a new species of diurnal frog in the genus crossodactylus ... · and bibron, 1841 (anura,...

9
Amphibia-Reptilia 26 (2005): 497-505 A new species of Diurnal Frog in the genus Crossodactylus Duméril and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1 , Carlos A.G. Cruz 2 , Renato N. Feio 3 Abstract. A new species of Crossodactylus, belonging to the C. gaudichaudii species group, was described from the Municipality of Santa Maria do Salto, in the Atlantic Rain Forest of the State of Minas Gerais, southeastern Brazil. The new species was characterised by its: small size; slender body; head being longer than its width; snout rounded in dorsal view and protruding in lateral view; skin very granulated on sacral region; males having four to six keratinized conical spines arranged in a circle on their thumb; inner tarsal fringe extensively developed; toes extensively fringed on both edges; and ventral body surface with brownish blotches and short stripes. The descriptions of advertisement calls and tadpoles are included. Resumo. Uma nova espécie de Crossodactylus, pertencente ao grupo de C. gaudichaudii, é descrita do município de Santa Maria do Salto, na Mata Atlântica do Estado de Minas Gerais, sudeste do Brasil. A nova espécie é caracterizada pelo corpo esbelto, cabeça mais longa que larga, focinho arredondado em vista dorsal e protuso em vista lateral, pele bastante granulada na região sacral, machos possuindo de quatro a seis espinhos córneos cônicos, dispostos em círculo no polegar, fimbria tarsal interna bastante desenvolvida, artelhos bastante fimbriados em ambas as margens e superfície ventral do corpo com manchas e pequenas faixas amarronzadas. Descrições do canto de anúncio e do girino são apresentadas. Introduction The genus Crossodactylus includes ten species distributed from northeastern to southern Brazil and northern Argentina (Carcerelli and Caram- aschi, 1992; Frost, 2004). Caramaschi and Saz- ima (1985) recognized three species groups in this genus: the C. gaudichaudii group, including C. aeneus Müller, 1924, C. bokermanni Cara- maschi and Sazima, 1985, C. caramaschii Bas- tos and Pombal, 1995, C. dantei Carcerelli and Caramaschi, 1992, C. gaudichaudii Duméril and Bibron, 1841, and C. lutzorum Carcerelli and Caramaschi, 1992, characterized by pro- truding snout and distinct canthus rostralis; the C. trachystomus group, including C. dis- 1 - Departamento de Ciências Biológicas, Museu de Ciên- cias Naturais, Pontifícia Universidade Católica de Minas Gerais, Av. Dom José Gaspar 290, Coração Eucarístico, 30535-610 Belo Horizonte, Minas Gerais, Brazil e-mail: [email protected] 2 - Departamento de Vertebrados, Museu Nacional, Univer- sidade Federal do Rio de Janeiro, Quinta da Boa Vista, 20940-040 Rio de Janeiro, RJ, Brazil e-mail: [email protected] 3 - Departamento de Biologia Animal, Universidade Fed- eral de Viçosa, 36561-000 Viçosa, Minas Gerais, Brazil e-mail: [email protected] par A. Lutz, 1925, C. grandis B. Lutz, 1951, and C. trachystomus (Reinhardt and Lütken, 1862 “1861”), characterized by short, rounded snout, and less evident canthus rostralis; and the monospecific group of C. schmidti Gal- lardo, 1961, characterized by very short snout and rounded canthus rostralis. Herein, we de- scribe a new species belonging to the C. gau- dichaudii species group, from the Atlantic Rain Forest of the State of Minas Gerais, southeast- ern Brazil. Descriptions of vocalizations, tad- poles, and notes on the species natural history are given. Materials and methods The examined specimens were deposited in the following collections: MNRJ (Museu Nacional, Rio de Janeiro, Rio Janeiro, Brazil), MZUSP (Museu de Zoologia, Universi- dade de São Paulo, São Paulo, São Paulo, Brazil), MC- NAM (Museu de Ciências Naturais, Pontifícia Universidade Católica de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil), and MZUFV (Museu de Zoologia “João Moojen de Oliveira”, Universidade Federal de Viçosa, Minas Gerais, Brazil). Measurements (in millimeters) of adult specimens were taken using a digital caliper, following the method of Du- ellman (1970) and Cei (1980). Abbreviations were as fol- lows: SVL (snout-vent length), HL (head length), HW (head © Koninklijke Brill NV, Leiden, 2005. Also available online - www.brill.nl

Upload: others

Post on 21-Jun-2020

1 views

Category:

Documents


0 download

TRANSCRIPT

Page 1: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

Amphibia-Reptilia 26 (2005) 497-505

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteriland Bibron 1841 (Anura Leptodactylidae) from Southeastern Brazil

Luciana B Nascimento1 Carlos AG Cruz2 Renato N Feio3

Abstract A new species of Crossodactylus belonging to the C gaudichaudii species group was described from theMunicipality of Santa Maria do Salto in the Atlantic Rain Forest of the State of Minas Gerais southeastern Brazil Thenew species was characterised by its small size slender body head being longer than its width snout rounded in dorsalview and protruding in lateral view skin very granulated on sacral region males having four to six keratinized conicalspines arranged in a circle on their thumb inner tarsal fringe extensively developed toes extensively fringed on both edgesand ventral body surface with brownish blotches and short stripes The descriptions of advertisement calls and tadpoles areincluded

Resumo Uma nova espeacutecie de Crossodactylus pertencente ao grupo de C gaudichaudii eacute descrita do municiacutepio de SantaMaria do Salto na Mata Atlacircntica do Estado de Minas Gerais sudeste do Brasil A nova espeacutecie eacute caracterizada pelo corpoesbelto cabeccedila mais longa que larga focinho arredondado em vista dorsal e protuso em vista lateral pele bastante granuladana regiatildeo sacral machos possuindo de quatro a seis espinhos coacuterneos cocircnicos dispostos em ciacuterculo no polegar fimbria tarsalinterna bastante desenvolvida artelhos bastante fimbriados em ambas as margens e superfiacutecie ventral do corpo com manchase pequenas faixas amarronzadas Descriccedilotildees do canto de anuacutencio e do girino satildeo apresentadas

Introduction

The genus Crossodactylus includes ten speciesdistributed from northeastern to southern Braziland northern Argentina (Carcerelli and Caram-aschi 1992 Frost 2004) Caramaschi and Saz-ima (1985) recognized three species groups inthis genus the C gaudichaudii group includingC aeneus Muumlller 1924 C bokermanni Cara-maschi and Sazima 1985 C caramaschii Bas-tos and Pombal 1995 C dantei Carcerelli andCaramaschi 1992 C gaudichaudii Dumeacuteriland Bibron 1841 and C lutzorum Carcerelliand Caramaschi 1992 characterized by pro-truding snout and distinct canthus rostralisthe C trachystomus group including C dis-

1 - Departamento de Ciecircncias Bioloacutegicas Museu de Ciecircn-cias Naturais Pontifiacutecia Universidade Catoacutelica de MinasGerais Av Dom Joseacute Gaspar 290 Coraccedilatildeo Eucariacutestico30535-610 Belo Horizonte Minas Gerais Brazile-mail lunapucminasbr

2 - Departamento de Vertebrados Museu Nacional Univer-sidade Federal do Rio de Janeiro Quinta da Boa Vista20940-040 Rio de Janeiro RJ Brazile-mail cagcruzuolcombr

3 - Departamento de Biologia Animal Universidade Fed-eral de Viccedilosa 36561-000 Viccedilosa Minas Gerais Brazile-mail rfeioufvbr

par A Lutz 1925 C grandis B Lutz 1951and C trachystomus (Reinhardt and Luumltken1862 ldquo1861rdquo) characterized by short roundedsnout and less evident canthus rostralis andthe monospecific group of C schmidti Gal-lardo 1961 characterized by very short snoutand rounded canthus rostralis Herein we de-scribe a new species belonging to the C gau-dichaudii species group from the Atlantic RainForest of the State of Minas Gerais southeast-ern Brazil Descriptions of vocalizations tad-poles and notes on the species natural historyare given

Materials and methods

The examined specimens were deposited in the followingcollections MNRJ (Museu Nacional Rio de Janeiro RioJaneiro Brazil) MZUSP (Museu de Zoologia Universi-dade de Satildeo Paulo Satildeo Paulo Satildeo Paulo Brazil) MC-NAM (Museu de Ciecircncias Naturais Pontifiacutecia UniversidadeCatoacutelica de Minas Gerais Belo Horizonte Minas GeraisBrazil) and MZUFV (Museu de Zoologia ldquoJoatildeo Moojen deOliveirardquo Universidade Federal de Viccedilosa Minas GeraisBrazil)

Measurements (in millimeters) of adult specimens weretaken using a digital caliper following the method of Du-ellman (1970) and Cei (1980) Abbreviations were as fol-lows SVL (snout-vent length) HL (head length) HW (head

copy Koninklijke Brill NV Leiden 2005 Also available online - wwwbrillnl

498 LB Nascimento CAG Cruz RN Feio

width) ED (eye diameter) TD (tympanum diameter) END(eye-nostril distance) IND (internarial distance) IOD (in-terorbital distance) UEW (upper eyelid width) THL (thighlength) TL (tibia length) FL (foot length including tarsallength)

Nomenclature and measurements (in millimeters) of tad-poles follow the methods of Altig and McDiarmid (1999)except for the interorbital and internarial distances whichwere taken between inner margins of eyes and nostrils re-spectively Tadpoles were fixed and preserved in 5 for-malin Vocalizations were recorded with a Panasonic taperecorder model RQ L31 and sonograms were analyzed us-ing the software Avisoft-Sonagraph Light 1 (version 27)and Cool Edit 2000 Vocalizations were digitalized andedited at a sampling frequency of 22050 Hz 16 bit reso-lution FFT with 256 points 100 frame FlatTop Windowand 50 overlap Advertisement call terminology followsDuellman and Trueb (1986)

Crossodactylus cyclospinus sp nov(fig 1A B)

Holotype MNRJ 34501 adult male collectedat Fazenda Duas Barras (1553primeS 4028primeW al-titude 800 m approximately) Municipality ofSanta Maria do Salto State of Minas GeraisBrazil on 15th October 2003 by RN FeioLB Nascimento CAG Cruz DP CabralMG Soares and PL Ferreira

Paratypes MNRJ 34500 34502-04 adultmales MNRJ 34505 juvenile collected withthe holotype MNRJ 35404 MNRJ 35406-07MCNAM 4813 and MZUFV 5609 adult malesMNRJ 35405 adult female MNRJ 36478 ju-venile collected at the type locality between5th to 9th January 2004 by LB NascimentoDS Fernandes DP Cabral and MG Soares

Diagnosis A species belonging to the Crosso-dactylus gaudichaudii species group character-ized by small size (males 232-244 mm SVLfemale 294 mm SVL) slender body headlonger than its width snout rounded in dorsalview and protruding in lateral view skin verygranulated on sacral region males having fourto six keratinized conical spines arranged in acircle on their thumb inner tarsal fringe exten-sively developed toes extensively fringed onboth edges ventral body surface with brown-ish blotches and short stripes advertisement callwith note duration ranging from 0003 to 004 sand fundamental frequency ranging from 700 to1200 Hz

Comparison with other species The newspecies was distinguished from all other species

Figure 1 Crossodactylus cyclospinus sp nov holotype (MNRJ 34501 SVL 239 mm) (A) Dorsal and (B) ventral view

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 499

of the C gaudichaudii species group by the cir-cular arrangement of conical spines (four to six)on their thumb (no circular distribution is ob-served in C gaudichaudii C aeneus C boker-manni C caramaschii C dantei and C lut-zorum) (Caramaschi and Sazima 1985 perobs) Crossodactylus cyclospinus sp nov dif-fered from C bokermanni C dantei C lut-zorum and C gaudichaudii by having minus-cule keratinized spines on its upper lip (absentin C bokermanni C dantei C lutzorum andC gaudichaudii) (Cochran 1955 Caramaschiand Sazima 1985 Carcerelli and Caramaschi1992 pers obs) The new species was furtherdistinguished from C caramaschii and C lut-zorum by the presence of a gland posterior to thebucal comissure (absent in C caramaschii andC lutzorum) (Carcerelli and Caramaschi 1992Bastos and Pombal 1995 pers obs) and fromC dantei by the elongated form of this gland(triangular in C dantei) (Carcerelli and Caram-aschi 1992 pers obs) A light stripe extend-ing from the snout to arm insertion was presentin the new species and in C bokermanni andC caramaschii (absent on C aeneus C dan-tei C gaudichaudii and C lutzorum) (Cara-maschi and Sazima 1985 pers obs) Addi-tionally Crossodactylus cyclospinus sp novC gaudichaudii and C aeneus differ from allother species of the C gaudichaudii group bythe presence of a lateral cream stripe on the pos-terior half of their flanks (absent in C boker-manni C caramaschii C dantei and C lutzo-rum) (Caramaschi and Sazima 1985 Carcerelliand Caramaschi 1992 Bastos and Pombal1995 pers obs)

Description of holotype Body slender headlonger than its width snout rounded in dorsalview and protruding in lateral view (fig 2AB) canthus rostralis distinct straight loreal re-gion concave weakly inclined nostrils smallrounded near the tip of snout directed later-ally tympanum large rounded approximatelytwo thirds of eye diameter supratympanic folddistinct labial spines minuscule keratinizedregularly distributed on upper lip premaxillary

Figure 2 Crossodactylus cyclospinus sp nov holotype(MNRJ 34501) (A) Dorsal and (B) lateral view of head(scale = 5 mm)

and maxillary teeth present vomerine teeth ab-sent tongue large ovoid distinct gland imme-diately posterior to the buccal comissure vo-cal sacs subgular paired weakly developedarms moderately robust six conical keratinizedspines arranged in a circle on dorsal surfaceof the thumb (fig 3A) fingers slightly fringedfinger lengths II sim= IV lt I lt III fingertips rounded (fig 3B) scutes weakly developedon fingers fingers without webbing subartic-ular tubercles rounded supernumerary tuber-cles present outer metacarpal tubercle ovoidinner metacarpal tubercle elliptical legs mod-

500 LB Nascimento CAG Cruz RN Feio

erately robust toes extensively fringed on bothsides tarsal fringe well-developed continuouswith toe I fringe toe lengths I lt II lt V lt

III lt IV toe tips truncated (fig 3C) outermetatarsal tubercle small protruding roundedinner metatarsal tubercle large elliptical dor-sal skin rugose very granulated on sacral re-gion some small granules along dorsolateralfold ventral skin of thigh tibia and around cloa-cal region gular region and belly smooth

Color in preservative Dorsum brown withdark spots more concentrated at sacral regiondark brown stripe extending dorsolaterally fromsnout to inguinal region cream stripe extend-ing laterally from snout to arm insertion im-mediately below the dorsolateral stripe lateralcream stripe beginning at the posterior half offlank reaching inguinal region tympanum lightbrown dorsal surfaces of thigh tibia and feetlight brown with darker transversal bars armslight brown with darker spots dorsally iris lightcopper gular and chest regions cream withbrownish scattered blotches and short stripesventral surfaces of upper arms inner surface offorearms thigh tibia and belly cream outersurface of forearms hands and feet dark brownventrally

Measurements of holotype SVL 239 HL114 HW 78 ED 30 TD 21 END 15 IND32 IOD 26 UEW 22 THL 121 TL 125FL 171

Variation Conical spines on the thumb var-ied in their degree of keratinization and number(four to six in males absent in the female) Inthe female the tarsal fringe was less well de-veloped and the lateral cream stripe on flankwas wider Some specimens showed scatteredbrownish blotches and short stripes on the bellyVariation in measurements can be seen in Ta-ble 1

Tadpole Measurements of four tadpoles indevelopmental stage 36 (Gosner 1960) can beseen in Table 2 Their body was elliptical indorsal view and ovoid in lateral view (fig 4A)

Table 1 Mean (x) standard deviation (SD) and rangeof measurements (mm) of nine adult males and one adultfemale of Crossodactylus cyclospinus sp nov

Male Female

X SD Range

Snout vent length 237 04 232-244 294Head length 98 09 91-114 100Head width 79 03 76-83 88Eye diameter 32 03 29-36 37Tympanum diameter 23 03 19-29 24Eye nostril distance 17 03 12-21 23Internarial distance 34 03 30-40 41Interorbital distance 28 03 25-33 36Upper eyelid 23 02 20-26 21Thigh length 118 06 107-127 122Tibial length 128 03 124-132 133Foot length 150 27 125-182 147

Table 2 Mean (x) standard deviation (SD) and rangeof measurements (mm) of four tadpoles of Crossodactyluscyclospinus sp nov in developmental stage 36 (Gosner1960)

X SD Range

Total length 443 12 427-455Body length 156 02 153-158Body width 104 04 100-110Body height 79 07 75-90Tail length 292 11 276-300Tail height 96 05 88-99Internarial distance 34 01 33-36Interorbital distance 30 02 29-33Eye diameter 19 01 19-20Eye-nostril distance 12 01 11-13Oral disc width 37 01 36-38

snout rounded in dorsal and lateral views bodywider than its height eyes directed laterallynostrils at the same distance between the snoutand the eyes located dorsally bean-shapedpores of lateral line system distributed on dor-sal and lateral surfaces of body spiracle shortsinistral free distally its opening slightly ante-rior to mid-body and postero-dorsally orientedoral disc ventral lips developed laterally emar-ginated bordered by one row of small papil-lae with a wide gap on anterior lip scatteredsubmarginal papillae distributed laterally toothrow formula 2(2)3(1) third posterior tooth rowshorter than others jaw sheath strongly devel-oped serrate upper jaw sheath wide-arched in-

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 501

Figure 3 (A) Dorsal view of finger I showing the keratinized spines (scale = 1 mm) ventral view of (B) hand and (C) foot(scale = 5 mm)

cised and lower jaw sheath V-shaped (fig 4B)cloacal tube dextral medium-sized with wideopening tail approximately twice body lengthslightly higher than body dorsal fin slightlyhigher than ventral fin and extending onto theposterior third of body dorsal and ventral fins

slightly arched tail tip nearly rectilineal tailmusculature higher than dorsal fin extending tothe tail tip In preservative dorsum was lightbrown with small scattered brown blotchesventral surface with fine brown reticulations andgrayish reflex tail with regular brown blotches

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 2: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

498 LB Nascimento CAG Cruz RN Feio

width) ED (eye diameter) TD (tympanum diameter) END(eye-nostril distance) IND (internarial distance) IOD (in-terorbital distance) UEW (upper eyelid width) THL (thighlength) TL (tibia length) FL (foot length including tarsallength)

Nomenclature and measurements (in millimeters) of tad-poles follow the methods of Altig and McDiarmid (1999)except for the interorbital and internarial distances whichwere taken between inner margins of eyes and nostrils re-spectively Tadpoles were fixed and preserved in 5 for-malin Vocalizations were recorded with a Panasonic taperecorder model RQ L31 and sonograms were analyzed us-ing the software Avisoft-Sonagraph Light 1 (version 27)and Cool Edit 2000 Vocalizations were digitalized andedited at a sampling frequency of 22050 Hz 16 bit reso-lution FFT with 256 points 100 frame FlatTop Windowand 50 overlap Advertisement call terminology followsDuellman and Trueb (1986)

Crossodactylus cyclospinus sp nov(fig 1A B)

Holotype MNRJ 34501 adult male collectedat Fazenda Duas Barras (1553primeS 4028primeW al-titude 800 m approximately) Municipality ofSanta Maria do Salto State of Minas GeraisBrazil on 15th October 2003 by RN FeioLB Nascimento CAG Cruz DP CabralMG Soares and PL Ferreira

Paratypes MNRJ 34500 34502-04 adultmales MNRJ 34505 juvenile collected withthe holotype MNRJ 35404 MNRJ 35406-07MCNAM 4813 and MZUFV 5609 adult malesMNRJ 35405 adult female MNRJ 36478 ju-venile collected at the type locality between5th to 9th January 2004 by LB NascimentoDS Fernandes DP Cabral and MG Soares

Diagnosis A species belonging to the Crosso-dactylus gaudichaudii species group character-ized by small size (males 232-244 mm SVLfemale 294 mm SVL) slender body headlonger than its width snout rounded in dorsalview and protruding in lateral view skin verygranulated on sacral region males having fourto six keratinized conical spines arranged in acircle on their thumb inner tarsal fringe exten-sively developed toes extensively fringed onboth edges ventral body surface with brown-ish blotches and short stripes advertisement callwith note duration ranging from 0003 to 004 sand fundamental frequency ranging from 700 to1200 Hz

Comparison with other species The newspecies was distinguished from all other species

Figure 1 Crossodactylus cyclospinus sp nov holotype (MNRJ 34501 SVL 239 mm) (A) Dorsal and (B) ventral view

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 499

of the C gaudichaudii species group by the cir-cular arrangement of conical spines (four to six)on their thumb (no circular distribution is ob-served in C gaudichaudii C aeneus C boker-manni C caramaschii C dantei and C lut-zorum) (Caramaschi and Sazima 1985 perobs) Crossodactylus cyclospinus sp nov dif-fered from C bokermanni C dantei C lut-zorum and C gaudichaudii by having minus-cule keratinized spines on its upper lip (absentin C bokermanni C dantei C lutzorum andC gaudichaudii) (Cochran 1955 Caramaschiand Sazima 1985 Carcerelli and Caramaschi1992 pers obs) The new species was furtherdistinguished from C caramaschii and C lut-zorum by the presence of a gland posterior to thebucal comissure (absent in C caramaschii andC lutzorum) (Carcerelli and Caramaschi 1992Bastos and Pombal 1995 pers obs) and fromC dantei by the elongated form of this gland(triangular in C dantei) (Carcerelli and Caram-aschi 1992 pers obs) A light stripe extend-ing from the snout to arm insertion was presentin the new species and in C bokermanni andC caramaschii (absent on C aeneus C dan-tei C gaudichaudii and C lutzorum) (Cara-maschi and Sazima 1985 pers obs) Addi-tionally Crossodactylus cyclospinus sp novC gaudichaudii and C aeneus differ from allother species of the C gaudichaudii group bythe presence of a lateral cream stripe on the pos-terior half of their flanks (absent in C boker-manni C caramaschii C dantei and C lutzo-rum) (Caramaschi and Sazima 1985 Carcerelliand Caramaschi 1992 Bastos and Pombal1995 pers obs)

Description of holotype Body slender headlonger than its width snout rounded in dorsalview and protruding in lateral view (fig 2AB) canthus rostralis distinct straight loreal re-gion concave weakly inclined nostrils smallrounded near the tip of snout directed later-ally tympanum large rounded approximatelytwo thirds of eye diameter supratympanic folddistinct labial spines minuscule keratinizedregularly distributed on upper lip premaxillary

Figure 2 Crossodactylus cyclospinus sp nov holotype(MNRJ 34501) (A) Dorsal and (B) lateral view of head(scale = 5 mm)

and maxillary teeth present vomerine teeth ab-sent tongue large ovoid distinct gland imme-diately posterior to the buccal comissure vo-cal sacs subgular paired weakly developedarms moderately robust six conical keratinizedspines arranged in a circle on dorsal surfaceof the thumb (fig 3A) fingers slightly fringedfinger lengths II sim= IV lt I lt III fingertips rounded (fig 3B) scutes weakly developedon fingers fingers without webbing subartic-ular tubercles rounded supernumerary tuber-cles present outer metacarpal tubercle ovoidinner metacarpal tubercle elliptical legs mod-

500 LB Nascimento CAG Cruz RN Feio

erately robust toes extensively fringed on bothsides tarsal fringe well-developed continuouswith toe I fringe toe lengths I lt II lt V lt

III lt IV toe tips truncated (fig 3C) outermetatarsal tubercle small protruding roundedinner metatarsal tubercle large elliptical dor-sal skin rugose very granulated on sacral re-gion some small granules along dorsolateralfold ventral skin of thigh tibia and around cloa-cal region gular region and belly smooth

Color in preservative Dorsum brown withdark spots more concentrated at sacral regiondark brown stripe extending dorsolaterally fromsnout to inguinal region cream stripe extend-ing laterally from snout to arm insertion im-mediately below the dorsolateral stripe lateralcream stripe beginning at the posterior half offlank reaching inguinal region tympanum lightbrown dorsal surfaces of thigh tibia and feetlight brown with darker transversal bars armslight brown with darker spots dorsally iris lightcopper gular and chest regions cream withbrownish scattered blotches and short stripesventral surfaces of upper arms inner surface offorearms thigh tibia and belly cream outersurface of forearms hands and feet dark brownventrally

Measurements of holotype SVL 239 HL114 HW 78 ED 30 TD 21 END 15 IND32 IOD 26 UEW 22 THL 121 TL 125FL 171

Variation Conical spines on the thumb var-ied in their degree of keratinization and number(four to six in males absent in the female) Inthe female the tarsal fringe was less well de-veloped and the lateral cream stripe on flankwas wider Some specimens showed scatteredbrownish blotches and short stripes on the bellyVariation in measurements can be seen in Ta-ble 1

Tadpole Measurements of four tadpoles indevelopmental stage 36 (Gosner 1960) can beseen in Table 2 Their body was elliptical indorsal view and ovoid in lateral view (fig 4A)

Table 1 Mean (x) standard deviation (SD) and rangeof measurements (mm) of nine adult males and one adultfemale of Crossodactylus cyclospinus sp nov

Male Female

X SD Range

Snout vent length 237 04 232-244 294Head length 98 09 91-114 100Head width 79 03 76-83 88Eye diameter 32 03 29-36 37Tympanum diameter 23 03 19-29 24Eye nostril distance 17 03 12-21 23Internarial distance 34 03 30-40 41Interorbital distance 28 03 25-33 36Upper eyelid 23 02 20-26 21Thigh length 118 06 107-127 122Tibial length 128 03 124-132 133Foot length 150 27 125-182 147

Table 2 Mean (x) standard deviation (SD) and rangeof measurements (mm) of four tadpoles of Crossodactyluscyclospinus sp nov in developmental stage 36 (Gosner1960)

X SD Range

Total length 443 12 427-455Body length 156 02 153-158Body width 104 04 100-110Body height 79 07 75-90Tail length 292 11 276-300Tail height 96 05 88-99Internarial distance 34 01 33-36Interorbital distance 30 02 29-33Eye diameter 19 01 19-20Eye-nostril distance 12 01 11-13Oral disc width 37 01 36-38

snout rounded in dorsal and lateral views bodywider than its height eyes directed laterallynostrils at the same distance between the snoutand the eyes located dorsally bean-shapedpores of lateral line system distributed on dor-sal and lateral surfaces of body spiracle shortsinistral free distally its opening slightly ante-rior to mid-body and postero-dorsally orientedoral disc ventral lips developed laterally emar-ginated bordered by one row of small papil-lae with a wide gap on anterior lip scatteredsubmarginal papillae distributed laterally toothrow formula 2(2)3(1) third posterior tooth rowshorter than others jaw sheath strongly devel-oped serrate upper jaw sheath wide-arched in-

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 501

Figure 3 (A) Dorsal view of finger I showing the keratinized spines (scale = 1 mm) ventral view of (B) hand and (C) foot(scale = 5 mm)

cised and lower jaw sheath V-shaped (fig 4B)cloacal tube dextral medium-sized with wideopening tail approximately twice body lengthslightly higher than body dorsal fin slightlyhigher than ventral fin and extending onto theposterior third of body dorsal and ventral fins

slightly arched tail tip nearly rectilineal tailmusculature higher than dorsal fin extending tothe tail tip In preservative dorsum was lightbrown with small scattered brown blotchesventral surface with fine brown reticulations andgrayish reflex tail with regular brown blotches

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 3: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 499

of the C gaudichaudii species group by the cir-cular arrangement of conical spines (four to six)on their thumb (no circular distribution is ob-served in C gaudichaudii C aeneus C boker-manni C caramaschii C dantei and C lut-zorum) (Caramaschi and Sazima 1985 perobs) Crossodactylus cyclospinus sp nov dif-fered from C bokermanni C dantei C lut-zorum and C gaudichaudii by having minus-cule keratinized spines on its upper lip (absentin C bokermanni C dantei C lutzorum andC gaudichaudii) (Cochran 1955 Caramaschiand Sazima 1985 Carcerelli and Caramaschi1992 pers obs) The new species was furtherdistinguished from C caramaschii and C lut-zorum by the presence of a gland posterior to thebucal comissure (absent in C caramaschii andC lutzorum) (Carcerelli and Caramaschi 1992Bastos and Pombal 1995 pers obs) and fromC dantei by the elongated form of this gland(triangular in C dantei) (Carcerelli and Caram-aschi 1992 pers obs) A light stripe extend-ing from the snout to arm insertion was presentin the new species and in C bokermanni andC caramaschii (absent on C aeneus C dan-tei C gaudichaudii and C lutzorum) (Cara-maschi and Sazima 1985 pers obs) Addi-tionally Crossodactylus cyclospinus sp novC gaudichaudii and C aeneus differ from allother species of the C gaudichaudii group bythe presence of a lateral cream stripe on the pos-terior half of their flanks (absent in C boker-manni C caramaschii C dantei and C lutzo-rum) (Caramaschi and Sazima 1985 Carcerelliand Caramaschi 1992 Bastos and Pombal1995 pers obs)

Description of holotype Body slender headlonger than its width snout rounded in dorsalview and protruding in lateral view (fig 2AB) canthus rostralis distinct straight loreal re-gion concave weakly inclined nostrils smallrounded near the tip of snout directed later-ally tympanum large rounded approximatelytwo thirds of eye diameter supratympanic folddistinct labial spines minuscule keratinizedregularly distributed on upper lip premaxillary

Figure 2 Crossodactylus cyclospinus sp nov holotype(MNRJ 34501) (A) Dorsal and (B) lateral view of head(scale = 5 mm)

and maxillary teeth present vomerine teeth ab-sent tongue large ovoid distinct gland imme-diately posterior to the buccal comissure vo-cal sacs subgular paired weakly developedarms moderately robust six conical keratinizedspines arranged in a circle on dorsal surfaceof the thumb (fig 3A) fingers slightly fringedfinger lengths II sim= IV lt I lt III fingertips rounded (fig 3B) scutes weakly developedon fingers fingers without webbing subartic-ular tubercles rounded supernumerary tuber-cles present outer metacarpal tubercle ovoidinner metacarpal tubercle elliptical legs mod-

500 LB Nascimento CAG Cruz RN Feio

erately robust toes extensively fringed on bothsides tarsal fringe well-developed continuouswith toe I fringe toe lengths I lt II lt V lt

III lt IV toe tips truncated (fig 3C) outermetatarsal tubercle small protruding roundedinner metatarsal tubercle large elliptical dor-sal skin rugose very granulated on sacral re-gion some small granules along dorsolateralfold ventral skin of thigh tibia and around cloa-cal region gular region and belly smooth

Color in preservative Dorsum brown withdark spots more concentrated at sacral regiondark brown stripe extending dorsolaterally fromsnout to inguinal region cream stripe extend-ing laterally from snout to arm insertion im-mediately below the dorsolateral stripe lateralcream stripe beginning at the posterior half offlank reaching inguinal region tympanum lightbrown dorsal surfaces of thigh tibia and feetlight brown with darker transversal bars armslight brown with darker spots dorsally iris lightcopper gular and chest regions cream withbrownish scattered blotches and short stripesventral surfaces of upper arms inner surface offorearms thigh tibia and belly cream outersurface of forearms hands and feet dark brownventrally

Measurements of holotype SVL 239 HL114 HW 78 ED 30 TD 21 END 15 IND32 IOD 26 UEW 22 THL 121 TL 125FL 171

Variation Conical spines on the thumb var-ied in their degree of keratinization and number(four to six in males absent in the female) Inthe female the tarsal fringe was less well de-veloped and the lateral cream stripe on flankwas wider Some specimens showed scatteredbrownish blotches and short stripes on the bellyVariation in measurements can be seen in Ta-ble 1

Tadpole Measurements of four tadpoles indevelopmental stage 36 (Gosner 1960) can beseen in Table 2 Their body was elliptical indorsal view and ovoid in lateral view (fig 4A)

Table 1 Mean (x) standard deviation (SD) and rangeof measurements (mm) of nine adult males and one adultfemale of Crossodactylus cyclospinus sp nov

Male Female

X SD Range

Snout vent length 237 04 232-244 294Head length 98 09 91-114 100Head width 79 03 76-83 88Eye diameter 32 03 29-36 37Tympanum diameter 23 03 19-29 24Eye nostril distance 17 03 12-21 23Internarial distance 34 03 30-40 41Interorbital distance 28 03 25-33 36Upper eyelid 23 02 20-26 21Thigh length 118 06 107-127 122Tibial length 128 03 124-132 133Foot length 150 27 125-182 147

Table 2 Mean (x) standard deviation (SD) and rangeof measurements (mm) of four tadpoles of Crossodactyluscyclospinus sp nov in developmental stage 36 (Gosner1960)

X SD Range

Total length 443 12 427-455Body length 156 02 153-158Body width 104 04 100-110Body height 79 07 75-90Tail length 292 11 276-300Tail height 96 05 88-99Internarial distance 34 01 33-36Interorbital distance 30 02 29-33Eye diameter 19 01 19-20Eye-nostril distance 12 01 11-13Oral disc width 37 01 36-38

snout rounded in dorsal and lateral views bodywider than its height eyes directed laterallynostrils at the same distance between the snoutand the eyes located dorsally bean-shapedpores of lateral line system distributed on dor-sal and lateral surfaces of body spiracle shortsinistral free distally its opening slightly ante-rior to mid-body and postero-dorsally orientedoral disc ventral lips developed laterally emar-ginated bordered by one row of small papil-lae with a wide gap on anterior lip scatteredsubmarginal papillae distributed laterally toothrow formula 2(2)3(1) third posterior tooth rowshorter than others jaw sheath strongly devel-oped serrate upper jaw sheath wide-arched in-

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 501

Figure 3 (A) Dorsal view of finger I showing the keratinized spines (scale = 1 mm) ventral view of (B) hand and (C) foot(scale = 5 mm)

cised and lower jaw sheath V-shaped (fig 4B)cloacal tube dextral medium-sized with wideopening tail approximately twice body lengthslightly higher than body dorsal fin slightlyhigher than ventral fin and extending onto theposterior third of body dorsal and ventral fins

slightly arched tail tip nearly rectilineal tailmusculature higher than dorsal fin extending tothe tail tip In preservative dorsum was lightbrown with small scattered brown blotchesventral surface with fine brown reticulations andgrayish reflex tail with regular brown blotches

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 4: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

500 LB Nascimento CAG Cruz RN Feio

erately robust toes extensively fringed on bothsides tarsal fringe well-developed continuouswith toe I fringe toe lengths I lt II lt V lt

III lt IV toe tips truncated (fig 3C) outermetatarsal tubercle small protruding roundedinner metatarsal tubercle large elliptical dor-sal skin rugose very granulated on sacral re-gion some small granules along dorsolateralfold ventral skin of thigh tibia and around cloa-cal region gular region and belly smooth

Color in preservative Dorsum brown withdark spots more concentrated at sacral regiondark brown stripe extending dorsolaterally fromsnout to inguinal region cream stripe extend-ing laterally from snout to arm insertion im-mediately below the dorsolateral stripe lateralcream stripe beginning at the posterior half offlank reaching inguinal region tympanum lightbrown dorsal surfaces of thigh tibia and feetlight brown with darker transversal bars armslight brown with darker spots dorsally iris lightcopper gular and chest regions cream withbrownish scattered blotches and short stripesventral surfaces of upper arms inner surface offorearms thigh tibia and belly cream outersurface of forearms hands and feet dark brownventrally

Measurements of holotype SVL 239 HL114 HW 78 ED 30 TD 21 END 15 IND32 IOD 26 UEW 22 THL 121 TL 125FL 171

Variation Conical spines on the thumb var-ied in their degree of keratinization and number(four to six in males absent in the female) Inthe female the tarsal fringe was less well de-veloped and the lateral cream stripe on flankwas wider Some specimens showed scatteredbrownish blotches and short stripes on the bellyVariation in measurements can be seen in Ta-ble 1

Tadpole Measurements of four tadpoles indevelopmental stage 36 (Gosner 1960) can beseen in Table 2 Their body was elliptical indorsal view and ovoid in lateral view (fig 4A)

Table 1 Mean (x) standard deviation (SD) and rangeof measurements (mm) of nine adult males and one adultfemale of Crossodactylus cyclospinus sp nov

Male Female

X SD Range

Snout vent length 237 04 232-244 294Head length 98 09 91-114 100Head width 79 03 76-83 88Eye diameter 32 03 29-36 37Tympanum diameter 23 03 19-29 24Eye nostril distance 17 03 12-21 23Internarial distance 34 03 30-40 41Interorbital distance 28 03 25-33 36Upper eyelid 23 02 20-26 21Thigh length 118 06 107-127 122Tibial length 128 03 124-132 133Foot length 150 27 125-182 147

Table 2 Mean (x) standard deviation (SD) and rangeof measurements (mm) of four tadpoles of Crossodactyluscyclospinus sp nov in developmental stage 36 (Gosner1960)

X SD Range

Total length 443 12 427-455Body length 156 02 153-158Body width 104 04 100-110Body height 79 07 75-90Tail length 292 11 276-300Tail height 96 05 88-99Internarial distance 34 01 33-36Interorbital distance 30 02 29-33Eye diameter 19 01 19-20Eye-nostril distance 12 01 11-13Oral disc width 37 01 36-38

snout rounded in dorsal and lateral views bodywider than its height eyes directed laterallynostrils at the same distance between the snoutand the eyes located dorsally bean-shapedpores of lateral line system distributed on dor-sal and lateral surfaces of body spiracle shortsinistral free distally its opening slightly ante-rior to mid-body and postero-dorsally orientedoral disc ventral lips developed laterally emar-ginated bordered by one row of small papil-lae with a wide gap on anterior lip scatteredsubmarginal papillae distributed laterally toothrow formula 2(2)3(1) third posterior tooth rowshorter than others jaw sheath strongly devel-oped serrate upper jaw sheath wide-arched in-

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 501

Figure 3 (A) Dorsal view of finger I showing the keratinized spines (scale = 1 mm) ventral view of (B) hand and (C) foot(scale = 5 mm)

cised and lower jaw sheath V-shaped (fig 4B)cloacal tube dextral medium-sized with wideopening tail approximately twice body lengthslightly higher than body dorsal fin slightlyhigher than ventral fin and extending onto theposterior third of body dorsal and ventral fins

slightly arched tail tip nearly rectilineal tailmusculature higher than dorsal fin extending tothe tail tip In preservative dorsum was lightbrown with small scattered brown blotchesventral surface with fine brown reticulations andgrayish reflex tail with regular brown blotches

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 5: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 501

Figure 3 (A) Dorsal view of finger I showing the keratinized spines (scale = 1 mm) ventral view of (B) hand and (C) foot(scale = 5 mm)

cised and lower jaw sheath V-shaped (fig 4B)cloacal tube dextral medium-sized with wideopening tail approximately twice body lengthslightly higher than body dorsal fin slightlyhigher than ventral fin and extending onto theposterior third of body dorsal and ventral fins

slightly arched tail tip nearly rectilineal tailmusculature higher than dorsal fin extending tothe tail tip In preservative dorsum was lightbrown with small scattered brown blotchesventral surface with fine brown reticulations andgrayish reflex tail with regular brown blotches

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 6: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

502 LB Nascimento CAG Cruz RN Feio

Figure 4 Tadpole of Crossodactylus cyclospinus sp nov (MNRJ 36479) stage 36 (Gosner 1960) (A) Lateral view (scale= 5 mm) and (B) oral disc (scale = 1 mm)

on dorsal fin and caudal musculature ventralfin poorly brown pigmented Three newly meta-morphosed individuals measured SVL 147-157 (x = 51) The tadpole of C cyclospi-nus sp nov was distinguished from all otherknown tadpoles of Crossodactylus by having anincised upper jaw sheath (concave in C boker-manni C gaudichaudii) (Caramaschi and Saz-ima 1985 Francioni and Carcerelli 1993)

Vocalization On 7th January 2004 the ad-vertisement call of one male of Crossodacty-lus cyclospinus sp nov was recorded this wasa long call (x = 433 s SD = 074 range357-625 s n = 25 calls) with many harmonicnotes (x = 633 notes SD = 119 range35-98 n = 25 calls) intercall interval varied(x = 1436 s SD = 1006 range 329-3380 sn = 16 intercall intervals) mean note dura-tion was 0028 s (SD = 0004 range 0003-004 s n = 1601 notes) mean note interval

was 004 s (SD = 0003 range 0029-0065 sn = 1576 note intervals) The dominant fre-quency was generally situated in the second andthird harmonic (x = 4981 Hz SD = 6266range 3488-5447 n = 25 calls) and funda-mental frequency ranged from 700 to 1200 Hz(fig 5A B) Two other call types were emittedwhen the male noticed the presence of the ob-server (herein called A and B) (fig 5C) Oneor two notes constituted call A when consistingof one note call duration ranged from 001 to0012 s (x = 0011 s SD = 0001 n = 4 calls)with two notes call duration ranged from 0031to 0089 s (x = 058 s SD = 0015 n = 56calls) first note duration ranged from 0004 to0062 s (x = 003 s SD = 0015 n = 56 calls)the interval between notes ranged from 001 to002 s (x = 0015 s SD = 0002 n = 56 calls)and the second note duration ranged from 0008to 0019 s (x = 0013 s SD = 0003 n =56 calls) In both cases dominant frequency

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 7: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 503

Figure 5 (A) Sonogram of part of the advertisement call (B) waveform of the advertisement call (C) waveform of theadvertisement call preceded by calls A (solid arrow) and B (dashed arrow) and (D) waveform of calls A (solid arrow) and B(dashed arrow) of Crossodactylus cyclospinus sp nov recorded at Fazenda Duas Barras Municipality of Santa Maria doSalto Minas Gerais Brazil on 07 January 2004 0130 PM air temperature 30C Voucher specimen MNRJ 35406

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 8: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

504 LB Nascimento CAG Cruz RN Feio

ranged from 4112 to 5297 Hz (x = 464083 HzSD = 31305 n = 61 calls) Call B con-sisted of two notes differing from A by a longerfirst note (fig 5D) Call B duration ranged from0397 to 049 s (x = 0432 s SD = 0051n = 3 calls) First note duration ranged from0363 to 0459 s (x = 0401 s SD = 0051n = 3 calls) the interval between notes rangedfrom 0018 to 0023 s (x = 0021 s SD =0003 n = 3 calls) and the second note dura-tion ranged from 0008 to 0011 s (x = 001 sSD = 0002 n = 3 calls) The dominant fre-quency of call B ranged from 4328 to 5103 Hz(x = 4823 Hz SD = 4299 n = 3 calls) Ad-vertisement calls of the C gaudichaudii speciesgroup are known for C gaudichaudii (Weygodtand Carvalho-e-Silva 1992) and C caramaschii(Bastos and Pombal 1995) (table 3) Weygodtand Carvalho-e-Silva (1992) presented four dif-ferent calls for C gaudichaudii recorded in ter-raria The first one was considered an aggres-sive encounter call and the other three adver-tisement calls The call named ldquolong range ad-vertisement callrdquo by Weygodt and Carvalho-e-Silva (1992) was the only one used herein forcomparisons since it was the most similar tothe advertisement call of C cyclospinus sp novBoth calls were similar in the number of notesper call (35-98 notes in C cyclospinus sp nov25-130 notes in the call of C gaudichaudii)

Table 3 Comparision between acoustic parameters of theadvertisement calls of species of the Crossodactylus gau-dichaudii group Data from Weygoldt and Carvalho-Silva(1992) and Bastos and Pombal (1995)

C cyclospinus C cara- C gaudi-sp nov maschi chaudii

Call duration (s) 357-625 471-609 2-13Intercall interval (s) 329-338 ndash ndashNotes per call 35-98 49-59 25-130Notes duration (s) 0003-004 ndash 004-005Intervals between 0029-0065 ndash 004-005

notes (s)Intercall interval (s) 329-338 ndash ndashDominant 3488-5447 5000 ndash

frequency (Hz)Fundamental 700-1200 1600 ndash

frequency (Hz)

and intervals between notes (0029-0065 s inC cyclospinus sp nov 004-005 s in C gau-dichaudii) However they differed in note du-ration (0003-004 s in C cyclospinus sp nov004-005 s in C gaudichaudii) and fundamen-tal frequency (700-1200 Hz in C cyclospinussp nov 2000-5500 kHz in C gaudichaudii)Advertisement call of C cyclospinus sp novwas similar to C caramaschii but differed infundamental frequency [700-1200 Hz in C cy-clospinus sp nov 1600 Hz in C caramaschii(Bastos and Pombal 1995)] The dominant fre-quency on the third harmonic in C cyclospi-nus sp nov and C caramaschi support the ideaof a relationship between the species of genusCrossodactylus and Hylodes as suggested byBastos and Pombal (1995)

Natural History Adults juveniles and tad-poles of Crossodactylus cyclospinus sp novwere diurnal and were collected during the wetseason in October 2003 and January 2004 Aswith other species in the C gaudichaudii groupexcept C bokermanni (Caramaschi and Saz-ima 1985) the new species occurred in shallowrivulets in forested areas in Atlantic Rain For-est Males were vocalizing from rocks or underplants in the middle of these rivulets The frogswere vigilant and when a male noticed the pres-ence of an observer he turned to this directionand emitted an A or B call When threatenedindividuals hid in the litter or entered the wa-ter The female was collected at the same site asthe males creamy white ovules were observedthrough transparent skin on her belly

Distribution The new species is only knownfrom the type locality at Fazenda Duas BarrasMunicipality of Santa Maria do Salto State ofMinas Gerais southeastern Brazil This localityis in the mountain range called Serra do Caririwhich separates the States of Minas Gerais andBahia and the Basin of Jequitinhonha and Bu-ranheacutem rivers

Etymology The specific name refers to thecircular distribution of keratinized spines onthe thumb Cyclo is a greek vernacular name

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005

Page 9: A new species of Diurnal Frog in the genus Crossodactylus ... · and Bibron, 1841 (Anura, Leptodactylidae) from Southeastern Brazil Luciana B. Nascimento 1,CarlosA.G.Cruz2, Renato

A new species of Diurnal Frog in the genus Crossodactylus Dumeacuteril and Bibron 505

meaning ldquocircle ringrdquo and -spinus is a Latinvernacular name meaning ldquospinerdquo

Additional specimens examined C aeneus MNRJ 33195-98 Teresoacutepolis Rio de Janeiro Brazil C caramaschiiMNRJ 31421 MNRJ 18675 Eldorado Satildeo Paulo BrazilC dantei MNRJ 4769 (holotype) MNRJ 4770-4811(paratypes) Murici Alagoas Brazil C gaudichaudiiMNRJ 27586-88 27610-11 27626-27 27706-08 MC-NAM 5109-5120 Rio de Janeiro Rio de Janeiro BrazilC lutzorum MNRJ 4753 (holotype) MNRJ 4754-4768(paratypes) Valenccedila Bahia Brazil

Acknowledgements We thank JP Pombal Jr and BVSPimenta for comments and suggestions on the manuscriptR Young for the English revision DP Cabral MG SoaresPL Ferreira and DS Fernandes for field and laboratoryassistance L Aleixo for logistical support PR Nasci-mento for the drawings Conselho Nacional de Desenvolvi-mento Cientiacutefico e Tecnoloacutegico (CNPq) for financial sup-port and CAGC fellowship This work is part of the projectldquoInventaacuterio de aacutereas prioritaacuterias para conservaccedilatildeo da bio-diversidade nos vales dos rios Jequitinhonha e Mucurirdquosupported by Project on the Conservation and SustainableUse of Brazilian Biological DiversityBrazilian Ministryof the Environment (PROBIOMMA) and coordinated byConservation International do Brasil (CI-Brasil) InstitutoBrasileiro do Meio Ambiente e Recursos Naturais Ren-ovaacuteveis (IBAMA) provided licenses to collect the speci-mens (1992002Fauna MINAS GERAIS 0201501883602and 23103 NUFASMINAS GERAIS 0201501883602-16)

References

Altig R McDiarmid RW (1999) Body plan Develop-ment and morphology In Tadpoles The Biology ofAnuran Larvae McDiarmid RW R Altig Eds p 24-51 Chicago The University of Chicago Press

Bastos R Pombal Jr JP (1995) New species of Crosso-dactylus (Anura Leptodactylidae) from the AtlanticRain Forest of southeastern Brazil Copeia 1995 436-439

Caramaschi U Sazima I (1985) Uma nova espeacutecie deCrossodactylus da Serra do Cipoacute Minas Gerais Brasil(Amphibia Leptodactylidae) Revista Brasileira de Zo-ologia 3 43-49

Carcerelli LC Caramaschi U (1992) Ocorrecircncia dogecircnero Crossodactylus Dumeacuteril and Bibron 1841 nonordeste brasileiro com descriccedilatildeo de duas espeacutecies novas(Amphibia Anura Leptodactylidae) Revista Brasileirade Zoologia 52 415-422

Cei JM (1980) Amphibians of Argentina Monitore Zoo-logico Italiano (NS) Monog 2 xii+609

Cochran DM (1955) Frogs of Southeastern Brazil UnitedStates National Museum Bulletin 206 xvi+423

Duellman WE (1970) The hylid frogs of Middle AmericaMonograph of the Museum of Natural History 21 1-372

Duellman WE Trueb L (1986) Biology of AmphibiansMcGraw Hill New York

Francioni E Carcerelli LC (1993) Descriccedilatildeo do girinode Crossodactylus gaudichaudii Dumeacuteril and Bibron1841 (Anura Leptodactylidae) Memoacuteria do InstitutoButantan 55 63-67

Frost DR (2004) Amphibian Species of the WorldAn Online Reference V30 (22 August 2004) Elec-tronic database accessible at lthttpresearchamnhorgherpetologyamphibiaindexhtmlgt American Museumof Natural History New York USA [captured on 10 Oc-tober 2004]

Gosner KL (1960) A simplified table for staging anuranembryos and larvae with notes on identification Her-petologica 16 183-190

Weygoldt P Carvalho e Silva SP (1992) Mating andoviposition in the hylodine frog Crossodactylus gau-dichaudii (Anura Leptodactylidae) Amphibia-Reptilia13 35-45

Received May 19 2005 Accepted July 13 2005