1. Variation in Oophaga pumilio
SALAMANDRA 44 4 225-234 Rheinbach, 20 November 2008 ISSN 0036-3375
Variation in Oophaga pumilio
(Amphibia: Anura: Dendrobatidae) in western Panama
Abel Batista & Gunther Köhler
Abstract. We studied variation in colour pattern and morphology in ten populations of Oophaga pumilio
from 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, variation
in the morphometric characters studied is small among these populations. There are examples of distinct
populations 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 ejemplos
de distintivas poblaciones cercanas sin aparentes barreras fisiográficas que no muestran evidencias de
hibridación. Interpretamos estas transiciones abruptas como una evidencia de ausencia o de reducido
flujo 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 observer
Schmidt, 857 (strawberry poison frog) was proves them to be identical, I propose to use
based 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 lost
cas del Toro, Panama, collected by Josef v. (Savage 968) so that direct comparisons of
Warszewicz. Schmidt (858) provided a type material can not be done. An additional
brief colour description and an illustration of nominal species, D. galindoi Trapido, 953
the dorsal view of this species, now in the ge- was described from the Province of Bocas del
nus 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 populations
ity: 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 et
O. Schmidt’s pumilio and argued that the il- al. (997, 2003) provided notes on the colour
lustration 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. 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 third
Fig. 1. Map showing the collecting sites of Oopha-
ga pumilio in this study. Localities are: (1) Cerro finger (3FD) at greatest width; width of third
Tebata; (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 greatest
and Summers et al. (997, 2003). However, width; body width (BW) at greatest width of
we claim the lack of a detailed study of the body; tympanum diameter (TD) horizontal
geographic variation of phenotypic patterns tympanum diameter, based on a estimated
and 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 capitalized
in 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 compared
Descriptions of populations are based on our material to the illustrations therein (their
specimens examined by the authors. Meas- Figs. 35, 46, 47, 48, 49).
urements and colouration data were taken
on 87 specimens of Oophaga pumilio from 0
localities (see Fig. ). Abbreviations for muse- Results
um collections follow those of Leviton et al. Description of the populations examined
(985), except MHCH (Museo Herpetologico
de 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 of
breviations 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 Orange
diagonally from angle of jaw to tip of snout; Yellow (8), Scarlet (4) or same colour of
head 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 dark
rior edge of eye to posterior corner of naris; greyish, with black flecks; venter pale grey;
internaris distance (IND) between cent- limbs black.
226

3. Variation in Oophaga pumilio
Fig. 2. Oophaga pumilio from Cerro Tebata (Loca- Fig. 3. Oophaga pumilio from Isla Bastimentos
lity 1 in Fig. 1), Bocas del Toro, Panama. (Locality 2 in Fig. 1), Bocas del Toro, Panama.
R emarks: Individuals from Costa Rica and
Nicaragua mostly agree well with this descrip-
tion. However, it has been reported that in
rare instances individuals from La Selva can
be 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 from
southern 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 a
C 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: Bocas
individuals 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 Flame
white 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 dark
timentos 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: Bocas
frogs 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. Abel Batista & Gunther Köhler
Tab. 1. Size and proportions of adult Oophaga pumilio from localities 1-10 as shown in Figure 1. Each
range 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 mm
SVL (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2
HL/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 0.373-0.427
FL/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 0.043-0.058
TD/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.23
EL/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.26
IND/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 0.258-0.282
FAL/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 0.244-0.27
HAL/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 .33-2.05
3FD/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-.562
4TD/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 0.37-0.5
BW/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 surfaces
our 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 head
R 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 or
dividuals found on the forest floor during a darker with Jet Black (89) spots; lips coloured
rainy afternoon. as ventral surfaces; tympanum Jet Black (89)
Sp e cimens examine d: PANAMA: Bocas or same colour as dorsum; ventral surfaces of
del 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. 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, males
darker 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 solid
Sp 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 during
82º2’30.7’’W, 20 m: SMF 8660-6. daytime in a mature secondary forest and in
disturbed areas. Most frogs were found on
Cerro Brujo (Locality 6 in Fig. ) the ground, one was observed climbing a tree
C 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. Abel Batista & Gunther Köhler
Fig. 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 Olive
Fig. 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 form
Río Uyama (Locality 7 in Fig. ) stripes, dorsal and ventral surfaces of limbs
C olourat ion in life: Dorsal ground as well as ventral surfaces of head and body
colour 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 same
form stripes; venter Light Sky Blue (68D) as dorsum; males with darker throat (Fig. 9).
with large black blotches. Sp e cimens examine d: PANAMA: Bocas
C olour af ter six mont hs in pre- del Toro: Quebrada la Gloria (9 km SW from
s 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 of
or black (Fig. 8). head and body uniform red; in SMF 86608
R emarks: This series was collected in the frontal and parietal region suffused with
late 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; in
the ground, one was observed climbing a tree SMF 86609 forelimbs Scarlet (4), hands Raw
230

7. Variation in Oophaga pumilio
Fig. 8. Oophaga pumilio from Río Uyama (Locality Fig. 10. Oophaga pumilio from Kusapin (Locality 9
7 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 surfaces
of 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 solid
brown to dark brown; lips Sky Blue (68C) dark greyish; venter almost black; limbs dark
or 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 during
edge of tympanum usually bordered by a daytime on the floor in a mature secondary
thin 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 of
and flanks (Fig. 0). head and body uniform Leaf Green (46)
231

8. Abel Batista & Gunther Köhler
to Parrot Green (260) or with Jet Black (89) Discussion
flecks; upper surfaces of limbs brown, Blue
Black (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 highly
part) 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 obvious
venter (Fig. ). physiographic barriers that show no evidence
C olour af ter s e ven mont hs in pre - of hybridization in terms of colouration. In
s er vat ive: Dorsal surfaces of head and the area of Almirante an abrupt transition is
body solid dark greyish, some specimens found between a population consisting of a
keep their black spots on dorsum and flecks black and white morph and red morph with
in the venter; limbs dark brown; venter grey. blue legs (Summers et al. 2003). In the area of
R emarks: The series was collected during Chiriquí Grande, three distinct phenotypes
daytime in a wetland forest, most specimens meet and abrupt transitions are reported
were found climbing on trees at less than two between them: a population consisting of a
meters 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 green
marca 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 interpret
86564-73. 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 for
trary, variation in morphometric characters O. pumilio “a static subspecies concept does
studied is comparatively small among these not seem well suited to the dynamics of the
populations. All morphometric characters situation”. However, the descriptions of the
show largely overlapping ranges among most phenotypes by these authors agree well with
populations (Table ). In respect of the mor- our observations indicating that over the
phometric characters examined, the Popa course of at least four decades the phenotypic
population (Locality 4 in Fig. ) is the most pattern had remained stable. The underlying
distinctive 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 that
lations. The series from Bastimentos (Local- females of O. pumilio preferentially choose
ity 2 in Fig. ) has the highest average values mates of their own colour morph (Siddiqi
for FAL/SVL and HAL/SVL indicating that et al. 2004). Under captive conditions it has
these 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 courtship
do overlap to some degree with the other behavior (mating calls) only towards females
populations. of their own phenotype but not towards pop-
ulations from Bastimentos Island (G. Var-
232

9. Variation in Oophaga pumilio
gas 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. Meike
the 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 instrumental
to potential predators but also to conspecif- in enabling Abel Batista and Marcos Ponce to
ics, 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 Ciencias
al. 2004). Fisher’s runaway process of sexu- Naturales y Exactas de la Universidad Autónoma
al selection is potentially an important force de Chiriquí (UNACHI), David, Panama, for their
generating character divergence between support. This paper is based on part upon work
closely 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 de
different colours morphs can produce fer- Chiriquí (UNACHI), David, Panama. We are
tile offspring. However, the survival rates of grateful to Alfred A. Schmidt for his financial
offspring 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 her
to 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 on
mtDNA data indicated that the studied pop-
ulations are phylogenetically closely related References
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Manuscript received: 29 November 2006
Authors’ 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