughout range (De Riva,

differ- note (240 ms) emitted sporadically, with a low num- unpublished data) leads us to believe that the

be the result ber of pulses and high dominant frequency (2870 ences in call descriptions reported may

Hz) (Fig. 2b). of differences in methods of analysis rather than

Duellman (1972) described the calls of Hyla actual taxonomiedifferences between populations. egleri (a junior synonym according to Hoogmoed &

Gruber, 1983). He described a call composed of Scinax parkeri(Gaige, 1929)

SVL: 23.3 4-7 notes, repeated at a rate of 39.5 notes/minute. This extremely small species (maximum

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are rubra Scinax rubra Scinax parkeri Scinax Scinax nasica Scinax garbei Scinax fus Scinax Scinax Scinax Scinax Table nebulosa I. 2nd 1st covaria chiquitana castroviejoi castroviejoi shown

note* 1st note 2nd next. Summary note note* When of of more numerical 11 7 5 61 19 25 12 39 38 5 calls Number than

Note one 29.5 162.9 185.7 240.6 57.5 947 162.2 243 26.1 27.8 parameters (ms) type (24-34.3) of (295-1469) (275) (24.6-32.3) is (3.7) (142.7-197.5) (16.2) (167.1-218.4) (17.6) (201.1-305) (43.6) (42.5-98.7) (10.3) (137.7-187.3) (13) (185.3-338.8) (48.6) (21.7-32.5) (2.6) (1.6) duration the considered, 1865.76 2096 2307.7 2374.6 972.9 1695.1 864.1 2172.4 891.1 905.6 frequency vocalisations. the (1776.9-2059.6) 2019.2-2140.4) (34.1) (2201-2402.9) (88.6) (2221.2-2564.4) 848.1-1110.6) (56.2) (1514.4-1776.9) (93.9) (827.9-928.8) (24.5) (2100-2261.5) (59.9) (848.1-928.8) (17.4) (868.3-1070.2) (53.1) Fundamental (111.1) (124.9) (Hz) note For each followed of 1865.76 2096 2777.9 2867.3 972.9 3715.4 864.1 2172.4 2848.7 2406 frequency the Dominant by (1776.9-2059.6) (2019.2-2140.4) (34.1) (2705.8-2826.9) (50.6) (2685.6-3129.8) (848.1-1110.6) (56.2) (3452.9-3957.7) (827.9-928.8) (24.5) (2100-2261.5) (59.9) (2766.3-2988.5) (47.1) (2301.9-2584.6) (43.1) an (111.1) (167.7) (136.9) (Hz) asterisk parameters,

1 1 1 1 1 1 1 0-2 1 1 /call Notes is the used mean 10 5.1 53.1 6.1 11.4 18.6 13.4 8.12 30 6 for (4-6) (7-9) 23-42 (5-7) (6-6) (0) /note Pulses (9-11) (0.7) (10-14) (17-21) (11-17) (0.9) (16-83) (6) (0.4) (1.3) (1.4) (2.6) (15.6) (0.6) the values are 69.7 342.1 55.6 91.1 122.1 234.3 216.1 100.2 56 50.12 construction followed (1) /second Pulses (291.5-375) (67.4-74.4) (1.983) (96.2-104.9) (2.9) (54.2-57.5) (1.3) (74.7-113.4) (15) (53.8-57.8) (46.6-56.5) (1.8) (114.2-128.4) (3.7) (157.7-276.5) (185.8-243.9) (33.7) (25.1) (11.8) by of the the 185.4 136.7 151.1 22.5 337.1 29.3 100.2 64.4 1504.2 1504.2 standard (83-206.7) (2.4-47.5) (11.2-49.4) (48.2-84.2) /minute Notes phenetic (164.7-206) (29.2) (103.6-156) (28.8) (54.6) (22.8) (223.8-457.8) (56.3) (8.9) (62.9-121.9) (15.7) (12.8) (1348.3-1685.4) (79.6) (1348.3-1685.4) (79.6) trees. deviations

33.6 33.6 151.1 22.5 337.1 29.3 100.2 64.4 80.6 80.6 /minute Calls between (16.6-46.8) (15.5) (16.6-46.8) (15.5) (83-206.7) (54.6) (2.4-47.5) (22.8) 223.8-457.8) (56.3) (11.2-49.4) (8.9) (62.9-121.9) (15.7) (48.2-84.2) (12.8) (6.9-115.6) (26.6) (6.9-115.6) (26.6)

0.708 0.612 0.641 0.828 0.663 0.678 0.596 0.842 Pulse parentheses. (0.45-0.81) (0.5-0.767) interval /inter-pulse (0.644-0.843) (0.065) (0.137) (0.567-0.707) (0.045) (0.724-0.912) (0.063) (0.543-0.848) (0.084) (0.074) (0.524-0.805) (0.075) (0.766-0.913) (0.047) duration Ranges Downloaded from Brill.com10/06/2021 07:12:18AM via free access 82 /. De la Riva et al. - Calls of Bolivian Scinax

member of the dubious S. and found them to be mm) presumably is a Hyla x-signata (Spix, 1824)

& It was different. Both the audiospectrogram and the nu- staufferi group (Duellman Wiens, 1992).

merical data for S. rubra in formerly considered by Lutz (1973) a junior syno- reported

a differ from our recordings of Bolivian S. nym of S. fuscomarginata (Lutz, 1925) position markedly

of the call of followed by several authors (Frost, 1985; De la rubra. However, the structure H. x-

Estado Venezuela 1 in Riva, 1990a) although Duellman & Wiens (1992) signata from Miranda, (fig.

resembles but it shows considered S. parker i as a valid species. It occurs at Rivero, 1969) our Fig. 2d, a

least in northern Bolivia and adjacent Brazil. It is slightly lower fundamental frequency. It would not

characteristic of and is a be that Rivero considered S. x-signata open habitats, probably unlikely

S. rubra vice since prolonged breeder during the whole rainy season. what actually was (and versa) the

of both is Recordings were obtained at the type locality, taxonomy species very complex.

Buenavista (17°27'S 63°40' W), where males called Duellman(1970b) described the mating call of S.

low in flooded The rubra from Santa Cecilia perched on grasses a large area. (then Hyla rubra) (Ecu-

call consistedof one short pulsed note (186 ms) with ador) as being a single short note (mean duration

high dominant and fundamental frequencies (ap- 130 ms) with dominant frequency centered at 1581

proximately 2300 and 2800 Hz) (Fig. 2c). Calling Hz and an average pulse rate of 63 pulses/second.

activity was sporadic. To the best of our knowl- Campbell (1971) later described the mating call of

from edge, no previous descriptions of the calls of Scinax Scinax rubra (then Hyla) Panama as one parkeri have been published. pulsed note (pulse frequency 66-72 pulses/second)

with a dominantfrequency of 1600 Hz and a dura-

Scinax rubra (Laurenti, 1768) tion of 140-150 ms. Whereas the duration and

This medium-sized species (maximum SVL: males pulse rate are somewhat comparable to the charac-

females 43.6 is in the teristics of the note described us 36.1 mm, mm) widespread longer by (duration

142-197 67-74 Neotropics from southern Panama throughout range ms, pulse rate range pulses/

of his does with northern South America east the Andes to cen- second), description not agree our

tral Bolivia, and southeastern Brazil. It is likely that recordings because it does not report the existence

fact that the a complex of species is currently being included of a secondary note. However, the

of the call located 1600 under this name (Frost, 1985). The of S. dominantfrequency was at

is reflected all in the of rubra includes both forests and open, disturbed Hz not at audiospectrogram

the call 2 At the areas. (fig. of Campbell, 1971). any rate,

characteristics of the call of the Panama- Recordings were obtained at Puerto Almacén, spectral

resemble where S. rubra is an explosive species breeding in nian frogs as depicted in this figure do

indi- temporal ponds. Males emitted their calls while those found in our analyses. Campbell (1971)

characteristics of his perching on both leaves and branches of bushes and cated that the call recordings

often similar those for this in small trees. Males formed large choruses that were to reported species

persisted until dawn. The call was emitted sporadi- Ecuador by Duellman (1970b).

of the cally and consisted of two pulsed notes (Fig. 2d). Duellman (1978) provided a description

The longer note (primary note) is often emitted in mating call of Scinax rubra (then Hy la) from Santa

isolation and is longer in durationand higher in fre- Cecilia (Ecuador) which coincided with that of the

and Panamanian calls of this described quency than the second (163 ms 2100 Hz, ver- species by

sus 29 ms and 1865 Hz in the second note). The Campbell (1971). On the other hand, Schlüter

the call of Scinax rubra second note can be repeated with a high number of (1979) described (then

from of two differ- pulses (342 pulses per second). Hyla) Peru as being composed

Rivero (1969) described the call of S. rubra (then ent types of calls, a long (200 ms) pulsed note and

Hyla rubra) from Estado Bolívar, Venezuela as a a shorter note (50 ms approximately). Similarly,

200 ms long pulsed call with dominantfrequency of our recordings from Bolivia show two note types as

about 1250 Hz. He compared this call with that of well, though with shorter durations (long 142-167

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Fig. 4. Phenetic tree resulting from the audiospectrographic Fig. 3. Phenetic tree resulting from the UPGMA analyses of 12

correlations of the notes of the calls of of numerical parameters ofthe calls of eight species ofScinax from primary eight species

Scinax from Bolivia. The scale represents correlations. Bolivia. The scale represents standard distances.

whose monophyly is not demonstrated (Duellman ms, short 24—34 ms). The dominant frequencies of

& Wiens, 1992). The advertisement call of S. par- our recordings appear to be slightly lower for both keriis similar to that of S. nebulosa, according to notes than those shown by Schlüter (1979), how- both Figs. 3 and 4. ever, the pulse rate of the longer note shown for the The audiospectrographic correlation phenogram Peruvian frogs (estimated to be 60 pulses/second) (Fig. 4) is rather consistent with the UPGMA is similar to that found by us (range 67-74 pulses/ phenetic tree, but does not show concordance with second). The discrepancy in mating calls between the tentative of the S. phylogeny genus. Again, cas- the descriptions of the calls from Panama and Ecu- troviejoi is markedly separated, but not to the same ador on the one hand, and from Peru and Bolivia degree as in Fig. 3. This species is grouped with S. the other on hand, suggests that the individuals chiquitana, and S. nebulosaand S. parkeri (the pair recorded in these locations may belong to two dif- most closely clustered). These four species probably ferent types or may have developed distinctly dif- belong to four different species groups. Although ferent dialects. A careful re-evaluation of the taxo- inboth Fig. 3 and Fig. 4, S. rubra and S. garbei, and nomie status of S. rubra in these locations be may S. fuscovaria and S. nasica, are clustered together, fruitful. the relative positions of the pairs are not the same.

The two phenograms cluster the species in a simi-

lar way. The affinity between the trees resulting Quantitative comparisons oftheadvertisement calls from the two methodologies is even more striking

if we consider that the audiospectrographic correla- The UPGMA phenetic tree (Fig. 3) has a co-phe- tion method does neither include any variables netic correlation coefficient of 0.897, which indi- related to the rate of emission of the utterances nor

cates a solid level of consistency. On the other to the number of different notes in a call (five out

tree hand, the consistency between the and the cur- in of twelve parameters included the UPGMA). rently accepted phylogenetic relationships within However, as we can see, there is no correlation be-

the Scinax (see Duellman & is genus Wiens, 1992) tween phylogeny and characteristics of the calls.

not the the high, although relationships among spe- The phenograms do not clearly reflect ecological

cies not clear taxonomists. few are to The estab- relationships either. Most of the species considered

lished affinities are not well reflected by the pheno- are mainly open habitat dwellers. Scinax chiquitana

S. nebulosa and S. are related gram. garbei closely lives in the forest, and S. garbei and S. rubra are

(both belong to the rostrata but their calls the group) euryoecious species found both in open and in

in the On the other are not grouped tree. hand, S. forest formations. These last two species are dis-

is parkeri clearly differentfrom the rest of the spe- tantly associated in the phenograms, and the differ-

cies, because of its small size, although it currently ence in habitat of S. chiquitana is not reflected in

is included with S. either of the nasica in the S. staufferi group, phenograms.

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to authors rubra in Panama J. 5: In summary, although according some group (Anura, Hylidae). Herpetol.,

similarities in their 52-55. closely related species may show Clark, C.W., P. Marler & K. Beeman, 1987. Quantitativeanaly- calls (Schiotz, 1973), the study of the genus Scinax sis of animal vocal phonology:an applicationto swamp spar- the does not conform to this trend and supports row song. Ethology, 76: 101 — 115. did find results of Duellman(1970b) who not great Donnelly, M.A. & C.W. Myers, 1991. Herpetological results of

coherence between the classification of the species the 1990 Venezuelan expedition to the summit of Cerro Guai-

quinima, with new Tepui reptiles. Amer. Mus. Novitates, of Hyla andthe similarities among mating calls (but 3017: 1-54. see Márquez et al., 1993). This lack of consistency Drewry, G.E. & A. Stanley Rand, 1983. Characteristics of an the rela- may be a result of the fact that intrageneric acoustic community of Puerto Rican frogs of the genus this far from tionships of groupare being clearly es- Eleutherodactylus. Copeia, 1983: 941-953.

tablished. On the other hand, the agreement be- Duellman, W.E., 1970a. Identity of the South American hylid

Garbeana garbei. Copeia, 1970: 534-538. tween the two methods of analysis suggests that the Duellman, W.E., 1970b. The hylid frogs of Middle America. holistic approach provided by the audiospectro- Monogr. Mus. nat. Hist. Univ. Kansas, 1: 1-753. for graphic correlation is a powerful technique Duellman, W.E., 1972. South American frogs of the Hyla

of sounds. This new Meded. comparison technique may rostrata group (Amphibia, Anura, Hylidae). Zool.

eventually become a more objective and agile alter- Leiden, 47: 179-195.

1978. The biology of an equatorial herpeto- native to the labor-intensivetechnique of extraction Duellman, W.E.,

fauna in Amazonian Ecuador. Univ. Kansas Mus. nat. Hist, of numerical variables for multivariate analysis. misc. Publ., 65: 1-352.

Duellman, W.E.& R.A. Pyles, 1983. Acoustic resourceparti-

tioning in anurancommunities. Copeia, 1983: 639-649.

Acknowledgments Duellman,W.E.& A.W. Salas, 1991. Annotated checklist of the

and reptiles of Cuzco Amazónico, Peru. Univ.

Kansas Mus. nat. Hist. occ. Pap., 143: 1-3. We are indebted to the Museo de Historia Natural "Noel

W.E.& J.J. Wiens, 1992. The status of the hylid frog Kempff Mercado", of Santa Cruz de la Sierra, Bolivia, for its Duellman, Ololygon and the recognition of Scinax Wagler, 1830. collaboration. Fieldwork in South America was supported by a genus

Univ. Kansas Mus. Hist. occ. 151: 1-23. Asociación de de Doñana I. De la nat. Pap., grant from the Amigos to 1977. Fouquette, M.J., Jr. & A.J. Delahoussaye, Sperm mor- Riva. Edwin Chacón allowed us to record calls in his ranch, phology in the Hyla rubra (Amphibia, Anura, Hyli- Puerto Almacén. We are particularly gratefulto A. Machordom group

dae), and its bearing on generic status. J. Herpetol., 11(4): who performed the phenetic analyses of the calls through

387-396. UPGMA and generatedthe phenetic trees from the audiospec-

D.R. 1985. of the world. A taxo- Additional for the Frost, (ed.), Amphibianspecies trogram correlation matrix. help analysis was nomie and geographicalreference: 1-732 (Allen Press Inc. & provided by S. Reig. Sound analyses were funded by project ASC, Lawrence, Kansas). CYCIT PB 89-0045C (PI: P. Alberch) Ministerio de Educación

1977. Taxonomie from the Madeira useful Heyer, W.R., noteson frogs y Ciencia, Spain. C.W. Clark and R. Cocroft provided and Purus rivers, Brasil. Papéis avuls. Zool. S. Paulo, 31: methodological advice for the analyses.

141-162.

Heyer, W.R., A. Stanley Rand, C.A. Goncalves da Cruz, O.L.

Peixoto & C.E. Nelson, 1990. Frogs of Boraceiá. Arq. Zool. References S. Paulo, 31: 237-410.

Hödl, W., 1977. Call differences and calling site segregation in

floating meadows. Barrio, A., 1965. El géneroPhysalaemus en la Argentina. Phy- anuran species from central Amazonian

sis, 25: 421-448. Oecologia, 28: 351-363.

Barrio, A., 1966. Divergencia acústica entre el canto nupcial de Hoogmoed, M.S. & U. Gruber, 1983. Spix and Wagler type

natural history Leptodactylus ocellatus (Linné) y L. chaquensis Cei (Anura, specimens of reptiles and amphibians in the

Leptodactylidae). Physis, 26: 275-277. musea in Munich (Germany) and Leiden (The Netherlands).

Blair, W.F., 1955. Differentiation ofmatingcall in spadefoots, Spixiana, Suppl. 9: 319-415.

and genus Scaphiopus. Texas J. Sei., 7: 183-188. Jameson, D.L., 1955. Evolutionary trends in the courtship

14: Blair, W.F., 1958. Mating call in the speciation of anuranam- mating behavior of Salientia. Syst. Zool., 105—119.

of phibians. Am. Nat., 92: 27-51. Johnson, C., 1959. Genetic incompatibilityin the call races

Copeia, 1959: 327-335. Blair, W.F., 1959. Call differences as an isolation mechanism in Hyla versicolor Le Conte in Texas.

southwestern toads. Texas J. Sei., 8: 87-106. Lutz, B., 1973. Brazilian species ofHyla: i-xviii, 1-260 (Univ.

Campbell,H.W., 1971. Observations on two species of the Hyla Texas Press, Austin).

Downloaded from Brill.com10/06/2021 07:12:18AM via free access Bijdragen tot de Dierkunde, 64 (2) - 1994 85

Márquez, R., I. De la Riva & J. Bosch, 1993. Advertisement Rohlf, F.J., 1992. NTSYS-pc: Numerical taxonomy system of

calls of Bolivian species of Hyla. Biotropica, 25(4). programs, version 1.70. (Exeter Publishing Co., Setauket,

Reynolds, R.P. & M.S. Foster, 1992. Four new species of frogs New York).

and one new species of snake from the Chapare region of Schiotz, A., 1973. Evolution of anuranmating calls. Ecological

other J.L. Vial of the Bolivia, with notes on species. HerpetologicalMonogr., aspects. In: (ed.), Evolutionarybiology anu-

6: 83-104. rans; contemporary research on major problems: 311-319

Riva, I. de la, 1990a. Lista preliminar de los anfibios de Bolivia (Univ. Missouri Press, Columbia).

con datos sobre su distribución. Boll. Mus. reg. Sei. nat. Tori- Schlüter, A., 1979. Bio-akustische Untersuchungen an Hyliden

no, 8: 261-319. in einem begrenzten Gebiet des tropischen Regenwaldes von

Riva, I. de la, 1990b. Una especienuevade Ololygon procedente Peru. Salamandra, 15: 211-236.

de Bolivia (Anura: Hylidae). Revta. esp. Herpetol., 4: 81-86. Sneath, P.H. A. & R.R. Sokal, 1973. Numerical taxonomy. The

Riva, I. de la, 1992a. Comentarios sobre el género Gastrotheca principles and practice of numerical classification: i-xv,

(Anura Hylidae) en Bolivia y descripción de una nuevaespe- 1—573 (W.H. Freeman, San Francisco).

Revta. 6: cie. esp. Herpetol., 15-22. Tosi, J.A., O. Unzueta, L.R. Holdridge & A. Gonzales, 1975.

Riva, I. de la, 1992b. A new species of Phrynopus from Bolivia Mapa ecológico de Bolivia (M.A.C.A., La Paz).

(Anura: Leptodactylidae). Herpetologica, 48: 111-114. Zimmerman, B.L., 1983. A comparison of structural features of

I. A Riva, de la, 1993. new species of Scinax (Anura: Hylidae) calls of open and forest habitat frog species in the Central

from Argentina and Bolivia. J. Herpetol., 27(1): 41-46. Amazon. Herpetologica, 39: 235-246.

1969. Rivero, J.A., Sobre la Hyla rubra Laurenti y la Hyla x-

signata Spix (Amphibia, Salientia). Mems. Soc. Cienc. nat. Received: 17 March 1993

La Salle, 29: 109-118. Revised: 16 July 1993

Downloaded from Brill.com10/06/2021 07:12:18AM via free access