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237
A new species of harlequin toad from Amazonian Ecuador
Open access at hps://www.salamandra-journal.com
© 2024 Deutsche Gesellscha für Herpetologie und Terrarienkunde e.V. (DGHT), Germany
15 November 2024 ISSN 0036–3375
SALAMANDRA 60(4): 237–253 SALAMANDRA
German Journal of Herpetology
A new species of harlequin toad (Bufonidae: Atelopus)
from Amazonian Ecuador
A P, A T-V, J C,, R B, D J. P,
A B. Q R, C H. H, J P. R-P,,,
D S-V, J M G, S L
1 Biogeography Department, Trier University, Universitätsring 15, 54296 Trier, Germany
2 Centro Jambatu de Investigación y Conservación de Anbios, Fundación Jambatu, Giovanni Farina 566 y Baltra,
San Rafael, Ecuador
3 Photo Wildlife Tours, Isla Isabela 1021 y Río Coca, 170501, Quito, Ecuador
4 Fundación Cóndor Andino, Tamayo N24-260 y Lizardo García, 170523, Quito, Ecuador
5 Laboratory Mecadev UMR 7179 CNRS/MNHN, Département Adaptations du Vivant, Bâtiment d’Anatomie Comparée,
55 rue Buon, 75005, Paris, France
6 Department of Biology, University of Dayton, 300 College Park, Dayton, Ohio 45469, USA
7 Fundación Ecominga, Mariscal Foch esquina, Juan León Mera 721, Quito 170524, Ecuador
8 Instituto Nacional de Biodiversidad (INABIO), Pasaje Rumipamba 341 y Av. de los Shyris, Quito, Ecuador
9 La Sapada Equipo Herpetológico-Finca La Argentina, Puyo, Pastaza Ecuador
10 Centro de Investigación de la Biodiversidad y Cambio Climático (BioCamb) e Ingeniería en Biodiversidad y Recursos Genéticos,
Facultad de Ciencias de Medio Ambiente, Universidad Indoamérica, Av. Machala y Sabanilla, Quito, Ecuador
11 Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales COCIBA,
Laboratorio de Biología Evolutiva, Calle Diego de Robles s/n y Pampite, Cumbayá, Quito, Ecuador
12 Tandayapa Cloud Forest Station, Universidad San Francisco de Quito USFQ, P.O. Box 17-1200-841, Quito, Ecuador
Corresponding author: A P, ORCID 0000-0001-8731-6312,
e-mail: amadeus.plewnia@t-online.de
Manuscript received: 14 October 2024
Accepted: 8 November 2024 by J K
Abstract. For nearly four decades, harlequin toads, genus Atelopus, have suered unparalleled population declines. While this also results
in limited understanding of alphataxonomic relationships, these toads face an urgent need for advances in systematics to inform conserva-
tion eorts. However, high intraspecic variation and cryptic diversity have hindered a comprehensive understanding of Atelopus diversity.
is is particularly exemplied among Amazonian populations related to A. spumarius, where decades of taxonomic work have not been
able yet to unravel relationships between the many forms, while the names coined so far have led to taxonomic confusion leaving numer-
ous lineages unnamed. A recent comprehensive phylogenetic study has revealed new insights into the systematics of harlequin toads with
an emphasis on Amazonian forms, identifying several unnamed lineages. We here describe one of these evolutionary lineages as a new
species, restricted to the Ecuadorian Amazon basin, in an integrative taxonomic approach using molecular, morphological, bioacoustic
and larval information. With this, we contribute to a better understanding of Atelopus diversity as the baseline of conservation action.
Key words. Amphibia, Anura, amphibian crisis, bioacoustics, cryptic diversity, Neotropics, osteology, integrative taxonomy.
Resumen. Durante casi cuatro décadas, los sapos arlequín, del género Atelopus, han sufrido descensos poblacionales sin precedentes.
Aunque esto también resulta en una comprensión limitada de las relaciones alfa-taxonómicas, estos sapos enfrentan una necesidad ur-
gente de avances en sistemática para orientar los esfuerzos de conservación. Sin embargo, la alta variación intraespecíca y la diversidad
críptica han dicultado una comprensión integral de la diversidad de Atelopus. Esto se ejemplica particularmente entre las formas ama-
zónicas relacionado con A. spumarius, donde décadas de trabajo taxonómico aún no han podido esclarecer las relaciones entre las muchas
formas, mientras que los nombres asignados hasta ahora han llevado a una confusión taxonómica que deja numerosos linajes sin nombre.
Un reciente estudio logenético integral ha revelado nuevos conocimientos sobre la sistemática de los sapos arlequín con énfasis en las
formas amazónicas, identicando varios linajes sin nombre. Aquí describimos uno de estos como una nueva especie restringida a la cuen-
ca amazónica ecuatoriana en un enfoque taxonómico integrativo utilizando información molecular, morfológica, bioacústica y larval.
Con esto, contribuimos a una mejor comprensión de la diversidad de Atelopus como base para las acciones de conservación.
Palabras clave: Amphibia, Anura, bioacústica, crisis de anbios, diversidad críptica, Neotrópicos, osteología, taxonomía integrativa.
238
A P et al.
Introduction
Unparalleled declines and possible extinctions have af-
fected harlequin toads, Atelopus D B, ,
over the last four decades with their conservation status
not having improved since (L M et al. , L-
et al. ). As a result, harlequin toads are oen con-
sidered the poster-child of the precipitous global amphib-
ian declines commonly framed as the amphibian crisis
(L et al. , L et al. ). With this, we
see an urgent need for advancing Atelopus systematics in
the context of conservation (cf. M et al. ,
V-T et al. ).
With more than species, many of which remain un-
described, harlequin toads are considered the most spe-
cies-rich bufonid genus (R-A et al. ,
L et al. ). High intraspecic plasticity coupled
with cryptic diversity has yet prevented a comprehensive
understanding of harlequin toad systematics, particularly
regarding forms from the Amazon basin (N G-
, L et al. , , ).
Since the descriptions of A. spumarius by C ()
from lowlands of western Amazonia and A. pulcher by
B () from the Andean foothills of Amazo-
nia, both names have been applied to many populations in
both the western and eastern basin, with additional names
coined for some of those as subspecies (reviewed by L-
et al. a, ).
In contrast to the earlier view, taxonomists have since
agreed that most western and eastern Amazonian popula-
tions related to A. spumarius or A. pulcher represent dis-
tinct taxa (e.g. L D R , L et
al. a,b, , R-A et al. , J
et al. ). However, it has remained dicult to draw
taxonomic conclusions, particularly because () access to
fresh material of A. spumarius sensu stricto for molecular
studies was hampered, () Atelopus remain overall poorly
sampled in Amazonia, and () at least for some nominal
species, repeated contact during Pleistocene cold phases
makes molecular delimitation challenging (L et al.
a, N G ). Action is further com-
plicated by the relatively low number of species included
in molecular phylogenetic approaches (e.g. L et al.
, R et al. ).
A recent study (L et al. in press) presents a
broadly updated phylogenetic reconstruction of harlequin
toads with vast taxonomic implications as a ‘roadmap’ for
future systematic work. e sampling is best for Amazo-
nian harlequin toads, revealing several formerly unnamed
lineages in western Amazonia, which are oen separated
by riverine barriers. Moreover, the rediscovery of A. spu-
marius sensu stricto by the authors close to its type locality
nally allows to thoroughly discriminate species from the
available names (L et al. in press). e purpose of
this paper is to describe one of the new species-level line-
ages indentied by L et al. (in press) as a new spe-
cies.
Material and methods
Nomenclatural acts
e electronic edition of this article conforms to the re-
quirements of the amended International Code of Zoo-
logical Nomenclature (ICZN). Hence, the new name given
herein is available under the code from the electronic edi-
tion of this article. is published work and the nomen-
clatural act it contains have been registered in ZooBank,
the online registration system for the ICZN. e LSID (Life
Science Identier) for this publication is: urn:lsid:zoobank.
org:pub:EEC-DCC--B-CFFEE. e
electronic edition of this work was published in a journal
with an ISSN, and has been archived and is available from
the following digital repositories: salamandra-journal.
com, zenodo.org.
Material examined
Specimens examined are housed in AMNH (American
Museum of Natural History, New York), BMNH (Brit-
ish Museum, London), ANSP (Academy of Natural Sci-
ences, Philadelphia), CBF (Colección Boliviana de Fauna,
La Paz), CJ (Centro Jambatu de Investigación y Conser-
vación de Anbios, San Rafael), DHMECN-INABIO (In-
stituto Nacional de Biodiversidad, Quito), FMNH (Field
Museum, Chicago), GOV (eld number, Gustavo Orcés,
at USNM), IAVH-Am (Instituto Alexander von Hum-
boldt – Colección de Anbios, Villa de Leyva), ICN (In-
stituto de Ciencias Naturales, Bogotá), KU (Natural His-
tory Museum, e University of Kansas, Lawrence), LG
(eld number, Jean Lescure, at MNHNP), MCZ (Muse-
um of Comparative Zoology, Harvard University, Cam-
bridge), MHNC (Museo de Historia Natural del Cusco,
Cusco), MHNG (Muséum d’histoire naturelle, Genève),
MNCN (Museo Nacional de Ciencias Naturales, Madrid),
MNHNP (Muséum National d’Histoire Naturelle, Paris),
MTD (Museum für Tierkunde, Dresden), MUBI (Museo
de Biodiversidad del Perú, Cusco), MUSM (Museo de His-
toria Natural, Universidad Nacional Mayor de San Mar-
cos, Lima), NHMW (Naturhistorisches Museum Wien,
Vienna), NHRM (Naturhistoriska Rijkmuseet, Stock-
holm), QCAZ (Museo de Zoología, Pontica Universidad
Católica del Ecuador, Quito), USNM (National Museum
of Natural History, Washington), UTA (University of Tex-
as at Arlington, Arlington), and ZFMK (Zoologisches For-
schungsmuseum Alexander Koenig, Bonn).
Morphology
Terminology and methods for the description of external
morphology and morphometrics follow L et al. (in
press) and references therein. Abbreviations are as follows:
239
A new species of harlequin toad from Amazonian Ecuador
SVL, snout–vent length; TIBL, tibia length; FOOT, foot
length; HLSQ, head length from the squamosal; IOD, in-
terorbital distance; HDWD, head width; EYDM, eye diam-
eter; EYNO, eye to nostril distance; ITNA, internarial dis-
tance; FAL, length of exed forearm; HAND, hand length;
THBL, thumb length; SW, sacrum width. Measurements
are provided in mm.
Osteology
We obtained skeletal data using a GE v|tome|x M mi-
cro-computed tomography system (microCT) at the Uni-
versity of Florida’s Nanoscale Research Facility in Gains-
ville, Florida. Specimens were scanned with a kV X-
ray tube and diamond-tungsten target with the voltage
and current set to kV and A, respectively. e raw
X-ray data were processed using the GE datos|x soware
to produce tomogram and volume les. e volume les
were imported into VG StudioMax (Volume Graphics,
version .), and the skull and skeletons were isolated
using VG StudioMax’s suite of segmentation tools. Com-
puted tomography data have been deposited in Morpho-
Source (https://www.morphosource.org). Osteological de-
scriptions follow L et al. (in press) and references
therein.
Bioacoustics
A recording of ms duration was available for analy-
sis. Calls were analysed using Audacity ... at a sampling
rate of . kHz and -bit resolution. Frequency informa-
tion was obtained using Fast Fourier Transformation (FFT
width at Hann window function). Terminology, de-
scription and visualization follow L et al. (in press)
and references therein. e analyzed call recording was de-
posited in full length in the Fonoteca Zoológica (www.fo-
nozoo.com) of the Museo Nacional de Ciencias Naturales
(CSIC), Madrid, and CJ sound archive. Calls were recorded
by AP from the life holotype in the laboratory on Sep-
tember , ca. : pm, with a Sennheiser ME direc-
tional microphone attached to an Olympus DM- digital
recorder. Distance to recorder was . m and ambient tem-
perature was ca. °C.
Tadpoles
Description format for larvae follows L et al. ().
Developmental stages are those of G (). In this
paper, we redescribe the tadpole of the new species based
on a single specimen from Río Puyo (Ecuador), formerly
described under the name Atelopus spumarius spumarius
by D L (), and on photos of laborato-
ry-reared larvae in life.
Distribution and natural history
We compiled information on species distribution based on
specimens from museum collections and eld observations.
Conservation status
We suggest a category based on the IUCN Red List Catego-
ries and Criteria Version ., Second edition (IUCN ).
We calculated the Extent of Occurrence (EOO) for the
species as the area contained by a minimum convex poly-
gon over all known localities and the Area of Occupancy
(AOO) as the area of inhabited grid cells when applying a
grid cell size of km by km, centered around the coor-
dinate in ESRI ArcGIS Pro ... (IUCN ). Additional
threats are discussed in this section.
Results
Based on the molecular results of L et al. (in press)
as well as comparative evidence from external and internal
morphological, bioacoustic and larval characters, we here
describe one of the undescribed species-level lineages, oc-
curring in the Ecuadorian Amazon basin, indentied by
L et al. (in press) as a species new to science.
Atelopus colomai sp. n.
Figs 1–2, S1
ZooBank LSID: urn:lsid:zoobank.org:act:DEFC--F-
-DFEE
Atelopus spumarius non C. – R-A et al. :
, R et al. : , C-B M-S
: , T et al. : .
Atelopus spumarius spumarius non C. – D L
: , L : .
Atelopus pulcher non B. – R : , G
: , P : (color illustration of paratype ZFMK
), L et al. b: .
Atelopus pulcher pulcher non B. – P : .
Atelopus a. spumarius. – C et al. : .
Atelopus sp. (a. A. pulcher). – L et al. a: .
Atelopus sp. (spumarius complex). – W et al. : .
A. sp. (spumarius complex from Puyo, Ecuador). – M-
L et al. : .
A. sp. Otoyacu (spumarius complex). – J et al. : .
A. sp. cf. spumarius Centro Oriente. – L et al. : Sup-
plementary Table .
Holotype: Ecuador: Provincia Pastaza: Río Pucayacu, Nof
Diez de Agosto (°’” S, °’” W, m a.s.l.), CJ
(Figs –), adult male, leg. September by
J C, A B. Q R, D
M and A P.
240
A P et al.
Paratopotypes: CJ , –, –,
(Fig. S), adult females, same data as holotype.
Paratypes: Ecuador: Provincia Pastaza: NW of El Triun-
fo (°’” S, °’” W, m a.s.l.), CJ –,
, adult males, leg. September by J C-
, A B. Q R, D M,
Z L and A P; Reserva Otoyacu
(°’” S, °’” W, m a.s.l.), CJ , , adult
males, and CJ , , adult females, leg. unknown;
Finca la Argentina, N of Puyo (°’.” S, °’” W,
m a.s.l.), DHMECN-INABIO , adult female, leg.
August by J P. R P, J I S-
L, P B, N K and P-
V; Río Cononaco, ZFMK , subadult fe-
male, leg. April by E P; Río Villano, near
Villano, BMNH .–. ( m a.s.l.), adult fe-
males, leg. (date unknown) by R O.
Referred material: Ecuador: Provincia Orellana: Yasuní
National Park, road from Pompeya Sur to Iro on km ,
QCAZ ; Yasuní National Park, road from Pompe-
ya Sur to Iro on km , QCAZ , , , ,
; Provincia Pastaza: Reserva Otoyacu, CJ , ,
, , –, ; Río Pucayacu, CJ ,
CJ ; . km SE of Puyo, USNM , , JAP
–; Río Puyo, km S Puyo, KU ; trail from
Unión Base to Rosario Yacu, SE of Puyo, MCZ Herp
A-–, A-–; Veracruz, ca. km E of
Puyo, USNM , KU –; Río Oglán, GOV
S; km on road from San Ramón to El Triunfo, Río
Pucayacu, QCAZ , , –, , ,
–, , , –, –,
–, –, –, , , ,
–, –, , ; Finca de la Co-
operativa Mariscal Sucre, Río Pucayacu, QCAZ ;
Parroquia Teniente Hugo Ortiz, QCAZ ; Conona-
co, UTA WWL ; Canelos, MCZ Herp A-–;
Baños and Canelos, AMNH –; Río Villano:
BMNH .–; Puyo, FMNH ; historic prov-
ince Santiago-Zamora (now provinces Zamora Chin-
chipe and Morona Santiago), NHRM RRB/..
(specimens).
Denition: Atelopus colomai sp. n. is placed in the genus
Atelopus based on having presacrals I and II fused, the
presence of a gastromyzophorous tadpole with a ventral
belly sucker and its mitochondrial and nuclear markers be-
Figure 1. Preserved male holotype of Atelopus colomai sp. n. in dorsal, ventral and lateral views (CJ 12198). Scale bar 10 mm. Photos
by J C.
241
A new species of harlequin toad from Amazonian Ecuador
ing nested within Atelopus (L et al. in press). e
species is dinstinguished from all other species by the com-
bination of the following characters: () A small-sized spe-
cies with mean SVL of adult males . ± . mm, N= ,
and adult females . ± . mm, N = ; () slender body
(SW/SVL = . ± . in males, . ± . in
females) with () long legs (TIBL/SVL = . ± . in
males, . ± . in females); () acuminate snout in
dorsal view, protruding beyond apex of lower jaw; ()tym-
panic membrane absent, tympanic annulus not visible,
columella present, ostia pharyngea present; () phalan-
geal formula of hand ---; webbing absent on hand;
() rst nger short (THBL/HAND = . ± . in
males, . ± . in females), rst nger in males
with keratinized dark brown spiculae forming the nup-
tial pad; () phalangeal formula of foot ----, web-
bing formula of toes I– to II to .–III. to –.
to IV. to –.V; () skin smooth, dorsally and laterally
covered with dense well-dened minute spiculae, result-
ing in a velvety appearance of the skin on respective sur-
faces; skin ventrally smooth to areolate; () vertebral col-
umn conspicuous, neural processes absent; () in life, dor-
sum black with few yellow-green asymmetric circles, dots
or lines; dorsolaterally yellow-green foam-like reticulation
forming a band from above eye to groin with numerous
black rounded irregular spots (smaller than EYDM); later-
ally a black band present from tip of snout almost to groin,
rarely interrupted by yellow-green lines; limbs and dorsal
surfaces of hand and foot with yellow-green foam-like re-
ticulation with black spots (smaller or larger than EYDM),
thumb and sometimes tips of ngers and toes dorsally or-
ange to vermillion red; in males, throat and venter cream,
posteriorly becoming yellowish, sometimes with few tiny
black dots (smaller than or equal to diameter of nger III)
on throat and black dots (smaller than EYDM) on later-
al and posterior vent; in females, throat and venter yellow
with numerous black dots (smaller or larger than EYDM);
in both sexes, palms, plants and an ovally-shaped blotch
on ventral surface of cloaca and adjacent areas of thighs
vermillion red, the latter being surrounded by black dots
(smaller than EYDM); iris black with a golden yellow ring
surrounding horizontally oriented pupil; () in preserva-
tive: above dark brown background color with cream pat-
tern, below whitish cream with brown dots; vermillion
red life markings faded to pale orange, yellow or cream;
() pulsed calls long for an Atelopus ( ± ms) with
dominant frequency ± Hz.
Diagnosis: Atelopus colomai sp. n. (Figs –, S) can be
readily distinguished from all other Atelopus species (as far
known) by molecular genetics (monophyly in a concate-
nated (S, S, CytB) mitochondrial phylogeny, support
by molecular species delimitation and lack of haplotype
sharing in POMC, L et al. in press). It can be mor-
phologically distinguished from all congeners by the com-
bination of small size, dorsal and lateral skin covered with
dense well-dened minute spiculae, ventral skin smooth to
slightly areolate, presence of a columella and dorsal pat-
tern, except from A. spumarius C, sensu stricto
and A. manauensis J, F L, . From
these species, it diers by longer call duration ( ±
ms vs. ± ms in A. spumarius sensu stricto, L-
et al. in press; ± ms in A. manauensis, J
et al. ), in female ventral coloration and pattern in life
(yellow with equally distributed black rounded dots on
vent smaller or larger than EYDM vs. throat and chest or-
ange or yellow, venter anteriorly cream yellow, posteriorly
orange or yellow, anteriorly with numerous brown or black
dots smaller than or equal to EYDM, sometimes forming
a continuous band on chest in A. spumarius sensu stricto,
L et al. in press; venter white or whitish cream with
spots in A. manauensis, J et al. ) and in having an
oval vermillion red blotch surrounded by small black dots
on ventral surface of cloaca and onto adjacent thighs in life
(vs. ventral surface of cloaca and thighs bright orange in
A. spumarius sensu stricto, and bright red in A. manauen-
sis, all lacking black dots). Further, A. colomai sp. n. diers
from both taxa in having a more pointed, acuminate snout
in lateral view in females (vs. subacuminate to truncate and
less protruding in A. spumarius sensu stricto; blunt in A.
manauensis). Moreover, from A. manauensis, the new spe-
cies diers in having a longer head (males HLSQ/HDWD
. ± ., N = , vs. . in A. manauensis, N = ,
J et al. ). e tadpole of the new species diers
from that of A. manauensis by having the lower beak about
twice as long as the upper one (vs. upper beak only slightly
shorter than lower, G under the name A. pul-
cher).
Atelopus colomai sp. n. is morphologically similar to its
congeners of the avescens-spumarius clade sensu L-
et al. (in press), as well as A. andinus R, ,
A.loettersi D R, C-F, C,
B P, , and the A. tricolor B,
complex (including the junior synonyms A. rugulo-
sus N, and A. willimani D-B, )
from the eastern Andean versant of Peru and Bolivia, and
to A. palmatus A, and A. planispina J-
E, from the eastern Andean versant
of Ecuador. With respect to these supercially similar spe-
cies, the new species diers as follows: from the nomi-
nal A. barbotini L, , A. avescens D
B, (including its junior synonym A. vermicu-
latus MD, ), A. franciscus L, and
the A. hoogmoedi L, complex (including the
available name A.hoogmoedi nassaui O J,
) in pattern and coloration in life (dorsally black with
few yellow-green asymmetric circles, dots or lines; dorso-
laterally yellow-green foam-like reticulation forming a
band from above eye to groin with numerous black round-
ed irregular spots; males below cream, posteriorly becom-
ing yellowish, sometimes with few tiny black dots and a
vermillion red shaped blotch on ventral surface of cloaca
and adjacent areas of thighs, females ventrally yellow with
numerous black dots and a vermillion red shaped blotch
on ventral surface of cloaca and adjacent areas of thighs vs.
dorsally black with red to pink markings, ventrally reddish
242
A P et al.
pink in A.barbotini; dorsally unicolored reddish, brown-
ish or yellow or brown and with olive, brownish reddish
and copper-like vermiculation, ventrally reddish pink in
males, pink in females in A. avescens; dorsally olive to
blackish brown, ventrally reddish in A. franciscus; dor-
sally dark brown or black with yellow, pink, bluish, white,
greenish or orange irregular spots and reticulated dorso-
lateral bands or solid bars, ventrally yellow, pink, bluish,
white, greenish or orange, in some specimens with dark
spots or blotches in the A. hoogmoedi complex). All these
forms, except some populations of the A. hoogmoedi com-
plex, lack red palmar and plantar surfaces in life (vs. pre-
sent in A.colomai sp. n.). All Guianan harlequin toads have
shorter call durations ( ± ms vs. ± ms in
A. barbotini; ± ms in A. avescens; ± ms
in A. franciscus; ± ms in the A. hoogmoedi com-
plex from French Guiana; ± ms in the A. hoog-
moedi complex from Amapá, Brazil; – ms in the
A. hoogmoedi complex from Pará, Brazil; L ,
C et al. , C-C C ).
Tadpoles of the new species dier from those of A. aves-
cens, A. franciscus and the A. hoogmoedi complex in hav-
ing the lower beak about twice as long as the upper (vs.
upper beak only slightly shorter than lower; B et al.
, L et al. ) and in coloration in life (black
to dark brown with dense irregular pale brown to golden
markings that consist of aggregated minute dots vs. black
with symmetrical golden spots anterior and posterior eye,
at the base of the tail and on tail n in A. avescens and
A.franciscus; ventrally entirely translucent with few whit-
ish cream dots in the A.hoogmoedi complex (B et al.
, L et al. ). e new species can be distin-
guished from A. pul cher B, by smaller male
SVL (. ± . mm, N = , vs. . ± . mm, N = ,
in A. pulcher; L et al. a), female ventral col-
oration in life (yellow with numerous black dots and a ver-
million red shaped blotch on ventral surface of cloaca and
adjacent areas of thighs vs. vermillion red with irregular
dark spots in A. pulcher, L et al. a) and longer
calls (, ± ms vs. , ± ms in A. pulcher, L-
et al. a). Tadpoles of A. colomai sp. n. dier from
those of A.pulcher in having the lower beak about twice
as long as the upper (vs. one third to one fourth as long)
and the nostrils positioned at about height of eye (vs. below
height of eye; L et al. a). Atelopus colo mai sp.
n. diers from A. seminiferus C, in smaller female
SVL (.± . mm, N = , vs. .mm, N = ), absence
of lateral warts (present in A.seminiferus) and dorsal col-
oration in preservative (dorsally black with few yellowish
cream asymmetric circles, dots or lines; dorsolaterally yel-
lowish cream foam-like reticulation forming a band from
above eye to groin with numerous black rounded irregu-
lar spots vs. dark brown without pattern in A. seminiferus).
From A. andinus, A. loettersi and the A. tricolor complex,
A. colo mai sp. n. is distinguished by the presence of a colu-
mella (vs. absence; L et al. , R. B unpubl.
data). From A. loettersi, it can be distinguished by female
ventral coloration and pattern in life (yellow with numer-
ous black dots and a vermillion red shaped blotch on ven-
tral surface of cloaca and adjacent areas of thighs vs. mostly
red, in some specimens with irregular brown blotches in
A. loettersi; D R et al. ). e new species can
be distinguished from A. andinus and the A. tricolor com-
plex by the absence of warts (vs. presence) and by presence
of dorsolateral reticulation (vs. absence). Tadpoles of the
new species dier from those of the A. tricolor complex in
having the upper beak shorter than the lower (vs. longer;
L et al. ). Atelopus colomai sp. n. can be distin-
guished from A. palmatus and A. planispina by coloration
and pattern in life (dorsally black with few yellow-green
asymmetric circles, dots or lines; dorsolaterally yellow-
green foam-like reticulation forming a band from above
eye to groin with numerous black rounded irregular spots;
males below cream, posteriorly becoming yellowish, some-
times with few tiny black dots and a vermillion red ovally-
shaped blotch on ventral surface of cloaca and adjacent ar-
eas of thighs, females below yellow with numerous black
dots and a vermillion red ovally-shaped blotch on ventral
surface of cloaca and adjacent areas of thighs vs. green
blotches on reddish brown dorsum, absence of reticula-
tion; ventrally unicolored orange in females of A. palmatus
and whitish to reddish in A. planispina; P ) and
skin texture (smooth, covered in minute spiculae vs. pres-
ence of small spiculae on anks, upper arm and from eye
fading towards groin; P ). Tadpoles of A. colomai
sp. n. dier from those of A. palmatus in exhibiting a gold-
en tail pattern in life (vs. absence; M-L et al.
) and in having the upper beak shorter than the lower
(vs. longer; M-L et al. ).
Description of holotype: Adult male. Body slender (SW/
SVL .); head slightly longer than wide (HLSQ/HDWD
.); in prole, snout protruding beyond apex of lower
jaw, acuminate in dorsal view, forming a triangle from
nostrils to tip of snout; nostrils directed laterally, mod-
erately protuberant, not visible in dorsal view, about two
thirds from eye to tip of snout in lateral view, situated an-
terior to apex of lower jaw; canthus rostralis well dened,
concave in dorsal view between eye and nostril; loreal re-
gion concave; lips not ared; top of snout depressed; head
plain in lateral view; eyelid ared, densely covered in min-
ute spiculae; tympanic membrane absent, tympanic an-
nulus not visible; supratympanic crest poorly developed;
choanae small, rounded; ostia pharyngea present; tongue
about twice as long as wide, broadest at its tip (situated
posteriorly); vocal slits present; limbs long and slender
(TIBL/SVL .); webbing absent on hand; ngers and
toes lack lateral fringes; palmar tubercle rounded, poor-
ly developed; supernumerary palmar and plantar tuber-
cles absent; thenar and subarticular tubercles indistinct;
tips of digits rounded; phalangeal formula of hand ---
; rst nger short (THBL/HAND .), dorsally covered
with keratinized dark brown spiculae forming the nuptial
pad; inner metatarsal tubercle indistinct, outer metatar-
sal tubercle rounded, poorly developed; relative length of
toes I<II<III<V<IV; phalangeal formula of foot ----,
243
A new species of harlequin toad from Amazonian Ecuador
webbing formula of toes I–II–III.–IV–.V; tar-
sal fold absent; foot roughly three quarters of tibia; skin
smooth on entire body, dorsally and laterally covered with
dense, well-dened minute spiculae, resulting in a velvety
appearance of the skin; skin smooth to slightly areolate on
ventral surfaces of cloaca and adjacent areas of thighs, ven-
ter and throat; cloacal opening in an inconspicuous tube,
directed posteriorly; vertebral column visible through the
skin, neural processes absent.
In life (Fig. ), dorsum black with few yellow-green
asymmetric circles, dots or lines; dorsolaterally yellow-
green foam-like reticulation, forming a band from above
eye to groin bearing numerous black rounded markings
(smaller than EYDM); lips bordered by thin black line; lat-
erally a black band from tip of snout almost to groin; ven-
trolaterally transition from yellow-green reticulation with
numerous black dots to unicolored cream throat and ven-
ter; throat and venter cream with few minute dark dots in
Figure 2. Life aspects of Atelopus colomai sp. n. from the type locality. Le column: dorsal (CJ 12406; top) and ventral (CJ 12409;
bottom) views of adult female paratopotypes. Right column: dorsal and ventral views of adult male holotype (CJ 12198). Scale bar
10mm. Photos by J C.
244
A P et al.
gular region (smaller than or equal to diameter of nger
III), venter posteriorly becoming yellowish with few black
dots (smaller than EYDM); limbs dorsally black, covered
in yellow-green bands, ventrally cream to yellow bearing
black dots or bars; thumb and tips of ngers and toes dor-
sally orange to vermillion red; palms, plants and an ovally-
shaped blotch on ventral surface of cloaca and adjacent ar-
eas of thighs vermillion red, the latter being surrounded by
black dots; iris black with a golden yellow ring surrounding
horizontally oriented pupil.
In preservative, above blackish brown with yellowish
cream pattern; below whitish cream with blackish brown
dots; vermillion red life markings faded to pale cream with
pale reddish remnants. Proximal (anterior) half of tongue
has black pigmentation.
Measurements: SVL ., TIBL ., FOOT ., HLSQ
., IOD ., HDWD ., EYDM ., EYNO ., ITNA .,
FAL ., HAND ., THBL ., SW ..
Variation: For meristic variation of characters see Table .
Paratypes (Fig. , S) correspond to the description above.
However, specimens from Reserva Otoyacu and Río Vil-
lano are slightly larger than topotypic individuals and
specimens from NW El Triunfo. Among the paratypes is
one malformed specimen (CJ ) with partial anoph-
thalmia, in which the right eye is not fully developed and
almost completely covered with skin.
Sexual dimorphism is apparent in A. colomai sp. n. with
females being larger than males (SVL . ± . mm, N=
, vs. . ± . mm, N = ). In addition, females lack
nuptial pads and vocal slits, dier in coloration by hav-
ing a yellow venter with equally distributed rounded black
dots (smaller or larger than EYDM) in the gular region, on
chest and venter (Fig. ).
Cranial osteology: General osteological features of a
male (CJ ; MorphoSource Media ID ) and
a female (CJ , paratype; MorphoSource Media ID
) are depicted in Figure . e skull of A. co-
lomai sp. n. is triangular in dorsal view. e skull length
is .mm, the skull width is . mm, and the skull height
is .mm in CJ ; the skull length is . mm, the skull
width is . mm, and the skull height is . mm in CJ .
e skull roof is smooth. In anterior view, the septomaxilla
is U-shaped with medial and lateral rami that extend poste-
riorly towards the vomer. e lateral ramus is much broad-
er than the medial ramus and bears a nasal process. e os-
sied sphenethmoid is smooth and underlies the posterior
and medial margins of nasals, as well as the anterior and
anterolateral margins of the frontoparietals. e posterior
limit of the sphenethmoid is about one-third the length of
the margin of the orbit. e prootics are fused to the pos-
terolateral edge of the frontoparietals. e exoccipitals en-
circle the foramen magnum and are fused to the fronto-
parietals dorsally, the prootics laterally, and the parasphe-
noid anteroventrally. e otic capsule is well-ossied. Col-
umellae are present, cylindrical in shape. e pars media
of the columella is slender and slightly bowed, and the pars
interna is not ossied. e operculum in the fenestra ova-
lis is mineralized and visible. e nasals are triangular and
bear an acuminate maxillary process that extends ventro-
laterally toward the maxilla and contacts the pars facialis of
the maxilla in CJ and nearly contacts the pars facialis
of the maxilla in CJ . e nasals are moderately sepa-
rated medially. e frontoparietals are rectangular and the
orbital edges are straight. Medial articulation is complete
in both specimens. A suture line is visible only in CJ .
A supraorbital ange is absent. Occipital groves, which
are canals for the carotid artery, are present and partially
roofed with bone. In both specimens, the grooves are cov-
Table 1. Measurements (in mm) and proportions of the type se-
ries of Atelopus colomai sp. n.; mean ± SD (range), excluding
subadult paratype ZFMK 44976. For abbreviations see Method
section.
Males (N = 6) Females (N = 10)
SVL 23.2 ± 1.46
(21.2–25.6)
29.8 ± 2.05
(26.4–33.7)
TIBL 10.0 ± 0.16
(9.9–10.3)
12.9 ± 0.47
(12.3–13.7)
FOOT 7.5 ± 0.37
(6.9–8.0)
10.6 ± 1.09
(8.8–12.0)
HLSQ 7.6 ± 0.33
(7.2–8.1)
9.0 ± 0.46
(8.4–9.9)
IOD 2.5 ± 0.37
(2.1–3.0)
3.2 ± 0.37
(2.7–3.9)
HDWD 6.6 ± 0.20
(6.3–6.8)
8.0 ± 0.27
(7.6–8.4)
EYDM 2.8 ± 0.09
(2.7–2.9)
3.1 ± 0.44
(2.4–3.9)
EYNO 1.9 ± 0.18
(1.6–2.0)
2.6 ± 0.27
(2.2–3.1)
ITNA 2.1 ± 0.08
(2.0–2.2)
2.6 ± 0.20
(2.3–3.0)
FAL 7.4 ± 0.34
(7.1–7.9)
9.4 ± 0.73
(8.1–10.6)
HAND 5.1 ± 0.05
(5.1–5.2)
7.0 ± 0.47
(6.4–7.9)
THBL 2.3 ± 0.13
(2.2–2.5)
3.0 ± 0.51
(1.8–3.6)
SW 6.0 ± 0.46
(5.5–6.8)
7.7 ± 0.48
(6.9–8.7)
TIBL/SVL 0.433 ± 0.028
(0.387–0.472)
0.443 ± 0.021
(0.407–0.466)
SW/SVL 0.261 ± 0.020
(0.230–0.289)
0.260 ± 0.016
(0.225–0.288)
HLSQ/SVL 0.328 ± 0.025
(0.289–0.350)
0.303 ± 0.010
(0.281–0.318)
FAL/SVL 0.318 ± 0.015
(0.301–0.338)
0.315 ± 0.026
(0.261–0.345)
HLSQ/HDWD 1.144 ± 0.067
(1.059–1.238)
1.136 ± 0.052
(1.036–1.238)
THBL/HAND 0.448 ± 0.024
(0.431–0.490)
0.425 ± 0.066
(0.281–0.507)
245
A new species of harlequin toad from Amazonian Ecuador
ered along the midpoint but open at the anterior and pos-
terior tips. Posteriorly, the frontoparietals are fused to the
prootics and exoccipitals. e vomers are small, edentate,
medially separated, crescent-shaped, and triradiate with an
anterior ramus, prechoanal ramus, and postchoanal ramus.
e anterior ramus is directed anterolaterally, pointed at
the anterior tip, and is roughly the same size as the pre- and
postchoanal ramus. e pre- and postchoanal rami form
the anteromedial margin of the choana. e prechoanal
ramus is directed towards the maxilla, and the postchoa-
nal ramus is directed posterolaterally. e postchoanal ra-
mus is fused to the overlying sphenethmoid. e neopala-
tine is elongate and triangular. It underlies, and is partially
fused to, the sphenethmoid and extends towards the max-
illa, nearly making contact with the preorbital process in
CJ and making contact in CJ . e neopalatine is
narrow and pointed at its medial tip, widens as it approach-
es the maxilla, and is cylindrical and pointed at its anterior
border. e parasphenoid has an inverted T-shape and the
cultriform process underlies the sphenethmoid in CJ
(but does not in CJ ). e cultriform process extends
beyond the midpoint of the orbit. e terminal end of the
cultriform process is rounded in shape. e parasphenoid
alae are directed posterolateral to the cultriform process
and are fused to the overlying otic capsule. e posterior
margin of the parasphenoid and a medial posterior process
are dicult to discern. e maxillary arcade is complete
and edentate. e paired premaxillae each possess a dorsal
alary process that is directed anterolaterally. e lower half
of the alary process is the same width as the upper half.
e pars palatina is biradiate with medial and lateral pro-
cesses. e lateral process is more than twice as long as the
medial process. e medial process is well developed and
triangular. A concave border is present where the lateral
and medial processes of the pars palatina come together.
e posterior tip of the pars dentalis is pointed and articu-
lates with the maxilla. e external surface of the maxil-
lae is smooth. e pars palatina, which extend along the
lingual margin of the maxilla, is narrow, and the posterior
half of this shelf articulates with the anterior ramus of the
pterygoid. e pars fascialis of the maxilla is directed; its
dorsomedial margin variably contacts, or nearly contacts,
the neopalatine, sphenethmoid, and maxillary process of
the nasal. e anterior end of the maxilla is truncate and
contacts the premaxilla. e posterior end of the maxil-
la is pointed and slightly overlaps the quadratojugal. e
quadratojugal is a small, slender, L-shaped bone that un-
derlies the ventral arm of the squamosal and contacts the
posterior process of the maxilla at is anterior margin. e
paired squamosals possess an otic ramus posterodorsally
and a ventral ramus. An anterodorsal zygomatic ramus is
not evident. e otic ramus is expanded dorsally and has a
broad articulation with the crista parotica of the otic cap-
sule, leaving only the posterior end of the prootic free. e
angle between the dorsal surface of the squamosal and the
anterior margin of the ventral ramus is nearly perpendicu-
Figure 3. Comparative skull osteology of western Amazonian Atelopus in dorsal, ventral and lateral views. Top row: A. spumarius sensu
stricto (female neotype, MNHNP 1979/8382, for details on scanning see L et al. in press). Bottom row: A. colomai (male CJ
8187 from Ecuador, Pastaza, Reserva Otoyacu). Not to scale.
246
A P et al.
lar. e ventral ramus is at and blade-like. A small squa-
mosal keel is present at the junction between the otic ra-
mus and the upper half of the ventral arm. e pterygoid
is triradiate, bearing anterior, medial, and posterior rami.
e anterior ramus articulates with the pars palatina of the
maxilla. e medial and posterior rami are of equal length
but the medial ramus is much broader. e medial ramus
nearly contacts the prootic. e posterior ramus is at and
blade-like. e palatoquadrate is cartilaginous and there-
fore not visible in the microCT dataset. e lower jaw is
composed of three ossied elements and Meckel’s carti-
lage, which is not visible in the microCT dataset. e men-
tomeckelian is the most anterior element that forms a car-
tilaginous symphysis at the midline of the jaws. Postero-
dorsally, the mentomeckelian is fused to the dentary. e
dentary is slender, thin, and edentate; this element overlays
the anterior half of the angulosplenial. e angulosplenial
is the largest mandible element and forms the lingual sur-
face of the lower jaws. e lower jaw length is . mm in CJ
and .mm in CJ , slightly shorter than the length
as the skull from the occipital condyle to the premaxilla.
Atelopus colomai sp. n. can be distinguished from A.spu-
marius sensu stricto by its smooth skull roof (vs. rugose;
Fig. ), having the frontoparietals completely fused (vs.
fused only over two posterior thirds of their total length;
Fig. ), the posterior end of the maxilla not being free (vs.
free), the pars glenoidalis and the pars jugularis of the
quadratojugal being subrectangular and elongated, respec-
tively (vs. pars glenoidalis thickend, pars jugularis short),
and a less straight orbital margin of the maxilla. However,
variation in these characters exists between the two popu-
lations of A. spumarius sensu stricto, which might be at-
tributed to imaging conditions or natural variation (L-
et al. in press).
e discriminatory power of osteological features needs
to be seen with some caution as intraspecic variation, sex-
ual dimorphism and taxonomic value remain poorly un-
derstood for harlequin toads and other anuran genera (see
e.g. K et al. ). For a comparative discussion of
osteological features in harlequin toads related to A. spu-
marius sensu stricto see Lötters et al. (in press).
Vocalization: e call recording (CJ ec.cj.aud.; Fonozoo
Sound Code ; Fig. ) contains pulsed calls with dura-
tion ± ms (– ms). Intervals between calls
last ± ms (– ms). Dominant frequency
is ± Hz (– Hz). Amplitude is generally as-
cending over the course of the call being interrupted by –
temporal decreases until it reaches its maximum followed
by a subsequent decrease over the last – pulses (Fig. ).
Call energy is also present in the frequency components re-
lated to the pulse rate, which shows weak modulation and a
minimal increase toward the end of the call. Each call con-
sists of ± pulses (–), that show a length of ± ms
(– ms) each. Pulses slightly increase in length over the
course of each call and are not modulated in frequency.
Figure 4. Oscillogram and corresponding spectrogram of two pulsed calls recorded from the holotype of Atelopus colomai sp. n.
247
A new species of harlequin toad from Amazonian Ecuador
Tadpole: D L () described a single
gastromyzophorous tadpole from Río Puyo in Provincia
Pastaza, Ecuador, under the name A. spumarius spumari-
us. We assign their specimen (KU ; Fig. ) to A.colo-
mai sp. n., based on the resemblance of living tadpoles
(Fig. ) reared from topotypic adults of A. colomai sp. n.
under laboratory conditions and based on being the only
harlequin toad known from the general area. Our exami-
nation of the tadpole in Stage corroborates most charac-
ters given by D L (). However, the il-
lustration provided with the description needs rectication
as the nostrils are actually located at the level of the eye,
and not below the eye. In addition, the upper beak appears
longer than the lower on the drawing, which is in error and
conicts the description. Our examination of the larva also
suggests the presence of a submarginal papilla on each side
at the end of the lowest row of labial teeth of the lower labi-
um (P), although this requires further corroboration due
to the poor status of preservation of the individual.
Based on this specimen and additional live specimens,
the tadpole of A. colomai sp. n. is characterised by: body de-
pressed in lateral view, attened ventrally, longer than wide,
widest between eye and spiracle, snout rounded in dorsal
view; eyes large, directed laterally to anterolaterally and
positioned dorsolaterally, not visible in ventral view; nares
ovoid, directed anterodorsally, situated closer to eye than to
anterior margin of snout, situated at the height of the lower
corner of eye; abdominal sucker circular to ovoid, anterior-
ly bordering the oral disc, posteriorly extended to height of
the opening of spiracle; spiracle sinistral, cylindrical, short
tube not attached to body wall, positioned ventrolaterally
and oriented posteriorly; anal tube medial, tubular; cau-
dal musculature massive, decreasing rapidly in distal half
of rounded tail; caudal ns low, dorsal n slightly round-
ed, highest in distal half of tail, not extending onto body;
margin of ventral n straight; oral disc ventral, almost as
wide as body, anteriorly and laterally bordered by one row
of short rounded papillae, submarginal papilla present on
each side of lower labium (P). Labial tooth row formula
/, tooth rows all abouth equal in length; upper beak of
jaw sheaths narrow, almost transverse; lower beak broadly
V-shaped, about twice as long as upper beak.
In life, back, anks and caudal muscle black to dark
brown with dense irregular pale brown to golden markings
that consist of aggregated minute dots, extending margin-
ally on transparent tail n; below, proximal half of body
whitish translucent with black dots, dots most dense be-
low suctorial disc, golden dots towards anks, distal half of
body black with scattered golden dots towards anks and
tail, tail cream with black and golden dots; pupil black sur-
rounded by cupreous to golden ring; towards metamor-
phosis, the pattern and coloration remain stable with a de-
crease of golden markings at late stages; limbs turn orange
in Stage (Fig. ). In preservative, the single specimen
used for the tadpole description by D L
() is entirely faded (Fig. ).
ere is limited variation in external morphology of
Atelopus tadpoles. e upper beak being shorter than the
lower is a larval character only known from the avescens-
spumarius clade (cf. M-L et al. , P-
G et al. , L et al. ). Tadpoles of
ve species from this clade have been described, A. aves-
cens, A. franciscus, the A. hoogmoedi complex, A. manau-
ensis and A. pulcher. Atelopus colomai sp. n. can be distin-
guished from them in having the lower beak about twice as
long as the upper. e upper beak is only slightly shorter
than the lower in the other species except A. pulcher (G-
, B et al. , L et al. ). It is
only one third to one fourth the length of the lower beak in
A. pulcher (L et al. a). From A. pulcher, the new
species is distinguished further by having the position of
the nostrils at about height of eye (vs. below height of eye;
L et al. a). In aspects of life coloration, larvae of
A. colomai sp. n. dier from those of A. avescens, A. fran-
ciscus and the A. hoogmoedi complex by being black to dark
brown with dense irregular pale brown to golden markings
that consist of aggregated minute dots (black with symmet-
rical golden spots anterior and posterior eye, at the base of
the tail and on tail n in A. avescens and A. franciscus; be-
low entirely translucent with few whitish cream dots in the
A. hoogmoedi complex; B et al. , L et al.
). e coloration of A.manauensis is barely described
(G under the name A.pulcher) so that compari-
son with the new species remains uncertain.
Figure 5. Preserved tadpole of Atelopus colomai sp. n. (KU 121385,
G Stage 37) in ventral, dorsal and lateral views (from top
to bottom). Scale bar 5 mm. Photos: KUDA 14472–14474, KU
Digital Archive.
248
A P et al.
Atelopus palmatus is the sole species with a described
tadpole which is not assigned to any of the dened clades
in L et al. (in press). Tadpoles of A. colomai sp. n.
can be distinguished from those of A. palmatus (cf. M-
-L et al. ) in having the upper beak shorter
than the lower (vs. longer), and in having golden tail pat-
tern in life (vs. lacking).
Distribution: Atelopus colomai sp. n. is known from the
Amazonian lowlands of Ecuador, ranging from ca. –
m a.s.l. Historically, A. colomai sp. n. was reported
from less than localities in Orellana and Pastaza prov-
inces, all situated between the Río Napo and the Río Pastaza
(Fig. ). However, specimens collected by R B-
in (NHRM RRB/..) originate from
Figure 6. Ontogenetic change in an individual of Atelopus colomai sp. n. from tadpole to subadult, reared under laboratory conditions
at CJ from parental stock collected at the type locality (specimens not preserved). Upper le: CJ (sc 10997), Stage 25, lateral, dorsal
and ventral views, total length = 8.3 mm; upper right: Stage 41, total length = 14.5 mm; Stage 42, total length = 14.1 mm; Stage 46,
71 days aer Stage 25, 6.8 mm SVL; bottom: subadult 244 days aer Stage 46, 18 mm SVL. Scale bar 10 mm. Photos by S
G S, CJ.
249
A new species of harlequin toad from Amazonian Ecuador
“Zamora”, which, by that time, was the name for large parts
of the southern Ecuadorian and adjacent Peruvian Ama-
zonian lowlands. us, the distributional limits south- and
eastward remain uncertain. roughout the last years
and despite intensive eorts to locate the species, A. colo-
mai sp. n. was conrmed only in few localities in Pastaza
province, NE and SE of Puyo.
Natural history: e species inhabits terra rme lowland
rain forest and can be found in close proximity to slow-
moving clear water streams and rivers (Fig. ). Adult males
were heard calling throughout the day with an activity
peak aer rainfalls, in the morning and late aernoon as
observed in August and September. Reproductive males
defend territories along the streams where entering males
are approached by the resident and pursued, accompanied
by intense calling, observed in the same months. Territo-
rial behavior was also observed by T et al. ().
Males were found perching on elevated positions such as
branches, fallen trees and rocks in the lower understory
up to . m above ground during the day. At night, males
were inactive, resting on leaves up to m above ground.
Females were found at night only, sleeping on leaves or in
bromeliad axils – m above ground. At the type locali-
ty, amplectant pairs were observed in March, April, July–
October and December (T et al. ; authors’ ob-
servations). Tadpoles were collected in July (D
L ) and metamorphs were observed in Sep-
tember suggesting that reproduction may take place year-
round.
Figure 7. Map of the central upper Amazon basin showing the known distributions of western Amazonian Atelopus related to A.spu-
marius. Red dots correspond to A. spumarius sensu stricto, yellow dots to A. pulcher, green dots to A. seminiferus, blue triangles to
A.colomai sp. n.; stars in the respective colors indicate type localities. White squares correspond to populations of unsolved taxonomic
status. Basemap shows major Amazonian river systems and relief.
250
A P et al.
Atelopus colomai sp. n. can be locally abundant with
up to individuals per person*hour found in nocturnal
visual encounter surveys at the type locality in September
. Northwest of El Triunfo, where some of the paratypes
were collected, the species is less common with individu-
als found in approx. person*hours in . T et
al. () observed individually identied specimens
through the course of a two-year capture-recapture study
in a population along Río Pucayacu, close to the type local-
ity. For information on syntopic anurans at the type locali-
ty see C et al. (). Monitoring studies conduct-
ed between and in Yasuní National Park (R
) identied this species as rare. Only ve individuals
were discovered at two locations within the park.
When disturbed, specimens of A. colomai sp. n. pre-
sent their colorful palms and soles while slowly walking
away (P et al. : Supplementary Video), simi-
lar to congeners from eastern Amazonia as described by
R et al. ().
Conservation status: We suggest listing Atelopus colomai
sp. n. as Endangered under criteria B and Ba,b of the
IUCN Red List of reatened Species as () it is known to
persist only at six threat dened localities, () all of which
are severely fragmented and () all of which are threatened
by ongoing deforestation, agriculture, road construction
and pollution (observed September ; August–Septem-
ber , June–August ). Including historic localities,
the EOO of the new species is roughly km² and AOO
km² while it is km² and km² for proven extant
localities only.
At the type locality, an almost complete deforestation
of remaining habitats occurred in late (Fig. ). Field-
work in August revealed the presence of few individu-
als in a remaining m wide fragment of forest along one
side of the stream. Deforestation has also been concluded
as the reason for declines in another population of the new
species by T et al. (). In the remaining four sites,
there is a constant reduction in the extent and quality of
remaining habitat. e species is known from one private
protected area, Reserva Otoyacu, and was formerly found
in Parque Nacional Yasuní from where it has not been con-
rmed for more than years.
Batrachochytrium dendrobatidis (Bd), considered a main
driver of extinctions in Atelopus (e.g. L M et al. ,
L et al. , L et al. ), has been repeatedly
detected in A. colomai sp. n. (L et al. ). However,
no declines related to Bd were noticed, similar to other low-
land Atelopus species (F et al. , , B
et al. ). T et al. () conducted a comprehensive
Bd sampling, documenting the pathogens’ absence in Atelo-
pus but its presence in syntopic anurans in their study site.
Atelopus colomai sp. n. is further likely to be threatened
by climate change in the near future, which might both im-
pact the species habitat as well as interfer and exacerbate
existing threats such as disease (L et al. ). e
species from the western Amazon are among the harlequin
toads likely to be most aected by climate change (L-
et al. ).
Atelopus colomai sp. n. is currently kept in a laboratory
survival-assurance colony in CJ, where it has already been
bred successfully in F generation. A backup colony is held
at Centro de Conservación de Anbios, Bioparque Amaru,
Cuenca.
Etymology: We dedicate this species to our friend and col-
league L A. C, who has continuously contribut-
ed to the study and protection of harlequin toads for dec-
ades. e specic name colomai is an eponym (i.e. a noun
in genitive case). As English common name, we propose
‘Coloma’s harlequin toad’.
Remarks: Paratypes CJ , , (Fig. S) remain
alive (and were measured in life) to build up a laboratory
survival-assurance colony.
Figure 8. Habitat of Atelopus colomai sp. n. at the type locality,
with deforestation in the foreground (August 2022). Adults were
found in the lower vegetation. Photo by A P.
Figure 9. Deforestation in the habitat of Atelopus colomai sp. n. at
the type locality with plantations of cassava on the right (August
2023). Photo by A B. Q R.
251
A new species of harlequin toad from Amazonian Ecuador
Acknowledgements
We thank A A, W L and M J
N for valuable discussions or for providing samples
to this study. We are thankful to S G S
(CJ) for providing photographs. P B, D
C, Y J F, A G, T-
H, L J, N K, Z L, D-
M, J R, J S, M T,
P V as well as Bloque de Conservación del Jam-
bato Amazónico, Centro Ecológico Zanja Arajuno, Equipo her-
petológico La Sapada Puyo, Finca La Argentina and Hyla Reserve
kindly supported eldwork. For providing information on and
granting access to material deposited in museum collections, we
are thankful to D F (AMNH), N G (ANSP),
S L and J S (BMNH), J A-
† (CBF), M Y M, M U, G-
L, D I and F P (DHEMCN-
INABIO), J M (FMNH), A A, K-
V. V and S P. G (IAvH), P M.
R-C† and M R-C (ICN), R
B, W E. D† and A M (KU), C-
A, F P. C, M I. D, S
R. M, L M, R F. Q and J A.
T (MUBI), R O (MHNC), A S
(MHNG), P-Y G and A O
(MNHNP), S S and G G (NHMW),
S O. K (NHRM), A W (USNM), G-
G. P (UTA), and W B and M
F (ZFMK). Our study was conducted under permits MAE-
DNBCM--, - IC-FAU-DNB/MA (collection) and
--FO-DPAP-MA, --FO-DPAP-MA (export), all
issued by the Ministerio de Ambiente de Ecuador. For funding,
we are grateful to a NSF Postdoctoral Research Fellowship in Bio-
logy (DBI ) to D.J.P. We thank an anonymous reviewer,
M V and the editor, J K, for valuable com-
ments on the manuscript.
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Material examined
Atelopus andinus: Peru: San Martín: upper Río Biabo valley,
AMNH A (paratype), A (holotype); Loreto: Río Pis-
qui, AMNH A (paratype); border area of San Martín and
Loreto: Río Cachiyacu (Tocachi), AMNH A , A , A
(paratypes).
Atelopus barbotini: French Guiana: Saül vicinities, MNHNP
/ (holotype), /–/, /, /,
/ (paratypes).
Atelopus avescens: French Guiana: Cayenne area, MNHNP
a (paralectotype), (lectotype), LG –, , , ZFMK
; Lac des Americains, ICN –; Kaw area, MHNG
., ., ., ZFMK , .
Atelopus franciscus: French Guiana: Crique Grégoire (Keren-
roch), MNHNP / (holotype), /–/ (para-
types).
Atelopus hoogmoedi complex: French Guiana: Haut Maroni:
Monts Atachi-Bacca, MNHNP A (holotype), A, A–
A (paratypes); Haut Oyapock, ICN , ; Guyana:
Karisparu, BM .–.; Potaro District, Echerak, BM
.–.; Upper Patavo River, BM ...; no
exact locality, BM ..–..; Iwokrama, MTD ; Mabu-
ra Hill, MTD ; Suriname: Brownsberg, AMNH , KU
, ; Kabo, MTD ; Nassau Mountains Plateau,
MTD (referable to A. hoogmoedi nassaui). Brazil: Amapá:
Serra do Navio, ZFMK , ; Pará: Sudam Floral Reserve,
km south-east of Santarém, KU –; no exact local-
ity, BM ...–..., NHMW /, /.
Atelopus loettersi: Peru: Cusco: km . from Puente Fortaleza
to Quincemil, MHNC (holotype), – (paratypes);
Madre de Dios: Tambopata, MUSM .
Atelopus manauensis: Brazil: Amazonas: N of Manaus,
NHMW /, /–.
Atelopus minutulus: Colombia: Meta: km – on road from
Guayabetal to Manzanares, ICN (holotype), –,
, –, (paratypes), IAvH-Am .
Atelopus palmatus: Ecuador: Pastaza: San Francisco on Río
Pastaza, NHRM NNN/.. ( syntypes); Río Negro,
SC.
Atelopus planispina: Ecuador: Napo: Mount Sumaco, near San
José Nuevo, MNCN – (syntypes); San Rafael, ICN .
Atelopus pulcher: Peru: San Martín: vicinities of Tarapoto,
KU –, , MUSM , , –, ZFMK
, –, , ; Loreto: Chayahuitas (“Chya-
vetas”), BM ... (lectotype), ...–... (pa-
ralectotypes).
Atelopus seminiferus: Peru: from between Balsapuerto and
Moyobamba, ANSP (holotype).
Atelopus spumarius sensu stricto: Peru: Loreto: Colonia (= Co-
lonia Ancón on the right shore of Río Zumún close to the junc-
ture with the Río Yaguasyacu, a tributary to the Río Ampiyacu),
MNHNP / (neotype); km NE Pebas, AMNH A –
, MUBI –.
Atelopus tricolor complex: Bolivia: La Paz: km from Car-
rasco, Serranía de Bellavista, CBF –; Río Ñeques, km ,
Charazani-Apolo road, Yungas de La Paz, CBF ; Pilon-La-
jas, CBF ; Cochabamba: Río Ronco, Chapare, CBF ;
“old” road from Villa Tunari to Cochabamba, Chapare, ZFMK
–; Peru: Cuzco: Marcapata valley, BM ...–
... (paralectotypes), MNHNP (paralectotype?),
ZFMK (lectotype); km south-west of Santa Isabel, Río
Cosñipata, KU ; Puno: vicinity of Juliaca, AMNH
(holotype of A. rugulosus).
Supplementary data
e following data are available online:
Supplementary Figure S1. Lateral, dorsal and ventral aspects of
living paratopotypes kept at CJ (in columns from le to right CJ
12425, 12406, 12409). Photos by S G S
(CJ).
