Taxonomic Review of Allobates (Anura, Aromobatidae) from ... Rodrigues_2007_Taxonomic Review... · Taxonomic Review of Allobates (Anura, Aromobatidae) from the Atlantic Forest, Brazil

Taxonomic Review of Allobates (Anura, Aromobatidae) from theAtlantic Forest, Brazil

VANESSA K. VERDADE1

AND MIGUEL T. RODRIGUES

Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Caixa Postal 11461, CEP 05422–970,

Sao Paulo, Brazil

ABSTRACT.—We present the results of a taxonomic review of the four species of Allobates endemic to the

Atlantic Forest Domain in Brazil. A total of 880 preserved specimens from 29 localities covering their range in

Atlantic Forest were studied based on external attributes. Characters formerly described as diagnostic in

original descriptions and others cited for dendrobatids in the literature were studied along the total

geographic range of the populations of these species, including the type localities. We found no discrete

characters, qualitative or quantitative, capable of differentiating the species. Most local and geographic

variation was limited to snout-vent length and color pattern. Considering the lack of evidence otherwise, we

use the results of our morphological analysis to allocate all Atlantic Forest species of Allobates to synonymy

with Allobates olfersioides.

Dendrobatids are a monophyletic group offrogs that are mostly known for their brightcolor and poisonous skin. Nevertheless, abouthalf of the species are cryptically and dullcolored, presumably nontoxic and until recently(Grant et al., 2006) placed in the genus Colos-tethus sensu lato. There had been about a hun-dred recognized species in the genus rangingfrom Nicaragua to southeastern Brazil (Frost,2006). The systematics of the family has recentlygone through significant changes (Grant et al.,2006) supported by a molecular and morpho-logical based phylogeny. The family Dendroba-tidae, as previously known, corresponds now toDendrobatoidea, which includes the familiesAromobatidae and Dendrobatidae. The fourspecies of Colostethus from Atlantic Forest,now allocated to the genus Allobates, are placedin the family Aromobatidae.

The Brazilian species of Allobates are leaf litterinhabitants of forests and can be found through-out the Amazon Basin, Brazilian Savannas, andAtlantic Forest. Most of the 17 currentlyrecognized species are Amazonian and de-scribed recently, probably as a consequence ofa vast and poorly sampled territory along withincreased knowledge of the natural history ofthese frogs (e.g., Morales, 2000; Lima andCaldwell, 2001; Caldwell and Lima, 2003). Thereare four species in eastern Brazil, endemic to theAtlantic Forest Domain (Ab’Saber, 1977): Allo-bates alagoanus (Bokermann, 1967), Allobatescapixaba (Bokermann, 1967), Allobates carioca(Bokermann, 1967), and Allobates olfersioides(Lutz, 1925).

Allobates olfersioides was described as Eupem-phix olfersioides Lutz, 1925 from the coastalregion of the state of Rio de Janeiro (RJ).Cochran (1955) considered it a synonym ofPhyllobates brunneus Cope, 1887. Bokermann(1966) restricted the type locality of A. olfer-sioides to Angra dos Reis (RJ) and, in 1967,described the other three Atlantic Forest species:Phyllobates alagoanus from Usina Sinimbu, Man-gabeiras, state of Alagoas; Phyllobates capixabafrom Lagoa do Macuco, Refugio Sooretama,Linhares, state of Espırito Santo; and Phyllobatescarioca from Represa Rio Grande, Jacarepagua,Rio de Janeiro, state of Rio de Janeiro. Boker-mann (1967) also considered P. olfersioides re-lated to P. alagoanus, P. capixaba and P. carioca.Edwards (1974) placed these four species inColostethus, as did Silverstone (1975). Grant et al.(2006) based on an extensive sampling andmolecular plus morphological based phylogenyallocated them to the genus Allobates.

The four species are very similar in externalattributes and were described based on smalland geographically remote samples. The char-acters used by Bokermann (1967) to separatethese species were basically color pattern,length of limbs and fingers, size of carpal andtarsal tubercles, shape of head, and snout-ventlength. All of these characters are presumablysubject to geographical variation, a possibilitythat could not be tested at that time.

Bokermann (1967) also presented informationon call variation, pointing out that the calls of A.capixaba and A. carioca were distinct from that ofA. olfersioides, but, except for saying that the callof A. capixaba was more spaced than those of A.olfersioides, no further description was given.Besides, the putative call differences between1 Corresponding Author. E–mail: [email protected]

Journal of Herpetology, Vol. 41, No. 4, pp. 566–580, 2007Copyright 2007 Society for the Study of Amphibians and Reptiles

species were reported by the collector (F. M.Oliveira) and not based on the authors’ fieldexperiences or knowledge. The main character-istics of the advertisement call of Allobates (a trillor an acute buzz) can vary in fundamentalfrequency, number of pulses and spacing,according to environmental conditions andsometimes, as a function of population density(Coloma, 1995; Junca, 1998; Grant and Rodri-guez, 2001; Caldwell et al., 2002a). Consideringthe subjective call differences reported and theirvariation, we think that the evidence alone isweak to keep the Atlantic Forest species ofAllobates separated. Moreover, the observationson vocalization pointed out in the speciesdescription refer only to the surroundings ofthe type locality, and geographical variation ofcall should be considered. Unfortunately, wewere unable to find available calling records incollections, either institutional or personal, andour attempts to get new ones failed. Becausethis problem will probably persist in the future,because the Atlantic Forest species of Allobatesare showing declines (Weygoldt, 1989; Izeck-sohn and Carvalho-e-Silva, 2001), specimensgathered in collections will be the main sourcefor studying species limits of this complex in theAtlantic Forest. Indeed, the samples collectedduring the last 40 years are a source of new dataon morphological variation that could clarifythe similarities and differences among the fourspecies of Allobates from Atlantic Forest.

We present, herein, a detailed study of themorphological variation of these populationsalong a latitudinal gradient that includes thetype localities of the presently recognizedspecies. We assume that external morphologyis the strongest available source of data sustain-ing the Atlantic Forest species of Allobates, and,if we found no discrete differences in morpho-logical characters among populations, theyshould be considered synonyms until newsources of data can be explored.

MATERIALS AND METHODS

To cover their total range, 880 preservedspecimens of Allobates from 29 localities werestudied (Fig. 1, Appendices 1 and 2). All speci-mens were observed regarding external mor-phology. We measured and classified 396individuals to be used in the statistical analysis.This number comprises all individuals fromlocalities with N below 100 and 100 randomlypicked up individuals from localities with Nabove 100. Twenty-two measurements wereobtained for the specimens included in theanalysis: snout-vent length (SVL); thigh length(THL) from the middle of the cloacal opening tothe outer edge of the flexed knee; tibia length

(TIL) from the outer edge of the flexed knee tothe heel; foot length (FOL) from the heel to thetip of toe IV; head width (HW) between theangle of jaws; head length (HL) perpendicularto the imaginary line linking the angle of jaws tothe tip of the snout; internarial distance (IND)between the inner edge of the nares; eye-nostrildistance (END) from the anterior corner of theeye to the outer edge of the nostril; eye diameter(ED) from posterior to anterior corners; tympa-num diameter (TD) from posterior to anterioredges; interorbital distance (IOD) between theinner edges of the eyelids; hand length (HAL)dorsally from the junction of radio-ulna andcarpal to the tip of the finger III; thumb length(TBL); second, third, and fourth fingers length(FL2, FL3, FL4) dorsally from the base to the tipof the finger; third finger diameter (TFD)dorsally at the distal edge of the second fingertubercle; greatest length of the inner and outercarpal tubercles (ICT, OCT); and greatest lengthof the inner and outer metatarsal tubercles(IMT, OMT).

All measurements were taken on the left sideof the specimens. The measurements wereobtained with a stereomicroscope except forSVL, taken by a dial caliper, to the nearest of0.01 mm. The axis of the greatest length at ICT,OCT, IMT, and OMT variables was the same forall specimens.

Qualitative characters studied were: shape ofthe snout (dorsal, ventral, and lateral views);pattern of the lateral dark stripe between the

FIG. 1. Geographic distribution of Atlantic ForestAllobates. Open circles represent the type localities ofthe four currently admitted species. Numbers are thelocalities sampled (see Appendix 2). The OTUsstudied are indicated by OTU1, 2, 3, and 4.

TAXONOMIC REVIEW OF ALLOBATES FROM THE ATLANTIC FOREST 567

nares and the eyes; color pattern of the upperlip; presence of a median lingual process (Grantet al., 1997, 2006); color pattern of the belly;color pattern on dorsal view; pattern of distri-bution of tubercles; pattern of dark stripes ongroin; pattern of the dorsolateral and laterallight stripes; pattern of the lateral dark stripe(Fig. 2); cloacal stains; color pattern of the thigh;presence/absence and extension of metatarsalfold; and toe webbing.

To make sure that there were no extralimitalspecimens belonging to other species in oursample, and that the Atlantic Forest specieswere restricted to the study area, we diagnosedthem from all other Brazilian species (SociedadeBrasileira de Herpetologia, 2005) previouslyplaced in the genus Colostethus (now in thegenera Allobates, Anomaloglossus, and Hyloxalus),based on literature and direct examinationwhen possible: Allobates brunneus (Cope, 1887);Allobates caerulodactylus, (Lima and Caldwell,2001); Allobates conspicuus (Morales, 2000); Allo-bates crombiei (Morales, 2000); Allobates fuscellus(Morales, 2000); Allobates gasconi (Morales,2000); Allobates goianus (Bokermann, 1975);Allobates granti (Kok et al., 2006); Allobatesmarchesianus (Melin, 1941); Allobates masniger(Morales, 2000); Allobates nidicola (Caldwell andLima 2003); Allobates sumtuosus (Morales, 2000);Allobates vanzolinius (Morales, 2000); Anomalo-glossus beebei (Noble, 1923); Anomaloglossusstepheni (Martins, 1989); Anomaloglossus tama-cuarensis (Myers and Donnelly 1997); andHyloxalus peruvianus (Melin, 1941). We decidedto diagnose them also from Allobates trilineatus(Boulenger, ‘‘1883’’ 1884) and Anomaloglossusdegranvillei (Lescure, 1975), considering theirdistribution close to Brazil (Grant and Rodri-guez, 2001; Frost, 2006).

The abbreviations for the herpetological col-lections consulted for preserved specimens andcall records are presented throughout the text asfollows: EI (Eugenio Izechson collection, Uni-versidade Federal Rural do Rio de Janeiro,UFRRJ); JJ (Jorge Jim collection, Universidade

Estadual de Sao Paulo, UNESP); MNRJ (MuseuNacional do Rio de Janeiro); MUFAL (Museu daUniversidade Federal de Alagoas); MZUSP(Museu de Zoologia da Universidade de SaoPaulo, currently including the Werner Boker-mann collection, WCAB); UEFS (UniversidadeEstadual de Feira de Santana); UFPB (Universi-dade Federal da Paraıba); UFRJ (UniversidadeFederal do Rio de Janeiro); USNM (UnitedStates National Museum); ZUEC (Museu deHistoria Natural, Universidade de Campinas,UNICAMP).

There were no obvious secondary sexualcharacters to differentiate males and femalesamong the four species studied. To avoiddamage caused by dissection we checked formorphometric differentiation between sexesonly in a sample from Tijuca, Rio de Janeiro.Forty-four adult individuals (checked by matu-rity of gonads) from a total of 400 weredissected, and 20 males and 20 females werecompared under a Student t-test at a 95%interval of confidence.

Three samples of tadpoles identified by thepresence of nursery adults or young meta-morphs were available for study—Ilheus (Ba),Una (Ba), and Tijuca (RJ). These samplescontained specimens in different developmentalstages and were not suitable for geographicalcomparison. Tadpoles from Tijuca (RJ) wereused to describe the larvae following Altig andMcDiarmid (1999) standard measurements anddescriptive terminology.

Statistical Analysis.—Specimens were firstobserved and compared within localities todetect possible cases of sympatry, and thenobserved and compared among localities fol-lowing a north-south transect. In addition, weran a preliminary stepwise Discriminant Anal-ysis to assess differences among localitiestreated as individual operational taxonomicunits (OTUs) and considering the only case ofsympatry suggested in the literature (Allobatesolfersioides and A. carioca; Bokermann, 1967) asindividual units. The observational and statisti-cal results were subsequently considered togroup the specimens in four OTUs, based onoverall similarity and geographical distribution.The OTUs are also congruent to the gapsobserved in the geographical distribution ofthe four species considered together (Fig. 1).The specimens from the states of Alagoas andSergipe were treated as OTU1 (N 5 57), speci-mens from the state of Bahia as OTU2 (N 5 68),specimens from the state of Espırito Santo asOTU3 (N 5 11), and specimens from the state ofRio de Janeiro as OTU4 (N 5 260; Fig. 1).Statistics were obtained for each OTU.

The absence of obvious sexual charactersmade it difficult to classify the specimens as

FIG. 2. Schematic drawing of an Allobates with thenomenclature used for the study of color pattern ofthe specimens.

568 V. K. VERDADE AND M. T. RODRIGUES

young or adults. To minimize deviations of themeans, we chose not to use very small speci-mens. As a criterion of SVL exclusion, we usedthe SVL average less the standard deviation ofthe OTU 1, which presented the smallestindividuals. The value obtained was 11.4 mm.All the specimens with SVL equal to or smallerthan 11.4 mm were excluded from the sub-sequent quantitative analysis, which left us with372 specimens to be statistically analyzed: OTU1 (N 5 44), OTU 2 (N 5 66), OTU 3 (N 5 10),and OTU 4 (N 5 252). None of the samplespresented normal distributions for all variables,even with data log-transformed, but deviationswere not severe enough to prevent parametrictests.

As an attempt to express differences in shapeamong individuals in different samples, somebody ratios were used in addition to the rawmeasurements. They are THL/SVL, HAL/SVL,FOL/SVL, HW/HL, END/HL, ED/HL, TD/HL, TD/ED, IND/HW, TBL/FL2, FL2/HAL,and FL4/HAL.

To classify the specimens, three stepwiseDiscriminant Analyses were run separatelybased on the morphological measurements,body ratios, and qualitative data. The numberof steps was unspecified, and all groups wereassigned to equal probabilities before analysis.Sexual dimorphism for the Tijuca sample wasassessed by the Student t-test on each variableand body ratio at a confidence interval of 95%.Means are given 6 1 SD.

RESULTS

All localities sampled are restricted to thecoastal region of eastern Brazil. The localityfarthest north is Novo Dino, state of Alagoasand the farthest south is Angra dos Reis, stateof Rio de Janeiro. The farthest west locality isBaixo Guandu in state of Espırito Santo. Thereare two major gaps along the north-southtransect. One between Areia Branca in the stateof Sergipe and Elısio Medrado, state of Bahia,and the other between Santa Teresa, state ofEspırito Santo and Teresopolis, state of Rio deJaneiro. These areas correspond to drier envir-onments. In the north, the coastal region iscomposed by fixed sand dunes, but there aremany forested regions that could conceal thespecies. The southern region is still covered byforests. It seems more probable that the absenceof samples of Allobates from these areas is anartifact of collection (Heyer, 1988).

Qualitative Data.—All individuals presenta rounded snout tip in dorsal and lateral viewswith nares slightly prominent and an upper jawprojecting slightly above the lower jaw. Thegeneral shape of the snout in dorsal view

resembles a trapezoid; the body is slender andelliptical (Fig. 3A, B). The dorsal skin is gener-ally smooth, but inconspicuous tubercles maybe present at the inguinal region or throughoutthe entire dorsum.

Dorsal pattern consists of a reddish-brownbackground over which are darker stripes thatcross once or more at the middle of the dorsum,suggesting intercrossing Xs. Dorsal pattern canvary from uniform to highly pigmented (Fig. 4;for summary, see Table 1). Individuals witha uniform dorsum are more frequent (about15%) in samples from the state of Bahia (OTU 2)but still occur in lower rates in other samples (,1% from the State of Rio de Janeiro, OTU 4).Individuals with a single X on the dorsum aremore common (74%) in the states of Alagoasand Sergipe (OTU 1), but specimens withmultiple X also occur in these samples (24%).The multiple-X pattern is the most common inOTU 2, 3, and 4, present in about 40, 80, and90% of the individuals, respectively (Table 1).There is some variation in this pattern. Some-times the largest X has longer arms that barelytouch each other. This variation is found inindividuals from samples from the states ofEspırito Santo (OTU 3) and Rio de Janeiro (OTU4) and is more common among the former(45%). The dorsal parts of the thigh and tibiahave two or three dark stripes bordered bylighter ones in almost all specimens examined.In some individuals, the dark stripes are muchnarrower, almost a line, in others the stripes areabsent.

Laterally, all individuals present a dark stripefrom the tip of the snout to the groin, involvingthe nares and partially the eyes along the side ofthe body (Fig. 5). Apparent north-south varia-tion exists in the width and definition of thisstripe. Following the north-south transect, thereis a tendency of the dark lateral stripe to becomewider with a crescent reticulation toward theventer and inguinal region (Fig. 6). Individualsfrom the state of Espırito Santo (OTU 3) are theones in which this characteristic is more pro-nounced. To the south, the dark lateral stripereverses to solid in individuals from northernRio de Janeiro (OTU 4), but it is still wide. At thesouthernmost localities, the lateral stripe isnarrower and solid, as observed in individualsfrom the state of Alagoas, Sergipe (OTU 1) andBahia (OTU 2).

The dark lateral stripe can be dorsallybordered by a narrow pale stripe that varies inlength and continuity. Most of the individuals(97%) from the state of Alagoas (OTU 1) do notpresent the pale dorsolateral stripe (Table 1).When it occurs, it is present from the eyes to thearms. The pale dorsolateral stripe is not presentin 57% of the individuals from the state of Bahia


(OTU 2) and can be observed from the eyes tothe arms in 36% of them and from the eyes tothe groin in 7%. In the state of Espırito Santo(OTU 3), all individuals present the paledorsolateral stripe. It is observed from the eyesto the arms in 9% of them and from the eyes tothe groin in 91% of them. To the south, only16% of the individuals from the state of Rio deJaneiro (OTU 4) present the pale dorsolateralstripe.

Individuals from the states of Alagoas andSergipe (OTU 1) commonly (62%) have venterpigmentation restricted to the throat, neverextending from throat to groin (Table 1). Sam-ples of state of Bahia (OTU 2) specimens with animmaculate venter are more frequent (57%).There are two equally common patterns ofventer pigmentation in individuals from thestates of Espırito Santo (OTU 3) and Rio deJaneiro (OTU 4). Venter pigmentation extends

to the pectoral or inguinal regions in about 30%of individuals in both states. The general colorpattern is highly variable, and we did not findany geographical consistency or discrete localvariation to justify partitioning of the samples.

Quantitative Data.—It was impossible to di-agnose populations studied and refer them toany of the recognized species based on bodyproportions or the following diagnostic char-acters pointed out in literature: SVL, ratio oflength of the thumb and the second finger,absolute length of the fourth finger, and shapeof the tarsal tubercle (Bokermann, 1967; Ed-wards, 1974; Rivero, 1988; Morales, 2000).Table 2 presents the descriptive statistics forthe main measurements taken for individuals inthe four OTUs.

Individuals from the states of Alagoas andSergipe (OTU 1) are smaller (t df 5 107 5 26.316,P 5 0.000) than those from the state of Bahia

FIG. 3. Allobates olfersioides (MNRJ 5094): (A) dorsal and (B) lateral views of head, ventral views of (C) righthand and (D) left foot.


(OTU 2; Table 2). Individuals from the state ofBahia (OTU2) are larger (t df 5 74 5 22.324, P 5

0.023) than individuals from the state of EspıritoSanto (OTU 3). Mean of SVL of individuals fromthe states of Espırito Santo (OTU3) and Rio deJaneiro (OTU 4) are not different (t df 5 260 5

20.042, P 5 0.389). Despite statistically signif-icant OTUs, differences in individuals SVLamong localities are not clear when visualizedin a graphic of dispersion (Fig. 7). The smallestmaximum sizes are found among individualsfrom the State of Espırito Santo, but it representsthe smallest OTU (N 5 10), with localitiespoorly sampled.

The results of the discriminant analysis basedon both raw measurements and body ratiosshow overlap among the four operational units.The results for the discriminant analysis based

on body ratios are presented in Figure 8. Thefirst and second discriminant functions explain98% of the differences among the OTUs. Themost important variable on function 1 is TBL/FL2 and on function 2 is FL4/HAL. Thevariation of the values of the TBL/FL2 andFL4/HAL for each locality from north to southcan be seen in Figure 9. The shape of the tarsaltubercle shows variation, and individuals fromthe same sample can have a lump, or a straight-or comma-shaped ridge.

We tried to rearrange the localities intodifferent OTUs, following, for example, the mainriver barriers (e.g., Sao Francisco, Jequitinhonha,Doce, and Paraıba do Sul), but the results werealways similar. None of the measurements orproportions studied presented evidence to sep-arate specimens into different groups.

TAXONOMIC CONCLUSION

Our study of Allobates from the AtlanticForest made it clear that the characters currentlyused to differentiate the recognized specieswere not useful in our sample (Bokermann,1967; Edwards, 1974; Rivero, 1988; Morales,2000). Variation in SVL throughout their dis-tributions is probably a result of restrictedsampling in some of the localities (Fig. 7). Thedata also show that the relative lengths of thethumb, second, and fourth fingers are highlyvariable, as already found for other species ofAllobates (Grant and Castro, 1998). Diagnosticdifferences between species are probably mis-interpretations based on limited series.

The only record of sympatry among theAtlantic Forest Allobates was published byBokermann (1967) from Represa Rio Grande,state of Rio de Janeiro, where A. olfersioides andA. carioca were found. This sample, presently atMZUSP, includes 19 specimens, three of whichwere identified by Bokermann as A. carioca (theholotype and the two paratypes). At first, thesespecimens look slightly different from thetypical A. olfersioides, presenting a larger sizeand uniform dorsum. However, difference insize detected visually is not significant when themean SVL is compared (t df 5 16 5 1.62, P 50.12). In addition, although very rare (less than1%), the uniform dorsum is present in othersamples of A. olfersioides from the state of Rio deJaneiro. Izecksohn and Carvalho-e-Silva (2001)have already suggested the possibility that thesetwo taxa are actually synonyms. When com-pared with individuals from other OTUs, theseindividuals are very similar to those from thestate of Bahia (OTU 2) presenting the samegeneral characteristics in size and color pattern.Unfortunately, vocalization, the only otherargument used to justify this species could not

FIG. 4. Dorsal view of adult Allobates olfersioidesshowing variation on dorsal color pattern.


be tested here because there are no availablecalling records and the attempt to get newrecords did not succeed.

According to the data presented above, thereis no external morphological evidence to keepthe four species of Allobates known from theAtlantic Forest as separate units. It is possiblethat new ecological, behavioral, cytogenetic, ormolecular evidence could contribute to ourunderstanding of the systematics of this group.Until such time as additional evidence is avail-able, we suggest a more conservative postureand treat A. alagoanus, A. capixaba, and A. cariocaas synonyms of Allobates olfersioides.

Allobates olfersioides (Lutz, 1925)Figure 3

Eupemphix olfersioides Lutz, 1925: 138.Holotype.—MNRJ 783 (currently exists only as

a piece of flesh), collected on the coast of thestate of Rio de Janeiro, Brazil; Angra dos Reis,state of Rio de Janeiro, according to Bokermann(1966).

TABLE 1. Frequencies of dorsal and ventral color patterns taken from adults of Atlantic Forest Allobates

included in the four operational taxonomic units (OTU; see text for definitions of OTUs).

OTU 1 OTU 2 OTU 3 OTU 4(N 5 57) (N 5 68) (N 5 11) (N 5 260)

Central dorsal patternUniform 1 (2%) 9 (13%) 0 1 (, 1%)Single X 42 (74%) 2 (3%) 0 0Multiple Xs 14 (24%) 57 (84%) 5 (45%) 240 (92%)Long arm X 0 0 6 (55%) 19 (7%)

Dorsolateral pale stripeAbsent 55 (97%) 39 (57%) 0 219 (84%)From eye to arm 2 (3%) 24 (36%) 1 (9%) 15 (6%)From eye to inguinal region 0 5 (7%) 10 (91%) 26 (10%)

Venter pigmentationAbsent 15 (26%) 29 (42%) 1 (9%) 50 (19%)Restricted to throat 35 (62%) 16 (24%) 2 (19%) 34 (14%)From throat to chest 7 (12%) 19 (28%) 4 (36%) 84 (32%)From throat to inguinal region 0 4 (6%) 4 (36%) 92 (35%)

FIG. 6. Lateral view of adult Allobates olfersioidesshowing variation on dark lateral stripe.

FIG. 5. Allobates olfersioides, live specimen fromMaceio, state of Alagoas (Photograph: G. Skuk).


Phyllobates capixaba Bokermann, 1967: 349.Holotype.—WCAB 19252 (MZUSP 73752), col-

lected at Refugio Sooretama, Linhares, state ofEspırito Santo, Brazil.

Phyllobates alagoanus Bokermann, 1967: 351.Holotype.—WCAB 2801 (MZUSP 73707), col-

lected at Mangabeiras, state of Alagoas, Brazil.

Phyllobates carioca Bokermann, 1967: 352.Holotype.—WCAB 38610 (MZUSP 76653), col-

lected at Represa Rio Grande, Jacarepagua, Riode Janeiro, state of Rio de Janeiro, Brazil.

Allobates olfersioides Grant, Frost, Caldwell,Gagliardo, Haddad, Kok, Means, Noonan,Schargel, and Wheeler, 2006.

TABLE 2. Descriptives for the main measurements taken from adults of Atlantic Forest Allobates included in thefour operational taxonomic units (OTU; see text for definitions of OTUs). Data are presented as mean 6 SD (N) range.

OTU 1 OTU 2 OTU 3 OTU 4

Snout–vent length (SVL) 14.5 6 1.4 (43) 16.5 6 1.7 (66) 15.1 6 1.5 (10) 15.5 6 1.4 (252)11.5–17.2 11.5–18.8 13.3–17.2 11.5–18.6

Thigh length (THL) 6.5 6 0.6 (43) 7.8 6 0.7 (65) 6.7 6 0.6 (10) 7.1 6 0.6 (249)5.0–7.7 5.5–8.8 5.7–7.5 3.8–8.5

Tibia length (TIL) 6.8 6 0.5 (42) 7.8 6 0.7 (65) 7.0 6 0.3 (10) 7.2 6 0.6 (249)5.5–7.5 5.2–8.8 6.5–7.6 4.8–9.0

Hand length (HAL) 2.9 6 0.4 (43) 3.4 6 0.4 (66) 3.0 6 0.2 (10) 3.3 6 0.4 (252)2.2–3.8 2.0–4.7 2.8–3.3 2.0–4.1

Thumb length (TBL) 1.5 6 0.2 (43) 1.8 6 0.3 (66) 1.5 6 0.2 (10) 1.6 6 0.2 (252)1.1–1.8 1.1–2.4 1.2–1.8 0.9–2.2

Head length (HL) 3.7 6 0.3 (43) 3.7 6 0.3 (66) 3.5 6 0.3 (10) 3.7 6 0.3 (252)3.1–4.4 2.8–4.7 3.1–4.1 2.9–4.9

Foot length (FOL) 5.8 6 0.7 (43) 6.9 6 0.7 (66) 6.1 6 0.3 (10) 6.6 6 0.6 (252)2.9–6.8 4.2–8.1 6.8–7.2 4.2–7.6

Second finger length (FL2) 1.1 6 0.1 (43) 1.4 6 0.2 (66) 1.2 6 0.2 (10) 1.4 6 0.2 (251)0.8–1.3 0.7–1.9 0.9–1.4 0.8–1.8

Head width (HW) 4.7 6 0.3 (44) 5.3 6 0.5 (66) 4.8 6 0.3 (10) 4.9 6 0.4 (252)3.7–5.4 3.8–6.2 4.3–5.2 3.9–5.7

Fourth finger length (FL4) 0.6 6 0.1 (43) 0.9 6 0.1 (66) 0.8 6 0.1 (10) 0.9 6 0.1 (252)0.4–1.0 0.6–1.2 0.7–0.9 0.4–1.2

FIG. 7. Values of the snout–vent length (SVL) measured in each sample of Atlantic Forest Allobates studied.Dots represent maximum SVL values, the squares are mean, and the bars, the standard error for each sample.Localities are presented with shortened names (see Appendix 2) from north (left) to south (right). The namesabove the graphic are the states to which the localities belong.


Diagnosis.—A small species of dendrobatid(maximum SVL 5 19 mm in males andfemales); thumb slightly longer than secondfinger (FL2 on average 85% of TBL); third fingernot swollen in adult males (TFD 0.3 mm inaverage); throat, chest, and belly light cream,almost unpigmented; toes III–IV basallywebbed; dorsum brown, generally adornedwith a pattern of intercrossing Xs; pale dorso-

lateral stripe absent or present; dark lateralstripe continuous extending from tip of snout togroin; ventrolateral stripe present, poorly de-fined; skin of dorsum smooth or slightlygranular in sacral region; skin of venter smooth;paracloacal ‘‘C’’ like light stains present; cloacaltubercles absent; median lingual process absent.Testis unpigmented in adult males.

Comparison with Other Species.—Given theiroverall similarity, in the following section wecompare Allobates olfersioides with all species ofAllobates, Anomaloglossus, and Hyloxalus re-corded from Brazil, and those likely to be foundin Brazilian territory. Allobates brunneus differsby presenting a wider and diffuse dark lateralstripe, vocal sacs visible in males, and a dorsumwith dark blotches; A. caerulodactylus differs bypossessing blue coloration on digits and toediscs, and basal web between toes II, III and IV;A. conspicuus, A. crombiei, and A. gasconi presentreduced fringes on toes. The abdomen isgranulated in A. conspicuus; A. crombiei havesecond and fourth fingers of same size (FL2 .FL4 in A. olfersioides) and dark blotches ondorsum; the males of A. gasconi and A. fuscelluspresent a swollen third finger and sexualdimorphism in color with venter darker inmales than females; A. goianus differs by theabsence of a conspicuous dark lateral stripe,toes fringed and webbed and dorsal colorpattern formed by four reddish-brown blotches;A. granti differs by presenting dorsal colorpattern uniform with brown flecks, anal flap,hind limbs with granular skin, reduced web-bing between toes II, III, and IV (between III andIV in A. olfersioides), narrow and longer head(HL 88% of HW, HL 30% of SVL; 80% and 24%in A. olfersioides), larger hands (HAL 30% ofSVL; 21% in A. olfersioides), and longer legs (TIL47% of SVL; 44% in A. olfersioides); A. marche-sianus differs from A. olfersioides by havinga uniform dorsum scattered with darker spots,by the dark lateral stripe which is wider at theinguinal region and can be seen dorsally, by thethumb distinctively longer than the secondfinger (FL2 about 70% of TBL; 85% in A.olfersioides), and by the gray throat in males; A.masniger has a more robust body, the dorsum isuniformly brown with some dark spots in twoparallel lines, and the males have a dark throat;A. nidicola differs from A. olfersioides by its largermaximum size (to 21.4 mm), presence of basalweb between toes II, III, and IV, presence ofa dark lateral stripe short and diffuse and throatand chest black to light gray in males; A.sumtuosus differs by presenting reduced fringeson toes II, III, and IV, and swollen third finger inmales; A. trilineatus differs from A. olfersioidesmainly by males presenting finger III and oftenfinger II strongly swollen, venter stippled with

FIG. 8. Graphic representation of the results of theDiscriminant analyses based on the Atlantic ForestAllobates body ratios divided in four operationaltaxonomic units (OTU). The OTUs are congruent tothe political limits of the states of Alagoas/Sergipe,Bahia, Espırito Santo, and Rio de Janeiro, Brazil (fordetailed distribution see Fig. 1).

FIG. 9. Ratios of thumb length by length of finger 2(upper graphic) and length of finger 4 by hand length(lower graphic) of Atlantic Forest Allobates studied foreach locality. Squares are mean and bars are thestandard errors for each sample. The localities arepresented with shortened names (see Appendix 2)from north (left) to south (right). The names above thegraphic are the states to which the localities belong.


gray and by the dark lateral stripe along flankthat becomes diffuse at the midbody to theinguinal region; A. vanzolinius differs from A.olfersioides by its larger maximum size (to22.9 mm) and by males presenting a darkerthroat than females; Anomaloglossus beebei bypresenting dorsum uniformly brown with scat-tered dark dots, by the presence of medianlingual process, swollen third finger, reducedweb between toes II, III, and IV, and by thepresence of a well defined tarsal fold; A.degranvillei has a pigmented and marbledventer, a more dorsally located lateral darkstripe with borders less defined, a dark cloacalregion, the median lingual process is present,the toes are fringed and a dermal fold is presentat the outer margin of the feet; A. stepheni differsby presenting a wider and robust body shape,median lingual process, a swollen digit III inmales, toes II and III webbed and a conspicuouslight dorsolateral stripe from eyelid to theinguinal region; A. tamacuarensis differs byreaching a larger size (to 25 mm), by presentingmedian lingual process, dorsal blotched pattern,inner tarsal fold and fringes along side offingers; Hyloxalus peruvianus differs by reachinga larger size (to 21 mm), by presenting finger IIand toe IV fringed, by the presence of an outertarsal fold, and toes II, III, and IV webbed.

Description.—Based on MNRJ 5094 from thetype locality Angra dos Reis, state of Rio deJaneiro. Head slightly wider than long (HL 80%of the HW); head length 24% of the SVL; snoutrounded in dorsal and lateral views, with naresslightly prominent located and opening lateral-ly; upper jaw slightly projected; canthus ros-tralis rounded; loreal region slightly concave,almost flat; internarial distance 39% of the HW;eye-nostril distance 62% of the eye diameter(ED); tympanum 42% of the ED with theanteroventral margin distinct and the poster-odorsal obscured by the depressor musculature;tongue attached anteriorly; median lingual pro-cess absent; vocal slits present; dorsal surfacesmooth with low, inconspicuous granules re-stricted to sacral region; dorsal surfaces of thighand shanks smooth; although discolored there isindication of a pattern of intercrossing exes ondorsum; a dark lateral stripe extends from thetip of snout, involving the nares and partiallythe eyes, to the groin; ventral surface smooth;belly light cream; no anal flap above vent; noenlarged tubercles near anus; forelimb slender,skin smooth; dark bands present mainly at theouter margin of arms; no ulnar fold, handlength 21% of SVL; appressed thumb slightlylonger than finger II (FL2 87% of TBL); finger IIlonger than IV (FL4 77% of FL2); relative fingersize when appressed III . II 5 I . IV; outermetacarpal tubercle nearly round, diameter 16%

of HAL; inner metacarpal tubercle elliptical,slightly smaller than the former; supernumerarytubercle absent; one subarticular tubercle onfingers I, II and IV and two on finger III; nofringes on fingers; finger III not swollen;webbing between fingers absent; tips of digitsexpanded (width of finger about 75% of thewidth of the disc); discs with distinct dorsalscutes; thighs and tibia brown with two darkstripes that form a continuum when hind limbsare in natural position; tibia length 44% of SVL;relative length of appressed toes IV . III . V .

II . I; discs on toes larger than the width of toes(as for fingers); rudimentary webbing presentonly between toes III and IV; inner metatarsaltubercle elliptical, outer metatarsal tuberclerounded; one subarticular tubercle on toes Iand II, two on toes III and V, and three on toeIV; toes not fringed; metatarsal fold absent;tarsus smooth with an inconspicuous tubercle;tarsal fold absent.

Measurements of the Specimen Described (mm).—SVL 16.9; HL 4.1; HW 5.1; ED 2.4; END 1.5; IOD2.2; HAL 3.5; TBL 1.5; FL2 1.3; FL4 1.0; THL 7.8;TIL 7.5; FOL 7.4.

Color Variation in Preservative (N 5 880).—Inmost specimens, the dorsum shows a pattern ofdark brown intercrossing Xs, with branches thatcan vary in length and width in a lighterbackground (Fig. 4). It is uniformly light brownin some individuals where intercrossing Xs areabsent. A dark lateral stripe extending from thetip of the snout, through the eyes to the inguinalregion is always present. This stripe is narrowerat the nares level, becoming gradually widerthrough the lateral of the body narrowing againat the inguinal region. The tympanum iscovered by the dark stripe dorsally (abouta third). The lateral stripe can be uniformlydark throughout the length or present scatteredwhite spots at the inguinal region, being diffusetoward the venter and inguinal region (Fig. 6).At the end of the stripe, there is an obliquenarrow light stripe that divides it from the shortdark stripes present at the dorsum. The belly islight cream, and some tiny spots of dark brownpigmentation of some individuals can be seen inthe throat and chest or through the entire belly.The arms and legs present the same backgroundcolor of the dorsum. The arms present somedark bands mainly at its outer margin, and thedorsum of the thigh and tibia generally showtwo or three dark brown stripes that vary inwidth and form a continuum when the leg isflexed. The palms of hands and sole of feet arepigmented as the general background color. Atthe end of the fingers and toes, there are lighttransversal stripes. There are C-like light creamstains at each side of the cloacal aperture. The


color pattern does not seem to vary ontogenet-ically.

Color in Life.—Based on MZUSP 132193–95from Una, state of Bahia, and photographs of anindividual from Porto Seguro, state of Bahia, anindividual from Maceio, state of Alagoas(Fig. 5), and an individual from Rio de Janeiro,State of Rio de Janeiro (Izecksohn and Carvalho-e-Silva, 2001:fig. 78). The iris is bronze. Thegeneral color pattern follows that described forpreserved specimens. The light cream parts lookmore bright and white in live specimens, and thecontrast between the background color and thedarker pattern is more evident.

Sexual Dimorphism.—The data obtained fromTijuca, state of Rio de Janeiro suggested nomarked sexual dimorphism. Females are slight-ly larger (SVL) than males (t df 5 39 5 22.15, P 50.04; female 5 16.4 6 0.7 mm [N 5 20]; male 516.0 6 0.6 mm [N 5 20]) and present animmaculate belly more frequently. Fifty percentof the females examined had an immaculatebelly, 35% a belly pigmented from throat tochest, and 15% a belly pigmented from throat toinguinal region. No males had an immaculatebelly, 15% had a belly pigmented from throat tochest, and 85% a belly pigmented from thethroat to the inguinal region. Males and femalesdid not differ in any other variable. Theseresults agree with the general pattern of colordimorphism observed for the genus (Savage,1968; Edwards, 1974; Coloma, 1995; Kok et al.,2006).

Advertisement Call.—The vocalization of Allo-bates olfersioides was described as an acute trill(Lutz, 1954; Izecksohn and Carvalho-e-Silva,2001), but unfortunately we were unable to findany recordings of their calls at the collectionsconsulted (WCAB, ZUEC, and UFRJ) or withcolleagues herpetologists.

Larval Definition (Fig. 10).—Based on 22 tad-poles (stages 25 to 41) and six metamorphs(stages 42 to 46) from Tijuca, state of Rio deJaneiro (MNRJ 23729), reared in captivity by B.Lutz. Larval definition follows Altig andMcDiarmid (1999), and the stages are identifiedfollowing Gosner (1960). All the qualitativecharacters studied do not vary among thetadpoles. The characteristics presented belongto a single stage-36 individual. The measure-ments taken were total length (TL); body length(BL); greatest body width (GBW), measuredbehind the eyes in dorsal view; greatest bodyheight (GBH), measured at the plane of thecenter of the coiled intestine; tail length (TAL);tail musculature height (TMH); tail musclewidth (TMW); maximum tail height (MTH);dorsal (DFH) and ventral (VFH) fin height,measured at 6mm from the body terminus; andventral internarial distance (IND); interorbital

distance (IOD); eye diameter (ED); distancebetween nostril and spiracle aperture (DNS);and length of the upper (LUJ) and lower (LLJ)jaw sheaths.

Body depressed (GBH 78% of GBW), ellipticalin dorsal view, oval in lateral view; snoutbroadly rounded in dorsal and ventral views,oval in lateral view; eyes located dorsally,oriented dorsolaterally; nares dorsolateral withrounded apertures, located twice as close to thetip of snout than to the eyes; spiracle single,lateral, sinistral, aperture lateral; vent tubemedial with lateral displacement, aperturedextral; tail long (TAL twice as long as BL); tailfins low (DFH plus VFH 47% of MTH); ventralfin near parallel to tail musculature, dorsal fin atan angle of 30u to tail musculature; tail tipnarrowly rounded, forming an internal angle ofapproximately 60u; origin of dorsal fin 0.5 mmanterior to the body terminus; origin of ventralfin 0.6 mm posterior to the junction of the bodywall and the ventral margin of the tail muscle;oral disc transversally elliptical, positionedanteroventrally, forming an angle of about 45uwith the midsagittal plan; lateral emarginationpresent; anterior labium with five papillae ineach side, separated by a large dorsal gap;posterior labium with 24 papillae with no gap;papillae in a uniserial row, with pointed tip; twostraight labial ridges at the anterior labium, thefirst with a central gap; three straight labialridges at the posterior labium, without gaps;each labial ridge with a single row of labialteeth; dental formula 2(1)/3.

The preserved larvae are faded, but somebrown reticulations over the body that becomerounded blotches toward the end of the tail arestill visible. Izecksohn and Carvalho-e-Silva(2001) described the life color pattern of larvaeas body blackish with a white spot at the end ofthe snout.

Metamorphosis appears to take place withindividuals measuring a SVL 7.4 6 0.09 mm onaverage (N 5 6), almost half the mean size ofadults. The hands of metamorphs presentsupranumerary tubercles, one at the base offinger II, one at the base of finger III, and one atthe base of finger IV. Dorsal color pattern ofhands, thighs, and shanks are already visiblebut not as contrasting as verified in adults.Although faded it is possible to see a moreintense pigmentation laterally on the body fromthe tip of snout to the inguinal region. Thetympanum is not visible.

Measurements of the Larvae Described (mm).—TL 23.3; BL 7.8; GBW 4.5; GBH 3.5; TAL 16.2;TMH 2.1; TMW 2.4; MTH 3.2; DFH 0.8; VFH 0.7;IND 1.7; IOD 2.1; ED 0.8; DNS 4.3; LUJ 0.8; LLJ0.4.


Natural History.—Allobates olfersioides is di-urnal and lives on the forest floor. Whendisturbed, it immediately jumps and dives intothe forest litter (Izecksohn and Carvalho-e-Silva,2001). The tadpoles hatch in humid terrestrialnests and are carried by their parents to puddlesof small rivulets on the forest floor where theyfeed until metamorphosis (Lutz, 1954; Boker-mann, 1967; Izecksohn and Carvalho-e-Silva,2001). Two nursing frogs captured at Una(MZUSP 132191, 132202) and two from Ilheus(MZUSP 93872–73), Bahia were examined andfound to be male. They carried tadpoles atGosner stage 25 agreeing with reports for otherdendrobatids (Coloma, 1995; Junca, 1998; Cald-well et al., 2002b). Unfortunately, we haveinformation on the number of tadpoles carriedfor only one of the specimens from Una (12).The frogs from Ilheus were in a bottle with 22tadpoles. Previous reports on Allobates foundeight to 48 tadpoles being carried by the parent(Caldwell et al., 2002b). We found 14 of the 20females dissected from Tijuca, municipality ofRio de Janeiro, state of Rio de Janeiro withmature eggs in the ovaries. The females hada maximum of 11 mature eggs (about 1.5 mmeach) and mean of 8 6 1.4. Based on these data,

we suggest that the tadpoles from Ilheusbelonged to two individuals. The presence ofmature eggs and developing follicles togetherindicate that females reproduce more than oncein the reproductive period, as observed forother species of Aromobatidae (Junca, 1998).Gravid females were observed in samples fromIlheus (MZUSP 93874–913) in August; fromTijuca (MZUSP 93914–94021) in February, Au-gust, and September; and from Represa RioGrande, State of Rio de Janeiro (MZUSP 93664–665) in April, suggesting reproduction through-out the year.

Distribution.—Allobates olfersioides occurs incoastal forests of the Atlantic Domain from sealevel to about 1000 m a.s.l. The species can befound from the north of the state of Alagoas tothe south of the state of Rio de Janeiro (between08u549S and 23u009S latitudinal degrees), witha probable occurrence in Parque Estadual doRio Doce, State of Minas Gerais.

DISCUSSION

The lack of information on the natural historyand molecular phylogeography of Allobates fromthe Atlantic Forest populations are important

FIG. 10. Tadpole of Allobates olfersioides (MNRJ 23729, Tijuca, Rio de Janeiro, state of Rio de Janeiro): (A)lateral and (B) dorsal views, and (C) detail of oral disc, ventral view.


barriers to obtaining a better picture of itsdifferentiation. Data on vocalization are anecdot-al, and reported declines in the states of EspıritoSanto and Rio de Janeiro (Weygoldt, 1989;Izecksohn and Carvalho-e-Silva, 2001) render itdifficult to obtain vocalizations and tissue sam-ples, delaying further discussions on possiblegeographical call and molecular variation.

As far as we know, the distribution ofAllobates olfersioides seems to be unrelated toany hydrographic basin, and no river in thecoastal region of Brazil appears to be a geo-graphic barrier. The presence of continuouscharacter gradients suggests uninterruptedgene flow between adjacent populations, atleast until recently. Other amphibian popula-tions widely distributed in the coastal area ofBrazil seem to present similar patterns ofgeographic variation. Pombal and Haddad(1992); studying populations of Phyllomedusaburmeisteri baihana and P. b. burmeisteri from thestates of Rio de Janeiro, Espırito Santo, andBahia; found a continuum in the color pattern ofthe posterior surface of the thigh of theindividuals, with more complex character fre-quency changes at the state of Espırito Santo(Silva-Filho and Junca [2006] recently attributedspecies status to these subspecies, but withoutconsidering variation in the state of EspıritoSanto). A similar pattern was observed byJackson (1978) while studying the pattern ofcharacter variation of lizards of the genusEnyalius (Leiosauridae). Again, the populationsfrom the state of Espırito Santo were thosepresenting the largest amount of variation(Jackson, 1978) compared to that along a north-south transect.

These congruent geographical patterns maybe a hint of historical events that occurred fromthe southern state of Bahia to the northern stateof Rio de Janeiro. It may be caused by thecontinuity of forest habitats along the coastbefore deforestation or by a residual effect ofa complex introgression caused by the floodingof the habitats along the coast prior to the lastmarine transgression (Suguio and Nogueira,1999). Molecular phylogeographic studiesmight provide much needed evidence for thediscussion of these hypotheses.

Acknowledgments.—We are indebted to thefollowing colleagues for the loan of specimens:E. Freire, E. M. Goncalves, F. Junca, F. M. Souza,H. Zaher, J. Jim, J. Pombal, M. Soares, O. L.Peixoto, S. P. Carvalho-e-Silva, U. Caramaschi,and W. R. Heyer. We thank Hussam Zaher andC. Castro-Mello for allowing the access andlogistics at the MZUSP; P. E. Vanzolini forproviding important data from specimens fromW. C. A. Bokermann collection; V. R. Morales for

providing literature, S. P. Carvalho-e-Silva and E.Izecksohn, for sharing their knowledge onAllobates populations at the state of Rio deJaneiro, G. Skuk for the picture of the livespecimen. We also thank the Brazilian environ-mental organization IBAMA, for collection per-mits. P. Narvaes, D. Pavan, and T. Grant read andcriticized the manuscript. This paper is a resultfrom the project ‘‘Revisao sistematica das espe-cies de Colostethus da Mata Atlantica’’ supportedby FAPESP-Fundacao de Amparo a Pesquisa doEstado de Sao Paulo process (97/13503–6) asfulfillment to the requirement of the master’sdegree in zoology by the Instituto de Biociencias,Universidade de Sao Paulo, Sao Paulo, Brazil.

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Accepted: 9 May 2007.

APPENDIX 1

Specimens Examined

Allobates brunneus: Brazil, Mato Grosso, Chapadados Guimaraes: MZUSP 121875–82. Allobates goianus:Brazil, Goias, Chapada dos Veadeiros: MZUSP 76652(holotype), 73706, 76651 (paratypes). Allobates march-esianus: Brazil, Amazonas, Reservas INPA–WWF:MZUSP 67823–25. Allobates masniger: Brazil, Para,Parque Nacional da Amazonia: MZUSP 69166–67(paratypes); SW of Itaituba, Rio Tapajos: MZUSP69168–71 (paratypes). Allobates olfersioides: BRAZIL,Alagoas, Maceio: MUFAL 98735–36, 98739–40; Man-gabeiras: MZUSP 73823–30 (A. alagoanus, paratypes),73707 (A. alagoanus holotype), 78191–78197; Manimbu:MZUSP 11994, 11997–33; Murici: MZUSP 76432; NovoDino: MNRJ 9910; Rio Largo: MUFAL 930809–810;Bahia, Cumuruxatiba: MZUSP 59433; Elısio Medrado:UEFS 81; Ilheus: MNRJ 23767–70, 23728, MZUSP81133, 93874–913, 93872–73; Porto Seguro: MZUSP63195–97, 126361–65, UFPB 866150, 866153–54, 866193,866196–97, 866212–13; Sao Jose: MZUSP 63495; Una:MZUSP 63719–21, 125848, 132188–206; Urucuca:MZUSP 26916–18, 132187; Espırito Santo, BaixoGuandu: MZUSP 35674; Linhares: MZUSP 93871;Refugio Sooretama: MZUSP 76628–76629 (A. capixaba,paratype), 73752 (A. capixaba, holotype); Santa Teresa:MZUSP 53559–60; USNM 208141, 208147, 200452–53;Rio de Janeiro, Angra dos Reis: MNRJ 5094; Araruama:MNRJ 4115, 15398–406, 4143, 15431–39; Imbiara: MNRJ23716–19; Itaguaı: EI 4321–56, JJ 7787, 7792–96, ZUEC2741–42, MNRJ 23724–27; Represa Rio Grande: MZUSP73714 (A. carioca, paratype), 73753 (A. carioca, paratype),76653 (A. carioca, holotype), 93651–65, 93671; SacraFamılia do Tingua: EI 8598–600, JJ 7788–91, ZUEC10824; Serra da Piedade: MNRJ 23730–34; MZUSP9801–9803; Teresopolis: MNRJ 23721–23, MZUSP


53448–51, USNM 208396–208401; Tijuca: JJ 224–231,MZUSP 76418, 76728–29, 93624–49, 93666–767, 93914–94377; Tingua: EI 4290, 4306; Sergipe, Areia Branca:MZUSP 88954. Allobates sumtuosus: Brazil, Para, Re-serva Biologica Rio Trombetas: MZUSP 69157–60(paratypes). Anomaloglossus beebei: French Guian, SautMais: MZUSP 58230. Anomaloglossus degranvillei:French Guian, Saul: MZUSP 58231. Anomaloglossusstepheni: Brazil, Amazonas, Manaus: MZUSP 64569(holotype), 64570 (paratype).

APPENDIX 2

Gazetteer (State, code number used at Fig. 1, specificlocality, approximate coordinate).—Alagoas, 4, Maceio:09u409S, 35u419W; 6, Mangabeiras: 09u569S, 36u059W; 5,Manimbu: 09u529S, 36u099W; 2, Murici: 09u189S,35u569W; 1, Novo Dino: 08u549S, 35u389W; 3, Rio

Largo: 09u299S, 35u509W; Bahia, 15, Cumuruxatiba:17u069S, 39u119W; 8, Elısio Medrado: 12u569S, 39u319W;10, Ilheus:14u489S, 39u039W; 14, Porto Seguro: 16u269S,39u059W; 11, Sao Jose: 15u099S, 39u189W; 12, Jussari,Serra do Teimoso: 15u099S, 39u319W; 13, Una: 15u179S,39u059W; 9, Urucuca: 14u339S; 39u189W; Espırito Santo,18, Baixo Guandu: 19u319S, 41u029W; 17, Linhares:19u239S, 40u049W; 16, Refugio Sooretama: 19u009S,40u009W; 19, Santa Teresa: 19u569S, 40u379W; Rio deJaneiro, 29, Angra dos Reis: 23u009S, 44u199W; 20,Araruama: 22u529S, 42u209W; 22, Imbiara: 22u429S,42u379W; 26, Itaguaı: 22u489S, 43u419W; Rio de Janeiro,24, Represa Rio Grande: 22u559S, 43u259W; 28, SacraFamılia do Tingua: 22u309S, 43u379W; Rio de Janeiro,27, Serra da Piedade: 23u009S, 43u399W; 21, Teresopo-lis: 22u269S, 42u299W; Rio de Janeiro, 25, Tijuca:22u489S, 43u329W; 23, Tingua: 22u339S, 43u269W;Sergipe, 7, Areia Branca: 10u469S, 37u199W.


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Taxonomic Review of Allobates (Anura, Aromobatidae) from ... Rodrigues_2007_Taxonomic Review... · Taxonomic Review of Allobates (Anura, Aromobatidae) from the Atlantic Forest, Brazil