BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. A New Species of Pseudopaludicola (Anura: Leptodactylidae: Leiuperinae) from Northeastern Brazil Author(s): Felipe de Medeiros Magalhães , Daniel Loebmann , Marcelo Nogueira de C. Kokubum , Célio Fernando Baptista Haddad , and Adrian Antonio Garda Source: Herpetologica, 70(1):77-88. 2014. Published By: The Herpetologists' League URL: http://www.bioone.org/doi/full/10.1655/HERPETOLOGICA-D-13-00054 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofitpublishers, academic institutions, research libraries, and research funders in the common goal of maximizing access tocritical research.
A New Species of Pseudopaludicola (Anura: Leptodactylidae:Leiuperinae) from Northeastern BrazilAuthor(s): Felipe de Medeiros Magalhães , Daniel Loebmann , Marcelo Nogueirade C. Kokubum , Célio Fernando Baptista Haddad , and Adrian Antonio GardaSource: Herpetologica, 70(1):77-88. 2014.Published By: The Herpetologists' LeagueURL: http://www.bioone.org/doi/full/10.1655/HERPETOLOGICA-D-13-00054
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in thebiological, ecological, and environmental sciences. BioOne provides a sustainable onlineplatform for over 170 journals and books published by nonprofit societies, associations,museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated contentindicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercialuse. Commercial inquiries or rights and permissions requests should be directed to theindividual publisher as copyright holder.
Herpetologica, 70(1), 2014, 77–88� 2014 by The Herpetologists’ League, Inc.
A NEW SPECIES OF PSEUDOPALUDICOLA (ANURA:LEPTODACTYLIDAE: LEIUPERINAE) FROM NORTHEASTERN
BRAZIL
FELIPE DE MEDEIROS MAGALHAES1, DANIEL LOEBMANN
2, MARCELO NOGUEIRA DE C. KOKUBUM3,
CELIO FERNANDO BAPTISTA HADDAD4, AND ADRIAN ANTONIO GARDA
5,6
1 Programa de Pos–graduacao em Sistematica e Evolucao, Laboratorio de Anfıbios e Repteis (LAR), Departamento deBotanica, Ecologia e Zoologia (DBEZ), Centro de Biociencias, Universidade Federal do Rio Grande do Norte, Campus
Universitario, Lagoa Nova, 59078–970, Natal, RN, Brazil2 Laboratorio de Vertebrados Terrestres, Instituto de Ciencias Biologicas, Universidade Federal do Rio Grande, Avenida
Italia, km 8, Vila Carreiros, 96.203–900, Rio Grande, RS, Brazil3 Laboratorio de Herpetologia, Unidade Academica de Ciencias Biologicas e Programa de Pos-graduacao em Ciencias
Florestais, CSTR/Universidade Federal de Campina Grande (UFCG), CEP 58708-110, Patos, PB, Brazil4 Laboratorio de Herpetologia, Departamento de Zoologia, Instituto de Biociencias, Universidade Estadual Paulista
Avenida 24 A, 1515, Bairro Bela Vista, 13506–970, Rio Claro, SP, Brazil5 LAR, DBEZ, Centro de Biociencias, Universidade Federal do Rio Grande do Norte, Campus Universitario, Lagoa
Nova, 59078–970, Natal, RN, Brazil
ABSTRACT: We describe a new species of Pseudopaludicola endemic to the Caatinga ecoregion ofnortheastern Brazil. The new species is characterized morphologically by small size, lack of T-shape terminalphalanges, smooth upper eyelids, presence of abdominal and vocal sac folds, light cream-colored vocal sac,and short hind limbs with tibia–tarsal articulation reaching the eye. The tadpole has a globular body shape,low tail fins, and a ventral oral disc bordered by a single row of marginal papillae with a wide anterior gap andtwo ventrolateral gaps; oral formula: 2(2)/2(1). The advertisement call consists of a well-defined sequence ofnotes comprising three nonconcatenated pulses, each with long interpulse intervals (111 6 21 ms) andaverage note duration of 238 6 31 ms. The mean dominant frequency is 5636 6 300 Hz, and increases fromthe first to the last pulse in each note. The advertisement call easily distinguishes the new species from allother congeners for which calls are known.
THE GENUS Pseudopaludicola comprises 18species of small frogs in South America,occurring from northern Argentina to Ven-ezuela east of the Andes (Frost, 2013). Thegenus forms the basal taxon in the leptodac-tylid subfamily Leiuperinae (Pyron andWiens, 2011). They are frequently foundcalling during the day (and occasionally atnight) around open areas in tropical forests(Amazon and Atlantic forests) and in openformations (Chaco, Pampas, Cerrado, andCaatinga). Several new Pseudopaludicola spe-cies have been described in recent years(Toledo, 2010; Toledo et al., 2010; Pansonatoet al., 2012; Roberto et al., 2013) and the callsand tadpoles of described species have beencharacterized (Pereira and Nascimento, 2004;Laufer and Barreneche, 2008; Giaretta andFacure, 2009). The number of recognizedspecies increased from 5 in the late 1980s
(Haddad and Cardoso, 1987) to 18 in 2013(Frost, 2013). Some of the recently describedspecies occur in heavily studied areas (such asMinas Gerais and Sao Paulo states; Toledo,2010; Toledo et al., 2010). The discovery ofnew species in well-studied regions suggeststhat additional undescribed species are likelyto occur in poorly known areas such as thesemiarid Brazilian Caatinga (Tabarelli andSilva, 2003; see Roberto et al., 2013).
The most recent compilation for the herpe-tofauna of the Caatinga lists 173 species (48anurans, 3 caecilians, 10 turtles, 3 crocody-lians, and 109 squamates; Rodrigues, 2003).These numbers are likely to increase substan-tially, however, as 40% of the biome has neverbeen surveyed, and about 80% remainsunderstudied until recently (Tabarelli andSilva, 2003). Indeed, several lizard (Arias etal., 2011a,b; Passos et al., 2011) and frog (e.g.,Pombal et al., 2012) species have beenrecently described from the region.6 CORRESPONDENCE: e-mail, [email protected]
77
Here we describe a new species of Pseudo-paludicola, which is widespread in the Caa-tinga ecoregion. We characterize the species,advertisement call and larval external mor-phology, and we compare it to allied species.
MATERIALS AND METHODS
We collected frogs at several localitiesthroughout northeastern Brazil (see geograph-ic distribution, Fig. 1). We euthanized spec-imens by rubbing lidocaine cream onto theirabdomens or through CO2 asphyxiation. Weindividually tagged each specimen, collectedtissue samples, fixed specimens in 10%formalin, and transferred them to 70%ethanol for permanent storage (collectingpermits to AAG SISBIO 32527–1, DL SIS-BIO 12545-2, and MNCK SISBIO 25267-1).We measured morphometric variables withthe use of a Digimess digital caliper (60.01mm) according to Pansonato et al. (2013) and
followed the terminology for external mor-phology from Duellman (1970). Specimensused in this description are housed in theCelio F.B. Haddad Collection at the Depar-tamento de Zoologia, Universidade EstadualPaulista, Campus de Rio Claro, Sao Paulo,Brazil (CFBH) and in the HerpetologicalCollection of Universidade Federal do RioGrande, Rio Grande, Rio Grande do Sul,Brazil (FURG). Tissue samples are housed inColecao de Anfıbios de Repteis da Universi-dade Federal do Rio Grande do Norte(CLAR–UFRN) and Laboratorio de Herpe-tologia da Universidade Federal de CampinaGrande (LHUFCG). Specimens examined forspecies comparisons are listed in Appendix I.
We described the advertisement call for thenew species based on recordings of nineindividuals from five localities (Table 1; alllatitude and longitude readings based onWGS84 data): Missao Velha (78140S, 39880W;
FIG. 1.—Geographic distribution of Pseudopaludicola pocoto sp. nov. in South America. The inset shows thenortheastern region of Brazil and the grey area represents the boundaries of the Caatinga biome. Symbols denote thetype locality (triangle) and additional localities where the new species was collected (circles). CEARA (CE): 1—SantaQuiteria, 2—Novas Russas, 3—Fortaleza, 4—Missao Velha, RIO GRANDE DO NORTE (RN): 5—Macaıba, 6—SerraNegra, PARAIBA (PB): 7—Patos, 8—Serra de Santa Catarina, PERNAMBUCO (PE): 9—Betania.
78 [Vol. 70, No. 1HERPETOLOGICA
363 m above sea level [asl]), Morada Nova(5860S, 388220W; 50 m asl), and Nova Russas(48420S, 408330W; 230 m asl) municipalities,Ceara State; Aguiar municipality (78000S,388130W; 380 m asl), Paraıba State andMacaıba municipality (58530S, 358220W; 11m asl), Rio Grande do Norte State. Frogswere recorded with the use of a MarantzPMD 661 coupled with a Sennheiser ME66directional microphone (Rio Grande doNorte; from 1800 to 2100 h), a Sony TCM5000 EV coupled with a Sennheiser ME66(Morada Nova, Ceara; at 2000 h), anOlympus S11 with a built-in microphone(Missao Velha, Ceara; at 1927 h), and aMicrotreck II coupled with a SennheiserME66 directional microphone (Paraıba;from 1800 to 2300 h). We analyzed adver-tisement calls (15 notes per recording) inRaven Pro v1.4 and constructed audiospectrograms with the following parameters:fast Fourier transform (FFT) window width¼512, frame¼100, overlap¼75, and flat topfilter. Variables used in comparisons arelisted in Table 1. We follow the terminologyfor frog call descriptions used by Duellmanand Trueb (1986) and McLister et al. (1995),who consider a ‘‘note’’ as a single unit ofsound, consisting of one or more pulses,produced during a single airflow cycle. Wedeposited call recording files in the followingcollections: Arquivos Sonoros da Universi-dade Federal do Rio Grande do NorteT
AB
LE
1.—
Ad
vert
isem
ent
call
par
amet
ers
ofP
seud
opal
udic
ola
poco
tosp
.n
ov.
Mea
n6
SD
(ran
ge).
Bra
zilia
nst
ate
acro
nym
s:R
N:
Rio
Gra
nd
ed
oN
orte
,C
E:
Cea
ra,
PB
:P
araı
ba.
Col
lect
ion
nu
mb
erN
ote
du
rati
on(m
s)P
uls
es/n
ote
Pu
lse
du
rati
on(m
s)In
terp
uls
ein
terv
al(m
s)D
omin
ant
freq
uen
cy(H
z)T
emp
erat
ure
(8C
)M
un
icip
alit
y/st
ate
AS
UF
RN
145
228
625
(190
–284
)3
56
1(4
–8)
106
615
(82–
136)
5608
687
(551
3–56
85)
24M
acaı
ba/
RN
AS
UF
RN
146
268
616
(226
–289
)3
46
0.6
(3–5
)12
76
20(9
4–16
6)55
816
100
(551
3–58
57)
24M
acaı
ba/
RN
AS
UF
RN
153
220
637
(126
–263
)3
46
0.6
(3–5
)10
46
26(4
9–16
0)53
406
142
(516
8–55
13)
24.3
Mac
aıb
a/R
NA
SU
FR
N22
426
66
15(2
24–2
81)
36
61
(4–8
)12
46
11(7
8–13
5)55
626
87(5
340–
5685
)–
Mis
sao
Vel
ha/
CE
AS
UF
RN
225
211
64
(204
–220
)3
56
1(4
–6)
976
6(8
1–10
9)62
786
125
(585
7–63
74)
–M
orad
aN
ova/
CE
DL
005
269
615
(237
–290
)3
56
1(3
–7)
126
616
(105
–156
)54
556
97(5
168–
5685
)–
Nov
asR
uss
as/C
EL
HU
FC
G45
320
76
14(1
76–2
27)
34
60.
5(3
–5)
926
18(4
3–12
8)58
726
155
(551
3–62
02)
26S
erra
de
San
taC
atar
ina/
PB
LH
UF
CG
452
245
626
(194
–286
)3
46
0.6
(3–5
)11
66
15(8
7–14
7)56
216
108
(534
0–58
57)
25.3
Ser
rad
eS
anta
Cat
arin
a/P
BL
HU
FC
G30
624
36
23(2
02–2
82)
2–3
36
0.5
(3–5
)11
66
14(8
5–14
6)53
766
94(5
186–
5513
)25
Ser
rad
eS
anta
Cat
arin
a/P
BM
ean
238
631
(126
–290
)3
56
1(3
–8)
111
621
(43–
166)
5636
630
0(5
168–
6373
)
TABLE 2.—Measurements (in millimeters) of Pseudopalu-dicola pocoto sp. nov. type series from municipalities ofFortaleza, Novas Russas, and Santa Quiteria, Ceara State,
(ASUFRN), Daniel Loebmann Sound Col-lection (DL), and Laboratorio de Herpeto-logia da Universidade Federal de CampinaGrande (LHUFCG).
We collected larvae in a pond at the gatesof Escola Agrıcola de Jundiaı (5853 0S,358210W; 11 m asl), Macaıba municipality,Rio Grande do Norte State, Brazil, in June2012, a week after amplectant pairs wereobserved in the same pond. No other speciesof frogs were observed reproducing at thissite and no other larvae were collectedduring this period. An extensive survey offrogs of the region has been published forthe region (Magalhaes et al., 2013), and a keyto anuran larvae of the area is beingcompleted by FMM. Although we were notable to rear larvae to metamorphosis toconfirm their identity in this study, ourknowledge of the species and their larvaein the region, and the characteristics of thesite where they were collected, enable us toattribute the larvae to the new speciesunequivocally. We anesthetized and pre-served larvae in 10% formalin and tookmorphometric measurements with the use ofMitutoyo digital calipers (60.01 mm), exceptfor eye diameter, interorbital and internarialdistance, and spiracle and cloacal tubelength, which were measured with an ocularmicrometer in a Leica-EZ4D stereomicro-scope. We described larvae with the use ofthe terminology and measurements fromAltig and McDiarmid (1999) and determineddevelopment stage according to Limbaughand Volpe (1957), as modified by Gosner(1960); variables used in this work are listedin Table 4. The specimen used for theillustration and description (AAGARDA9438) and the remaining lot with 11specimens (AAGARDA 8825) were alsodeposited in the CLAR–UFRN.
Holotype.—CFBH 26842, adult male,Santa Quiteria Municipality, Ceara State,Brazil (048190S, 408100W; 20 m asl), collect-ed on 25 March 2009, by D. Loebmann.
TA
BL
E3.
—A
dve
rtis
emen
tca
llp
aram
eter
sof
Pse
udop
alud
icol
asp
ecie
sth
atem
itca
llsw
ith
non
con
cate
nat
edp
uls
es.
Mea
n6
SD
(ran
ge).
Sp
ecie
sN
ote
du
rati
on(m
s)In
terp
uls
ein
terv
al(m
s)P
uls
es/n
ote
Dom
inan
tfr
equ
ency
(Hz)
Ref
eren
ce
P.
ameg
hini
706
10(4
0–11
0)20
66
(1–3
0)4.
36
0.3
(3–6
)39
896
247
(318
7–43
99)
Pan
son
ato
etal
.(2
013)
P.
falc
ipes
4030
242
00–5
800
Had
dad
and
Car
dos
o(1
987)
P.
min
eira
406
420
62
245
936
263
Per
eira
and
Nas
cim
ento
(200
4)P
.m
urun
du99
69
(87–
112)
36
4(0
–12)
4.93
60.
70(3
–6)
5720
632
0(5
170–
6370
)T
oled
o(2
010)
P.
poco
tosp
.n
ov.
238
631
(126
–290
)11
16
21(4
9–16
6)3
5636
630
0(5
168–
6373
)T
his
stu
dy
P.
salt
ica
606
10(3
0–10
0)20
64
(3–4
0)3.
36
0.8
(1–4
)45
476
531
(335
9–54
69)
Pan
son
ato
etal
.(2
013)
P.
salt
ica
766
19(3
0–90
)19
610
(0–3
2)2.
086
0.51
(1–3
)46
006
180
(439
0–49
10)
Tol
edo
(201
0)P
.se
rran
a85
622
(25–
115)
126
7(0
–26)
3.07
60.
80(1
–4)
5750
611
0(5
510–
6030
)T
oled
o(2
010)
80 [Vol. 70, No. 1HERPETOLOGICA
Paratypes.—CFBH 26843–26847, all adultmales, collected with the holotype: CFBH20185–20287 adult males, Novas RussasMunicipality, Ceara State, Brazil (048410S,408330W; 270 m asl), collected on 16 March2008, by D. Loebmann: FURG 2179–2184,adult males, Fortaleza Municipality, CearaState, Brazil (038460S, 388390W; 17 m asl),collected on 25 January 2009, by D. Loeb-mann and I.J. Roberto.
Diagnosis.—The new species is assigned tothe genus based on the small size andpresence of hypertrophied antebrachial tu-bercles (Lobo, 1995). It is distinguished fromall other species of the genus by the followingcombination of characters: (1) smaller size(SVL range ¼ 11.5–14.5 mm in adult males);(2) light cream-colored vocal sac with alongitudinal fold; (3) absence of T-shapedterminal phalanges (phalange shape similar toP. facilpes; see Fig. 2B in Cardozo and Suarez,2012); (4) short hind limbs (tibio-tarsalarticulation reaching eye); (5) abdominal foldscomplete; (6) advertisement call emitted as aseries of well-defined sequences of threenonconcatenated pulses per note with longintervals (111 6 21 ms; range 43–166 ms)between each pulse; (7) mean note duration238 6 31 ms (range 126–290 ms); (8) average
dominant frequency 5636 6 300 Hz (range5168–6374 Hz).
The new species is distinguished morpho-logically from all congeners belonging to theP. pusilla species group (P. boliviana, P.ceratophyes, P. llanera, and P. pusilla) by theabsence of either T-shaped terminal phalan-ges (see Cardozo and Suarez, 2012) orexpanded toe tips (disks or pads). We didnot include P. canga as a member of P. pusillagroup based a on recent reassessment of itsosteological features (Cardozo and Suarez,2012). Also, the new species is differentiatedfrom P. boliviana, P. ceratophyes, and P.llanera by the absence of palpebral tubercles(tubercles are present on the upper eyelids ofthese three species; Lynch, 1989).
Compared to other species of Pseudopalu-dicola that are not part of P. pusilla group (P.ameghini, P. canga, P. facureae, P. falcipes, P.giarettai, P. hyleaustralis, P. mineira, P.murundu, P. mystacalis, P. parnaiba, P. salt-ica, P. serrana, and P. ternetzi), the newspecies differs morphologically from P. fal-cipes by the presence of a continuousabdominal fold (abdominal fold interrupted
FIG. 2.—Pseudopaludicola pocoto sp. nov., CFBH26842 (holotype) from Santa Quiteria Municipality, CearaState. (A) Dorsal and (B) lateral views of head, (C) lefthand, and (D) left foot.
TABLE 4.—Measurements (in millimeters) of Pseudopalu-dicola pocoto sp. nov. larvae at Gosner Stages 29–31 fromEscola Agrıcola de Jundiaı, Macaıba municipality, RioGrande do Norte State. n ¼ number of measured
or absent in P. falcipes; Lobo, 1994); from P.canga, by the absence of dorsal folds and smallsize with SVL reaching 14.5 mm in males(dorsal folds are present in P. canga and maleSVL range ¼ 14.6–16.2 mm; Giaretta andKokubum, 2003); from P. murundu, P. saltica,and P. serrana by shorter hind limbs withtibio-tarsal articulation reaching the eyes(these three species have very long hind limbswith tibio-tarsal articulation reaching beyondthe end of the snout; Toledo, 2010) and by thepresence of a light cream-colored vocal sac(vocal sac is dark gray in P. serrana and P.murundu; Toledo, 2010). Compared to P.ameghini (males SVL range ¼ 14.1–19.3 mm;Pansonato et al., 2013) and P. giarettai (malesSVL range ¼ 16.2–18.0 mm; Carvalho, 2012)has a smaller size and less robust body.
Even though we found no reliable charac-ters that morphologically distinguish adults ofP. pocoto from P. facureae, P. hyleaustralis, P.mineira, P. mystacalis, P. parnaiba, and P.ternetzi, the distinctive advertisement calldistinguishes the new species from thesespecies and from all other congeners forwhich call descriptions are available. The callof P. pocoto is characterized by well-definedsequences of three nonconcatenated pulsesper note. This distinguishes the new speciesadvertisement call from P. canga (Pansonatoet al., 2012), P. facureae (Andrade andCarvalho, 2013), P. giarettai (Carvalho,2012), P. hyleaustralis (Pansonato et al.,2012), and P. parnaiba (Roberto et al.,2013), which emit calls with single pulsednotes. Additionally, the call structure (threenonconcatenated pulses/note) distinguishesthe calls of P. pocoto from P. boliviana (callswith sets of five notes each one formed by 3–6concatenated pulses; Dure et al., 2004) and P.mystacalis (12–14 concatenated pulses/note;Pansonato et al., 2013). A longer interpulseinterval and note duration distinguish theadvertisement call of P. pocoto from all othercongeners that emit calls with nonconcaten-ated pulses (Table 3). Such parameters alsodistinguish calls of P. pocoto from those of P.ternetzi (note duration and interpulse intervalare below 100 and 10 ms, respectively,Cardozo and Toledo, 2013).
The marginal papillae configuration (singlerow of marginal papillae with one anterior and
two ventrolateral gaps) distinguishes the larvaof P. pocoto from those of P. mineira, P.murundu, and P. serrana (gaps on theposterior labium absent, Pereira and Nasci-mento, 2004; Toledo, 2010; Toledo et al.,2010), P. boliviana (only one central gap onthe posterior labium; Kehr and Schaefer,2005), P. falcipes, and P. facureae (twoventrolateral and one central gap on theposterior labium; Giaretta and Facure,2009). Also, the labial tooth row formula[2(2)/2(1)] separates the larva of P. pocotofrom the tadpoles of P. facureae and P.falcipes [2(2)/3 in these species; Giaretta andFacure, 2009].
Description of the holotype.—Body ellipti-cal and robust. Head elliptical, as wide as long.Snout subelliptical in dorsal view and roundedin profile (Fig. 2A,B). Nostrils protuberant,directed dorsolaterally. Canthus rostralisrounded. Loreal region slightly concave.Choanae rounded. Eye not protuberant as aresult of fixation, and its diameter representsabout 57% of the interorbital distance; inter-orbital area flat. Tympanum not visibleexternally. Vocal sac singular, slightly expand-ed externally with longitudinal folds; vocal slitsdeveloped, laterally on mouth floor. Vomerineteeth absent. Tongue elliptical, longer thanwide. Finger length I,II,IV,III. Toe lengthI,II,V,III,IV. Finger and toe tips withoutdisks, slightly expanded and round (Fig.2C,D). Thumb with keratinized brown nuptialpad. Finger webbing absent and toe webbingreduced I–II 2þ–3 III 3þ–4 IV–V. Fringesdeveloped on all toes (mainly II, III, and IV)and less developed on fingers. External fringeon Toe V continues almost to the externalmetatarsal tubercle. Fingers and toes withconical, singular subarticular tubercles. Largesubconical ulnar tubercle. Internal metacarpaltubercle elliptical; external metacarpal tuber-cle ovoid. Hind limbs robust and long withtibia-tarsal articulation reaching eye. Thighlength similar to tibia length; foot slightlylonger than thigh. Toes long. Supernumerarytubercles absent on the sole of foot. Metatar-sal tubercles present, internal elliptical andlarger than the external; external more protu-berant and conical in shape compared tointernal. Well-developed fold from internalmetatarsal tubercle to the mid-ventral tarsus,
82 [Vol. 70, No. 1HERPETOLOGICA
ending in a tarsal tubercle less protuberantthan the external metatarsal tubercle. Abdom-inal folds are also present; belly skin smooth.Ventral surface of thigh granular. Skin texturesmooth. Measurements of the holotype arepresented in Table 2.
Coloration of the holotype.—In preserva-tive, dorsal surfaces of dorsum and limbs arepale brown, belly is while, and vocal sac is palecream-colored. White spots on upper lipsextending to the flanks. Dorsal surface of legswith brown stripes.
Color in life.—There are no data aboutcolor in life of the holotype. Therefore, colorin life and its variation are described based onparatypes and specimens from other localities(Fig. 3). As with most congeneric species,individuals of P. pocoto vary in whether theyhave a vertebral line (white, orange or red).Usually, dorsal surfaces of body and limbs arelight brown. In some individuals, well-defineddark brown stripes are present on the dorsalsurface of thighs (a few), the interorbitalregion (single), and postcranial region (singleW-shaped stripe). Scattered orange blotchesare also present on the dorsal surface of thebody and thighs. The upper jaw has whitespots, extending to the flanks; the belly andvocal sac are light cream-colored.
Variation in the type series.—The typeseries comprises only males (n ¼ 14, Table2); therefore, comparisons between males andfemales are not possible. Males have nuptialpads well developed, as well as a singlerounded vocal sac. The ratio of head tosnout–vent length ranges from 0.34 to 0.39(mean¼0.37). The tympanic membrane is notvisible (or is difficult to discern) and does notexhibit circumferential grooves. The head isslightly wider or equal to its length; the ratioof head width to head length ranges from 0.81to 1.00 (mean¼ 0.92). Hand length comprisesapproximately one-half the foot length (theratio of hand length to foot length ranges0.45–0.55; mean ¼ 0.49). The ratio of footlength to SVL ranges from 0.54 to 0.65 (mean¼ 0.59), whereas the ratio of hand length toSVL ranges from 0.26 to 0.33 (mean ¼ 0.29).The forearm length to hand length ratioranges from 0.50 to 0.63 (mean ¼ 0.56). Theratio of internarial distance to interorbitaldistance ranges from 0.33 to 0.46 (mean ¼
0.39). The ratio of eye diameter to eye–snoutdistance ranges from 0.70 to 0.90 (mean ¼0.77). The ratio of snout width to head widthranges from 0.34 to 0.52 (mean ¼ 0.44).Specimens with a vertebral line, a featureusually present in other species of the genus,were recorded within the type series of P.pocoto.
Etymology.—The specific epithet refers tothe similarity of the advertisement call to thesound of a horse trotting, which in Portugueseis expressed by the onomatopoeia pocoto.
Distribution.—Pseudopaludicola pocoto hasbeen collected in three adjacent states inBrazil at sites within the Caatinga ecoregion:(1) in Rio Grande do Norte State it was foundin Escola Agrıcola de Jundiaı (MacaıbaMunicipality) and Estacao Ecologica doSerido, Serra Negra Municipality (6839 0S,378240W; 178 m asl); (2) in Paraıba State atSerra de Santa Catarina, Aguiar Municipalityand Patos Municipality (078010S, 378160W;250 m asl); and (3) in Ceara State at FortalezaMunicipality, semipermanent ponds betweenCrateus and Nova Russas municipalities andat Santa Quiteria, Morada Nova and MissaoVelha municipalities (Fig. 1). Additionally, thenew species has been documented in ReservaParticular do Patrimonio Nacional MaurıcioDantas, Betania municipality, PernambucoState, where it was identified as Pseudopalu-dicola sp. 2 (Borges-Nojosa and Santos, 2005).
Natural history notes.—The new speciescan be found in open areas within theCaatinga ecoregion. Males can be foundcalling at dusk (1730 h) and are active mostly
FIG. 3.—Unvouchered male of Pseudopaludicola pocotosp. nov. in life from Estacao Ecologica do Serido, SerraNegra Municipality, Rio Grande do Norte State, Brazil.
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during the night (the last recording was takenat 2300 h). We observed that males preferen-tially call from the margins of shallowtemporary ponds (depth » 5 cm ) with sandyor muddy bottoms, actively call during theentire rainy season, and maintain their activityfor as long as the reproductive sites remainflooded (usually from March–September). Wealso found males of Dendropsophus branneri,Dermatonotus muelleri, Elachistocleis cesarii,Physalaemus albifrons, P. cuvieri, Pseudopa-ludicola mystacalis, Rhinella granulosa, andScinax x–signatus calling in and around thesame habitats. The larvae of P. pocoto can befound along the margins of shallow semiper-manent ponds (about 2–5 cm in depth) withsparse vegetation and they hide in the mudwhen disturbed. The larvae of the previouslymentioned species can also be found in thesame habitats as P. pocoto.
Advertisement call description.—We ana-lyzed the advertisement calls of nine speci-mens (15 notes per recording) of P. pocotofrom five localities encompassing its entireknown distribution (Table 1). All males were
recorded at night calling on the ground at themargins of temporary ponds. The advertise-ment call of P. pocoto consists of a well–defined sequence of notes, each one formedby sets of three nonconcatenated pulses (Fig.4) with mean note duration of 238 6 31 ms(range ¼ 126–290 ms) emitted at intervals of220 6 59 ms (range 136–507 ms) betweeneach note. Also, P. pocoto may occasionallyemit notes with one, two (more frequent), orfour nonconcatenated pulses. One or twopulse notes may be emitted prior to startingthe three-pulsed note sequences, but theseand the four-pulsed note are rarely emitted.Mean duration of each pulse is 5 6 1 ms(range¼3–8 ms). Each pulse is followed by aninterpulse interval of 111 6 21 ms (range ¼43–166 ms). The average rate of emission is 136 2 pulses/s (range ¼ 10–18) and 124 6 16notes/min (range ¼ 94–150). Mean dominantfrequency is 5636 6 300 Hz (range ¼ 5168–6374 Hz). The call exhibits frequency modu-lation, increasing from the first to the lastpulse in each note. A single note is emitted
FIG. 4.—Advertisement call of Pseudopaludicola pocoto sp. nov. from Escola Agrıcola de Jundiaı, MacaıbaMunicipality, Rio Grande do Norte State. (A) Amplitude of the waveform; (B) Spectrogram. Air temperature¼ 24.08C.
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during a single airflow (personal observationsof FMM and AAG).
Description of larva.—Body is dorsoventral-ly compressed with a globular appearance(Fig. 5). Snout is rounded in all views. Eyesare small and positioned dorsally. Nares areoval and located closer to eyes than to snout,without projection on marginal rim. Thespiracle is sinistral, short and narrow, openingon the middle third of the body bellow thelateral line with proximal wall fused to it. Thecloacal tube is medial, narrow, and fused tothe fin. The dorsal fin slightly arched,emerging on the last third of the body; ventralfin with margin parallel to the longitudinal tailaxis; both fins are lower than body and similarin height. The tail ends in a rounded tip. Theoral disc is ventral and laterally emarginated;single row of marginal papillae with a widegap on the anterior labium and two ventrolat-eral gaps on the posterior labium; submarginalpapillae absent. Labial tooth row formula:2(2)/2(1); A–1, A–2; P–1 with similar lengthwhereas P–2 is slightly shorter; A–2 is widelyinterrupted and P–1 is narrowly interrupted.
Jaw sheaths are narrow with triangular serra-tion; upper jaw sheath is arc-shaped and lowerjaw sheath v-shaped. In life, body and tailmusculature are dark brown, and fins aretranslucent with few scattered brown-pig-mented blotches along its extension.
Remarks.—Pseudopaludicola pocoto iswidely distributed across the Caatinga andseveral populations were found within pro-tected and disturbed areas. Therefore, weconsider the new species not endangered andrecommend it be listed as Least Concernfollowing IUCN categories. Because we onlycollected the new species at sites within theCaatinga or in ecotonal areas, it is likelyendemic to this ecoregion, and therebyprovides additional evidence that the speciesrichness of this region is likely underestimat-ed.
DISCUSSION
The conservative external morphology andsmall size of species of Pseudopaludicolahamper the assessment of species diversityof the genus. Nevertheless, advertisement call
FIG. 5.—Schematic representation of Pseudopaludicola pocoto sp. nov. tadpole from Escola Agrıcola de Jundiaı,Macaıba Municipality, Rio Grande do Norte State, Brazil. (A) Lateral, dorsal, and ventral views; (B) detail of the oraldisk.
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parameters and tadpole morphology can aid inthe comparison and delimitation of species inthe genus. Indeed, several recently describedand revalidated species are diagnosed primar-ily through advertisement calls (Carvalho,2012; Pansonato et al., 2012; Andrade andCarvalho, 2013; Pansonato et al., 2013).
Despite the importance of advertisementcalls in the delimitation of species of Pseudo-paludicola, the terminology used for callparameters in different studies is not alwayscompatible. For example, for P. saltica theterm note (Pansonato et al., 2013) was used torefer to the same structure previously namedpulse (Toledo, 2010), whereas the term call(Pansonato et al., 2013) was applied to thestructure previously named note (Toledo,2010). Likewise, the call of P. canga wasdescribed as notes composed of one–ninenonconcatenated pulses (Giaretta and Koku-bum, 2003), and later considered as a series ofsingle pulsed notes (Pansonato et al., 2012).We recommend that authors consider a noteas the sound emitted during a single airflowcycle (McLister et al., 1995). In Pseudopalu-dicola this note may be formed by singlepulses (P. canga, P. facureae, P. hyleaustralis,P. giarettai, and P. parnaiba), by concatenatedpulses (P. mystacalis and P. boliviana), or bysets of nonconcatenated pulses (species inTable 3).
Ecological and behavioral characteristicscan also help to distinguish syntopic species.For instance, we found P. pocoto in syntopywith P. mystacalis, and both species aremorphologically indistinguishable. Neverthe-less, P. mystacalis calls mostly during the day,whereas P. pocoto calls mostly during thenight, and the advertisement calls are easilydistinguished. Other researchers have alsofound different species of Pseudopaludicolacalling during different hours of the day atdifferent sites (Loebmann and Haddad, 2010;Pansonato et al., 2013). Finally, the use ofmolecular tools will be important for under-standing species diversity and evolutionaryrelationships within the genus, as relativelyfew species have been included in previousmolecular phylogenetic analyses (e.g., Pyronand Wiens, 2011).
Acknowledgments.—We thank I. Joventino and R. Avilafor data on geographic distribution and advertisement
calls, and L.F. Toledo for information on Pseudopaludi-cola calls. R.A. Pyron and D.B. Shepard kindly revised theEnglish. FMM and AAG thank the crew of Laboratorio deAnfıbios e Repteis–UFRN for help in fieldwork. FMMthanks Capes for his Masters fellowship. Parts of thisresearch were supported by two grants from CNPq toAAG (552031/2011–9 and 558317/2009-0). CFBH isgrateful for grant 2008/50928-1, Sao Paulo ResearchFoundation (FAPESP), and the research fellowship ofCNPq.
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Accepted: 26 October 2013Associate Editor: Christopher Raxworthy
