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Source : MNHN, Paris

DLP 01-03-11 - 66097

AIVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

July 2011 __ Volume 27, N° 4 27, N°4

——— Bibliothèque Centrale Muséum

| LL ULIILL

330910056084 7

Description of the advertisement calls

of nine species of

Fejervarva Bolkay, 1915 and

Minervarya Dubois, Ohler &

Biju, 2001 from China, India and Nepal

Stéphane GROSJEAN & Alain DUBoIS

Reptiles et Amphibiens, UMR 7205 OSEB,

Département Systématique & Evolution, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<sgrosjea@mnhn.fr>, <adubois@mnhn.fr>

The advertisement calls of eight species of the genus Fejervarya and of Minervarya

sahyadris (Dicroglossidae) are described in detail. This includes the advertisement calls of the

holotypes of F pierrei, F. teraiensis and M. sahyadris, as well as of a paratype of F

nepalensis. For each species, temporal and frequency parameters are provided, as well as an

oscillogram and a spectrogram. The particular calls emitted before the advertisement call

proper are discussed. The advertisement calls described herein are compared to previous call

descriptions of the same species and some calls of unidentified Fejervarya species reported in

literature are tentatively assigned to calls described herein. Calls of species which could occur

in sympatry are compared to each other. The call of M. sahyadris is compared with the call of

the recently described second species of the genus, M. chilapata.

INTRODUCTION

The genus Fejervarya is a taxonomically complex group of morphologically similar frogs

comprising 32 currently recognized species distributed in Pakistan, India, Nepal, southern

China, Indochina to theislands of the Sunda Shelf, and Japan and Papua New Guinea (FROST,

2010fhttp://research.amnh.org/vz/herpetology/amphibia/amphibia/], 4 October, 2010).

Though the members of this genus inhabit open areas such as paddy fields and are

usually quite common when present, their taxonomy is far from being fully resolved. Indeed

this group of frogs is composed of morphologically very similar species, generally named

Fejervarya limnocharis, which was so considered as a very largely distributed species. Even

before molecular data (VEITH et al., 2001; KURABAYASHI et al., 2005; DJONG et al., 2007; .

KURAMOTO et al., 2007), advertisement calls turned out to be very useful in discriminating

several new species of Fejervarya in the small country of Nepal, within the group that had

been previously named Rana limnocharis (DuBois, 1975). The usefulness of advertisement

Source : MINHN, Paris

118 ALYTES 27 (4)

calls in discovery of new species or in the discrimination of sibling species is no longer debated

(BOGERT & SENANAYAKE, 1966; LITTLEJOHN & OLDHAM, 1968; KURAMOTO, 1980; NARINS,

1983: MaTsuI et al., 1986; SCHNEIDER & SINSCH, 1992; SCHNEIDER et al., 1993; NARINS

et al., 1998). After the discovery of several species in Nepal (Dugois, 1975), Roy & ELEP-

FANDT (1993) recorded the call of three different Fejervarya species in a nearby region,

northeastern India, thus confirming the existence of a species complex related to Fejervarya

limnocharis.

Several field trips done by one of us (AD) from 1972 to 1992 permitted to record the calls

of eight species of Fejervarya (F nepalensis, F. pierrei, F. rufescens, FE. multistriata, EF.

syhadrensis, F. sp. 1, nd Æ sp. 2, among which the calls of the holotypes of Æ

pierrei and E teraiensis and of one paratype of FÆ nepalensis), and of the holotype of

Minervarya sahyadris. Four of these Fejervaryan calls have been briefly described previously

(Dusois, 1975) and the call of F. rufescens used in this work has been recently described

(KuRAMOTO & DuBois, 2009). The call of F rufescens had been previously described by

KADADEVARU et al. (2000). Despite the importance of advertisement calls in species recogni-

tion and the abundance of the species of this genus in open and anthropized habitats, the calls

of only a few additional Fejervarya species have been described until know: a species of

Thailand referred to as FÆ. limnocharis (HEYER, 1971), Æ. cancrivora from Kalimantan, Borneo

(Maïsui, 1982), three unidentified Fejervarya species from northeastern India (ROY & ELEP-

FANDT, 1993), ecies from southwestern India referred to as Æ limnocharis (KANAMADI et

al., 1995), a species from Bali, Indonesia tentatively referred to as Æ limnocharis (MARQUEZ &

EEKHOUT, 2006), FE granosa, F. kudremukhensis and F caperata from southwestern India

(KuRAMOTO et al., 2007; the call of Æ kudremukhensis has been described as Æ cf. keralensis in

KURAMOTO & JosHy (2001) and the call referred to as Æ limnocharis in the same paper could

belong to Æ. caperata). The call of M. sahyadris has been described recently (KURAMOTO &

Jos, 2001; KADADEVARU et al., 2002; KURAMOTO et al., 2007; in the two first references as

Limnonectes syhadrensis, KURAMOTO et al., 2007).

eraiensis

The goals of this paper are (1) to describe the calls of the specimens recorded during the

fieldtrips of AD, (2) to provide the call characteristics of the holotypes of Æ pierrei, F

teraiensis and M. sahyadris, these data being particularly useful in integrative taxonomy of

Amphibians (e.g. Vierres et al., 2009), and (3) to compare them with the previously published

call descriptions of Fejervarya species.

MATERIAL AND METHODS

The advertisement calls of nine species of Dicroglossidae of the genera Fejervarya and

Minervarya were recorded in the field in India, Nepal and China from 1972 to 1992 by one of

us (AD) (tab. 1, fig. 1-2).

Recordings were made using either an Uher Report 4000 or a Sony TCDM-S$ tape

recorders with two microphones LEM, and Scotch magnetic 215 and TDK SA-X90 tapes.

Oscillograms and spectrograms were prepared with the Canary 1.2.4 software from the

Cornell Laboratory of Ornithology (CHARIF et al., 1995). The sampling rate used to convert

the signals to digital format was 22.254 Hz with a 16-bit precision. A filter bandwidth of

Source : MNHN, Paris

GROSJEAN & DUBoIS 119

Fig. 1. - Recording localities and type localities of the Fejerrarya and Minervarya species recorded in

India and Nepal. Black stars represent type localities where advertisement calls have been recor-

ded, white stars represent type localities. 1: Srirangapatna, recording site of Fejervarya sp. 1: 2:

Udhagamandalim, recording site of jersurru sp. 2 3: Gundik “pe local and'recording site of

Coma ty pe Eat of

ensis and r

ï sy'hadrens

of Fejervar

11: Ham, recording site of Fejervai S

e, corn ite of Fejervarya teraiensis: 13: Patnitop, Jammu & Kashmir, north Indi

recording sites of several Æ syhadrensis displaying the same temporal and frequency cha

as the conspecific populations described here.

Source : MNHN, Paris

Table 1. List of the species studied with information on the place and date of recording. Air T°: air temperature.

Fejervarva sp. | Lost 8930 India IS/VIL1984 124136°N, 76.7042°E 740-800 m No data

{field number 7489.D) (21h00-01h20)

Lost 8931-8932 | India IS VIL 1984 ngapatna 124136, 76.7042E 740-S00 m No data

{field number 7559.D) {21h00-01h20)

Fejervarva on 8933 India VIL1984 Gundia, 13.0833°N. 76.1167°E 200-220 m No data

rufescens (20h30-00h15) | Kempholey forest

& 8051 | india Gandi. BBA 7enerE | 200220m | Nodaa

{20h30-00h15) | Kempholey forest

Fejervarva sp. 2 @ 8935 India TVI9S4 Udhagamandalam TAN, 76.7042E 2200-2210 m No data

22h10-25h00)

Minervarva 20003031 holotype | 8936-8938 | India VIT1984 Gundia, TH OSSEN, 6LIOTE 200-220 m No data

sahyadris Kempholey forest

2000.3053 paratype 8939 India 2511/1984 Gundia, 13.0833N, 76.1167E | 200-20m No data

Kempholey forest

Fejervarva 1975.1628 paratype 5930 Nepal OVA1973 Kunga About 27.5°N, ST9SE 1580 m 210 °C

nepalensis (9h 10-22h45)

1975.1676 paratype s941 Nepal 12 V11973 Sitda Pokhari 27.32N, 87.38E 1530-1550 m | __No data

pierrei 0h1$-01h15)

1975.1706 paratype 8043 Nepal 22/1973 Biramode About 26.9167°N, 86.8°E 200 m No data

{19h45-23h10)

Faerere FETES BH NT | 2vior Biramode | Aou DENIERNIGNE | 00m Fo

sYhadrensis (9h45-23h10)

1995.2534 8945-8946 Nepal 6/VI/1973 Lumkua 27.37N. 87.0: 890 m No data

(20h20-24h00)

1978.1751 (3) 8947 Nepal 8/IX/1972 (20h30) Kaneagats About 28.2°N, 84.2°E S50m 22.7 °C

jervarva 1975.1763 holotype ET Nepal 2UVA973 Biramode About 26.9167°N, S6.8E 200 m 24.0 %

1eraiensis (0h15-01h15)

1976.1147 paratype 8949 Nepal 29/1V/1973 am 26.9167N,87.9167E | 1180-1210m

ei s Nepal 2IV/1973 Sanichare 26.68°N. S7.O8E 250m

{20h00-24h00)

Fes a BST Ce OV | ame | AMEN NRRRE | im ÉXES

multistriata {field number T.2062) (21h55) Province

(1) 19842346, 1984.2348—2350 and 1984.2356; (2) 7426.D, 7429.D, 7454.D, 7499.D, V3 and V6: (3) a single calling male

the recorded one: (4) 1976.1128—1130, 1976.1132—1135.

was caught but not necessarily

OTI

(+) LT SALATV

Source : MNHN, Paris

GROSJEAN & DUBOIS 21

Fig. 2. - Recording locality and type locality of Fejervarya multistriata from China. Black star represents

the type locality (Lantau Island, Hong Kong), the black dot represents the site where the

advertisement call has been recorded (Tunxi).

349.70 Hz and frame length of 512 points were used for both spectrogram and spectrum

analyses.

A call is defined as a series of notes emitted consecutively and separated from another

series of notes by a much longer interval than the interval separating two consecutive notes. In

the case of Fejervarya species a call lasts from about ten seconds to several minutes, with the

exception of Æ rufescens for which a call is composed of only a long note. These long calling

periods are named “calling bouts” hereafter. A note is an individual unit of sound composed

of impulsions called pulses. A pulse is an energetic impulse. In the case of Fejervarya species,

a note is composed of a series of pulses. The frequency bands are the different components

visible on the spectrogram, the lowest being the fundamental, and the other the different

harmonics. The band with the greatest emphasis is defined as the dominant frequency.

The following temporal parameters were scored from oscillograms: duration of the call

(dc) and intercall duration (the silence interval between two consecutive calls, dic), duration of

the notes (dn) and internote duration (the interval of silence between two consecutive notes,

din), number of notes per call (nn) and note rate (the number of notes per second, nns),

duration of pulses (dp), number of pulses per note (np), interpulse duration (dip) and pulse

rate (number of pulses per second, nps). Frequency measurements were scored from the

spectrum of a few notes within the signal and given as the mean of the frequency values of

these notes for the same frequency band. The visible frequency bands are noted fIb to f3b

from the fundamental frequency to the highest harmonic.

Source : MNHN, Paris

122 ALYTES 27 (4)

Most of the samples treated in this study are entire calls. However, the capacity of the

software being reduced, we had to choose calls of short duration (lower to 5 seconds).

Therefore the durations of calls (dn) are not typical values of the species.

Al specimens are deposited in the herpetological collections of the Muséum national

d'Histoire naturelle of Paris (MNHN); see tab. 1 for details. The calls described herein are

deposited in the Fonoteca Zoolôgica of the Museo National de Ciencias Naturales de Madrid

(FZ collection numbers 8930-8951). À sample for each species is available in the web checklist

of Frog Calls of the World, at <www.FonoZoo.com>.

AIT males of this genus have the same calling site. They call from the ground or from

grasses on the ground, very close to the water edge but never in the water itself, only sometimes

with the rear part of the body slightly in the water. Even if the males are within a flooded area,

they call sitting on emergent mounds, or perched on emergent plants as in the case of Æ

nepalensis from Sitda Pokhari. So as all recorded males were outside water, only air tempera-

ture is given when available (tab. 1). All the Fejervarya species call in choruses and as such are

heard from several dozen of meters by à human listener.

Itis not possible to provide a collection number to each animal because some individuals

could not be caught or were lost as a result of logistic problems. The calling male of Æ

rufescens Was lost during storing before transportation. In these cases, the snout-vent length

given was obtained by the mean of the snout-vent length of males caught in the population of

the calling male. The collection numbers of individuals taken into consideration for calcula-

tion of the means are the following (tab. 1):

(1) Fojervarya rufescens: 1984.2346, 1984.2348-2350 and 1984.2356.

2: 7426.D, 7429.D, 7454.D, 7499.D, V3 and V6.

G)A single calling male of Fejervarya syhadrensis has been caught but not nec

recorded one.

(4) Fejervarya teraiensis: 1976.1128-1130, 1976.1132-1135.

OBSERVATIONS

INDIA

Féjervarva sp. 1

The three calls studied here are entire sequences. The call of Fejervarya sp. 1 (fig. 3a,

tab. 2) is composed of a series of 9 to 16 multi-pulsed notes and lasts 3.24 s in average. The

duration of notes averages 0.11 s and the intervals between them 0.15 s. The notes are emitted

at a rate of 4.2 per second. Each note (fig. 3b) consists of about 20 pulses of very short

duration (about 5 ms) and lacks pulse intervals. The pulses are emitted at a rate of 181 pulses

per second on average. The duration of notes increases during the call. So, the number of

s increases with the duration of notes (from 14 to 28 for the largest range). The notes have

Source : MNHN, Paris

GROSJEAN & DUBOIS 123

8-8 0 D. De

3 ss 56

5 00 os m1 15 20 25 30 35 4 4

Fig. 3. — (a) Spectrogram and oscillogram of an advertisement call of Féjervarva sp. 1 (field number

D.7489) and (b) detailed oscillogram of the 7!" note.

a very fast (abrupt) rise time and decrease just as fast. The amplitude of pulses increases

abruptly at the beginning of the note and decreases slowly at the end.

The fundamental frequency lies at about 1724 Hz. The dominant frequency corresponds

to the second frequency band (the first harmonic) and lies at about 3417.6 Hz. A slight

upwards frequency modulation of the dominant frequency band is observed at the beginning

of notes.

Fejervarya rufescens (Jerdon, 1854)

The advertisement call of Fejervarya rufescens (fig. 4a, tab. 2) is a long noisy note with

increasing intensity. In this case, the call is synonym of note. The duration of the call is

relatively short with a mean of 0.349 second, separated by ence of 1.59 s in average and

emitted at a rate of 0.74 notes per second. The calls are formed of about 60 pulses (fig. 4b).

The amplitude of pulses increases greatly from the beginning of the note until the last pulses,

the amplitude of which decreases abruptly. The pulses are emitted at a high rate, without

silence between them.

Source : MNHN, Paris

ptl

Table 2. - Characteristics of the advertisement calls of several Fejervarya species and Minervarva sahyadris from India and of Fejervarya

multistriata from China. MNHN: inventory number of MNHN: svl: snout-vent length of the recorded male in mm (when a mean is

given, it refers to the mean of all caught males in the population of the recorded male: see Marerial and methods section); de:

duration of the call from first to last note: dic: duration of the interval between two consecutive calls; dn: duration of notes; din:

duration of the interval between two consecutive notes; nn: number of notes per call; nns: number of notes per second; dp: duration

of pulses: np: number of pulses per note; nps: number of pulses per second: fib: frequency of the band i. Time measurements

expressed in seconds; frequency in Hz. Values are given as: mean + standard deviation, minimum-maximum, (number of

measurements).

Fojervarsa sp. 1] 7489.D = 428 = Joinis207/02072365] 14 | 33 [about|157232/ 1605257 | 1807 F5 | {or

0.080-0.134 | 0.154-0.305 0006| 14-22 |164.0-175.9 5

ui) u3 u4) (ES =

Fejerarsa sp. 1] 7559.D - Jonosiéfonez483{ 125 | 47 labour] 207240 [188.72 81 |1555-1802/3017-3788 w

186-558 0.073-0.143 | 0.083-0.297 | 9-16 | 4.548 | 0.005 | 14-28 S

@) e3) @) à

Fejervana 0.349 4 0.72 | 1.59 + 0.46) x = 0.7 = 1117-1420[2410-3027| - 2

rufescens 0267-0422 0708

En @ @)

Fejervarsa sp.2 137 = Joossooxfoss200) 4 | 25 [ovoofs2s+00] 552 SIS7 | 10685 | -

0.068-0.128 | 0.371-0.417 +6

ct) G) ES

Minervarva | 2000.3031 07762010] - 0034: 0.006/0.071 0.055 O004-|6.63 + 141 365-370] - -

sahyadris 0.668-0.872 0.023-0.047 | 0.026-0.245 0007| 5-10

6) En en es

Minervarva | 2000.3055| 180 0.998 = [0.052 2 0:005/0.076 + 0.049 about [7002115] 2185 | 36934 - E

sahvadris 0.027-0.044 | 0.050-0.203 œ001| 6-9

uo ) 0)

Fejerarsa | 1.206 13 0381 133 [ooso-oon) 0210 53 [oo 911279 | 1371 3017 [4007

mulristriata 0.367-0.394 0.070-0.094 | 0.191-0.229 83.9-90.6

@) ei @) «)

Source : MNHN, Paris

GROSIEAN & DUBoIS 125

m5 300 50 0 650 ro Fo 00 650 300 #50

Fig. 4. — (a) Spectrogram and oscillogram of two advertisement calls of Féjervarya rufescens and (b)

detailed oscillogram of the 1* call.

Two frequency bands are present, the fundamental lying at 1117-1420 Hz whereas the

dominant lies at 2410-3027 Hz. The frequency of the note increases during its emission,

which is especially visible in the dominant frequency. This change of frequency within the

note is coupled with an intensity modulation to the note (increase in this case).

Fejervarya sp. 2

The call of Fejervarya sp. 2 (fig. Sa, tab. 2) is composed of short multi-pulsed notes. The

notes last 0.095 s in average, are separated by silences of 0.398 s in average and are emitted at

a rate of 2.5 notes per second. The maximum amplitude of the note is reached in the middle

of the note. The notes are composed of two or three groups of two pulses (fig. 5b). These

groups are separated by durations comprised between 0.012 and 0.047 second. There are no

silences inside the groups. Despite the variation of the duration of notes and of the number of

pulse groups, the call appears homogenous to the listener.

Two frequency bands are present, the fundamental lying at about 820 Hz and the

dominant at about 1070 Hz.

Source : MNHN, Paris

126 ALYTES 27 (4)

ékr tt ht ‘ k te

es Fo es æ ps ps 13 ES rs rs

Dr] co p3 ET ET ps eA

Fig. 5. — (a) Spectrogram and oscillogram of an advertisement call of Féjervarva sp. 2 and (b) detailed

oscillogram of the 2"! note.

The sample described here is only a short part of a calling bout. It does not represent an

entire call.

Minervarya sahyadris Dubois, Ohler & Biju, 2001

The specimen recorded is the holotype of Minervarya sahvadris.

The call of M. sahyadris (fig. 6a, tab. 2) lasts less than 1 second and is composed of a

series of 7-10 multi-pulsed notes. The notes which last in average 0.033 s are emitted very

quickly (about 10 notes per second) and are high-pitched. The duration of internote intervals

is relatively short (0.073 s in average). The first note is always longer, with a greater amplitude,

composed of more pulses, and separated from the second one (mean din = 0.190 s) by a longer

interval than all the following notes between them. The second note, though of same

amplitude and duration as the following ones, is frequently separated from the third one by à

slightly longer silence than those between the following notes (mean din = 0.096 s). The mean

duration of the internote intervals between the following notes is 0.048 s. Consequently, the

Source : MNHN, Paris

GROSJEAN & DUBOIS

104

m ls r

ms 0 200 400 600 800

1000

ms 0 200 400 600 800

1000

ms 540 550 séo 570 580 530

610

Fig. 6. — (a) Spectrogram and oscillogram of an advertisement call of the holotype of Minervarya

sahyadris (MNHN 2000.3031) and (b) detailed oscillogram of the 3" note.

Source : MNHN, Paris

8cI

Table 3. - Characteristies of the advertisement calls of four species of Fejervarva from Nepal. For legends, see Table 2. dip: interpulse

duration.

Species | MNHN de dic dn Din mn} nns dp flb | Pb | Bb

Fojervarsa EX] 3.76 : 013 | 0.460 2 0.036 13 | 000 171

nepalensis 0370-0406 | 0.430-0.50$

& en

Fjenara | 19754167 | 289 430 = [07320008 10] 22 | 0007 | - 140245 [1883/5613] -

nepalensis O.164-0.181 N7S-ULS

ao «oi uo

Fjervarsa | 19751680 | 295 296 = [os 20010 [0078200068 [9 0004 | <0001 760 [issoc|4212.3 | 85815

te | 0.247-0. 0.072-0.08$ >

| ww) «si w) C

Fojervarva 1706 | 300 = [o2s720.027 | 0.106 20.064 | 10 | 2 0.003 | <0001 74 1176[41143| - e)

picrrei 0.221-0.302 | 0.072-0. pl

ao w) 4

Faenana loraew | 271 T Toosiz 0007 loosisowor [26] o1 | 000 | - 193.7260 | is18 [677] - Ni

srhadrensis 0.0$4-0.081 | 0:039-0.067 1804-2017 &

6) es) 6 6) +

Fejervarsa | 199: 29.5 204 = [ooss+000 | 0059-0011 [27] 92 3 199.72 8.77

svhadrensis 291-297 0045-0066 | 0035-0226 91-93 183.6-219.1

ei (54) sr || « us)

Fejervana | 19781781 | 286 141 = [0082202004 [oser <oo21 [13 | 02 | 000 | - 2107 [17005 | 600 | 56227

sthadrensis 0046-0058 | 0.046-0.125

a ui?

Fcjervarsa mél 416 = [oo7i2000s [0232220007 | 9 | 56 | abou | - TS mir | 2430 | 38703

teraiensis 0.066-0.079 | 0.158-0.438 0014

Ferana | 1976.1147| 351 107 [ozes| 10520021 [03722017 | 3 bout DO 694 MAETIE"T

teraiensis 1.06-1.09 061-0119 | 0179-0557

En © «

Fcjervarsa = [css 137 [| 00810009 [oxi72oo7t | 4 | 32 hbouoo] - 69.6 1058 | 2116 |27166

eraiensis 319-440 0067-0094 | 0.189-0.374 |3-

ui) a) 4) 46) e)

Source : MNHN, Paris

GROSIEAN & DuBoiIs 129

emission rate of the notes increases during the call. The notes are composed of 5 to 10 very

densely arranged pulses lasting from 4 to 7 ms and without silence between them, and emitted

at a rate of 200 pulses per second (fig. 6b). The largest pulse is usually the first or the second

of each note, then their amplitude decreases. So the maximum amplitude of the note is

reached very quickly after what the amplitude of the note decreases more slowly. The

structure of the pulses resembles that of the note.

Only the fundamental frequency, which is also the dominant frequency, is visible. It lies

at about 3650 Hz.

NEPAL

Fejervarya nepalensis (Dubois, 1975)

Contrary to the other species, F nepalensis (fig. 7) is a forest species. During day time, it

is often found on the forest floor. It is also found near ponds at the edge of forests.

The advertisement call of Fejervarya nepalensis is a sequence of long multi-pulsed notes.

It is remarkable, however that this is the only characteristic shared by the calls of the two

specimens recorded from two different localities. The recording person noticed differences

between the two frogs in his fieldnotes. As these two calls differ in all respects, they are

described independently.

The call of Æ nepalensis from Kunga (paratype), which is an entire call sequence, is

composed of long notes (0.385 s in average) separated by long intervals (0.460 s in average)

- “

Source : MNHN, Paris

130 ALYTES 27 (4)

and emitted at a slow rate of 1.3 note/s (fig. 8a, tab. 3). The duration of the notes and of the

intervals between notes are the largest among the species described herein whereas the note

rate is the slowest of all species. The notes are composed of a large number of pulses (56-63)

of 3 ms of duration with a group of greatest amplitude at the beginning of each note (fig. 8b).

Three frequency bands are visible, the fundamental frequency lying at about 1710 Hz, the

dominant at about 2830 Hz and the third band of energy at about 4300 Hz. The dominant

frequency shows an upwards frequency modulation within each note.

D 0

5 00 05 10 ‘5 20 25 30 35 mi 5 50

s 2 30 En #2 #3

Fig. 8. — (a) Spectrogram and oscillogram of an advertisement call of one paratype of Fejervarya

nepalensis (MNHN 1975.1628) from Kunga and (b) detailed oscillogram of the 3 note. The notes

of a second calling male are visible between the notes of the described call.

The call of Æ nepalensis from Sitda Pokhari (paratype) is composed of shorter notes of

0.173 s in average separated by intervals of 0.300 s in average and emitted at a rate of 2.2

notes/s (fig. 9a, tab. 3). This is only a part of call which can last until two minutes. The notes

are composed of 21-22 pulses of 7 ms of duration arranged in three groups per note, the

pulses of each group increasing then decreasing in amplitude (fig. 9b). Two frequency bands

are visible, the first one at about 1830 Hz and the second at about 3630 Hz. The dominant

frequency is the fundamental.

In summary, the calls of these two specimens differ in the duration of the notes, the

duration of the intervals between them, the number of pulses per note, and the pulse and note

Source : MNHN, Paris

GROSJEAN & DUBoIS 131

ns 1550 1600 1650 1700 ET 1800

Fig. 9. - (a) Spectrogram and oscillogram of an advertisement call of one paratype of Fejervarya

nepalensis (MNHN 1975.1676) from Sitda Pokhari and (b) detailed oscillogram of the 4 note.

rate. The frequency bands are higher in individual from Kunga and the dominant frequency

is not represented by the same harmonic in the two individuals. These two calls probably

belong to two different species.

Fejervarya pierrei (Dubois, 1975) (fig. 10)

One of thecalls described herein was emitted by the holotype(MNHN 1975.1628, fig. 11).

The call of Fejervarya pierrei (fig. 12a, tab. 3) is a series of long multi-pulsed notes lasting

0.258 s in average. The durations of internote intervals are short, 0.092 s in average, in

comparison to the calls of the other Fejervarya species. However, due to the relatively

important duration of notes, the note rate (2.9 notes per second) is not affected. These notes

are composed of five or six groups of three to five large pulses separated by seven or eight

small pulses (fig. 12b) giving the call a trill-like sound. The first or the second group of pulses

has the greatest amplitude which decreases slightly in the subsequent pulse groups.

Two or three frequency bands are present, the fundamental lying at about 1800 Hz, the

second and dominant frequency is about 4200 Hz and the third one, when present, at about

8600 Hz.

This call sounds like a cricket song.

Source : MNHN, Paris

132 ALYTES 27 (4)

Fejervarya syhadrensis (Annandale, 1919)

The calls analysed herein (from Birtamode, Lumkua and Kaneagats, Nepal) are whole

sequences but chosen among the shortest for the convenience of the software limitations (the

maximum length capacity of the software is 14 s). The call of Fejervarya syhadrensis (fig. 13

and 14a, tab. 3) is a fast series of short multi-pulsed notes (0.055 s on average). The durations

of internote intervals are very reduced (din = 0.057 s, n = 89). This is the highest call rate

among calls studied here (9.17 notes per second on average) due to both short note and

internote intervals. Each note is composed of 9-15 pulses (about 0.005 s) without interpulse

Fig. LL. - The holotype of Füjervarya pierret (MNHN 1975. 60 from Birtamode, the 21° May 1973.

Source : MNHN, Paris

GROSJEAN & DuBoIs 133

14119

id j ER ï sur

RO ul # ul Le de È + vw ERA HE

"00 LE 19 20 30

| a

6 00 05 10 15 20 25 30

1000 b

6 1400 1450 1500 1550 1600 1650

Fig. 12. — (a) Spectrogram and oscillogram of an advertisement call of the holotype of Féjervarya pierrei

(MNHN 1975.1680) and (b) detailed oscillogram of the 5!” note.

interval (fig. 14b). The amplitude of pulses increases slightly at the beginning of call and

decreases in the same way at the end. The last pulse is always shorter than the previous ones.

These calls typically have one note then two groups of two notes before a rapid and

continuous emission of notes. Although the duration and the number of pulses of any note in

the call is noticeably the same, the silence duration between each of these groups is three to

four times bigger (0.171 + 0.037, n = 6) than the silences within both the groups of two notes

and the following succession of notes (0.048 + 0.007, n = 83).

The fundamental frequency is at about 1634 Hz and the dominant frequency peak (the

first harmonic) lies at about 3440 Hz. The spectrogram is characterized by a wide range with

up to six harmonie bands.

Fejervarya teraiensis (Dubois, 1984) (fig. 15)

The call of the specimen MNHN 1975.1763, which is the holotype of FÆ teraiensis

(fig. 16), described here, is only a small sample of a calling bout lasting about 1.5 minute. The

calls of the two other specimens analysed herein are whole calls.

The call of Fejervarya teraiensis (fig. 17a, tab. 3) is a continuous series of short multi-

pulsed notes. The duration of notes averages 0.085 s and the intervals between them, 0.31 s.

The duration of the intervals between the notes is highly variable, even within a call (range

0.158-0.557 s, SD = 0.118, n = 18). So the notes are emitted at an irregular rate. The note rate

Source : MNHN, Paris

134 ALYTES 27 (4)

ee.

Fig, 14. - (a) Spectrogram and oscillogram of an advertisement call of Féjervarya syhadrensis (MNHN

1995.2534) from Lumkua and (b) detailed oscillogram of the 16!” note.

Source : MNHN, Paris

GROSJEAN & DUBoIS 135

is about 3.2 notes per second. Each note consists of 5-8 large pulses (fig. 17b) without

interpulse intervals (dp = 0.014 s). The amplitude of the note increases quickly (the second

pulse is the largest) and decreases more slowly.

The dominant frequency is about 2450 Hz for the two first calls and 1040 Hz for the third.

We can notice here a change in the dominant frequency: the dominant frequency is equal to

f2b for the two first calls and f1b for the last one.

The spectrogram shows a wide frequency range with up to seven harmonic bands.

SL APS CAES

Fig. 15. A male paratype of Fejervarpa teraiensis (MNHN 1975.1764) from Burimorang, Nepal, the 271

July 1973.

Source : MNHN, Paris

136 ALYTES 27 (4)

es 00 05 10 15 20 25 M

tb dt +

Li 10 vo 140 vo rs 130 1% 130

Fig. 17. — (a) Spectrogram and oscillogram of an advertisement call of the holotype of Fejervarya

teraiensis (MNHN 1975.1763) and (b) detailed oscillogram of the 5!" note.

Table 4. - Comparison of the frequency ranges of each frequency band among the four species of Fejervarva recorded from Nepal and the

three species recorded from southwestern India.

Species recorded from Nepal Species recorded from southwestern India

FE teralemsis Eshadrenis | Fnepalensis FE pierrel Foenanasp.2 | Fenfscns | Foenanaspl

ne Lo 1547 EAN EAN 0x 11414 1518

es 2124 3236 2836 4142 10 24-30 30-38

sb 2739 53-56 43 86 5

CHINA

Fejervarya multistriata (Hallowell, 1861)

The call of Fejervarya multistriata (fig. 18a, tab. 2) is a short call (0.380 s) composed of

groups of two multi-pulsed notes. These are only a few notes emitted before a longer call.

These two groups of two notes are separated by an interval of 1.53 s. The duration of notes

averages 0.08 s and the intervals between them, 0.21 s. The note repetition rate is relatively

high (5.26 notes per second). Each note consists of seven or eight large pulses (10-12 ms)

which have a decreasing amplitude throughout the note (fig. 18b). There is no silence interval

between two consecutive pulses.

The dominant frequency is 1371 Hz. This is the only case where the dominant frequency

is equal to the fundamental among the Fejervarya species studied here (except for Æ nepalensis

Source : MNHN, Paris

GROSJEAN & DUBOIS 137

from Sitda Pokhari). The frequency of the first pulse of each note is higher than the second

resulting in frequency modulation.

al ÿ

DR me fn - EUR

Hu LI We ds

[he AMEN

+ s 00 LE 10 15 20

ss 00 LE 10 15 20

LOPRE 15 15 156 13e 200 202

Fig. 18. — (a) Spectrogram and oscillogram of an advertisement call of Fejervarya multistriata (T.2062)

and (b) detailed oscillogram of the 1° note of the second call. The notes of a second calling male

are visible between the notes of the described call.

DISCUSSION

The calls described herein are all, with the exception of FÆ rufescens, a long and regular

succession of trilled notes. These calls can last more than one minute. The same structure has

been reported for the calls of other species in northeastern and southwestern India (Roy &

ELEPFANDT, 1993; KANAMADI et al., 1995; KURAMOTO et al., 2007).

The calls of Fejervarya species are often introduced by a few notes, often grouped by two.

These notes are present before each advertisement call and in all calling males. These small

calls sound like timid attempts of advertisement calls and are emitted until the group of

calling males “feels at ease”. Then, as soon as a male is “sufficiently reassured”, it begins

emitting the typical advertisement call, immediately followed by all the other males in its

vicinity. If the frogs are “inhibited”, these notes can be emitted for a long period (GROSIEAN

& Duois, pers. obs.). These notes emitted alone or in pairs with a longer internote duration

Source : MNHN, Paris

138 ALYTES 27 (4)

before and after the typical advertisement call were reported by DuBois (1975), who reported

larger internote durations but larger note duration too unlike our results and interpreted these

notes as territorial call. In the same paper, Dumois (1975) illustrated a territorial call of F.

teraiensis (under the name Æ “limnocharis”) which was much longer and structurally different

from the notes of the advertisement call of this species. Furthermore, KANAMADI et al. (1995)

observed the territorial behaviour of a Fejervarya species and described the territorial call

whose notes have a different structure from the notes of the advertisement call. The few

tentative notes at the beginning of the calls reported here have, on the contrary, exactly the

same structure and the same duration as the following notes of the advertisement call. For

these reasons we do not consider these notes as the territorial call.

The advertisement calls of F nepalensis, F pierrei, F. syhadrensis and F. teraiensis were

briefly described elsewhere (Dugois, 1975, 1976).

The call of the specimen of Æ rufescens studied here was described by KURAMOTO &

Dunois (2009). The measurements of the call parameters reported by these authors and those

described here are of course very similar except for the number of pulses which is reported to

be 25 in average by KURAMOTO & DuBois (2009) and about 60 here (reflecting perhaps

differences in detectability of the pulses). The advertisement call of this species has also been

described previously from the Western Ghats (KADADEVARU et al., 2000). In this population

the number of pulses was 49 in average. The waveform of the call of Æ rufescens shown by

these authors was very similar to that of the present study. However, the temporal values of

our sample are bigger than those of Æ rufescens from the Western Ghats. For instance the

notes last 349 ms and contain 60 pulses in average in our samples vs 242 ms and 49 pulses, and

the interval between each note lasts 1.6 s in our samples vs 0.556 s. KADADEVARU et al. (2000)

reported five frequency bands whereas only two are visible in our specimens. The dominant

frequency matches in the two samples 2400-3000 Hz in this study and 2500-3520 Hz reported

by KADADEVARU et al. (2000). These differences could be due to temperature differences

(unfortunately, no data are available for our recording), geographic variation or different

hormonal state.

Roy & ELEPFANDT (1993) described the call of three unidentified species of Fejervarya

from northeastern India. All the values of the temporal parameters (note duration, interval

duration between notes and number of pulses) of the species referred to as Æ limnocharis are

higher than those of FÆ nepalensis from Kunga, the species from Nepal and northern India

which exhibits the greatest temporal value. On the contrary, the dominant frequency is lower

in “E limnocharis” than in F nepalensis. However, the general waveform of the notes of the

two species resembles greatly. The species named sp. 1 in the same paper fits the call

parameters of our Æ teraiensis, a species which also oceur in this region whereas the call

parameters of the species named sp. 2 have no equivalent in our sample.

The distribution of Æ syhadrensis, if actually this taxon is not a species complex, is large,

spreading from eastern Pakistan to Bangladesh, northern India and Nepal and most probably

to northeastern India. One of us (AD) identified and recorded a species close to Æ syhadrensis

reported here as Æ sp. 1. However, except for the frequency parameters and the high pulse rate,

the two calls share no similarity and sound differently. KANAMADI et al. (1995) described the

advertisement call of a seemingly undescribed Fejervarya species. The call parameters (both

temporal and frequency) reported by these authors perfectly fit the measurements of our Æ

sp. las does the structure of the note. These two populations are likely conspecific. On the

Source : MNHN, Paris

GROSJEAN & DUBOIS 139

other hand, AD recorded also the calls of several FÆ syhadrensis from a distant population, at

Patnitop, Jammu & Kashmir, northern India (about 1000 km distance: fig. 1). The calls of

these specimens (data not shown) are identical to those described here, confirming the

conspecificity of the two populations as well as the wide range of this species.

Among the eight Fejervarya species whose calls are described herein, three have been

recorded in southwestern India (FE sp. 1, EF rufescens and F sp. 2), four in Nepal (F nepalensis,

Æ pierrei, F syhadrensis and F. teraiensis) and one in China. Although the four Nepalese

species inhabit different zoogeographic areas, they can occur in syntopy, particularly Æ

syhadrensis which can be found together with any of the three other species (DUBoIs, 1975; fig.

1). Thus, their advertisement call could play an important role as a premating isolating

mechanism. The fundamental frequencies of three out of the four species overlap (tab. 4).

Only the frequency bands of FÆ teraiensis are clearly outside the range of the three other

species. Fejervarya nepalensis and F. pierrei have the same range of values for the fundamental

frequency, but the dominant frequency plays probably the most important role. However,

Roy & ELEPFANT (1993) suggested that a marked different dominant frequency between

individuals might be useful for interindividual recognition. Fejervarya syhadrensis and F

nepalensis from Sitda Pokhari present the same dominant frequency but the calls differ clearly

by the length of notes, the longest notes of Æ syhadrensis remaining always shorter than the

shortest notes of F nepalensis. Fejervarya syhadrensis which can be found in syntopy with the

three other species possesses the most divergent call. It has the shortest note durations, the

shortest intervals between notes and the fastest call (which is expressed by more notes, high

note rate and high pulse rate).

Fejervarya rufescens, F. sp. 1 and F. sp. 2 were all described from southwestern India

(Karnataka and Tamil Nadu States). So they could potentially be syntopic. Here again, the

advertisement call could be an important premating isolating mechanism. The values of the

first two frequency bands do not overlap (tab. 4). Furthermore, Æ rufescens has a very

different call relative to the two other southwestern India species. Its notes are the longest as

well as the intervals between them (which last more than one second). Due to the length of the

notes, the note rate is particularly slow and the number of pulses high. The call of Æ sp. 2 is

remarkable by its notes containing a low number of pulses.

The advertisement call of Minervarya sahyadris described herein is of importance as it is

the call of the holotype. The type locality is near the village of Gundia, Kempholey forest,

Hassan District, Karnataka, India (13.0833°N, 76.1167°E: altitude about 200 m). The

advertisement call of M. sahyadris has already been reported from southwestern India:

Mangalore and the Western Ghats (KURAMOTO & JosHy, 2001: KADADEVARU et al., 2002:

KURAMOTO et al., 2007; in the two first references as Limnonectes syhadrensis according to

KURAMOTO et al., 2007). The three descriptions from the literature differ substantially

between them, and the calls from the Western Ghats (KADADEVARU et al., 2002) are most

similar to the calls of the holotype. One of the most striking features of this callis the first note

which is longer than the following ones (including more pulses) and separated from the second

one by a longer interval than all the following note intervals of the call. This first longer note

has also been reported by KADADEVARU et al. (2002) but the first small pulse of the first note

mentioned by these authors has not been observed in our calls. The three previously described

calls and the calls described here share a mean dominant frequency of 3.6-3.8 kHz, a short

call duration (0.668-1.21 s) and a number of notes per call ranging from 7 to 28.

Source : MNHN, Paris

140 ALYTES 27 (4)

The calls reported by KURAMOTO & JosHY (2001) differ in structure from the other calls

assigned to this species (KADADEVARU et al., 2002; KURAMOTO et al., 2007; this paper) with

respect to the notes which increase in length during the emission of the call and which tend to

be divided in pulse groups, ultimately reaching a duration of 300 ms which is about ten times

the duration reported here and in KADADEVARU et al. (2002). The numerous and fine spectral

bands are not found in the calls described in the other papers. The three frequency bands

observed by KADADEVARU et al. (2002) were not found here, nor in KURAMOTO et al. (2007).

Although the description of KURAMOTO et al. (2007) was very short, the structure of the call

and the few parameters given fit well with the call of the holotype (although the notes are

emitted at a faster rate in the call of KURAMOTO et al. (2007), i.e. 15.7 notes/s vs. 9-11 notes/s

in the call of the holotype).

A second species of Minervarya has recently been described from West Bengal as M.

chilapata (Ourer et al., 2009) and a short part of its advertisement call described. The

structure of the call of this species is basically the same as that of M. sahyadris, consisting of

a rapid succession of pulsed notes. Three frequency bands are found in M. chilapata vs. only

onein M. sahyadris, lying at about 3.5, 7.0 and 10.5 kHz. Nevertheless the duration of the calls

is greater (from 0.9 to 1.9 s) in M. chilapata and their calls include more notes (11-27),

the duration of the notes is longer (70 ms in average), the notes are emitted at a quicker rate

(13.6 notes/s) and there is no interval between two consecutive notes. Furthermore, the first

longer note with a greater interval relative to the following characteristic of M. sahyadris was

not found in M. chilapata.

The genus Fejervarya is a conservative genus composed of morphologically similar

species. The taxonomy of this group is at present not fully resolved and new species will

probably be described with increasing population samplings and data set collections (espe-

cially of molecular and acoustic data). In this paper, we described the advertisement call of the

holotype of Æ pierrei, F. teraiensis and M. sahyadris, of paratypes of Æ nepalensis, and of

topotypes of Æ rufescens. These data should be very useful in species allocation (and

taxonomy in general) of newly collected (and recorded) specimens. On the other hand, the

advertisement calls attributed to F syhadrensis and F. multistriata must be taken with caution

as the recorded specimens were caught quite far from their respective type localities. In

particular a recent study based on molecular data (KoTAkI et al., 2010) showed that at least

some Chinese populations of Fejer 1 belong actually to Fejervarya limnocharis. Further

research is needed to confirm their specific allocation.

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Source : MNHN, Paris

Alytes, 2011, 27 (4): 142-150.

Description of the advertisement call

of the Moroccan midwife toad

(Alvtes maurus Pasteur & Bons, 1962)

Rafael MARQUEZ*, Juan Francisco BELTRAN**, Tahar SLIMANI***,

Mohamed RaDi***, Diego LLUSIA* & El Hassan EL MOUDEN***

* Fonoteca Zoolôgica, Departamento de Biodiversidad y Biologia Evolutiva,

Museo Nacional de Ciencias Naturales (CSIC).

José Gutiérrez Abascal 2, 28006 Madrid, Spain

<rmarquezmnen.csic.es>, <diego_Ilusia@mnen.csic.es>

** Departamento de Fisiologia y Zoologia, Facultad de Biologi

Universidad de Sevilla, Avda. Reina Mercedes 6, 41012 Sevilla

<beltran@ >

*#* Laboratoire Biodiversité et Dynamique des Écosystèmes, Département de Biologie,

Faculté des Sciences Semlalia, Université Cadi Ayyad,

Avenue Prince Moulay Abdellah, BP 511, 40000 Marrakech, Morocco

<slimani@ucam.ac.ma>, <radibam@hotmail.com>, <elmouden@ucam.ac.ma>

The Moroccan midwife toad is to date the only species of Alytes whose

advertisement call remains undescribed. We describe the calls of this

species from recordings obtained in a population of the Parc National de

Tazzeka (Middle Atlas range) on 21-24 April 2010. We sampled the

population calls with two methods: (1) we recorded advertisement calls of

focal animals at close range with al audio equipment (Sound Devices

722 & Sennheiser MKH70 or Olympus LS-10), and some of the recorded

individuals were captured after recording and their weight and size was

measured; and (2) we also placed an automated timed recording station

(Song Meter SM1) in the population, for long-term monitoring of calling

activity. Recordings obtained were analyzed using Raven Pro 1.3 software.

Males call at dusk and at night during rainy nights. They may call from

underground or under rocks or on the surface. Both males carrying eggs

and males without them were observed calling. We did not hear female calls.

Soil temperature during the focal recordings was between 13 and 18.9°C.

The mean snout-vent length and weight of the seven males recorded and

captured subsequently were 40.2 mm and 6.91 g (ranges 38-43 mm and

6-8 g). A total of 7 focal recordings of individual males including 202 calls

were obtained for analy: The call is a short tonal burst of sound, like a

ief rise time (attack) and a longer fall time, and

no additional salient amplitude modulation in the call. The total call

duration of the focal recordings was short (mean 97.3 ms, range 77-217

ms). The call has a very simple spectral structure with one emphasized

frequency (mean 1324 Hz, range 1218-1406 Hz) and no harmonics nor

frequency modulation. Overall, both adult sizes and call parameters are well

within the range of variation of those of Alytes obstetricans. The calls are

shorter in duration than those of Alytes cisternasii and Alytes dickhilleni

at similar temperatures.

Source : MNHN, Paris

MARQUEZ et al. 143

INTRODUCTION

The Moroccan midwife toad (Alytes maurus) is the least known of all the species of

Alytes. After the initial debate about its full specific status (ARNTZEN & SZYMURA, 1984;

ARNTZEN & GARCiA-PARIS, 1995; BONS & GENIEZ, 1996; SALVADOR, 1996; ALTABA, 1997), an

effort has been made to obtain information about its basic biology such as distribution,

phenology and co-occurrence with other amphibians (Lists, 1985; MELLADO & MATEO, 199

DONAIRE-BARROSO & BOGAERTS, 2003; MATEO et al., 2003; DONAIRE-BARROSO et al., 2006;

PLEGUEZUELOS et al., 2010). However, basic ecological information and more specifically

behavioral information is still not available for 4. maurus.

The species of Alytes studied to date have a courtship that involves male emitting

advertisement calls to attract females (CREsPo, 1981; BUSH, 1993, 1996, 1997; MARQUEZ &

BosCH, 1995; BUSH et al., 1996) and vocalizing females approaching the calling male (MAr-

an elaborate amplexus occurs on land (DE

HÉRON-ROYER, 1886; BOULENGER, 1912; LOPEZ-JURADO et al., 1979;

E , 1984; MÂARQUEZ & VERRELL, 1991; BUSH, 1993; Bush & BELL, 1997),

where males twine the strings with fertilized eggs around their ankles and keep them on land

for several weeks until the tadpoles are fully formed. The male then walks to the water and

releases the egg masses and the tadpoles hatch.

In this paper we contribute to the knowledge of 4. maurus by describing its advertise-

ment call, a characteristic that can be of use to detect the presence of this secretive species in

unexplored parts of its potential range and that can also be used to establish acoustic

monitoring protocols of key populations of this species (e.g., determining frequency ranges

of the species advertisement calls for automated recorders, etc.).

MATERIAL AND METHODS

Males were recorded in Tazzeka National Park, Taza (Morocco) at night on 21-24 April

2010 (34°08°23.12"N; 04°04°09.09”W; 1283 m a.s.1.) in a rocky area adjacent to permanent

springs known as Aïn Khabab. Focal recordings of localized males were obtained with a

Sound Devices 722 digital recorder with a Sennheiser MKH70 microphone or with an

Olympus LS-10 digital recorder on the nights of 21 and 24. Recordings were made at 96 KHz

and 24 bits. When the recorded individual could be captured after recording, the air and soil

temperature near the recorded individual were subsequently measured to the nearest 0.1°C

with a Fluke K 72 thermocouple thermometer. In addition, snout-vent length (SVL) to the

nearest millimeter was measured by pressing the male flat (ventrally) against a ruler, and

weight was measured with a Pesola spring scale to the nearest 0.5 g. If the males were carrying

eggs, the total number of eggs visible from the top plus the number visible from the bottom of

the animal was considered an estimate of the clutch size. AI animals collected were released

in the site to prevent any impact on the population.

In addition to the focal recordings, additional recordings were obtained with an auto-

mated recording system (Song Meter SMI, by Wildlife Acoustics, Concord, MA. USA),

Source : MNHN, Paris

144 ALYTES 27 (4)

which was left in the calling area from 21 to 24 April with a recording protocol of 55 minutes/

hour. Automated recordings were made at 44.1 KHz and 16 bits. Air and soil temperature (to

the nearest 0.3°C) and relative humidity (to the nearest 2.8 %) were recorded with Hobo

Pro-V2 and Hobo Pendant 64 Kb dataloggers adjacent to the automated recording station.

Recordings were analyzed with Raven Pro 1.3 software (CHARIF et al., 2008) on Apple

Macintosh computers in the facilities of Fonoteca Zoolôgica (Museo Nacional de Ciencias

Naturales, Madrid). For spectral analyses, a window size of 4000 samples was used (7.93 Hz

precision). Recordings were deposited in the collection of Fonoteca Zoolôgica (FZ Collec-

tion numbers 8994-9000). À sample recording is available on line on the web checklist

<www.FonoZoo.com/frogcallsoftheworld>.

RESULTS

Males call at dusk and at night during rainy weather. They may call from underground or

under rocks or on the surface. Both males carrying eggs and males without them were

observed calling. Only one male was observed (and recorded) carrying eggs (estimate 35 eggs:

fig. 1). We did not hear female calls (fig. 2). Only one type of call was recorded or heard during

the study period. The advertisement call of Alytes maurus simple pure tone with a fast rise

time and a slow and continuous (not amplitude-modulated) fall time (fig. 3).

Recordings were obtained from seven calling males, which were subsequently captured,

measured and weighted (the weight of one male could not be obtained accurately because it

was carrying a clutch of eggs). Average calling male SVL was 40.2 mm (SD 1.92, range 38-43,

n 7); average calling male weight was 6.91 g (SD 1.01, range 6-8, n 6). Average air temperature

during recording was 16.0°C (range 15.2-17.4°C); average soil temperature was 15.9°C (range

13.1-18.9°C). A total of 202 calls were analyzed, the minimum number of calls analyzed per

male was 17 and the average was 28.8.

The values measured for call duration and frequency obtained from the focal recordings

are shown in tab. 1. Data on call interval were not calculated because the recordings were

made while stimulating males’ responses with the emission of vocalisations and whistles by

the experimenters. No measurements of Sound Pressure Level were obtained, but the overall

call intensity resembled that of the Iberian species.

Calls were recorded with the automated recording system from 4 a.m. to 6 a.m. (GMT)

on 23 April (air temperature 8.1-8.3°C, soil temperature 11.1-11.2°C, relative humidity 96 %)

and 8 pm. to 2 a.m. on 23-24 April 2010 (air temperature 9.6-10.8°C, soil temperature

11.6-12.5°C, relative humidity 95.2-95.8 %). In both cases, noise of heavy rainfall was

recorded before the . À total of 466 male calls were recorded and analyzed (tab. 2), which

included calls in 13 discernible frequency categories. Given that male call frequency does not

change within calling nights in other species of A/ytes (MARQUEZ & BosCH, 1995), we estimate

that this was à minimum number of calling males in these recordings. In 9 cases, individual

calling males could be reasonably identified and inter-call intervals of the (likely) same

individual could be measured. Given that these call intervals were not affected by artificial

calls produced by the res hers, they are included in tab. 2. Call frequencies were similar to

those obtained in the focal recordings. On the other hand, call durations were longer, probably

a consequence of lower calling temperatures (see MARQUEZ & BoscH, 1995).

Source : MNHN, Paris

MARQUEZ et al. 145

Fig. 2. — Female of A/ytes maurus. © R. Märquez.

Source : MNHN, Paris

146 ALYTES 27 (4)

Fig. 3. — Oscillogram (top) and audiospectrogram (bottom) of a male advertisement call of Alpes

maurus (SVL 40 mm, weight 6 g), recorded at à soil temperature of 18.9°C. Sound parameters: 96

KHz, 24 bits. Spectrogram: FFT 512, Hanning filter.

DISCUSSION

Overall, the advertisement call of 4. maurus closely resembles the calls of the congeneric

species from the European continent, being a short pure tone, with no harmonics, no

frequency modulation (HEINZMANN, 1970; CRespo, 1981; CRESPoO et al., 1989; MARQUEZ &

BoscH, 1995) and substantially lower in frequency than the Balearic insular endemic, A/ytes

muletensis (BUSH, 1997). The relationship between call duration and temperature has been

shown to differentiate between the continental species and subspecies of Alytes (MÂARQUEZ &

BoscH, 1995). If we compare the results obtained with 4. maurus, the measurements from A.

maurus are well within the range of values obtained for 4. obstetricans in Iberia and are

shorter in duration than those of À. cisternasit and À. dickhilleni. The similarity of the call of

A. maurus With À. obstetricans is consistent with the closer phylogenetic affinity between these

two species relative to À. cisternasi, which occupies an ancestral position in the genus

(MARTINEZ-SOLANO et al., 2004).

The recordings obtained have a low standard deviation and range (less than 200 Hz).

This is admittedly a limited sample size, but if the recordings from the automated device

obtained over a complete season confirm this trend, this may indicate that the calling

individuals are homogeneous in size (because size and call frequency are correlated in other

Source : MNHN, Paris

MARQUEZ et al. 147

Table 1.- Values of call frequency and call duration from the seven focal recordings made.

Mean and standard deviation (SD) are the average of seven individual means. Range is

maximum and minimum of all measured calls.

Call frequency (Hz) Call duration (ms)

n males 7 7

n calls 202 202

Mean 1324.24 97.34

Average SD (between males) 33.95 0.01

Range ë 1218.8-1406.2 77.0-217.0

Table 2. - Values of call frequency, call duration and call interval from the recordings

obtained by the automated recording system. Mean and standard deviation (SD) are

the average of all calls recorded (not grouped by individual). Range is maximum and

minimum of all measured calls. Call interval was measured for nine clearly discernible

individuals.

Call frequency (HZ) | Call duration (ms) Call interval (s)

n males 13 13 9

n calls 466 466 243

Mean 1286.16 130.12 8.50

Average SD (between 26.89 0.02 3.04

males)

Range 1238.0-1367.2 62.0-190.0 0.4-19.9

species of 4/) see MARQUEZ, 1993). This homogeneity of size can be related to the

possibility that only some cohorts reach sexual maturity (those produced in more favorable

years). Of course this possibility needs to be confirmed by more continuous monitoring of the

population.

The population of À. maurus analyzed was the only one where we could record calling

activity in visits to more than seven different sites over five years where the species was

previously reported in the Rif and Atlas Mountains of Morocco. In all of them the larvae

were scarce or absent, and the sites all appeared to be encroached by human use of water

resources. The other species of A4lytes are known to be vulnerable to pathogens that cause the

extinction of populations (BoscH et al., 2001) and there are important conservation efforts in

place in other countries (BoscH, 2010). We believe that this species deserves a serious effort of

monitoring (SLIMANI et al., 2010) and that its conservation status is likely to be more

worrisome than currently believed (PLEGUEZUELOS et al., 2010).

Source : MNHN, Paris

148 ALYTES 27 (4)

RESUMEN

El sapo partero marroqui es hasta la fecha la ünica especie del género Alytes cuya

Ilamada de apareamiento aûn no ha sido descrita. En este articulo se describe la Ilamada de

apareamiento de esta especie a partir de grabaciones obtenidas en una poblaciôn del Parque

Nacional Tazzeka (Atlas Medio, Marruecos) entre el 21 y el 24 de Abril de 2010. Para el

muestreo de las vocalizaciones se utilizaron dos métodos: (1) grabaciôn de animales focales a

corta distancia con grabadoras digitales (Sound Devices 722 y Sennheiser MKH70 u Olym-

pus LS-10); y (2) grabaciôn de coros con un sistema automätico de grabaciôn con tempori-

zador (Song Meter SM1) con el que se realizarä el seguimiento a largo plazo de la actividad

acüstica de la poblacién. Los machos vocalizaron enterrados, bajo piedras o en la superficie,

al atardecer y durante la noche de dias Iluviosos. La temperatura del suelo durante las

grabaciones oscilé entre 13 y 18.9°C. La media de tamaño corporal y peso de siete machos fue

de 40.2 mm y 6.91 g (rangos 38-43 mm y 6-8 g). La Ilamada del sapo partero marroqui es un

tono puro de corta duracién, parecido a un corto silbido. Su estructura temporal se caracte-

riza por tener una fase de ataque muy breve, seguida de una larga caida sin modulaciones de

amplitud notables adicionales. La duraciôn total de la Ilamada fue de media 97.3 ms (rango

77-217 ms). La Ilamada tiene una estructura espectral simple y una sola frecuencia enfatizada

(media 1324 Hz, rango 1218-1406 Hz), careciendo de arménicas y de modulacién de

frecuencia. En conjunto, tanto el tamaño corporal como los parämetros de la Ilamada de

apareamiento del sapo partero marroqui se encuentran dentro del rango de variacién de 4.

obstetricans. Las Hlamadas son mäs cortas en duraciôn que las de A. cisternasit y À. dickhilleni

à temperaturas similares.

RÉSUMÉ

Le crapaud accoucheur du Maroc est la seule espèce du genre A/ytes dont le chant

d'accouplement n'a pas encore été décrit. Dans cet article le chant d'accouplement de cette

espèce est décrit à partir d’enregistrements obtenus dans une population du Parc National de

Tazzeka (Moyen Atlas, Maroc) entre le 21 et le 24 avril 2010. Deux méthodes ont été utilisées

pour obtenir les enregistrements: (1) enregistrements focaux d'animaux localisés à courte

distance avec un équipement audio (Sound Devices 722 et Sennheiser MKH70 ou Olympus

LS-10), avec capture et notation de poids et taille corporelle après enregistrement, et (2)

installation d’un système automatique d'enregistrement temporisé (Song Meter SMI) qui

sera utilisé pour suivre la phénologie acoustique de la population. Les enregistrements

obtenus ont été analysés avec le logiciel Raven 1.3. Les mâles émettent leurs chants enterrés,

cachés sous des pierres ou sur la surface du sol, le soir et la nuit après des journées pluvieuses.

La température du sol pendant les enregistrements était de 13-18.9°C. La taille et le poids

moyens des sept mâles enregistrés et capturés est de 40,2 mm et 6,91 g (extrêmes 38-43 mm et

6-8 g). Le chant d'accouplement du crapaud accoucheur du Maroc est un ton pur de courte

durée, semblable à un court sifflement. Sa structure temporelle ou enveloppe se caractérise par

une attaque très brève et une descente plus longue sans avoir d’autres modulations d’ampli-

tude appréciables. La durée moyenne des chants enregistrés est de 97,3 ms (extrêmes 77-217

ms). Le chant a une structure spectrale très

simple centrée sur une seule fréquence (moyenne

Source : MNHN, Paris

MARQUEZ et al. 149

1324 Hz, extrêmes 1218-1406 Hz), ne présentant ni harmoniques ni modulation de fréquence.

En général, aussi bien la taille que les paramètres du chant d’accouplement du crapaud

accoucheur du Maroc se trouvent dans le domaine de variation d’Alytes obstetricans. Les

chants sont plus courts que ceux d’A. cisternasii et A. dickhilleni à des températures similaires.

ACKNOWLEDGEMENTS

This work was funded by the Comité Mixto Hispano-Marroqui (AECID, Spain), Acciones Integra-

das A/016244/08 and A/023074/09 (PIs, T. Slimani & J. F. Belträn) and by the project ACOURA,

CGL2008-04814-C02 Ministerio de Ciencia e Innovaciôn (PL. R. Märquez). It was authorized by the

Haut Commissariat aux Eaux et Forêts et à la Lutte contre la Désertification (Maroc), Scientific Permits

N°98/2009 et 15/2010 HCEFLCD/DLCDP/DPRN/CFF. We thank M. Benitez, M. Chirosa, S. Fahd, J.

Barbadillo and D. Donaire for their help in localizing populations.

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Source : MNHN, Paris

Alytes, 2011, 27 (4): 151-153. 151

Subgeneric taxonomy and nomenclature

of the genus Hypselotriton

Wolterstorff, 1934

(Amphibia, Urodela)

Alain DuBois* & Jean RAFFAËLLI**

* Reptiles & Amphibiens, UMR 7205 OSEB.

Département Systématique & Evolution, Muséum national d'Histoire naturelle,

CP 30, 25 rue Cuvier, 75005 Paris, France

dubois@mnhn.fr>

** Penclen, 56420 Plumelec. France

<jean.raflaelli@laposte.net>

Two subgenera were recently recognized in the newt genus Hypselotri-

ton Wolterstorff, 1934. The nucleospecies of the subgenus Pingia Chang,

1935, described as Pachytriton granulosus Chang, 1933, was recently

shown not to be a member of this genus but a synonym of Pachytriton

labiatus (Unterstein, 1930). À new nomen is therefore proposed for this

subgenus which includes three species: Hypselotriton fudingensis (Wu,

Wang, Jiang & Hanken, 2010), Hypselotriton orientalis (David, 1875) and

Hypselotriton orphicus (Risch, 1983).

In a previous paper (DuBois & RAFFAËLLI, 2009), we proposed a new taxonomy of the

urodelan family Sazasanpripar Goldfuss, 1820, in which we restricted the former genus

Cynops Tschudi, 1838 to the Japanese species, whereas we placed all Chinese species in the

genus Hypselotriton Wolterstorff, 1934. Within this latter genus, we recognized two subge-

nera, corresponding 10 the two species-groups identified by ZHAO & HU (1984, 1988). The

subgeneric hyponymous (nominotypical) nomen Hypselotriton, the nucleospecies (type-

species) of which is Molge iolierstorfft Boulenger, 1905 by original designation, applies to the

subgenus including the species Hypselotriton chenggongensis (Kou & Xing. 1983), Hrpselo-

triton cyanurus (Liu, Hu & Yang, 1962) with its two subspecies, and Hypselotriton wolterstorffi

(Boulenger, 1905). For the second subgenus, including the species Hrpselotriton orientalis

(David, 1875) and Hypselotriton orphicus (Risch, 1983), we used the generic nomen Pingia

Chang, 1935. This nomen is based on the nucleospecies Pachytriton granulosus Chang, 1933,

the holotype of which was lost and the taxonomic status of which was long disputed. HOU et

al. (2009) had designated a neotype and provided a redescription of this species, which they

considered close to, although distinct from, Hyrpselotriton orientalis, so that we recognized

three species in the subgenus Pingia of Hypselotriton

Shortly alter however, NisHiKAWA et al. (2009) provided a detailed study of morpholo-

gical variation in the species Pachvtriton labiatus (Unterstein, 1930) and concluded that the

Source : MNHN, Paris

152 ALYTES 27 (4)

lost holotype and the neotype of Pachytriton granulosus belonged in fact in the latter species.

The confusion was due to the fact that the juvenile phenotype of Pachytriton labiatus is very

different from its adult phenotype. The specific nomen Pachytriton granulosus then becomes

an invalid junior synonym of Pachytriton labiatus, and the generic nomen Pingia an invalid

junior synonym of Pachytriton.

This finding leaves the second subgenus of Hypselotriton unnamed. Besides the two

species Hypselotriton orientalis and Hypselotriton orphicus, it includes the species Hypselotri-

1on fudingensis recently described by Wu et al. (2010). As no genus-series nomen is available

for it, we hereby provide a nomen for this subgenus. The entexognosis, the diagnosis and the

idiognosis we give to this taxon follow the same plans and should be compared with those of

the subgenus Æypselotriton in DUBoIs & RAFFAËLLI (2009: 45-48).

Subgenus Cynotriton nov.

Nucleospecies. — Triton ( Cynops) orientalis David, 1875, by present designation.

Etymology. — From the Greek Aunos, genitive of kuon (“dog”) and generic nomen Triton

Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. - The most inclusive holophyletic taxon including the species Hypselotriton

orientalis (David, 1875) and excluding the species Hypselotriton wolterstorfii (Boulenger,

1905).

sis, — (1) Frontal process of premaxillary short. (2) Parotoid glands well developed.

cules on external side of hands and feet absent. (4) Skin slightly to very granular.

(5) Adaptability in terrarium high. (6) Altitudinal distribution low (0-1000 m).

Idiognosis. — (1) TL 70-90 mm. (2) Habitus stout. Trunk almost quadrangular. Parotoids well

developed. Skin slightly to very granular. (3) Dorsal coloration dull, sometimes with bright

(red) spots or bands. Ventral coloration very bright, red. (4) Sex dimorphism strong, male

small. (5) Mainly aquatic, in lentic habitat. (6) Adaptability in terrarium high, with tolerance

of a large gradient of temperature (5-25°C). (7) Eastern China. (8) No tubercules on the

external side of hands and feet.

Content. — Hypselotriton (Cynotriton) fudingensis (Wu, Wang, Jiang & Hanken, 2010);

Hypselotriton (Cynotriton) orientalis (David, 1875); Hypselotriton (Cynotriton) orphicus

(Risch, 1983).

LITERATURE CITED

Dunons, A. & RarraëlLLi, J., 2009. - À new ergotaxonomy of the family Salamandridae Goldfuss, 1820

(Amphibia, Urodela). A/rtes, 26 (1-4): 1-85.

-C. Li, P.-P. & LÜ, S.-Q., 2009. - Rediscovery of Pingia granulosus (Chang, 1933) and

description of the neotype specimens [sie]. Sichuan Journal of Zoologr, 28 (1: 15-18,3 pl.

Source : MNHN, Paris

Dusois & RAFFAËLLI 153

NisHikawWa, K., JIANG, J.-P., MATSUI, M. & CHEN, C.-S., 2009. - Morphological variation in Pachytriton

labiatus and a re-assessment of the taxonomic status of 2 granulosus (Amphibia: Urodela:

Salamandridac). Current Herpetology, 28 (2): 49-64.

Wu, Y. WanG, Y., JIANG, K. & HANKEN, JL, 2010. — A new newt of the genus Cynops (Caudata:

Salamandridae) from Fujian Province, southeastern China. Zootaxa, 2346: 42-52.

Zuao, E. & Hu, Q. 1984. - Studies on Chinese tailed amphibians. Chengdu, Sicuan Scientific & Technical

Publishing House: [i-v] + 1-68, 3 pl.

_ 1988. - The classification and evolution of the Chinese Salamandridae. Jr: E. ZHAO, Q. HU, Y. JIANG

& Y. YANG (ed.), Studies on Chinese Salamanders, in: Contributions in Herpetology, 4, SSAR,

Oxford (Ohio): 12-26.

Corresponding editor: Stéphane GROSJEAN.

©ISSCA 2011

Source : MNHN, Paris

Alytes, 2011, 27 (4): 154-160.

The authorship and

date of the familial nomen Ranidae

(Amphibia, Anura)

Alain DuBois & Roger BOUR

Reptiles & Amphibiens, UMR 7205 OSEB,

Département Systématique & Evolution, Muséum national d'Histoire naturelle,

CP 30,25 rue Cuvier, 75005 Paris, France

<adubois@mnhn.fr>, <bour@mnhn.fr>

The anuran familial nomen RANIDAE has been credited with several

authorships and dates in the recent decades. Some of these changes were

due to the rediscovery of older works, and some to modifications in the

Rules of the Code. The rediscovery of the work of BaTscH (1796) brings a

new change in this respect, as this nomen was indeed created in this book.

This is the first familial nomen of amphibians created, whereas TESTUDINIDAE

Batsch, 1788 is the first familial nomen of reptiles.

The zoological Code (ANONYMOUS, 1999) recognizes three “groups of names” or better

“‘nominal-series” (DuBois, 2000): the species-, genus- and family-series. Family-series nomen-

clature (i.e., nomenclature of taxa at ranks family, superfamily, subfamily, tribe, subtribe and

additional intermediate ranks) is regulated by the Code but these Rules have regularly been

ignored by some zootaxonomists. Some believe that no Principle of Priority applies to these

nomina and that the valid nomen of a family-series taxon is fixed by “usage” or “consensus”.

Others imagine that any such nomen should be credited to the first publication where it was

used with its now correct spelling, e. : for the family including the genus Rana

Linnaeus, 1758. Still others think that, to be valid, a family-series nomen must be based on a

generic nomen considered valid, and must be changed when the latter becomes invalid (e.g..

for being discovered to be a junior synonym). AI of this is wrong. The valid nomen of a

family-series taxon is the senior one, among all of those potentially available for the taxon

(i.e., based on generic nomina now referred to the taxon), except if it is a junior homonym or

based on à generic nomen which is itself a junior homonym, and irrespective of the validity of

the generic nomen on which it is based (type genus or nucleogenus); and the author and date

of any family-series nomen are those of the work where a nomen based on this nucleogenus

coined for a taxon of any rank in the family-series, whatever its ending, under the

condition that this nomen was clearly in the nominative plural. The rationale for these Rules

was discussed in detail by Myers & LEVITON (1962) and Dumois (1984, 1987b, 2005a, 2011).

Another rather frequent mistake in zoological nomenclature consists in ignoring the

Principle of Coordination, which states that all family-series nomina based on the same

Source : MNHN, Paris

Duois & BOUR 155

generic nomen (e.g.. family RanD4r, subfamily RanIN4E, superfamily Ra oIDEA, tribe RANINI,

subtribe RAnA, etc.) have the same author and date, that of the first nomen ever proposed for

a family-series taxon containing the genus at stake. This Rule is too often ignored, even in

works by professional zoologists (e.g.: FROES, 1957; ALBOUY & CAUSSANEL, 1990; Gasc, 1990:

MEN & GiNsBURG, 1997) and, at least for some entries, in databases dealing with zoological

phylogeny, taxonomy and nomenclature (ANONYMOUS, 2011; BRANDS, 2011).

The familial nomenclatures of rather few zoological groups have been surveyed extensi-

vely for the valid nomina of taxa and especially for their valid authorships and dates. In many

groups, nomina are used following some kind of consensus, and no authors and dates are

given to nomina. Exceptions include the mammals (WiLsoN & REri 2005), the birds

(Bock, 1994), some mollusks (BOUCHET & ROCROI, 2005, 2010) and crustaceans (NG et al.,

2008). In herpetology, the only groups to have been exhaustively and seriously treated are: (1)

the recent amphibians, that were covered by DuBois (1983, 1984, 1985, 2005b), followed, but

for some mistakes (see DuBois, 19874), by FROST (1985, 2011); (2) the chelonians, treated by

Bour & Dusois (1985, 1986), followed, but for some mistakes, by RHODIN et al. (2008, 2009,

2010); (3) some snakes (McDiarMib et al., 1999). No such comprehensive treatment has been

published so far for the other groups of “reptiles”, which explains that regularly correctio:

have to be published concerning the authorships and dates of some family-s: omina (e.g..

Dusois & BouR, 2010b).

Even in the groups that have been seriously surveyed, it is not rare that earlier uses of

family-series nomina are discovered for well-known groups. This is easy to understand. The

earliest recognized family-series taxa were based on Linnaean generic nomina made available

in the 1758 and 1766-1767 editions of LINNAEUS’ Spstema Naturae. The latter works having

been well-known to all zoologists since their publication, any author could coin a family-

series nomen based on a generic nomen for a taxon including this well-known genus. Linnaeus

did not use the rank family (Dusois, 2007) and there is no official starting date for the use of

family-series nomina based on generic nomina in zoology. The e: st publication using this

system that we know of is that of BaTsCH (1788, 1789), where this author coined 17 familial

nomina that are duly available under the current Rules of the Code and that should be credited

to him, not to subsequent authors who used the same nomina (Dugois & BouR, 2010b).

In the early days of zoological nomenclature, authorship and date of nomina were not

strictly regulated and were often ignored because of a widespread “mihilism” (BRUUN, 1950;

Dusois, 2008), i.e., a propensity of some authors to claim authorship for some nomina

although they had not coined them but just “redefined” them. Now we have strict Rules, and

nomina are created once and for all and cannot be “redefined”, because of the existence of

what has been called the Principle of Nomenclatural Foundation (DuBois, 2011). In the three

nominal-series covered by the Code, nomina are “defined” only through their onomatophore,

not by any intensional or extensional definition (DuBois, 2011). Therefore, if two different

authors create independently (i.e., in the ignorance of the others work) two family-series

based on the same generic nomen, the latter is the onomatophore of both and they are strict

objective synonyms or isonyms (DUBoIs, 2000). It could be argued that in such a case the two

nomina are indeed independent homonymous and isonymous nomina, but this would have no

practical consequence on the valid author and date of the family-series nomen, which would

in all cases remain that of the first published one. Practically, it would be very difficult, if not

Source : MNHN, Paris

156 ALYTES 27 (4)

impossible in some cases, to ascertain whether the second author had used the nomen

proposed in an earlier work by the first author, or whether he/she thought he/she had indeed

coined a “new” nomen, because most authors of that time (and in fact still nowadays - even

in taxonomic revisions, faunistic lists, catalogues of specimens or taxa, etc.) just mentioned

family-series nomina but not their authors and dates. It is therefore much simpler, and without

any nomenclatural consequence, to consider that any family-series nomen based on a given

generic nomen has been coined only once, in the first publication where it appeared, and that

any subsequent appearance of this nomen, either identical or modified in its ending (e.g.,

—INAE instead of -14£), is not a new nomen but respectively a chresonym or an aponym of the

protonym used in the first publication (for the definitions of these terms, see DuBois, 2000).

This guideline was used in all our previous works (e.g., DuBois, 1984; Dugois & BoUR, 2010)

and we use it here.

The case of the family Ravip4r is a very enlightening one. Although this nomen, under

various spellings, has been used continuously by all authors since 1825 for a family including

the genus Rana Linnaeus, 1758, its authorship and date have changed regularly, in part

because of incomplete bibliographic surveys by authors, and in part because inappropriate

changes implemented in the Code in 1985 (for a discussion of this point, see DUBoIs, 1987b,

2011). The authorship and date traditionally credited to this nomen (e.g., DOWLING &

DUELLMAN, 1974-1978) was BONAPARTE (1831), because this author was thought (in error) to

have been the first one to use the correct spelling RawsDar for this family. DuBois (1984)

pointed out that the spelling Ranipar had been used already by Bot (1828), but, as established

by Dugois (1981), the nomen had in fact been created by GRAY (1825) under the spelling

Rawabar. However, an earlier nomen, Ravaripia Rafinesque-Schmaltz, 1814, based on Rana-

ria Rafinesque-Schmaltz, 1814, an invalid neonym (nomen novum) for Rana Linnaeus, 1758,

was also available for this family. According to the Code then in force (ANONYMOUS, 1964), the

family had to be named Ranpar Gray, 1825 (1814) and its eponymous taxa (Ra orp£4,

RanINAE, RANINI) had to be credited with the same authorship and date (Dumois, 1981, 1984).

The subsequent discovery (DUBois, 1985) that GoLbruss (1820) had already recognized a

family Ravaz should have led to a new authorship and date for the family, as Ramipaz

Goldfuss, 1820 (1814), but the third edition of the Code (ANONYMOUS, 1985) modified the

Rules in force in such c« so that the valid nomen became Rawipar Rafinesque-Schmaltz,

1814 (for details, see Dugois, 1985). Here we report on a new discovery that again modifies the

authorship and date of this familial nomen.

The work of BATSCH (1788) has never been completely forgotten (see e.g. STEINE

1907), but it was not until the resurrection by Bour & DuBois (1985) of the famil

nomen 7ksruninNes Batsch, 1788 as the valid nomen for the family T£srupinip4s that it was

used as the first identified source of available family-series nomina in zoology. DuBois & BOUR

(2010b) confirmed this fact and pointed to 16 other available and valid familial nomina over

the whole of zoology coined by BATSCH (1788, 1789) that had been ignored until then.

BaTsCH (1788, 1789) was not consistent in the derivation of his familial nomina. These

nomina belong in three categories (Dusois & BoUR, 2010b). Some were clearly based on an

available generic nomen recognized by Batsch as designating a valid genus: these rhizonyms

(Dusois, 2006b) are correctly formed and they are available nomina under the Code or

hoplonyms (Dusois, 2000). Others were not based on generic nomina: such arhizonpms

Source : MNHN, Paris

Dusois & BOUR 157

(Dusois, 20064) are incorrectly formed and are unavailable nomina under the Code or

anoplonyms (Dusois, 2000). Finally, others are based on available generic nomina, but the

latter were not used by Batsch as valid in the new family-series taxon; these nomina can be

called cenorhizonyms — from the Greek xevés (kenos), “empty, vain”, ét (rhiza), “root” and

ävoua (onoma), “name”. The status of such nomina is addressed in Article 11.7.1.1 of the

Code, which states that, for a new family-series nomen to be available it must be based on a

generic nomen “then used as valid in the new family-group taxon”. The meaning of “then” in

this Article is questionable, but we here follow the interpretation that we presented elsewhere

(Dusois & BouR, 2010b), according to which the generic nomen must be used as valid in the

new family-group taxon as recognized in the work where its nomen is created. According to this

interpretation, cenorhizonyms are also anoplonyms under the Code.

The 17 family-series rhizonyms in BATSCH (1788, 1789) are doubtless nomenclaturally

available. They have priority over all other nomina proposed later on for the same taxa and

should replace them. As these changes in authorships and dates do not imply any change in

the nomina of the family themselves, they can and must be implemented without delay in all

the zoological groups concerned (DuBois & BOUR, 2010).

Another work of BATSCH (1796) seems to be still rarer than his first two books. It was

mentioned in catalogues in the 19!" century, but forgotten in the 20°", It was “rediscovered” in

the 21" century (CHANDRA, 2005; RHODIN et al., 2008) by authors who mentioned it but did

not discuss its nomenclatural implications in zoology. This work was recently digitalized and

made available to all interested zoologists by “Die Bayerische Staatsbibliothek München”

(<http///reader.digitale-sammlungen.de/resolve/display/bsb10074788.html>), so we had

access to it. It so happens that this work contains two new available familial nomina, not

mentioned in Batsch's first work.

BaTsCH (1796: 73) recognized a family Orina. This nomen, based on Ovis Linnaeus,

1758, has priority over Bovipar Gray, 1821 for the family and its subordinated taxa including

the genus Ovis. As the nomen Bovipar has had a widespread use in zoology, it should be

validated against Oviva: we will address this question in a distinct paper.

BATSCH (1796: 179) also recognized a family Ravia, based on Rana Linnaeus, 1758. The

situation is clearer in this case. In 1788, Batsch had recognized a family “ Barracur”, With the

four frog genera Bufo Laurenti, 1768, Hyla Laurenti, 1768, Pipa Laurenti, 1768 and Rana

Linnaeus, 1758. Although a generic nomen Batrachus Schaeffer, 1760 was available at that

time (Dusois & BouR, 20104), it was a genus of “fishes” and it cannot be at stake here: the first

frog generic nomen Batrachus was created by RAFINESQUE-SCHMALTZ (1814) and cannot be

the basis for the stem of “Barracur” Batsch, 1788. The latter is therefore an arhizonym and

anoplonym (Dugois & BouR, 2010). In contrast, the nomen Raniva, which BATSCH (1796)

substituted for his nomen “Barracur”, being based on the generic nomen Rana Linnaeus,

1758 that he recognized as valid, is a rhizonym and hoplonym. It has priority over the nomen

Ranaipra Rafinesque-Schmaltz, 1814. This change is a very fortunate one, because it allows

to credit again the family-series nomen Rawip4E (widely used, and at many different ranks) to

a publication where it was indeed based on the generic nomen Rana Linnaeus, 1758 - and not

on a neonym of the generic nomen nowadays considered valid for the taxon (for details and

discussion, see DuBois 1987b, 2010).

Source : MNHN, Paris

158 ALYTES 27 (4)

Therefore Ravip4r Batsch, 1796 becomes the valid nomen of the family and of its

eponymous taxa (RANOIDEA, RANINAE, RANINI, RANINA) if recognized as valid taxa.

In herpetology, BATSCH (1788) recognized four families in his classis AMPHIBIA. The

nomen of one of them (Tesruives Batsch, 1788, for the single genus Zestudo Linnaeus, 1758)

was à rhizonym (based on the root of a generic nomen considered valid) and a hoplonym

(available) and should now be used as the valid nomen of this family, as Tesrupinipar Batsch,

1788. The other three (“Barracur” Batsch, 1788; “LacerTar” Batsch, 1788; SERPENTES"

Batsch, 1788) are cenorhizonyms (based on the root of a generic nomen considered invalid)

and anoplonyms (unavailable), and are therefore without nomenclatural existence.

Eight years later, BATSCH (1796) kept the same taxonomy for his classis AMPHIBIA but

modified the nomenclature of the families. He changed his family nomen 7ësrunines into

TESTUDINE s not a new nomen but just an aponym (derived form of nomen) of the

protonym (original form of nomen) 7£srunives, which therefore keeps the same authorship

and date, i.e., Batsch, 1788. He also modified his family nomina “LaC£RTAE" and SERPENTES"

into respectively “LacerriNa" and “S£RPENTINA": these are also aponyms, but as they remain

cenorhizonyms they remain anoplonyms and have therefore no nomenclatural existence.

Finally, he replaced his nomen “Barkacui” by the new nomen RanI4, which is a rhizonym

and hoplonym, and becomes the valid nomen of the family.

Ranipar Batsch, 1796 is the first familial nomen of amphibians ever created, whereas

Tesruniipar Batsch, 1788 is the first familial nomen of reptiles.

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2e 0

Corresponding editor: Stéphane GROSIFAN.

Source : MNHN, Paris

AIVTES

International Journal of Batrachology

published by ISSCA.

EDITORIAL BOARD

Chief Editor: Stephane GROSIEAN (Reptiles et Amphibiens, UMR 7205 OSEB, Département de Systématique &

Evolution, Muséum national d'Histoire naturelle, CP 30, 25 rue Cuvier, 75005 Paris, France;

<sgrosjea@mnhn.fr>).

Deputy anco ANDREONE (Museo Regionale di Scienze Naturali, Via G. Giolitti 36, 10123 Torino, ltaly;

<f'andreone@libero.it>).

Alytes Editorial Board: Ariadne ANGULO (Toronto, Canada); David C. BLACKBURN (Kansas, USA); Lauren E.

BROWN (Normal, USA); Angus I. CARPE TER (Norwich, UK): Ignacio DE LA Riva (Madrid, Spain); Rafa

MÂRQUEZ (Madrid, Spain); Masafumi MaTsuI (Kyoto, Japan); Annemarie OHLE €

Oliver Rë (Berlin, Germany): Miguel VENCES (Braunschweig, Germany) : Ché WELDON (Potchefstroon,

South Africa)

Amphibia Mundi Editorial Board: Alain Dumois, Chief Editor (Paris, France); Ronald 1. CROMBIE (San

Francisco, USA); Stéphane GROSIEAN (Paris, France); W. Ronald HEYER (Washington, USA); JIANG

Jianping (Chengdu, China); Esteban O. LAVILLA (Tücumän, Argentina): Jean-Claude RAGE (Paris,

France), David B. Wake (Berkeley, USA):

Technical Editorial Team (Pa rance): Alain DUBOIS (texts); Roger BOUR (tables); Annemarie OHLER (figures).

Book Review Editor: Annemarie OHLER (Paris, France).

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions to Authors, see Alvtes, 1997, 14: 175-200)

Alptes publishes original papers in English, French or Spanish, in any discipline dealing with amphibians,

side articles and notes reporting results of original research, consideration is given for publication to synthetic

review articles, book reviews, comments and replies, and to papers based upon original high quality illustrations

{such as colour or black and white photographs), showing beautiful or rare species, interesting béhaviours, etc.

The title should be followed by the name(s) and address(es) of the author(s). The text should be typewritten

or printed double-spaced on one side of the paper. The manuscript should be organized as follows: English

abstract, introduction, material and methods, results, discussion, conclusion, French or Spanish abstract,

acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or tab. 4. Figures should not excced 16

X* 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures and tables

should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & POLLS PELAZ, 1989; INGER

et al., 1974). References in the Literature Cited section should be presented as follows:

BourRer, R., 1942. - Les batraciens de l'Indochine. Hanoï, Institut Océanographique de l'Indochine: i-x +

1-547, pl. 1-4.

Gkar, J-D. & POLLS PELAZ, M. 1989. - Evolutionary genetics of the Rana esculenta complex. In: R. M. DAWLEY

& I. P. BoGaRT (ed.),

289-3

IxGëR, RE. Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some Southeust

Asian frogs of the genera Bufo and Rana. Biochem. Genet.. 12: 121-145

volution and ecology of unisexual vertebrates, Albany, The New York

e Museum:

Manuscripts should be submitted either as attached document by e-mail (pdf file including all figures and

word file without the figures if it is 100 big) or in paper form by mail but then in triplicate, cither to Stéphane

GROSIEAN (address above) if dealing with amphibian morphology, anatomy, systematics, biogeography, evolu-

tion, genetics, anomalies or developmental biology, or to Franco ANDREONE (address above) if dealing with

amphibian population genetics, ecology, ethology, life history or conservation biology, including declining

amphibian populations or pathology. Acceptance for publication will be decided by the editors following review

by at least two referees.

After acceptance, should be sent to the Chief Editor, as attachment by e-mail, the corrected text in word file

as well as solution figures.

Page charges are requested only from authors having institutional support for this purpose. The publication

of colour photographs à charged. For each publi ed paper, a free pdf or 25 free reprints are offered by ISSCA

to the author(s). Additional reprints may be purchased

Published with the support of AALRAM

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum National d'Histoire Naturelle, Paris, Fran

Directeur de la Publication: Alain DUBOIS.

Numéro de Commission Paritaire: 64851.

A 2011 Source : MNHN, Paris

Alytes, 2011, 27 (4): 117-141.

Contents

Stéphane GROSIEAN & Alain DuBoIs

Description of the advertisement calls of nine species of

Fejervarya Bolkay, 1915 and Minervarya Dubois, Ohler & Biju, 2001

from China, India and Nepal... 117-141

Rafael MARQUEZ, Juan Francisco BELTRAN, Tahar SLIMANI,

Mohamed RaDi, Diego LLUsIA & El Hassan EL MOUDEN

Description of the advertisement calls of the Moroccan midwife toad

(Alytes maurus Pasteur & Bons, 1962)............................... 142-150

Alain Duois & Jean RAFFAËLLI

Subgeneric taxonomy and nomenclature of the genus

Hypselotriton Wolterstorf, 1934 (Amphibia, Urodela) .................. 151-153

Alain Dugois & Roger BOUR

The authorship and date of the familial nomen Ranidae

PAROI DIG ANUTA) RTL E MAR en SRE ALERTE PRE Er ie MEET 154-160

Alytes is printed on acid-free paper.

Alytes is indexed in Bio

is, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 3° trimestre 2011.

©ISSCA 2011

Source : MINHN, Paris