Variation in Oophaga pumilioSALAMANDRA            44             4           225-234        Rheinbach, 20 November 2008   ...
Abel Batista & Gunther Köhler                                                      ers of naris; snout length (SL) from an...
Variation in Oophaga pumilioFig. 2. Oophaga pumilio from Cerro Tebata (Loca-   Fig. 3. Oophaga pumilio from Isla Bastiment...
Abel Batista & Gunther KöhlerTab. 1. Size and proportions of adult Oophaga pumilio from localities 1-10 as shown in Figure...
Variation in Oophaga pumilio   Cerro Brujo         Río Uyama         Quebrada la Gloria       Kusapin        Rio Krikamola...
Abel Batista & Gunther KöhlerFig. 5. Oophaga pumilio from Isla Popa (Locality   Fig. 7. Oophaga pumilio from Cerro Brujo (...
Variation in Oophaga pumilioFig. 8. Oophaga pumilio from Río Uyama (Locality   Fig. 10. Oophaga pumilio from Kusapin (Loca...
Abel Batista & Gunther Köhlerto Parrot Green (260) or with Jet Black (89)                       Discussionflecks; upper sur...
Variation in Oophaga pumiliogas pers. comm). In accordance with this,          Fuenmayor, Panama City, provided valuable a...
Abel Batista & Gunther KöhlerGuyer, G. & M.A. Donnelly (2005): Amphib-               Schmidt, O. (857): Diagnosen neuer F...
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  1. 1. Variation in Oophaga pumilioSALAMANDRA 44 4 225-234 Rheinbach, 20 November 2008 ISSN 0036-3375 Variation in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) in western Panama Abel Batista & Gunther KöhlerAbstract. We studied variation in colour pattern and morphology in ten populations of Oophaga pumiliofrom the Province of Bocas del Toro, western Panama. Our field work documented several distinct phe-notypes of colouration and pattern that are highly correlated geographically. On the contrary, variationin the morphometric characters studied is small among these populations. There are examples of distinctpopulations in close proximity without any obvious physiographic barriers that show no evidence of hy-bridization. We interpret these abrupt transitions as evidence of lack of or minimal gene flow.Key words. Anura, Dendrobatidae, Oophaga pumilio, colouration, distribution, morphology, variation,Panama.Resumen. Estudiamos la variación y morfología de varias poblaciones de Oophaga pumilio en la Pro-vincia de Bocas del Toro, oeste de Panamá. Nuestro trabajo de campo documenta varios fenotipos dis-tintivos de coloración y patrones que están altamente correlacionados geográficamente. Por el contrario,la variación morfométrica de caracteres estudiados es pequeña entre estas poblaciones. Hay ejemplosde distintivas poblaciones cercanas sin aparentes barreras fisiográficas que no muestran evidencias dehibridación. Interpretamos estas transiciones abruptas como una evidencia de ausencia o de reducidoflujo genético. Introduction to the species here described as typographi- cus. Until a comparison of the types of pumil-The description of Dendrobates pumilio O. io and typographicus by a competent observerSchmidt, 857 (strawberry poison frog) was proves them to be identical, I propose to usebased on material from the trail between Bo- the Keferstein name” for Costa Rican spec-cas del Toro and Volcan Chiriquí (5000-7000 imens. Unfortunately, the type of D. pumil-Polish feet = 50-60 masl), Province of Bo- io, originally in the Krakow Museum, is lostcas del Toro, Panama, collected by Josef v. (Savage 968) so that direct comparisons ofWarszewicz. Schmidt (858) provided a type material can not be done. An additionalbrief colour description and an illustration of nominal species, D. galindoi Trapido, 953the dorsal view of this species, now in the ge- was described from the Province of Bocas delnus Oophaga (Grant et al. 2006). Two more Toro, Panama, a name placed in the synony-names were proposed for this species, dif- my of O. pumilio by subsequent authors (Sav-fering in colouration and currently referred age 968, Silverstone 975).to as synonyms of O. pumilio: Dendrobates In accordance with most previous authors,typographus Keferstein, 867 (type local- Daly & Myers (967) treated all populationsity: Costa Rica) and D. ignitus Cope, 874 of strawberry poison frog from Nicaragua(type locality: Machuca, Nicaragua). Taylor to northwestern Panama as a single species,(952) examined material from Costa Rica O. pumilio, suggesting that it is a polymor-and was uncertain regarding the identity of phic species. These authors and Summers etO. Schmidt’s pumilio and argued that the il- al. (997, 2003) provided notes on the colourlustration in O. Schmidt (858) (Tafel II, Fig. variation in O. pumilio. We go along with the3) “could scarcely be regarded as belonging taxonomic opinion by Daly & Myers (967)© 2008 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT)http://www.salamandra-journal.com 225
  2. 2. Abel Batista & Gunther Köhler ers of naris; snout length (SL) from anterior edge of eye to tip of snout; forearm length (FAL) from proximal edge of palmar tuber- cle to outer edge of flexed elbow; hand length (HAL) from proximal edge of palmar tuber- cle to tip of third finger; shank length (SHL) from outer edges of flexed knee to heel; foot length (FL) from proximal edge of outer met- atarsal tubercle to tip of fourth toe; width of third finger (3FW) at penultimate phalanx just anterior to disc; width of disc of thirdFig. 1. Map showing the collecting sites of Oopha-ga pumilio in this study. Localities are: (1) Cerro finger (3FD) at greatest width; width of thirdTebata; (2) Bastimentos Island; (3) Solarte Island; toe (3TW) at penultimate phalanx just ante-(4) Popa Island; (5) Loma Partida Island; (6) Cerro rior to disc; width of disc of third toe (3TD)Brujo; (7) Río Uyama; (8) Quebrada La Gloria; (9) at greatest width; width of fourth toe (4TW)Kusapin; (10) Río Krikamola. at penultimate phalanx just anterior to disc; width of disc of fourth toe (4TD) at greatestand Summers et al. (997, 2003). However, width; body width (BW) at greatest width ofwe claim the lack of a detailed study of the body; tympanum diameter (TD) horizontalgeographic variation of phenotypic patterns tympanum diameter, based on a estimatedand morphometrics in O. pumilio. Here we circular tympanum.report upon the results of our study on the Colour descriptions refer to live speci-variation in colouration and morphometrics mens if not stated otherwise. The capitalizedin O. pumilio in western Panama. colours and colour codes (the latter in paren- theses), as applied here, are those of Smithe (975-98). In descriptions of colouration in Materials and methods preservative we used the terminology pro- posed by Grant et al. (2006) and comparedDescriptions of populations are based on our material to the illustrations therein (theirspecimens examined by the authors. Meas- Figs. 35, 46, 47, 48, 49).urements and colouration data were takenon 87 specimens of Oophaga pumilio from 0localities (see Fig. ). Abbreviations for muse- Resultsum collections follow those of Leviton et al. Description of the populations examined(985), except MHCH (Museo Herpetologicode Chiriquí, Davíd, Panama). Cerro Tebata (Locality 1 in Fig. 1) All measurements were taken on adult C olourat ion in life: Dorsal ground col-specimens (males and females not distin- our varies from Spectrum Orange (7), Scar-guished) using digital calipers and were let (4) to Yellowish Olive Green (50); dor-rounded to the nearest 0.0 mm. The fol- sum usually uniform, rarely with black spots;lowing measurements were taken (with ab- flank coloured as dorsum; dorsal surfaces ofbreviations indicated): length from snout to hind limbs Dusky Brown (9), Jet Black (89),vent (SVL); head length (HL) was measured rarely same colour of dorsum; venter Orangediagonally from angle of jaw to tip of snout; Yellow (8), Scarlet (4) or same colour ofhead width (HW) between angles of jaws; dorsum (Fig. 2).eye length (EL) from anterior to posterior C olour af ter f ive mont hs in pres er v -edge; eye to naris distance (END) from ante- at ive: Dorsal surfaces of head, body darkrior edge of eye to posterior corner of naris; greyish, with black flecks; venter pale grey;internaris distance (IND) between cent- limbs black. 226
  3. 3. Variation in Oophaga pumilioFig. 2. Oophaga pumilio from Cerro Tebata (Loca- Fig. 3. Oophaga pumilio from Isla Bastimentoslity 1 in Fig. 1), Bocas del Toro, Panama. (Locality 2 in Fig. 1), Bocas del Toro, Panama.R emarks: Individuals from Costa Rica andNicaragua mostly agree well with this descrip-tion. However, it has been reported that inrare instances individuals from La Selva canbe uniform dark blue with black spots (Guy-er & Donnelly 2005). Based on our material,individuals from northern populations (e.g.,from Nicaragua) are smaller than those fromsouthern Costa Rica and western Panama.Specimens examined: PANAMA: Bo-cas del Toro: Cerro Tebata, 9º33’37.3’’N, 82º5’ Fig. 4. Oophaga pumilio from Isla Solarte (Locality8.8’’W, 480 m: MHCH 578, SMF 86595-605. 3 in Fig. 1), Bocas del Toro, Panama.Isla Bastimentos (Locality 2 in Fig. ) uals from Red Frog and Wizard Beach have aC olourat ion in life: Dorsal ground col- uniform red dorsum or with dark spots, ven-our varies from Opaline Green (62D), Pale ter same colour as dorsum or dirty white. In-Horn Colour (92), Flesh Ocher (32D) and dividuals from Bahia Honda are orange uni-Flame Scarlet (5) to Spectrum Red (), usu- form.ally with Dusky Brown (9) blotches (some Sp e cimens examine d: PANAMA: Bocasindividuals immaculate); venter immaculate del Toro: Bastimentos Island: 09º2’4.’’N,dirty white with a suggestion of Pale Horn 82º’ 26.5’’W, 0 m: SMF 85359-36, 8663-34,Colour (92) or of Opaline Green (62D), or 86636.Pearl Gray (8) (Fig. 3).C olour af ter ten mont hs in pres er va - Isla Solarte (Locality 3 in Fig. )t ive: Dorsal surfaces of head, body and legs C olourat ion in life: Dorsum Flamewhite to pale grey with brown blotches; ven- Scarlet (5) or Spectrum Orange (7), immac-ter almost white. ulate; rarely with small Dusky Brown (9)R emarks: We collected our series on Bas- spots; venter same as dorsum, without darktimentos Island in the area of the cemetery sp ots (Fig. 4).in the late afternoon during light rain. The Sp e cimens examine d: PANAMA: Bocasfrogs were active on the ground and on tree del Toro: Solarte Island, Hospital Point, 0 m:trunks up to .5 m above the ground. Individ- SMF 85349-58. 227
  4. 4. Abel Batista & Gunther KöhlerTab. 1. Size and proportions of adult Oophaga pumilio from localities 1-10 as shown in Figure 1. Eachrange is followed by the mean and standard deviation in parentheses; for abbreviations see text.Character Cerro Tebata Bastimentos Solarte Island Popa Island Loma Partida Island 2.06 -23.89 mm 8.03-22.45 mm 7.33-9.56 mm 4-7 mm 5.98-9.84 mmSVL (22.09 ± 0.90) (20.306 ± .5) (8.8 ± 0.66) (5.7 ± 0.84) (8.6 ± .342) 0.28-0.334 0.26-0.38 0.27-0.294 0.270-0.306 0.26-0.30HW/SVL (0.30 ± 0.08) (0.283 ± 0.05) (0.28 ± 0.007) (0.286 ± 0.0) (0.28 ± 0.05) 0.272-0.324 0.265-0.34 0.282-0.34 0.269-0.300 0.28-0.32HL/SVL (0.3 ± 0.04) (0.283 ± 0.04) (0.3 ± 0.02) (0.286 ± 0.008) (0.3 ± 0.04) 0.420-0.500 0.48-0.5 0.43-0.5 0.408-0.463 0.43-0.48SHL/SVL (0.46 ± 0.02) (0.45 ± 0.02) (0.455 ± 0.02) (0.43 ± 0.020) (0.45 ± 0.07) 0.375-0.452 0.273-0.44 0.332-0.404 0.39-0.386 0.373-0.427FL/SVL (0.407 ± 0.02) (0.37 ± 0.036) (0.37 ± 0.022) (0.354 ± 0.07) (0.4 ± 0.02) 0.040-0.067 0.04-0.06 0.035-0.06 0.045-0.063 0.043-0.058TD/SVL (0.047 ± 0.008) (0.05 ± 0.0) (0.05 ± 0.0) (0.053 ± 0.006) (0.05 ± 0.006) 0.3-0.40 0.-0.4 0.04-0.22 0.23-0.5 0.-0.23EL/SVL (0.24 ± 0.007) (0.22 ± 0.0) (0.5 ± 0.005) (0.34 ± 0.00) (0.4 ± 0.0)SL/SVL 0.087-0.00 0.077-0.0 0.08-0. 0.082-0.04 0.085-0.05 (0.090 ± 0.003) (0.086 ± 0.007) (0.087 ± 0.005) (0.09 ± 0.006) (0.093 ± 0.006) 0.07-0.23 0.-0.3 0.02-0.7 0.095-0.26 0.04-0.26IND/SVL (0.6 ± 0.005) (0.5 ± 0.0) (0. ± 0.005) (0.2 ± 0.008) (0.6 ± 0.008) 0.243-0.285 0.233-0.33 0.25-0.27 0.244-0.28 0.258-0.282FAL/SVL (0.258 ± 0.04) (0.262 ± 0.022) (0.256 ± 0.005) (0.26 ± 0.02) (0.27 ± 0.0) 0.232-0.275 0.25-0.3 0.23-0.27 0.84-0.247 0.244-0.27HAL/SVL (0.250 ± 0.04) (0.27 ± 0.04) (0.25 ± 0.02) (0.222 ± 0.08) (0.258 ± 0.007) .44-2.40 .6-2.34 .6-2.452 .486-2.000 .33-2.053FD/3FW (.83 ± 0.273 ) (2.07 ± 0.85) (2.076 ± 0.264) (.740 ± 0.82) (.74 ± 0.246)3TD/3TW .22-.8 .2-.69 .8-.69 .000-.30 .06-.42 (.45 ± 0.82) (.4 ± 0.30) (.47 ± 0.45) (.046 ± 0.092) (.26 ± 0.6) .35-.86 .3-.84 .264-.947 .03-.862 .074-.5624TD/4TW (.55 ± 0.4) (.5 ± 0.8) (.604 ± 0.2) (.364 ± 0.243) (.36 ± 0.2) 0.400-0.478 0.4-0.52 0.405-0.5 0.346-0.46 0.37-0.5BW/SVL (0.437 ± 0.025) (0.46 ± 0.04) (0.45 ± 0.035) (0.395 ± 0.035) (0.458 ± 0.04)Isla Popa (Locality 4 in Fig. ) Isla Loma Partida (Locality 5 in Fig. )C olourat ion in life: Dorsal ground col- C olourat ion in life: Dorsal surfacesour Greenish Olive (49); legs Ultra Marine of head, body and limbs Olive Green (48),Blue (70A); posterior thigh Yellowish Ol- Brownish Olive (29), Indigo (73) or Tur-ive Green (50); venter immaculate Sky Blue quoise Green (64), uniform or with Jet Black(68C) (Fig. 5). (89) flecks or spots; lateral surfaces of headR emarks: Our series was collected in a ma- and body Light Sky Blue (68D); upper sur-ture secondary forest on Popa Island, all in- faces of limbs same colour as dorsum ordividuals found on the forest floor during a darker with Jet Black (89) spots; lips colouredrainy afternoon. as ventral surfaces; tympanum Jet Black (89)Sp e cimens examine d: PANAMA: Bocas or same colour as dorsum; ventral surfaces ofdel Toro: Popa Island, 9º3’4’’N, 82º08’28’’W, head, body and limbs uniform Light Sky Blue5 m: SMF 85338-48. 228
  5. 5. Variation in Oophaga pumilio Cerro Brujo Río Uyama Quebrada la Gloria Kusapin Rio Krikamola 7.2-8.88 mm 6.86-9.78 mm 6.6-8.84 mm 4.62-6.42 mm 6.22-8.84 mm (7.78 ± 0.56) (7.96 ± .02) (7.66 ± 0.93) (5.4 ± 0.55) (7.26 ± 0.76) 0.275-0.3 0.27-0.33 0.265-0.32 0.264-0.306 0.26-0.3 (0.285 ± 0.0) (0.286 ± 0.04) (0.29 ± 0.02) (0.288 ± 0.04) (0.284 ± 0.05) 0.277-0.32 0.274-0.39 0.283-0.34 0.268-0.337 0.283-0.33 (0.284 ± 0.0) (0.295 ± 0.03) (0.3 ± 0.04) (0.304 ± 0.08) (0.306 ± 0.07) 0.46-0.466 0.424-0.475 0.44-0.48 0.425-0.466 0.43-0.466 (0.44 ± 0.06) (0.45 ± 0.08) (0.46 ± 0.07) (0.45 ± 0.03) (0.442 ± 0.08) 0.35-0.46 0.35-0.42 0.344-0.394 0.332-0.392 0.363-0.422 (0.37 ± 0.022) (0.386 ± 0.024) (0.38 ± 0.02) (0.364 ± 0.02) (0.385 ± 0.07) 0.044-0.055 0.05-0.062 0.037-0.057 0.05-0.067 0.045-0.073 (0.049 ± 0.005) (0.055 ± 0.003) (0.044 ± 0.008) (0.06 ± 0.005) (0.053 ± 0.0) 0.2-0.42 0.5-0.4 0.05-0.25 0.3-0.45 0.05-0.35 (0.28 ± 0.006) (0.28 ± 0.008) (0.5 ± 0.0) (0.25 ± 0.0) (0.5 ± 0.0) 0.085-0. 0.082-0. 0.08-0.094 0.08-0. 0.077-0. (0.09 ± 0.005) (0.09 ± 0.003) (0.089 ± 0.006) (0.087 ± 0.006) (0.087 ± 0.007) 0.08-0.28 0.0-0.2 0.096-0.8 0.-0.2 0.-0.5 (0.2 ± 0.006) (0. ± 0.006) (0. ± 0.009) (0. ± 0.006) (0. ± 0.006) 0.254-0.265 0.237-0.275 0.248-0.277 0.238-0.27 0.25-0.28 (0.26 ± 0.004) (0.26 ± 0.05) (0.26 ± 0.0) (0.256 ± 0.0) (0.266 ± 0.0) 0.235-0.267 0.247-0.288 0.248-0.272 0.23-0.25 0.245-0.27 (0.253 ± 0.02) (0.264 ± 0.02) (0.262 ± 0.0) (0.24 ± 0.006) (0.255 ± 0.007) .49-2.022 .42-2.25 .545-.744 .5-3.0 .36-2.0 (.822 ± 0.2) (.82 ± 0.257) (.62 ± 0.76) (2.5 ± 0.474) (.684 ± 0.24) .0-.65 .06-.9 .84-.625 .06-.5 .0-.4 (.406 ± 0.83) (.4 ± 0.245) (.42 ± 0.8) (.22 ± 0.35) (.2 ± 0.32) .7-.84 .25-.64 .-.53 .05-.7 .093-.674 (.5 ± 0.24) (.42 ± 0.3) (.344 ± 0.55) (.4 ± 0.77) (.362 ± 0.66) 0.34-0.467 0.367-0.464 0.4-0.47 0.393-0.462 0.38-0.473 (0.43 ± 0.03) (0.428 ± 0.032) (0.442 ± 0.027) (0.423 ± 0.026) (0.42 ± 0.028)(68D) or Robin’s Egg Blue (93) male with with Light Sky Blue (68D) throat, malesdarker throat (Fig. 6). with darker throat (Fig. 7).R emarks: These frogs were very abundant C olour af ter ten mont hs in pres er v -in some areas of Loma Partida Island. at ive: Dorsal surfaces of head, body solidSp e cimens examine d : PANAMA: Bocas dark greyish; venter grey blue; limbs black.del Toro: Loma Partida Island, 9º0’37.2’’N, R emarks: The series was collected during82º2’30.7’’W, 20 m: SMF 8660-6. daytime in a mature secondary forest and in disturbed areas. Most frogs were found onCerro Brujo (Locality 6 in Fig. ) the ground, one was observed climbing a treeC olourat ion in life: Dorsal surfaces of ten meters above the ground.head, body and limbs Indigo (73); Vandike Sp e cimens examine d: PANAMA: Bo-Brown (22), uniform or with Jet Black (89) cas del Toro: Cerro Brujo, 9º’6.4’’N,spots; venter Venetian Blue (68B); females 82º’25.4’’W, 0 m: SMF 85329-37. 229
  6. 6. Abel Batista & Gunther KöhlerFig. 5. Oophaga pumilio from Isla Popa (Locality Fig. 7. Oophaga pumilio from Cerro Brujo (Loca-4 in Fig. 1), Bocas del Toro, Panama. lity 6 in Fig. 1), Bocas del Toro, Panama. two meters above the ground. At Río Uyama we observed one population of O. pumilio with red dorsum with black spots and blue legs in very close vicinity (more or less 800 m) to the black and white populations de- scribed here. Sp e cimens examine d: PANAMA: Bocas del Toro: Río Uyama; 9º08’55’’N, 82º9’28’’W, 35 m; SMF 8532-28. Quebrada la Gloria (Locality 8 in Fig. ) C olourat ion in life: Dorsal surfaces of head and body Orange Yellow (8) or OliveFig. 6. Oophaga pumilio from Isla Loma Partida Yellow (52), rarely Light Sky Blue (68D),(Locality 5 in Fig. 1), Bocas del Toro, Panama. with Sepia (9 or 29) or Sayal Brown (223C) blotches that have the tendency to formRío Uyama (Locality 7 in Fig. ) stripes, dorsal and ventral surfaces of limbsC olourat ion in life: Dorsal ground as well as ventral surfaces of head and bodycolour dirty white with large Sayal Brown Light Sky Blue (68D) with Sepia (29) blotch-(223C) blotches that have the tendency to es; some individuals with ventral colour sameform stripes; venter Light Sky Blue (68D) as dorsum; males with darker throat (Fig. 9).with large black blotches. Sp e cimens examine d: PANAMA: BocasC olour af ter six mont hs in pre- del Toro: Quebrada la Gloria (9 km SW froms er vat ive: Dorsal surfaces of head, body Chiriquí Grande Road): SMF 86626-30.dark greyish to brown, with the white are-as scarcely visible; venter almost black with Kusapin (Locality 9 in Fig. )white blotches scarcely visible; limbs brown C olourat ion in life: Dorsal surfaces ofor black (Fig. 8). head and body uniform red; in SMF 86608R emarks: This series was collected in the frontal and parietal region suffused withlate morning at the egde and within a mature Pratt’s Rufous (40); upper surfaces of fore-secondary forest. Most frogs were found on limbs same colour as dorsum or brown; inthe ground, one was observed climbing a tree SMF 86609 forelimbs Scarlet (4), hands Raw 230
  7. 7. Variation in Oophaga pumilioFig. 8. Oophaga pumilio from Río Uyama (Locality Fig. 10. Oophaga pumilio from Kusapin (Locality 97 in Fig. 1), Bocas del Toro, Panama. in Fig. 1), Comarca Ngöbe Bugle, Panama.Fig. 9. Oophaga pumilio from Quebrada la Gloria Fig. 11. Oophaga pumilio from about 2 km SW(Locality 8 in Fig. 1), Bocas del Toro, Panama. mouth of Río Krikamola (Locality 10 in Fig. 1), Comarca Ngöbe Bugle, Panama.Umber (223) and fingers Sky Blue (68C) suf-fused with Raw Umber (223); dorsal surfacesof hind limbs same colour as upper surfaces C olour af ter six mont hs in pres er v -or darker; posterior surfaces of hind limbs at ive: Dorsal surfaces of head, body solidbrown to dark brown; lips Sky Blue (68C) dark greyish; venter almost black; limbs darkor coloured as dorsum; tympanum brown brown, regions of ellbows and knees brown.to black or same colour as dorsum; upper R emarks: The series was collected duringedge of tympanum usually bordered by a daytime on the floor in a mature secondarythin black stripe, about two times the tym- forest on a hill near Kusapin town.panum length; transition of colours in flanks Sp e cimens examine d: PANAMA: Co-(red to sky blue) with dark flecks in some in- marca Ngöbe Bugle: Kusapin, 9º0’54.9’’N,dividuals; ventral surfaces of head and body 8º53’22.’’W, 8 m: SMF 86584-94.and limbs uniform Sky Blue (68) in females,male with Medium Plumbeus (87) throat; Rio Krikamola (Locality 0 in Fig. )some specimens with dark flecks on dorsum C olourat ion in life: dorsal surfaces ofand flanks (Fig. 0). head and body uniform Leaf Green (46) 231
  8. 8. Abel Batista & Gunther Köhlerto Parrot Green (260) or with Jet Black (89) Discussionflecks; upper surfaces of limbs brown, BlueBlack (90) or same colour as dorsum; tym- Oophaga pumilio exhibits a distinctive pat-panum Jet Black (89) (usually only at lower tern of phenotypic variation which is highlypart) or same colour as dorsum; ventral sur- correlated geographically; even several phe-faces of head and body uniform Olive Yellow notypically distinct populations occur in(52) to Lime Green (59), usually with black sympatry or parapatry (Myers & Daly 983,spots, rarely with Sky Blue (68C) blotches; Summers et al. 2003, Saporito et al. 2007).male with darker throat; ventral surfaces of This includes examples of distinct popula-limbs grey blue or dark blue; lips coloured as tions in close proximity without any obviousventer (Fig. ). physiographic barriers that show no evidenceC olour af ter s e ven mont hs in pre - of hybridization in terms of colouration. Ins er vat ive: Dorsal surfaces of head and the area of Almirante an abrupt transition isbody solid dark greyish, some specimens found between a population consisting of akeep their black spots on dorsum and flecks black and white morph and red morph within the venter; limbs dark brown; venter grey. blue legs (Summers et al. 2003). In the area ofR emarks: The series was collected during Chiriquí Grande, three distinct phenotypesdaytime in a wetland forest, most specimens meet and abrupt transitions are reportedwere found climbing on trees at less than two between them: a population consisting of ameters above the ground. black and yellow morph, one with a red dor-Sp e cimens exam ine d: PANAMA: Co- sum and bluish flanks, and one with a greenmarca Ngöbe Bugle: Kankintú, Río Kri- ground colour (Myers & Daly 983, Sum-kamola, 8º58’38.6’’N, 8º55’0.7’’W, 7 m: SMF mers et al. 2003, this study). We can interpret86564-73. these abrupt transistions as evidence of lack of or remarkably reduced gene flow. Never- theless, minimal gene flow may exists since Comparisons at Pueblo Nuevo and at Chiriquí Grande we found individuals that appear to be interme-Our study documented several distinct phe- diate in colouration between those popula-notypes of colouration and pattern that are tions (Fig. 2).highly correlated geographically. On the con- Daly & Myers (967: 973) stated that fortrary, variation in morphometric characters O. pumilio “a static subspecies concept doesstudied is comparatively small among these not seem well suited to the dynamics of thepopulations. All morphometric characters situation”. However, the descriptions of theshow largely overlapping ranges among most phenotypes by these authors agree well withpopulations (Table ). In respect of the mor- our observations indicating that over thephometric characters examined, the Popa course of at least four decades the phenotypicpopulation (Locality 4 in Fig. ) is the most pattern had remained stable. The underlyingdistinctive one. In several characters (FAL/ mechanisms and barriers that prevent mix-SVL, HAL/SVL, 3TD/3TW) it has barely or ing of the phenotypes in the natural popula-non-overlapping ranges with the other popu- tions are poorly understood. It is known thatlations. The series from Bastimentos (Local- females of O. pumilio preferentially chooseity 2 in Fig. ) has the highest average values mates of their own colour morph (Siddiqifor FAL/SVL and HAL/SVL indicating that et al. 2004). Under captive conditions it hasthese frogs possess relative long hind feet and been observed that certain males of O. pumil-toes, although the ranges of these characters io, i.e. red with blue legs, display courtshipdo overlap to some degree with the other behavior (mating calls) only towards femalespopulations. of their own phenotype but not towards pop- ulations from Bastimentos Island (G. Var- 232
  9. 9. Variation in Oophaga pumiliogas pers. comm). In accordance with this, Fuenmayor, Panama City, provided valuable as-Siddiqi et al. (2004) provided evidence that sistance with acquisition of these permits. Meikethe colours displayed by the various colour Piepenbring, Botanisches Institut J.W. Goethe-morphs are effective visual signals, not only Universität, Frankfurt, provided logistic support for our studies in Panama and was instrumentalto potential predators but also to conspecif- in enabling Abel Batista and Marcos Ponce toics, making sympatric speciation driven by visit Frankfurt, for a three months research stay.sexual selection possible. Thus, colouration We thank Pedro Caballero (Director of Istitu-may be important in maintaining the iden- to de Ciencias Ambientales y Desarrollo Sosteni-tities of the various populations (Siddiqi et ble) and Boris E. Sanjur, Facultad de Cienciasal. 2004). Fisher’s runaway process of sexu- Naturales y Exactas de la Universidad Autónomaal selection is potentially an important force de Chiriquí (UNACHI), David, Panama, for theirgenerating character divergence between support. This paper is based on part upon workclosely related populations (Pomiankowski supported by the Deutscher Akademischer Aus- tauschdienst (DAAD) to Abel Batista and to& Iwasa 998). Captive cross-breeding ex- Gunther Köhler through the Partnership Pro-periments in O. pumilio, including several gram between the J.W. Goethe-Universität Frank-populations from the Bocas del Toro Islands furt am Main, and the Facultad de Ciencias Nat-and mainland, demonstrated that frogs of urales y Exactas der Universidad Autónoma dedifferent colours morphs can produce fer- Chiriquí (UNACHI), David, Panama. We aretile offspring. However, the survival rates of grateful to Alfred A. Schmidt for his financialoffspring were relatively low (Summers et support of this study. We thank Marcos Ponce,al. 2004). These observations are supported José Torres and Ramon Luna for field assist-by the fact that breeding success appeared ence, Rigoberto Lopez for guide and facilities at his farm at Cerro Tebata, Alicia Cáceres for herto be higher if only captive frogs from the support to Abel Batista in this work. We appre-same geographical origin were put together ciate the help and comments by Stefan Lötters(Salewski 2005). and two anonymous reviewers who significantly Perhaps in contrast to all these observa- improved this paper.tions, molecular genetic studies based onmtDNA data indicated that the studied pop-ulations are phylogenetically closely related Referencesand most probably are members of the samespecies (Summers et al. 997, 2003). Summers Cope, E.D. (874): Description of some species ofet al. (2004) suggested that colour pattern is reptiles obtained by Dr. John F. Bransford,under single locus control with dominance, assistant surgeon United States Navy, while at-whereas colouration may be under polygenic tached to the Nicaragua surveying expedition in 873. – Proc. Acad. Nat. Sci. Philadelphia,control, or may represent a single locus sys- 26: 64-72.tem with incomplete dominance. Most of the phenetically distinct popu- Daly, J.W. & C.W. Myers (967): Toxicity of pana- manian poison frogs (Dendrobates): some bi-lations described in the present study have ological and chemical aspects. – Science, 56:very restricted geographical distributions. Ig- 970-973.noring the distinctness of these populations Dunn, E.R. (93): New frogs fromPanama andpresents a latent threat to them in the light of Costa Rica. – Occ. Pap. Bost. Soc. Nat. Hist.,the ongoing alterations and contaminations 5: 385-40.of their habitats. Grant, T., D.R. Frost, J.P. Caldwell, R. Gagliardo, C.F.B. Haddad, P.J.R. Kok, D.B. Acknowledgements Means, B.P. Noonan, W.E. Schargel & W.C. Wheeler (2006): Phylogenetic systematics ofCollecting and export permits were provided dart-poison frogs and their relatives (Amphib-by Yariela Hidalgo, Autoridad Nacional del ia: Athesphatanura: Dendrobatidae) – Bull.Ambiente (ANAM), Panama City. Querube D. Amer. Mus. Nat. Hist., 299: -262. 233
  10. 10. Abel Batista & Gunther KöhlerGuyer, G. & M.A. Donnelly (2005): Amphib- Schmidt, O. (857): Diagnosen neuer Frösche des ians and reptiles of La Selva, Costa Rica, and zoologischen Cabinets zu Krakau. – Sitzber. the Caribbean slope: a comprehensive guide. Math.-Natwiss. Akad. Wiss. Wien, 24: 0-5. – Berkeley and Los Angeles California (Uni- Schmidt, O. (858): Deliciae herpetologicae Mu- versity of California press): 299 pp. sei Zoologici Cracoviensis. Beschreibung derJungfer, K.H., P. Weygoldt & N. Juraske Im K.K. Museum zu Krakau befindlichen, Von (996): Dendrobates vicentei, ein neuer Pfeilg- J. v. Warszewicz in Neu-Granada und Boliv- iftfrosch aus Zentral-Panama. – Herpetofauna, ia gesammelten ungeschwänzten Batrachier. 8(03): 7-26. – Denkschr. Math.-Natwiss. Cl. Kaiserl. Akad.Leviton, A.E., R.H. Gibbs Jr., E. Heal & C.E. Wiss. Wien, 4: 237-258. Dawson. (985): Standards in Herpetology and Siddiqi, A., T.W. Cronin, E.R. Loew, M. Voro- Ichthyology: part I. Standard symbolic codes byev & K. Summers (2004): Interspecific and for institutional resource collections in herpe- intraspecific views of colour signals in the tology and ichthyology. – Copeia, 985: 802- strawberry poison frog Dendrobates pumilio. 832. – J. Exper. Biol., 207: 247-2485.Keferstein, W. (867): Über einige neue oder Silverstone, P.A. (975): A revision of the poi- seltene Batrachier aus Australien und dem tro- son-arrow frogs of the genus Dendrobates pischen Amerika. – Nachr. Gesell. Wissen. G. Wagler. – Nat. Hist. Mus. Los Angeles Co, Sci. A. Univ., Göttingen, 8: 34-36. Bull., 2: -55.Myers, C.W. & J.W. Daly (983): Dart-poison Smithe, F.B. (975-98): Naturalist’s colour guide. frogs. – Sci. Amer., 248: 20-33. Part I. Colour guide. 82 colour swatches. –Myers, C.W., J.W. Daly & V. Martinez (984): Amer. Mus. Nat. Hist., New York, USA. An arboreal poison frog (Dendrobates) from Summers, K., E. Bermingham, L. Weigt, S. Mc- western Panama. – Amer. Mus. Novit., 2783: cafferty & L. Dahlstrom (997): Phenotypic -20. and genetic divergence in three species of dart-Pomiankowski, A. & Y. Iwasa (998): Runaway poison frogs with contrasting parental behav- ornament diversity caused by Fisherian sexual ior – J. Heredity, 88: 8-3. selection. – Proc. Natl. Acad. Sci. Usa, 95: 506- Summers, K., T.W. Cronin & T. Kennedy (2003): 5. Variation in spectral reflectance among popu-Salewski, S. (2005): Erfolgreiche Haltung und lations of Dendrobates pumilio, the strawberry Zucht verschiedener Farbvarianten des Erd- poison frog, in the Bocas del Toro Archipelago, beerfröschchens, Dendrobates pumilio (Dend- Panama. – J. Biogeogr., 30: 35-53. robatidae). – Amphibia, 4: 2-8. Summers, K., T.W. Cronin & T. Kennedy (2004):Saporito, R.A., M.A. Donnelly, P. Jain, H.M. Cross-breeding of distinct colour morphs Garraffo, T.F. Spande, & J.W. Daly (2007): of the strawberry poison frog (Dendrobates Spatial and temporal patterns of alkaloid var- pumilio) from the Bocas del Toro Archipelago, iation in the poison frog Oophaga pumilio in Panama. – J. Herpetology, 38: -8. Costa Rica and Panama over 30 years. – Toxi- Summers, K., R. Symula, M. Clough & T.W con, 50: 757-778. Cronin (999): Visual mate choice in poisonSavage, J.M. (968): The dendrobatid frogs of frogs. – Proc. Biol. Sci., 266: 24-5. Central America. – Copeia, 968: 745-776. Taylor, E.H. (952): The frogs and toads of CostaSavage, J.M. (2002): The amphibians and reptiles Rica. – Univ. Kansas Sci. Bull., 35: 577-942. of Costa Rica: a herpetofauna between two Trapido, H. (953): A new frog from Panama, continents, between two seas. – Chicago and Dendrobates galindoi. – Fieldiana Zool., 34: London (Univ. Chicago Press): 934 pp. 8-87. Manuscript received: 29 November 2006Authors’ addresses: Abel Batista, Instituto de Ciencias Ambientales y Desarrollo Sostenible, Universi-dad Autonoma de Chiriquí, Panama, E-Mail: abelbatista@hotmail.com; Gunther Köhler, Forschungs-institut und Naturmuseum Senckenberg, Senckenberganlage 25, D-60325 Frankfurt a.M., Germany,E-Mail: gkoehler@senckenberg.de. 234

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