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Acta Herpetologica 16(2): 63-87, 2021
ISSN 1827-9635 (print) © Firenze University Press
ISSN 1827-9643 (online) www.fupress.com/ah
DOI: 10.36253/a_h-10742
A new species of the genus Noblella (Amphibia: Strabomantidae) from
Ecuador, with new information for Noblella worleyae
C R-P,,,,*, J M. G,, C K, D B-Z, M H-
, M C, D F. C-H,,
1 Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas & Ambientales COCIBA & Instituto de Diversidad Biológi-
ca Tropical iBIOTROP, Museo de Zoología/ Laboratorio de Zoología Terrestre, Campus Cumbayá, Quito, Ecuador
2 Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales COCIBA, Instituto BIOSFERA, Campus
Cumbayá, Quito, Ecuador
3 Instituto Nacional de Biodiversidad INABIO, Unidad de Investigación, Quito, Ecuador
4 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto, Instituto de Ciências Agrári-
as de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal
5 Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales COCIBA, Laboratorio de Biología Evolutiva,
Campus Cumbayá, Quito, Ecuador
6 Zoologisches Forschungsmuseum Alexander Koenig ZFMK, Leibniz-Institut zur Analyse des Biodiversitätswandels, Bonn, Germany
7 Southern Cross University, School of Environment, Science and Engineering, Lismore, Australia
8 University of New Brunswick, Department of Biology, Fredericton, Canada
*Corresponding autor. E-mail: creyesp@usfq.edu.ec
Submitted on: 2021, 31st March; revised on: 2021, 22nd July; accepted on: 2021, 23rd July
Guest Editor: Aaron M. Bauer
Abstract. We describe a new species of terrestrial-breeding frog of the genus Noblella from the northwestern slopes of
the Andes of Ecuador, in the province of Pichincha, Ecuador, and report a new locality for the recently described N.
worleyae. We include a detailed description of the osteology of both species and discuss their phylogenetic relation-
ships. e new species is dierentiated from other species of Noblella by having discs of ngers rounded, without
papillae; distal phalanges only slightly T-shaped; toes slightly expanded and rounded distally, without papillae; dorsum
uniform brown with irregular suprainguinal dark brown marks; venter yellowish cream, ventral surfaces of legs and
thighs reddish to brownish cream; and dark brown throat. e new locality for N. worleyae is located in Los Cedros
Reserve, an area highly threatened by mining. We highlight the importance of protecting endemic species of small
vertebrates in northwestern Ecuador.
Keywords. Frog, Los Cedros Biological Reserve, endemism, Imbabura, Mindo, Pichincha, phylogeny.
INTRODUCTION
e amphibian diversity in the tropical Andes is
outstanding (Duellman, 1988; Myers et al., 2000; Hut-
ter et al., 2013, 2017). Each year, several species are
described from montane forests of this biodiversity hot-
spot (e.g., Rojas-Runjaic et al., 2018; Guayasamin et al.,
2019; Paez and Ron, 2019; Reyes-Puig et al., 2019b; San-
ta-Cruz et al., 2019; Yanez-Muñoz et al., 2019; Acevedo
et al., 2020; Ospina-Sarria et al., 2020; Lehr et al., 2021).
Most described species from Ecuador belong to the
hyper-diverse genus Pristimantis (Paez and Ron, 2019;
Reyes-Puig et al., 2020a), but diversity in other anuran
taxa has also increased considerably (e.g., Osornophryne,
Hyloscirtus, Noblella, Centrolenidae; Mueses-Cis-
neros et al., 2010; Cisneros-Heredia and Gluesenkamp,
64 C. Reyes-Puig et alii
2010; Yánez-Muñoz et al., 2010a; Páez-Moscoso and
Guayasamin, 2012; Almendáriz et al., 2014; Guayasamin
et al., 2017a, 2019; Reyes-Puig et al., 2019c).
Terrestrial-breeding frogs of the genus Noblella
Barbour 1930 are minute-size anurans (SVL < 22 mm),
morphologically dierentiated by having terminal discs
on digits not or barely expanded, discs and circum-
ferential grooves present distally (except in N. duellm-
ani), terminal phalanges narrowly T-shaped, pointed
tips of at least Toes III‒IV, and an inner tarsal tubercle
(De La Riva et al., 2008; Hedges et al., 2008; Duellman
and Lehr, 2009). However, phylogenetic relationships
of Noblella are not fully resolved and its monophyly
is uncertain (De la Riva et al., 2017; Santa-Cruz et al.,
2019). As currently dened, Noblella includes 16 spe-
cies, fourteen distributed in the Andes of Ecuador, Peru,
and Bolivia, and two (N. losamigos and N. myrmecoides)
in the Amazonian lowlands from southeastern Colom-
bia, Ecuador, Peru, Bolivia, and western Brazil (Frost,
2021). During the last 15 years, the number of species in
the genus has doubled; and four new species have been
described since 2019 (Catenazzi and Ttito, 2019; Reyes-
Puig et al., 2019c, 2020b; Santa-Cruz et al., 2019). Cur-
rently, the total number of species of the genus Noblella
is 16, distributed in ten species in Peru, seven in Ecua-
dor, three in Bolivia, and one in Colombia and Brazil
(Frost, 2021).
Andean species of the genus Noblella show a high
level of endemicity, with very restricted distributions.
While some species of Noblella may apparently be
able to survive in environments modied by humans
(e.g., N. duellmani, N. losamigos, N. lochites, N. nature-
trekii; Duellman and Lehr, 2009; Reyes-Puig et al.,
2019c; Santa-Cruz et al., 2019); most species (e.g., N.
coloma, N. heyeri, N. personina, N. pygmaea; Lynch,
1986; Guayasamin and Terán-Valdez, 2009; Harvey et
al., 2013) seem to depend on undisturbed forest. ree
species of Noblella have been described from western
Ecuador, all from mature mountain forests: Noblella
heyeri (Lynch, 1986) occurs in southwestern Ecua-
dor and extreme northwestern Peru; Noblella coloma
Guayasamin and Terán-Valdez, 2009 is known from
its type locality and surroundings (Rio Guajalito and
Chiriboga area; Ron et al., 2019); and Noblella worleyae,
a recently described species is known just from seven
specimens, all found in mature forest in the Río Man-
duriacu Reserve, province of Imbabura, Ecuador (Reyes-
Puig et al., 2020b).
While the Ecuadorian Andes have suered serious
habitat destruction and fragmentation caused by expan-
sion of deforestation, agriculture, mining, among others
(Castellanos et al., 2011; Roy et al., 2018; Guayasamin et
al., 2019; Lessmann et al., 2019; Ortega et al., 2021), there
are still some areas with mature forests that have not
been exploited due to their complex topography, dicult
access, private protection, or preservation for touristic
activities. Unfortunately, all such sites are under strong
anthropogenic pressure, including mining concessions
and the expansion of agricultural boundaries, among
others (Cuesta et al., 2017; Roy et al., 2018; Guayasamin
et al., 2019; Ortega et al., 2021). ese privileged areas
have proven to keep an extremely high cryptic diver-
sity of small vertebrates and contain the last remnant
populations of numerous threatened species (Cisneros-
Heredia and Yanez-Muñoz, 2010; Reyes-Puig et al.,
2010, 2019a, 2019b; Yánez-Muñoz et al., 2010b, 2018;
Guayasamin et al., 2018, 2019, 2020; Sánchez-Nivicela et
al., 2018; Barrio-Amorós et al., 2020).
During the last ve years, we have carried out sur-
veys on the western slopes of the Andes in the provinces
of Imbabura and Pichincha, Ecuador. As a result of this
continuous eort, we found a new species of leaf-litter
frog of the genus Noblella, which we describe herein
based on a combination of morphological, molecular,
and osteological features. We also document new infor-
mation on distribution, external morphology and osteol-
ogy for the recently described Noblella worleyae, infor-
mation that was not described in detail in the original
description. We also include intraspecic variation that
will allow complete full with members of the same genus
in the future.
MATERIALS AND METHODS
Taxonomy
We followed the family taxonomy proposed by Heinicke et
al. (2018) and, also we revised De la Riva et al. (2017) and Bar-
rietos et al. (2021). For identifying species, we assumed the uni-
ed species concept (De Queiroz, 2005, 2007). Information for
species comparisons was extracted from the original descrip-
tions and cited once at the beginning of the comparison.
Study area and eldwork
Over the last three years (i.e., 2018–2020), we have car-
ried out eld surveys at several localities in montane forests of
northwestern Ecuador, mainly in the provinces of Imbabura
and Pichincha. Specimens of two dierent species of Noblella
were found in Mindo (province of Pichincha) and Los Cedros
Biological Reserve (province of Imbabura). Mindo is a small
town renowned for its adventure and nature-based touris-
tic activities; thus the area has numerous reserves that protect
cloud forests (Arteaga-Navarro et al., 2013). Los Cedros Bio-
logical Reserve is a protected area that contains 6,879 hectares
65
A new species of the genus Noblella from Ecuador
of premontane humid tropical forest and cloud mountain forest.
is reserve is located south of the Cotocachi-Cayapas Ecologi-
cal Reserve (state protected area), and is also recognized by its
endemic microfauna (Hutter and Guayasamin, 2015). Collected
specimens were euthanized with benzocaine, xed in 8% forma-
lin, and preserved in 75% ethanol. Liver and leg muscle tissue
samples were collected from all individuals prior to preserva-
tion. Tissues were preserved in 95% ethanol and stored at -20°C
at the Laboratorio de Biología Evolutiva USFQ. Specimens were
deposited in the Museo de Zoología, Universidad San Francisco
de Quito, Ecuador (ZSFQ).
DNA extraction, amplication, and sequencing
We obtained new DNA sequences of Noblella sp. n ov.
(ZSFQ 050–051). DNA was extracted from muscle or liver tis-
sue following the protocol by Peñael et al. (2019). Standard
polymerase chain reaction (PCR) was performed to amplify a
fragment of the mitochondrial gene 16S rRNA, using a combi-
nation of the following primers: 16L10, 16H36E, 16L34, 16H47
(Heinicke et al., 2007). Amplicons were sequenced in both
directions by the Macrogen Sequencing Team (Macrogen Inc.,
Seoul, Korea).
The new sequences were assembled and edited with
Geneious 7.1.7 (GeneMatters Corp). After assemblage, the
sequences were combined with sequences from GenBank for
all species of Noblella and representatives of the genera within
the Terrarana clade (sensu Hedges et al., 2008), including Bary-
cholos Heyer 1969, Bryophryne Hedges, Duellman & Heinicke
2008, Craugastor Cope 1862, Haddadus Hedges, Duellman &
Heinicke 2008, Holoaden Miranda-Ribeiro 1920, Ischnocnema
Reinhardt & Lutken 1862, Lynchius Hedges, Duellman & Hein-
icke 2008, Niceforonia Goin & Cochran 1963, Oreobates Jimé-
nez de la Espada 1872, Qosqophryne Catenazzi, Mamani, Lehr,
von May 2020, Phrynopus Peters 2873, Pristimantis Jiménez de
la Espada 1870, Psychrophrynella Hedges, Duellman & Heinicke
2008, and Microkayla De la Riva, Chaparro, Castroviejo-Fisher,
Padial 2017. GenBank codes are shown in our inferred phyloge-
netic tree (Fig. 1).
Phylogenetic analyses
Phylogenetic relationships were inferred using maximum
likelihood as the optimality criterion. e nal matrix, with
52 terminals, was aligned with MAFFT v.7 (Multiple Align-
ment Program for Amino Acid or Nucleotide Sequences: http://
mafft.cbrc.jp/alignment/software/), with the Q-INS-i strat-
egy. MacClade 4.07 (Maddison and Maddison, 2005) was used
to visualize the alignment, which contained a total of 492 bp.
Phylogenetic analyses were performed under the ML criteria
in GARLI 2.01 (Genetic Algorithm for Rapid Likelihood Infer-
ence; Zwickl, 2006) for the mitochondrial gene 16S. GARLI uses
a genetic algorithm that nds the tree topology, branch lengths,
and model parameters that maximize lnL simultaneously
(Zwickl, 2006). Individual solutions were selected aer 10,000
generations with no signicant improvement in likelihood, with
the signicant topological improvement level set at 0.01. en,
the nal solution was selected when the total improvement in
likelihood score was lower than 0.05, compared to the last solu-
tion obtained. Default values were used for other GARLI set-
tings, as per recommendations of the developer (Zwickl, 2006).
Bootstrap support was assessed via 1,000 pseudoreplicates
under the same settings used in tree search. Pairwise genetic
distances between species (uncorrected-p) for gene 16S were
calculated with PAUP 4a (Swoord et al., 1996).
External morphology
Diagnosis and description of the new species follow for-
mats proposed by Duellman and Lehr (2009) and Lynch and
Duellman (1997). For comparisons, we examined specimens
of other species of Noblella (see Appendix I). We followed the
sequence of characters proposed by Guayasamin and Terán-Val-
dez (2009). We measured preserved specimens using digital cal-
ipers to the nearest 0.01 mm. ese measurements are: snout to
vent length (SVL), from the tip of the snout to the cloaca; head
length (HL), measured from tip of snout to anterior edge of
tympanum; head width (HW), measured at midorbital region;
horizontal diameter of the eye (ED); eye–nostril distance (EN),
from anterior ocular angle to posterior edge of nostril; horizon-
tal diameter of tympanum (TD); minimum interorbital distance
(MIOD); minimum eyelid width (MWE); hand length (LH),
from posterior edge of palmar tubercle to tip of third digit;
shank length (LS), from the tip of the ankle to the knee; and
foot length (LF), from posterior edge of external metatarsal
tubercle to tip of Toe IV. We determined sexual maturity by the
presence of vocal slits or extended vocal sacs in males and by
the presence of eggs or convoluted oviducts in females. Detailed
illustrations of the head, hands and feet were done with Adobe
InDesign ©.
Osteology
Osteological descriptions were based on one specimen
of the new species (ZSFQ 050) and one of Noblella worleyae
(MZUTI 1709). Both specimens were scanned using a high-
resolution micro-computed tomography (micro-CT) desktop
device (Bruker SkyScan 1173, Kontich, Belgium) at the Zoolo-
gisches Forschungsmuseum Alexander Koenig (ZFMK, Bonn,
Germany). To avoid movements during scanning, specimens
were placed in a small plastic container and mounted with sty-
rofoam. Acquisition parameters comprised: An X-ray beam
(source voltage 43 kV and current 114 µA) without the use of
a lter; 800 projections of 500 ms exposure time each with a
frame averaging of 5 recorded over a 180° continuous rotation
(rotation steps of 0.3 degrees), resulting in a scan duration of
49 min; a magnication setup generating data with an isotrop-
ic voxel size of 19.16 µm (MZUTI 1709) and 14.55 µm (ZSFQ
050), respectively. The CT-dataset was reconstructed with
N-Recon soware (Bruker MicroCT, Kontich, Belgium) and
rendered in three dimensions using CTVox for Windows 64 bits
version 2.6 (Bruker MicroCT, Kontich, Belgium). Osteological
66 C. Reyes-Puig et alii
terminology follows Trueb (1973), Duellman and Trueb (1994),
Fabrezi and Alberch (1996), Guayasamin and Terán-Valdez
(2009), Scherz et al. (2017), and Suwannapoom et al. (2018).
Cartilage structures were omitted from the osteological descrip-
tions because micro-CT does not render cartilage.
RESULTS
Phylogenetic relationships and genetic distances (Fig. 1)
The inferred phylogeny shows that the new spe-
cies described herein is part of a clade composed of
taxa distributed along the western slopes of the Ecua-
dorian Andes. is clade is composed by the new spe-
cies Noblella sp. nov., Noblella coloma Guayasamin
and Terán-Valdez, 2009, and N. worleyae Reyes-Puig,
Maynard, Trageser, Vieira, Hamilton, Lynch, Culebras,
Kohn, Brito and Guayasamin, 2020. Uncorrected p
genetic distances are as follow: N. coloma (QCAZ 40579)
and the new species (ZSFQ 050) = 5.1%; N. coloma
(QCAZ 40579) and N. worleyae (ZSFQ 550–551) = 8.3%;
N. worleyae (ZSFQ 550–551) and the new species (ZSFQ
050) = 1.2%.
Generic placement
We place the new species in the genus Noblella
based on morphological and molecular evidence (Fig.
1). Morphologically, we assign the new species to the
genus Noblella, as dened by Hedges et al. (2008), based
on possession of the following traits: head not wider
than body; cranial crests absent; tympanic membrane
differentiated (undifferentiated in N. duellmani, N.
naturetrekii and N. madreselva); dentigerous processes
of vomers absent; terminal discs on digits not or barely
expanded; discs and circumferential grooves present dis-
tally (absent in N. duellmani); terminal phalanges nar-
rowly T-shaped; Finger I shorter than, or equal in length
to, Finger II; Finger IV containing two phalanges in
Noblella carrascoicola (De la Riva and Köhler, 1998), N.
lochites (Ly nch, 1976b), N. losamigos (Santa-Cruz et al.,
2019), N. myrmecoides (Ly nch, 1976b), N. naturetrekii
(Reyes-Puig et al., 2019c), and N. ritarasquinae (Köhler,
2000) and three phalanges in N. coloma (Guayasamin
and Terán-Valdez, 2009), N. duellmani (Lehr, Agu ilar,
and Lundberg, 2004), N. heyeri (Lynch, 1986), N. sp.
nov., N. lynchi (Duellman, 1991), N. madreselva (Cat-
enazzi, Uscapi, and von May, 2015), N. personina (Har-
vey, Almendáriz, Brito-M., and Batallas-R., 2013), N.
peruviana (Nob le, 19 21), N. pygmaea (Lehr and Cat-
enazzi, 2009), and N. thiuni (Catenazzi and Ttito, 2019);
Toe III shorter than Toe V (except in N. naturetrekii and
N. worleyae); tips of at least toes III–IV acuminate; sub-
articular tubercles not protruding; dorsum pustulate or
shagreen; venter smooth; SVL less than 22 mm.
Systematic accounts
Noblella mindo new species.
Noblella coloma Arteaga et al. (2013).
Figs. 2–8
LSID urn:lsid: lsid:zoobank.org:act:3B7741EF-BF26-
4589-B231-73F198AA1218
Proposed standard English name. Mindo Leaf Frog
Proposed standard Spanish name. Rana Noble de Mindo
Holotype. ZSFQ 050 (Fig. 2–6), adult female, col-
lected in El Cinto, 11 Km E from Mindo town, Mindo
(0.09022°S, 78.818581°W; 1,673 m; Fig. 2), province of
Pichincha, República del Ecuador, by Melissa Costales,
Matthijs Hollanders and Emilia Peñaherrera on 08 July
2017.
Paratypes (2 females, 2 males). Adult males (ZSFQ 049,
051) and adult females (ZSFQ 304–305) collected at the
type locality (same data as holotype), by Melissa Cos-
tales on 04 October 2015.
Etymology
e specic name “mindo” is a word of unknown
meaning in Panzaleo, an extinct pre-Columbian lan-
guage of northern Ecuador (Jijón y Caamaño 1940). It is
used as a noun in apposition, and alludes to the valley
of Mindo, where the type locality of the new species is
located. e remnant forests of this emblematic valley
protect several species of endemic amphibians and rep-
tiles such as Pristimantis mindo, Noblella mindo, and
Anolis proboscis.
Diagnosis
Noblella mindo sp. nov. (Figs. 3–8) presents the fol-
lowing characteristics: (1) skin of dorsum nely sha-
green, skin on venter smooth, discoidal fold slightly
dened, discoidal and thoracic folds absent; (2) tympan-
ic annulus and membrane visible externally, supratym-
panic fold inconspicuous (Figs. 3, 4); (3) snout short
(eye-to-nostril distance 57% of eye diameter), rounded
in dorsal and lateral views (Fig. 3); (4) eyelids without
tubercles; (5) dentigerous processes of vomers absent; (6)
vocal slits and sac present, nuptial pads not visible; (7)
ngers not expanded distally, nger tips rounded, with-
out papillae (Fig. 3); Finger I shorter than Finger II (Fig.
3); (8) distal phalanges slightly T-shaped, phalangeal for-
mula of hands: 2-2-3-3 (Fig. 7); (9) supernumerary pal-
mar tubercles present (slightly visible) mostly at the base
67
A new species of the genus Noblella from Ecuador
of the digits, ulnar tubercles diminutive and rounded,
subarticular tubercles rounded; circumferential grooves
absent; (10) one tarsal tubercle elongated and subconi-
cal (Fig. 3); two prominent metatarsal tubercles (inner
tubercle 3–4 times size of external); toes slightly expand-
ed and rounded distally, without papillae; (11) Toe V
shorter than Toe III, supernumerary plantar tubercles
absent, distal portions of circumferential grooves not
visible; (12) phalangeal formula of feet: 2-2-3-4-3 (Fig.
7); (13) in life, uniform brown dorsum, cream middor-
sal, longitudinal line distinct and present in all individu-
als, dark brown suprainguinal marks, white rictal gland,
Noblella thiuni (MK072732, CORBIDI 18723)
Noblella madreselva (MN056356, CORBIDI 15770)
Noblella madreselva (MN064565, CORBIDI 15769)
Psychrophrynella usurpator (KY652662, AC186-09)
Psychrophrynella usurpator (KU884559, CORBIDI 16495)
Psychrophrynella glauca (MG837565, CORBIDI 18729)
Noblella losamigos (MN100040, —)
Noblella losamigos (MN366392, MVZ 292687)
Noblella losamigos (KY652644, RvM3-12)
Noblella pygmaea (KY652645, MUSM 24536)
Noblella sp (KY652646, MUSM 27582)
Bryophryne bustamantei (KT276293, MHNC6019)
Bryophryne cophites (EF493537, KU173497)
Bryophryne phuyuhampatu (MF419259, —)
Bryophryne tocra (MF186398, MUBI5419)
Barycholos pulcher (EU186709, KU 217781)
Barycholos ternetzi (KU495150, MALCX17P10)
Noblella coloma (MT710932, QCAZ 40579)
Noblella coloma (MT710931, QCAZ 32702)
Noblella mindo sp. nov. (MT710934, ZSFQ-050)
Noblella mindo sp. nov. (MT710935, ZSFQ-051)
Noblella worleyae (MT710935, ZSFQ 551)
Noblella worleyae (MT710934, ZSFQ 550)
Noblella worleyae (MT710933, ZSFQ 2502)
Noblella lochites (EU186699, KU177356)
Noblella personina (MK838465, QCAZ 58818)
Noblella heyeri (JX267463, QCAZ 31471)
Noblella myermecoides (JX267464, QCAZ 40180)
Noblella naturetrekii (MK838462, DHMECN 13307)
Ischnocnema bilineata (JX267324, MNRJ 46476)
Holoaden bradei (EF493366, USNM 207945)
Holoaden luederwaldti (EU186710, MZUSP 131872)
Ischnocnema colibri (MH538418, CFBH_41810)
Ischnocnema parnaso (MH538421, CFBH 41812)
Niceforonia brunnea (EF493357, KU178258)
Niceforonia peraccai (EF493710, KU178266)
Niceforonia dolops (EF493394, —)
Lynchius parkeri (MK423937, QCAZ 61015)
Lynchius simmonsi (JF810005, QCAZ 41640)
Oreobates barituensis (JF810000, MCN1360)
Oreobates quixensis (QCAZ 31186, JF810003)
Phrynopus barthlenae (AM039653, —)
Phrynopus inti (MF651906, UMMZ 245218)
Phrynopus kauneorum (AM039655, —)
Phrynopus sp (AM039660, —)
Phrynopus pesantesi (AM039656, —)
Phrynopus mariellaleo ( MH538299, CORBIDI 11658)
Pristimantis cedros (KT210170, MZUTI 1710)
Pristimantis ornatissimus (KU574613, MZUTI 4329)
Pristimantis subsigillatus (KU999217, MZUTI 1999)
Haddadus aramunha (KF740845, UFBA 283)
Haddadus binotatus (KF740846, USP/TCX51ST09)
Craugastor aenigmaticus (MK211617, UCR 22737)
Craugastor tarahumaraensis (EU186702, —)
Craugastor_longirostris (EF493395, KU 177803)
0.05 length units
Microkayla +
Qosqophryne Clade
“Noblella”
Southern Clade
Noblella
Northern Clade
58
95
98
58
74
76
100
59
93
99
100
95
74
80
96
98
84
53
70
68
50
78
100
68
76
56
95
78
67
68
100
Fig. 1. Phylogeny of Noblella (light gray boxes) showing the relationships of N. mindo sp. nov. e phylogeny was inferred based on mito-
chondrial (16S) DNA sequences (16S; 52 terminals, 492 bp) and under the Maximum likelihood criterion. For each individual, museum
catalog number or, if unavailable, GenBank accession number is shown.
68 C. Reyes-Puig et alii
anks with dark brown band narrowing towards groin,
dotted with white, light groin, with low concentration
of melanophores, dark brown throat, chest and ventral
surfaces of arms with a white cross formed by a longi-
tudinal, ne line running from chin to chest crossing a
similar line departing from midventral surface of each
forelimb, yellowish-cream venter, reddish-copper iris
with minute turquoise scattered dots (Fig. 4); (14) SVL in
adult males 16.5–17.0 mm (mean 16.7 mm, n = 2), SVL
in adult females 18.3–19.5 mm (mean 19.0 mm, n = 3).
Comparisons (Fig. 6, Table 1)
Noblella mindo sp. nov. diers from its congeners by
the presence of rounded ngertips, without papillae; dis-
tal phalanges slightly T-shaped; toes slightly expanded
and rounded distally, without papillae; dorsum uniform
brown with middorsal cream line, suprainguinal marks
dark brown, rictal gland white, light groin, throat and
chest dark brown with white cross, and venter yellow-
ish cream. Noblella mindo sp. nov. is most similar and
closely related to N. coloma and N. worleyae (Fig. 1), but
they dier as follows (characters of N. mindo sp. nov. in
parentheses): Noblella coloma has all nger tips acumi-
nate (all rounded), dark middorsal line (light), dark ric-
tal gland (white), orange to reddish-venter (yellowish-
cream), dark groin (light), uniform dark brown throat,
chest and ventral surfaces of arms (dark brown with
white cross), ulnar tubercles absent (diminutive and
rounded), and smaller body size of 16.0 mm SVL in adult
female (18.3–19.5 mm SVL in adult females); the new
species (characters of N. mindo sp. nov. in parentheses)
is distinguished from Noblella worleyae has nger tips
slightly acuminate on Fingers I and IV and acuminate on
Fingers II and III (rounded), T-shaped distal phalanges
(slightly T-shaped), prootic and exoccipital fused to form
otoccipital (separated), sphenethmoid well-ossied and
ventrally fused at midline (moderately ossied, ventrally
fused at midline in posterior half and separated in anteri-
or half). For more comparison’s information see Table 1.
Noblella mindo sp. nov. has three phalanges on Fin-
ger IV like N. duellmani (Lehr, Aguilar and Ludenberg,
2004), N. heyeri (Lynch, 1986), N. lynchi (Duellman,
1991), N. madreselva (Catenazzi, Uscapi and von May,
2015), N. personina (Harvey, Almendáriz, Brito, and
Batallas, 2013), N. peruviana (Noble, 1921), N. pygmaea
(Lehr and Catenazzi, 2009), and N. thiuni (Catenazzi
and Ttito, 2019), but they dier as follows (characters
of N. mindo sp. nov. in parentheses): Noblella duellmani
has dorsal skin pustular (nely shagreen), tympanum
membrane and annulus absent (present), upper eyelid
bearing small tubercles (absent), ulnar tubercles coa-
lesced into low folds (diminutive and round, not form-
ing a fold), outer edge of tarsus bearing row of low and
elongate tubercles (absent), tips of Fingers I-II slightly
expanded and tips of Fingers III–IV slightly acuminate
(all nger tips rounded), venter brown with tan mottling
(yellowish-cream), and larger body size of 20.0 mm SVL
in adult female (18.3–19.5 mm SVL in adult females);
Noblella heyeri has dorsal skin weakly pustulate (nely
shagreen), snout subacuminate in dorsal view (round),
ulnar tubercles distinct and round (diminutive, round),
toe tips slightly acuminate (round), venter brown with
cream eck (yellowish-cream), and smaller body size of
13.1–15.9 mm SVL in adult females (18.3–19.5 mm SVL
in adult females); Noblella lynchi has dorsal skin pus-
tular (nely shagreen), snout subacuminate in dorsal
view (round), ulnar tubercles coalesced into low folds
(diminutive and rounded, not forming a fold), outer
edge of tarsus bearing row of low and elongate tubercles
(absent), toe tips weakly acuminate (round), and venter
brown with ne cream ecks (yellowish cream); Noblella
madreselva has dorsal skin with small tubercles (nely
shagreen), tympanic membrane not dierentiated and
tympanic annulus barely visible below skin (well-dier-
entiated), upper eyelid with minute tubercles (absent),
toe tips weakly acuminate (rounded), venter black with
large and irregularly shaped white mark (yellowish-
cream), and smaller body size of 17.6 mm SVL in adult
female (18.3–19.5 mm SVL in adult females); Noblella
Fig. 2. Distribution of Noblella mindo sp. nov and N. worleyae in
Ecuador.
69
A new species of the genus Noblella from Ecuador
personina has dorsal skin smooth with pustules (nely
shagreen), snout subtruncate in prole (round), nger
and toe tips acuminate with papillae (round, lacking
papillae), venter white (yellowish-cream), and smaller
body size of 15.6–17.9 mm SVL in adult females (18.3–
19.5 mm SVL in adult females); Noblella peruviana has
tympanic membrane not dierentiated (dierentiated),
toe tips slightly acuminate (round), and venter tan (yel-
lowish cream); Noblella pygmaea has dorsal skin tuber-
cular (nely shagreen), thoracic fold present (absent),
dorsolateral fold on anterior half of body present
(absent), upper eyelid bearing small tubercles (absent),
minute tubercle on heel present (absent), toe tips pointed
(round), venter pale grayish brown (yellowish-cream),
and smaller body size of 11.3–12.4 mm SVL in adult
females (18.3–19.5 mm SVL in adult females); Noblella
thiuni has thin dorsolateral folds visible on anterior half
of body (absent), tympanic membrane not dierentiated
(dierentiated), ngertips bulbous (round), ulnar tuber-
cles absent (present, diminutive and round), venter cop-
per reddish with a profusion of silvery spots (yellowish
cream), and smaller body size of 11.0 mm SVL in male
(16.5–17 mm in adult females). Noblella carrascoicola
(De la Riva and Köhler, 1998), N. lochites (Lynch, 1976b),
N. losamigos (Santa-Cruz et al., 2019), N. myrmecoides
(Lynch, 1976b), N. naturetrekii (Reyes-Puig et al., 2019c),
and N. ritarasquinae (Köhler, 2000) are easily dierenti-
ated from N. mindo sp. nov. by having two phalanges on
Finger IV instead of three.
Description of the holotype
Adult female (ZSFQ 050); head narrower than body,
its length 40.8% of SVL; head longer than wide; head
width 31.1% of SVL; snout round in dorsal and lateral
Fig. 3. Noblella mindo sp. preserved holotype, ZSFQ 050, adult
female, SVL = 18.3 mm. (A) Foot in ventral view. (B) Hand in ven-
tral view. (C) Head in dorsal view. (D) Head in lateral view. Illus-
trations by Carolina Reyes-Puig.
Fig. 4. Color patterns of Noblella mindo sp. nov. in life. (A, C) Dorso-lateral and ventral patterns of holotype, ZSFQ 050, adult female, SVL
= 18.3 mm. (B, D) Dorsolateral and ventral patterns of paratype, ZSFQ 051, adult male, SVL = 16.9 mm. Photos by Matthijs Hollanders.
70 C. Reyes-Puig et alii
views; canthus rostralis straight, slightly concave in pro-
le; loreal region slightly concave; upper eyelid 45.6% of
interorbital distance; eye-nostril distance 54.8% of eye
diameter; tympanum visible externally, tympanic mem-
brane dierentiated from surrounding skin; supratym-
panic fold indistinct. Dentigerous processes of vom-
ers absent and vomerine teeth absent; choanae laterally
oriented; tongue longer than wide, elongated, partially
notched posteriorly.
Skin of dorsum nely shagreen, evident tubercles
absent; skin on anks smooth; venter smooth; discoidal
fold slightly visible, dorsolateral folds and thoracic folds
absent; diminutive rounded ulnar tubercles; palmar
tubercle oval, about 2 times the size of the thenar tuber-
cle; supernumerary palmar tubercles present, mainly at
the base of the digits; proximal subarticular tubercles
prominent, rounded; phalangeal formula 2-2-3-3; ngers
not expanded distally, nger tips rounded, circumferen-
tial grooves absent; relative lengths of ngers: I < II < IV
< III; forearm lacking evident tubercles.
Hindlimb lengths moderate, tibia length 49.3%
of SVL; foot length 46.1% of SVL; dorsal surfaces of
hindlimbs shagreen; tubercles on the heel absent; one
prominent elongated tarsal tubercle on ventral surface
of tarsus; two metatarsal tubercles, inner elongated con-
spicuous, outer subconical; proximal and distal subar-
ticular tubercles well-dened; supernumerary tubercles
absent. Toes slightly expanded and rounded distally;
distal portions of circumferential grooves not visible;
phalangeal formula 2-2-3-4-3; relative lengths of toes: I <
II < V < III < IV.
Measurements of holotype (in mm)
SVL= 18.3, HL= 7.5, HW= 5.7, ED= 2.4, EN= 1.3,
MWE= 1.5, TD= 0.9, MIOD= 3.4, LH= 3.4, LS= 9.0, LF=
8.4. For measurements of the type series (mm) see Table 2.
Color of holotype in life (Fig. 4)
Dorsum brown, grayish brown towards the anks;
well-defined cream middorsal stripe, extending from
interparietal region to cloaca and continuing along pos-
terior surfaces of hindlimb. Loreal region black, extend-
ing as homogeneous dark band to upper insertion of arm
and into body anks, narrowing towards groin and limit-
ed dorsally with a lighter brown line; anks strongly light
ecked; groin dark. Rictal gland white. Venter and ven-
tral surfaces of hindlimbs yellowish cream; throat dark
brown with large irregular yellowish cream marks and
Fig. 5. Color variation of preserved Noblella mindo sp. nov. in (A–E) dorsal and (F–J) ventral views: (A, F) ZSFQ 305, paratype, adult
female, SVL = 19.5 mm; (B, G) ZSFQ 304, paratype, adult female, SVL = 19.2 mm; (C, H) ZSFQ 050, holotype, adult female, SVL = 18.3
mm; (D, I) ZSFQ 051, paratype, adult male, SVL = 17.0 mm; (E, J) ZSFQ 049, paratype, adult male, SVL = 16.5 mm. Photos by David
Brito-Zapata and Carolina Reyes-Puig.
71
A new species of the genus Noblella from Ecuador
medium longitudinal line. Forelimbs ventrally yellowish
cream with dark brown marks, dorsally light brown with
dark brown marks; iris reddish copper with minute scat-
tered turquoise dots. Hindlimbs like dorsum.
Color of holotype in ethanol (Fig. 5)
Dorsum brown, darker towards middorsum, well-
defined middorsal line cream, extending from inter-
parietal region to cloaca where stripe continues along
posterior surface of thighs and pes. Dorsal surfaces of
forelimbs brown with black spots. Labial bars absent;
rictal gland light brown. Loreal region black, extend-
ing as homogeneous dark band to upper insertion of
arm and into body anks, narrowing towards groin;
anks strongly light ecked; groin dark. Dorsal surfaces
of hindlimbs lighter brown than dorsum to cream with
dark eck and spots. roat dark brown with cream
irregular marks and medium longitudinal stripe. Chest,
venter and ventral surfaces of thigh and crus cream.
Variation of color patterns and external morphology
(Figs. 4–5)
Adult females ZSFQ 050, 304–305 and the adult
male ZSFQ 051 exhibit a cream middorsal stripe extend-
ing from interparietal region to cloaca; stripe of ZSFQ
305 is thinner and faintly dened. Dark suprainguinal
marks are faint in ZSFQ 049. roat, chest and ventral
surfaces of forelimbs are dark brown with a white cross
formed by a longitudinal, ne line running from chin to
chest, crossing a similar line departing from midventral
Fig. 6. Ventral views of (A–D) hands and (E–H) feet from three species of Noblella: (A, E) Noblella coloma, extracted from Guayasamin and
Terán-Valdez (2009); (B, F) N. worleyae (holotype); (C, G) N. worleyae (ZSFQ 3851); (D, H) N. mindo sp. nov. (holotype). Scale bars = 1
mm. Illustrations by Carolina Reyes-Puig.
72 C. Reyes-Puig et alii
Table 1. Main diagnostic characters of three species of Noblella from northwestern Ecuador.
Species
Characters
Source
Finger tips Toe tips Distal
phalanges
Toes
papillae
Ulnar
tubercles
Venter and throat
coloration
Prootic
and
exoccipital
Sphenethmoid
Length of
transverse
processes of
Presacrals
Neural arch of
Presacrals
Noblella
coloma Acuminate Slightly expanded and
acuminate distally T-shape d Absent Absent
Venter orange with
minute white and brown
spots
Separated
Well-ossied,
ventrally
not fused at
midline
II, VIII<V–
VII<IV<III
Presacrals
III–V with
raised medial
ridge
Terán-
Valdez and
Guayasamin,
2009
N.
worleyae
Slightly
acuminate
on Fingers I
and IV and
acuminate on
Fingers II and
III
Slightly expanded and
slightly acuminate
on Toes I and V, and
cuspidate tips on Toes
II–IV
T-shape d Absent Present
Venter yellowish cream
with minute speckling;
throat with irregular
brown marks to
homogeneously brown
Fused
to form
otoccipital
Well-ossied,
ventrally fused
at midline
II<V–
VIII<IV<III
With raised
medial ridge in
all presacrals
Reyes-Puig et
al., 2020
and this paper
N. mindo
sp. nov. Rounded Slightly expanded and
rounded distally
slightly
T-shape d Absent Present
roat, chest and ventral
surfaces of arms dark
brown with a white cross,
venter yellowish cream
Separated
Moderately
ossied,
ventrally fused
at midline in
posterior half
and separated
in anterior half
V–VIII<II–
IV<III
Presacrals
III–VIII with
raised medial
ridge
is paper
73
A new species of the genus Noblella from Ecuador
surface of each forelimb (ZSFQ 050, 304, 774), longitu-
dinal line on throat of ZSFQ 773 is faint, while cross is
almost unnoticeable in holotype due to extensive light
color of throat. Venter and ventral surfaces of hindlimbs
are cream (ZSFQ 305), dirty cream with diused irregu-
lar brown marks (ZSFQ 304), or pinkish cream (ZSFQ
049–051). e throat in females is cream (ZSFQ 304–
305) or pinkish cream (ZSFQ 050) with large irregular
dark brown marks; meanwhile in males it is homoge-
nously dark brown with a slightly dened medium lon-
gitudinal stripe (ZSFQ 049, 051).
Osteology
Osteological description of Noblella mindo sp. nov.
is based on micro-CT images of the adult female holo-
type (ZSFQ 050). Details of skull morphology and osteo-
logical aspects of hand and foot are presented in Fig. 7
and main skeletal features are shown in Fig. 8.
Skull (Fig. 7)
Skull slightly longer than wide; widest part is at
about where quadratojugal meets maxilla and is 89% of
skull length. Rostrum short; distance from anterior edge
of frontoparietals to anterior face of premaxilla is 16%
of skull length. At level of midorbit, braincase is about
38% of maximum skull width. Braincase combines well-
and poorly ossied elements. Frontoparietals are well-
developed bones, distinctly longer than broad, slightly
narrower anteriorly than posteriorly; narrowly separated
along most of their length and only fused in anterior
region. Boarder between frontoparietals and prootics not
well-resolved in micro-CT scan. Ventrally, prootics in
contact with parasphenoid alae. Prootics well-separated
from each other. Exoccipitals approximate one another
ventromedially and dorsomedially but still clearly sepa-
rated with about same distance ventrally and dorsally;
separated from frontoparietals. Anterolaterally, fron-
toparietals in contact with sphenethmoid. Sphenethmoid
ventrally at midline separated in anterior half and fused
in posterior half; posterior margin does not reach mid-
point of orbit and is broadly separated from prootic and
in ventral contact with parasphenoid. Cultriform process
of parasphenoid well-ossied posteriorly, thinning ante-
riorly, and about 28% width of braincase at mid-orbit.
Lateral margins of process approximately parallel. Par-
asphenoid alae long but poorly ossied at their lateral
ends. Neopalatines very thin and long, articulate with
sphenethmoid and approximate but not contact maxilla.
Columella (or stapes) large and well-ossied. Due to tiny
size and ne structure, septomaxilla is not well-resolved
in micro-CT scan. Dorsal investing bones moderate-
ly developed. Nasals thin and broadly separated from
one another, posteriorly in contact with anterior end of
frontoparietals and posterolaterally in thin contact with
maxilla. ey curve ventrally towards their lateral edges.
Small prevomers broadly separated from one another
medially, their anterior edge almost contacts a long and
thin posterior projecting ramus of septomaxilla. Maxil-
lary arcade bears many small, poorly resolved teeth on
premaxillae and maxillae. Premaxillae separated medi-
ally, and their anterodorsal alary processes rise divergent
from midline but still distinctly separated from nasals.
Premaxilla and maxilla in lateral contact, with ante-
rior edge of maxilla slightly overlapping lateral edge of
premaxilla. Pars palatina of premaxilla broad, with two
well-dened processes: medial process thin and acumi-
nate, running about parallel to its counterpart, being
distinctly separated from it; lateral process about the
same length, but slightly broader, especially at its trun-
cate posterior ending. Maxilla long, its posterior end
acuminate and in contact with quadratojugal. Triradiate
pterygoid bears a long, curved anterior ramus oriented
anterolaterally toward maxilla, with which it articulates
at ventral boarder slightly anterior to midline of orbit.
Posterior ramus of pterygoid about same length as medi-
al ramus and both about half length of anterior ramus;
however, posterior ramus more robust than other two.
Edge of medial ramus overlaps lateral edge of prootic.
Quadratojugal slender, almost straight and articulating
anteriorly with maxilla and posterodorsally with ventral
ramus of squamosal. Squamosal T-shaped, with a long
laminar otic ramus; zygomatic ramus much shorter and
slender; ventral ramus about same length as otic ramus,
laminar and broad, increasing in width ventrally. Man-
dible slim and edentate. Mentomeckelians small, medial-
Table 2. Measurements (in mm) of type series of Noblella mindo sp.
nov. Ranges followed by mean and standard deviation in parenthe-
ses.
Characters Noblella mindo sp. nov.
Females (n = 3) Males (n = 2)
SVL 18.3–19.5 (19.0 ± 0.6) 16.5–17.0 (16.7 ± 0.3)
HL 7.1–7.5 (7.2 ± 0.2) 6.2–7.0 (6.6 ± 0.6)
HW 5.7–6.7 (6.3 ± 0.5) 5.37–5.4 (5.38 ± 0.02)
ED 2.2–2.5 (2.4 ± 0.2) 1.9–2.0 (1.95 ± 0.1)
EN 1.26–1.29 (1.28 ± 0.01) 1.21–1.22 (1.215 ± 0.01)
MWE 1.2–1.5 (1.3 ± 0.2) 1.1–1.2 (1.15 ± 0.01)
TD 0.8–1.2 (0.9 ± 0.2) 0.78–0.82 (0.8 ± 0.02)
MIOD 3.3–3.4 (3.3 ± 0.1) 3.0–3.4 (3.2 ± 0.3)
LH 3.5–3.9 (3.7 ± 0.2) 3.0–3.2 (3.1 ± 0.2)
LS 9.0–9.3 (9.1 ± 0.2) 7.7–8.0 (7.8 ± 0.2)
LF 8.4–9.0 (8.6 ± 0.4) 7.3–7.5 (7.4 ± 0.1)
74 C. Reyes-Puig et alii
ly, and laterally slightly broadened, and separated medi-
ally by a narrow gap. Dentary long and thin, reaching to
about anterior corner of orbit; it is posteriorly acuminate
and overlapping angulospenial, seeming to be in contact
with this bone for about the posterior half of its length;
anteroventrally it contacts mentomeckelian bones.
Angulosplenial long and arcuate. Coronoid process is a
moderately long and slightly raised ridge. e only ossi-
ed portions of hyoid apparatus are two posteromedial
processes, which are anteriorly slightly and posteriorly
moderately expanded, approaching each other at ante-
rior ends but being still moderately separated.
Postcranium (Fig. 8)
Eight presacral vertebrae. All presacrals non-imbri-
cate. Presacral I longer than posterior vertebrae. All
except Presacral I bear well-developed diapophyses.
Transverse processes of Presacrals V–VIII similar in size,
being the shortest and thinnest, those of Presacrals II
and IV also about similar in size and being slightly larg-
er, and those of Presacral III being the longest and wid-
est of all transverse processes. Transverse processes of
Presacrals II and VIII have slightly anterolateral orienta-
tion, those of Presacral III are laterally oriented and the
others are slightly posterolaterally oriented. Neural arch
of Presacrals III–VIII bears a raised medial ridge. Sacrum
bears slightly expanded diapophyses. Urostyle long, slen-
der, slightly shorter than presacral portion of vertebral
column and bearing a well-pronounced dorsal ridge
along most of its length, beginning at its anterior end.
e bone has a bicondylar articulation with the sacrum.
Pectoral girdle with well-ossified coracoids, clavicles,
scapulae and cleithra. Suprascapular and sternum unos-
sied and not visible in micro-CT scan, and omosternum
hardly visible. Clavicles long and slim, oriented antero-
medially, slightly curved, with medial tips touching each
other. Laterally, clavicles rmly articulating with scapu-
lae. Coracoids stout and glenoidal and sternal ends about
equally expanded. Anterior edges of coracoids slightly
curved, posterior edges almost straight. Medial tips of
coracoids broadly separated from another. Scapula long,
with a prominent pars acromialis not separated from
pars glenoidalis. Cleithrum long, broader, and thicker at
scapular boarder, thinning posteriorly. In pelvic girdle,
long, slender iliac shas bearing conspicuous dorsolater-
al ridges along most of their length, except anteriormost
region. Ilia are fused posteriorly with ischium and pubis.
Ischium stout, whereas pubis is thin and blade-like.
Manus and pes (Fig. 7)
All phalanges are ossied with a phalangeal formula
for ngers and toes: 2-2-3-3 and 2-2-3-4-3, respectively.
Order of nger length: I < II < IV < III, and that of toes:
I < II < V < III < IV. Distal knobs present on terminal
phalanges of all ngers and toes. Terminal phalanges of
all toes and ngers narrower than penultimate phalan-
ges of all toes and ngers, respectively. Carpus and tar-
sus not well-resolved in micro-CT scan. However, carpus
seems to be composed of a radiale, ulnare, Element Y,
ossied prepollex element, Carpal 2 and a large post-axi-
al element probably representing a fusion of Carpals 3–5.
Tarsus seems to be composed of two tarsal elements:
Tarsal 1 and Tarsal 2 + 3, with latter being distinctly
larger than Tarsal 1. A moderately large centrale and
small ossied prehallux are also present. In ventral view,
three sesamoids of subequal sizes are overlaying proxi-
mal end of Metatarsals IV–V, a further smaller sesamoid
is overlaying parts of Tarsal 1.
Distribution and Natural History
Noblella mindo sp. nov. is only known from El Cin-
to (0.09022°S, 78.81858°W; 1,673 m), Mindo, province
of Pichincha, Ecuador (Fig. 2). Noblella mindo sp. nov.
inhabits secondary cloud forests, with the presence of
palmito (Bactris gasipaes) plantations and trees that have
emerged aer the massive logging of forests in the area.
ese forests have a high humidity index, dense leaf lit-
ter layer, and abundant epiphytes. It has a restricted dis-
tribution; sampling activities were carried out in a range
up to 3km around the type locality, and no individuals
nor calls of N. mindo sp. nov. were recorded. e gecko
Lepidoblepharis conolepis was found in sympatry. e
locality is surrounded by livestock areas and within the
type locality forest, there are trails used by farmers to
move their livestock. e population of Noblella mindo
sp. nov. could be impacted if livestock activity or defor-
estion expands. ree individuals (ZSFQ 049–051) were
found active during the day between 10:00 and 11:00
am; all frogs were on the ground in a 2-meter depth
hole.
Noblella worleyae Reyes-Puig, Maynard, Trageser,
Vieira, Hamilton, Lynch, Culebras, Kohn, Brito, and
Guayasamin 2020: New locality
Figs. 2, 9–12
New records (3 females, 1 male). All individuals were
collected at dierent localities inside Los Cedros Bio-
logical Reserve: ZSFQ 3851 (Figs. 9–10), adult female
and ZFSQ 3852, adult male, collected at 0.31501°N,
78.77943°W, 1,612 m (WGS84; Fig. 2), García Moreno,
Cotacachi, province of Imbabura, by David Brito-Zapa-
ta and Martín Obando on 26 October 2019. MZUTI
1708, adult female, collected at 0.31125°N, 78.78095°W,
1,417 m, by Giusseppe Gagliardi and JMG on 13 March
75
A new species of the genus Noblella from Ecuador
Fig. 7. Details of (A–D) skull morphology and osteological aspects of (E–F) hand and (G–H) foot of Noblella mindo sp. nov., ZSFQ 050,
holotype, adult female. e skull is shown in (A) dorsal, (B) ventral, (C) frontal, and (D) lateral views. alary p = alary process, angspl =
angulosplenial, col = columella, dent = dentary, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian bone, nas = nasal, npl = neo-
palatine, occ con = occipital condyle, otoc = otoccipital (fused prootic and exoccipital), pmax = premaxilla, prsph = parasphenoid, prvom
= prevomer, pter =pterygoid, qj = quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal. e right hand is shown
in (E) dorsal, and (F) palmar aspects; and the le foot in (G) dorsal, and (H) plantar aspects. Digits numbered I–V. antbr = os antebrachii
(radius + ulna), carp d = carpale distale, cent = centrale, b = bulare, mtc = metacarpalia, mtt = metatarsalia, ph d I–IV = nger phalanges
F1–F4, ph d I–V = toe phalanges F1–F5, prhl =prehallux, prpl = prepollex, rad = radius, tar d = tarsale distale, tib = tibiale, uln = ulnare.
Images prepared by Claudia Koch.
76 C. Reyes-Puig et alii
2012; MZUTI 1709, adult female, collected at 0.3184°N,
78.7837°W, 1,790 m, by Jaime Culebras and JMG on 15
March 2012.
Diagnosis
Specimens collected in Los Cedros reserve show
morphological characters described for Noblella worleyae
(Reyes-Puig et al., 2020; Figs. 6, 9–10) as follow (varia-
tion from original description in bold): (1) skin of dor-
sum nely shagreen, skin on venter smooth; (2) tympanic
annulus and membrane visible externally, supratympanic
fold inconspicuous; (3) snout, rounded in dorsal and lat-
eral views; (4) eyelids without tubercles; (5) dentigerous
processes of vomers absent; (6) vocal slits and vocal sac
present, nuptial pads not visible; (7) ngers not expand-
ed or slightly expanded distally, tips of Fingers I and IV
rounded and tips of Fingers II and III slightly acuminate
(originally described as tips of Fingers I and IV slightly
acuminate, Fingers II and III acuminate), without papil-
lae (Fig. 6), Finger I shorter than Finger II,(8); distal
phalanges T-shaped (originally described as slightly
T-shaped), phalangeal formula of hands: 2-2-3-3 (Fig.
11); (9) supernumerary palmar tubercles few but present,
ulnar tubercles diminutive and round (decreased by pres-
ervation eects), subarticular tubercles rounded, discs
lacking circumferential grooves; (10) one tarsal tubercle,
elongated and subconical, two metatarsal tubercles (inner
tubercle 2 times size of external); toes slightly expanded
distally and rounded on Toes I and V, cuspidate tips on
Toes II–IV, papillae present on Toes II–IV (Fig. 6); (11)
Toe V shorter than Toe III distal portions of circum-
ferential grooves present on Toes II–V, supernumerary
tubercles absent (12) phalangeal formula of feet: 2-2-3-4-
3 (Fig. 11); (13) in life, dorsum brown, with two suprain-
Fig. 8. Osteology of Noblella mindo sp. nov., ZSFQ 050, holotype, adult female. e full skeleton is shown in (A) dorsal, (B) ventral, and (C)
lateral views. antbr = os antebrachii (radius + ulna), clav = clavicle, cle = cleithrum, cor = coracoid bone, crur = os cruris (tibia + bula),
fem = femoral bone, b = bulare, hm = humeral bone, il = ilium, isch = ischium, pr p-m = processus postero-medialis, prsac v = presacral
vertebrae, pub = pubis, sac v = sacral vertebra, sc = scapula, ur = urostyle, tib = tibiale. Images prepared by Claudia Koch.
77
A new species of the genus Noblella from Ecuador
guinal dark brown marks; middorsal, longitudinal line
faint cream; rictal gland dark brown; anks with dark
brown band narrowing towards groin and with clusters
of turquoise specks towards ventral side; groin dark, with
high concentration of melanophores; throat, chest and
ventral surfaces of arms dark brown; venter yellowish
cream, with brownish-orange tones on ventral surfaces of
legs and thighs, iris reddish copper; (14) SVL in one adult
male 16.1 mm; in adult females 19.1–20.4 mm (mean 19.8
mm, n = 3) (range originally described 18.1–19.1 mm).
Variation of color patterns and external morphology
(Figs. 9–10)
Specimens of Noblella worleyae from Los Cedros
Reserve vary from brown (MZUTI 1708–1709) to dark
brown (ZSFQ 3851–3852). In preservative, specimen
ZSFQ 3852 has a grayish-brown dorsal coloration. Black
suprainguinal spots vary in size and may be diused but
are always present. All specimens exhibit a faint, cream
middorsal stripe extending from the head to cloaca, but
only in one specimen (MZUTI 1709) this line contin-
ues onto posterior surfaces of thigh, disappears in crus,
and reappears in posterior surfaces of pes. Specimens
MZUTI 1708 and ZSFQ 3852 have a homogenously dark
brown throat like the holotype, but in MZUTI 1709 the
throat is brown with scattered irregular cream marks.
Ventral surfaces of thighs and crus of ZSFQ 3852 retain
pinkish-cream color in preservative. Male ZSFQ 3852
exhibits an evident discoidal fold.
Osteology description
Osteological description of Noblella worleyae is
based on micro-CT images of an adult female (MZU-
TI 1709). Details of skull morphology and osteological
aspects of hand and foot are presented in Fig. 11 and
main skeletal features are shown in Fig. 12.
Fig. 9. Color pattern of Noblella worleyae in life: (A, C) Dorso-lateral and ventral patterns of ZSFQ 3851, adult female, SVL = 19.1 mm; (B,
D) Dorsal and ventral patterns of ZSFQ 3852, adult male, SVL = 16.1 mm. Photos by David Brito-Zapata.
78 C. Reyes-Puig et alii
Skull (Fig. 11)
Skull almost as wide as long; widest part is at about
where quadratojugal meets maxilla and is 97% of skull
length. Rostrum short; distance from anterior edge of
frontoparietals to anterior face of premaxilla is 18% of
skull length. At level of midorbit, braincase is about 34%
of maximum skull width. Braincase combines well- and
poorly ossied elements. Prootic and exoccipital seem
to be fused to form otoccipital. Frontoparietals are well-
developed bones, distinctly longer than broad, slightly
narrower anteriorly than posteriorly; narrowly separated
along most of their length and only fused in anterior
region. Posterior portion of braincase seems to be fully
enclosed by partial fusion of frontoparietals with otoc-
cipitals. However, there might still exist some traces of
boarders between bones, but these parts are not well-
resolved in micro-CT scans. Ventrally, otoccipitals are
in contact with parasphenoid alae. Prootic part of otoc-
cipitals are well-separated from each other. Exoccipital
parts approximate one another ventromedially and dor-
somedially but are still clearly separated with a broader
ventral than dorsal gap between them. Anterolater-
ally, frontoparietals are in contact with sphenethmoid.
Sphenethmoid is well-ossified and ventrally fused at
midline; posterior margin almost reaches midpoint of
orbit but is still broadly separated from prootic part of
otoccipitals and is in ventral contact with cultriform
process of parasphenoid. Cultriform process of paras-
phenoid is well-ossied posteriorly, thinning anteriorly,
and about 31% of width of braincase at mid-orbit. Lat-
eral margins of process are approximately parallel. Par-
asphenoid alae are long and well-ossied. Neopalatines
are very thin and long, articulating with sphenethmoid
dorsomedially and maxilla laterally. Columella (or sta-
pes) is large and well-ossied. Because of tiny size and
fine structure, septomaxilla is not well-resolved in
micro-CT scan. Dorsal investing bones are moderately
developed. Nasals are thin and broadly separated from
one another, posteriorly in contact with anterior end of
frontoparietals and posterolaterally in very thin contact
with maxilla. ey curve ventrally towards their lateral
edges. Small prevomers are broadly separated from one
another medially, their anterior edge approximates a
long and thin posterior projecting ramus of septomax-
Fig. 10. Color variation of preserved Noblella worleyae in (A–D) dorsal and (E–H) ventral views: (A, E) MZUTI 1708, adult female, SVL =
20.4 mm; (B, F) MZUTI 1709, adult female, SVL = 19.9 mm; (C, G) ZSFQ 3851, adult female, SVL = 19.1 mm; (D, H) ZSFQ 3852, adult
male, SVL = 16.1 mm. Photos by David Brito-Zapata and Carolina Reyes-Puig.
79
A new species of the genus Noblella from Ecuador
Fig. 11. Details of (A–D) skull morphology and osteological aspects of (E–F) hand and (G–H) foot of Noblella worleyae, MZUTI 1709. e
skull is shown in (A) dorsal, (B) ventral, (C) frontal, and (D) lateral views. alary p = alary process, angspl = angulosplenial, col = columella,
dent = dentary, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian bone, nas = nasal, npl = neopalatine, occ con = occipital con-
dyle, otoc = otoccipital (fused prootic and exoccipital), pmax = premaxilla, prsph = parasphenoid, prvom = prevomer, pter =pterygoid, qj =
quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal. e right hand is shown in (E) dorsal, and (F) palmar aspects;
and the le foot in (G) dorsal, and (H) plantar aspects. Digits numbered I–V. antbr = os antebrachii (radius + ulna), carp d = carpale distale,
cent = centrale, b = bulare, mtc = metacarpalia, mtt = metatarsalia, ph d I–IV = nger phalanges F1–F4, ph d I–V = toe phalanges F1–F5,
prhl =prehallux, prpl = prepollex, rad = radius, tar d = tarsale distale, tib = tibiale, uln = ulnare. Images prepared by Claudia Koch.
80 C. Reyes-Puig et alii
illa. Maxillary arcade bears many small, poorly resolved
teeth on premaxillae and maxillae. Premaxillae are sepa-
rated medially, and their anterodorsal alary processes
rise divergent from midline but are still distinctly sepa-
rated from nasals. Premaxilla and maxilla are in lateral
contact, with anterior edge of maxilla slightly overlap-
ping lateral edge of premaxilla. Pars palatina of premax-
illa is broad, with two well-dened processes: medial
process acuminate, and runs about parallel to its coun-
terpart, being distinctly separated from it; lateral pro-
cess is slightly shorter and broader. Maxilla is long, its
posterior end acuminate and in contact with quadrato-
jugals. Triradiate pterygoid bears a long, curved anterior
ramus that is oriented anterolaterally towards maxilla,
with which it articulates at ventral boarder anterior to
midline of orbit. Posterior ramus of pterygoid is about
same length as medial ramus and both are about half
length of anterior ramus; however, posterior ramus is
more robust than the other two. Edge of medial ramus
overlaps lateral edge of prootic part of otoccipital. Quad-
ratojugal is slender, slightly curved and articulates ante-
riorly with maxilla and posterodorsally with ventral
ramus of squamosal. Squamosal is T-shaped with long
laminar otic ramus; zygomatic ramus is much shorter
and more slender; ventral ramus is about same length
as otic ramus, laminar and broad, increasing in width
ventrally. Mandible is slim and edentate. Mentomeckeli-
ans are small, medially and laterally slightly broadened,
and medially contacting each other. Dentary is long and
thin, reaching to about anterior corner of orbit; posteri-
orly acuminate and overlapping angulospenial, seeming
to be in contact with this bone for most of its length,
except the most anterior part; anteroventrally it contacts
mentomeckelian bones. Angulosplenial is long and arcu-
Fig. 12. Osteology of Noblella worleyae, MZUTI 1709, adult female. e full skeleton is shown in (A) dorsal, (B) ventral, and (C) lateral
views. antbr = os antebrachii (radius + ulna), clav = clavicle, cle = cleithrum, cor = coracoid bone, crur = os cruris (tibia + bula), fem =
femoral bone, b = bulare, hm = humeral bone, il = ilium, isch = ischium, pr p-m = processus postero-medialis, prsac v = presacral verte-
brae, pub = pubis, sac v = sacral vertebra, sc = scapula, ur = urostyle, tib = tibiale. Images prepared by Claudia Koch.
81
A new species of the genus Noblella from Ecuador
ate. Coronoid process is a moderately long and strongly
raised ridge. e only ossied portions of hyoid appara-
tus are two posteromedial processes, which are anteri-
orly slightly more expanded than posteriorly, approach-
ing each other at anterior ends but being still distinctly
separated.
Postcranium (Fig. 12)
Eight presacral vertebrae. All presacrals are non-
imbricate. First presacral vertebra is longer than poste-
rior vertebrae. All except Presacral I bear well-developed
diapophyses. Transverse processes of Presacrals V–VIII
similar in size and being the thinnest and second short-
est, those of Presacral II being the shortest, and those of
Presacral III being the longest and widest of all trans-
verse processes. Transverse processes of Presacrals II
and VIII have slightly anterolateral orientation, those
of Presacrals III and VII are laterally oriented and oth-
ers are slightly posterolaterally oriented. Neural arch of
all presacrals bears a raised medial ridge. Sacrum bears
slightly expanded diapophyses. Urostyle is long, slender,
about similar in length as presacral portion of vertebral
column and bearing a well-pronounced dorsal ridge
along about two-thirds of its length, beginning at its
anterior end, with a lateral foramen in anterior region.
e bone has a bicondylar articulation with the sacrum.
Pectoral girdle with well-ossied coracoids, clavicles,
scapulae and cleithra. Suprascapular, omosternum, and
sternum unossied and not visible in micro-CT scan.
Clavicles are long and slim, oriented anteromedially,
slightly curved, with medial tips approaching but not
touching each other. Laterally, clavicles rmly articulat-
ing with scapulae. Coracoids are stout and glenoidal and
sternal ends are about equally expanded. Anterior edges
of coracoids are curved, the posterior edges are almost
straight. Medial tips of coracoids are broadly separated
from another. Scapula is long with a prominent pars
acromialis that is not separated from pars glenoida-
lis. Cleithrum is long, broader and thicker at scapu-
lar boarder, thinning posteriorly. In pelvic girdle, long,
slender iliac shas bear conspicuous dorsolateral ridges
along most of their length, except the anterior most
region. Ilia is fused posterirly with ischium and pubis.
Ischium is stout, whereas pubis is thinner and blade-like.
Manus and pes (Fig. 11)
All phalanges are ossied, with phalangeal formula
for ngers and toes: 2-2-3-3 and 2-2-3-4-3, respective-
ly. Order of nger length: I < II < IV < III, and that of
toes: I < II < V < III < IV. Distal knobs seem to be absent
on Finger I but are present on terminal phalanges of all
other ngers and toes. Nevertheless, they are not well-
resolved in micro-CT scans and are sensitive to thresh-
olds used during reconstruction. Terminal phalanges of
all toes and ngers are narrower than penultimate pha-
langes of all toes and ngers, respectively. Carpus and
tarsus are not well-resolved in micro-CT scan. However,
carpus seems to be composed of a radiale, ulnare, Ele-
ment Y, ossied prepollex element, Carpal 2 and a large
post-axial element probably representing a fusion of Car-
pals 3–5. Tarsus seems to be composed of two tarsal ele-
ments: Tarsal 1 and Tarsal 2 + 3, with latter being larg-
er than Tarsal 1. A moderately large centrale and small
ossied prehallux are also present. In ventral view, three
sesamoids of subequal sizes overlaying proximal end of
Metatarsals II–IV, a further smaller sesamoid overlaying
parts of Tarsal 1 and proximal end of Metatarsal I.
Natural History
We report a new locality for Noblella worleyae:
Los Cedros Biological Reserve, province of Imbabura,
Ecuador (Fig. 2), at approximately 8.4 km in a straight
line between the type locality of N. worleyae (i.e., Man-
duriacu Reserve). e individuals were found at several
point-localities between 1,417–1,790 m of elevation. e
reserve presents an important track of mature Low Mon-
tane Evergreen Forest. All specimens were collected at
night between 7:00 and 11:50 pm and were found on the
ground, in areas covered with abundant leaf litter. Indi-
viduals appeared inactive, because they were located by
movement only when litter was removed. Syntopic spe-
cies were Pristimantis mutabilis (Guayasamin et al.,
2017b), P. crenunguis, (Lynch, 1976a) and Alopoglossus
viridiceps (Torres-Carvajal and Lobos, 2014). Noblella
worleyae seems to be a rare species at Los Cedros Bio-
logical Reserve. Only two individuals were found in two
dierent surveys with known sampling eort. e rst
sampling was carried out between 22–25 August 2019,
with ve people for approximately nine hours per day.
e second sampling was carried out between 26–30
October 2019, with two people for about nine hours
per day. Individuals were found in the lower forest stra-
tum, camouaged extremely well in leaitter and hav-
ing an evasive behavior, similar to other Noblella species
(Reyes-Puig et al., 2019c).
DISCUSSION
Our phylogenetic analyses (Fig. 1) agree with pre-
vious studies that have shown that southern species
of Noblella (N. losamigos, N. madreselva, N. pygmaea,
N. thiuni) are more closely related to species of Psy-
chrophrynella, rather than to northern species of Noblel-
82 C. Reyes-Puig et alii
la (N. coloma, N. heyeri, N. lochites, N. mindo sp. nov., N.
myrmecoides, N. personina, N. worleyae) (Reyes-Puig et
al., 2019c, 2020; Santa-Cruz et al., 2019). Noblella peru-
viana is the type species of the genus, which, based on
geography, is most likely part of the Southern Clade.
However, since there are no sequences of N. peruviana,
we refrain from proposing a new generic arrangement.
Species richness of the genus Noblella has increased
dramatically over the last decade (Frost, 2020). About a
decade ago, only three species of Noblella were known
in Ecuador and no species had been described from the
northwestern slopes of the Andes of Ecuador (Cisneros-
Heredia and Reynolds, 2007). Nowadays, there are eight
species of Noblella reported from Ecuador, including
three from the western Ecuadorian Andes (N. coloma,
N. mindo sp. nov., and N. worleyae) that form a distinct
clade among the northern Noblella (Fig 1). e diversi-
ty of the genus Noblella has also increased in the Peru-
vian Andes, where in recent years three species been
described, forming a clade with the previously described
Psychrophrynella and Noblella (Catenazzi et al., 2015;
Santa Cruz et al., 2019; Catenazzi and Ttito, 2019).
Our new records of Noblella worleyae from Los
Cedros reserve add important intraspecic variation to
the original description, in terms of its body size, col-
oration and shape of tips of the digits. In particular,
variation in the ngertips was found, with tips of Fin-
gers I and IV varying from slightly acuminate (original
description) to rounded (data presented herein), and tips
of Fingers II and III from acuminate (original descrip-
tion) to slightly acuminate (data presented herein). We
also strengthen the original publication (Reyes-Puig et
al., 2020) with a detailed description of the osteology of
the species.
Diversification in Noblella seems be related with
the linearity of the Andes, with allopatric and parapa-
tric populations being separated by ecological and geo-
graphic barriers. In Ecuador, all species of Noblella are
allopatric and most of them are restricted to very spe-
cic geographic areas. Noblella mindo sp. nov. occurs in
low montane forest in the valley of Mindo, in the Nam-
billo River watershed, western slopes of the Pichincha
Massif, northwestern Andes, at 1,673 m; while N. worley-
ae inhabits low montane forest in the Manduriacu-Los
Cedros watersheds, southern slopes of the Toisan mas-
sif; and N. coloma is restricted to the cloud forests of the
Río Guajalito watershed, western slopes of the Atacazo
volcano. Noblella worleyae is separated from N. coloma
and N. mindo sp. nov. by the Guayllabamba River Valley
(Fig. 2), an important biogeographic barrier, especially
for frog species with low vagility (Hillman et al., 2014).
Although the valley of Mindo (type locality of N. mindo
sp. nov.) and the valley of Guajalito (type locality of N.
coloma) are ca. 20 km apart in straight line, they are in
dierent watersheds, separated by the Nambillo River
and complex orogeny caused by the Pichincha massif
and the Atacazo volcano.
All species currently recognized under the genus
Noblella are miniaturized frogs, among the smallest
Neotropical vertebrates (Duellman and Lehr, 2009).
ey are cryptic and adapted to live amidst or under
leaf litter in forests, where they are oen overlooked by
amphibian visual surveys, being easier to locate by their
calls or through pitfall traps (Reyes-Puig et al., 2019c).
Although some species may be abundant locally, most
species appear to have low densities. For example, year-
ly surveys between 2000–2012 at the type locality of
N. coloma produced only three records, while up to 20
individuals of N. lochites were found in 2014 at a single
locality in the province of Zamora-Chinchipe (D. F. Cis-
neros-Heredia pers. obs.).
Most remnants of mature forests in the Andes of
Ecuador are nowadays either inside public or private pro-
tected areas or persist due to their inaccessibility. Private
conservation initiatives have become extremely impor-
tant in Ecuador (Betancourt et al., 2018; Guayasamin et
al., 2018, 2019; Reyes-Puig et al., 2019a, 2019b, 2019c),
where public protected areas do not cover all critically
important regions for biodiversity (Lessmann et al., 2014;
Cuesta et al., 2017; Reyes-Puig et al., 2017).
Unfortunately, habitat loss due to unsustainable
expansion of the agricultural frontier, mining and infra-
structure projects have placed a heavy burden on sev-
eral private reserves (Roy et al., 2018; Guayasamin et
al., 2019). In the early 2000s, the region of Mindo was
heavily threatened by the construction of an oil pipeline
(Oleoducto de Crudos Pesados OCP). Fortunately, local,
national and international protests managed to promote
some actions to mitigate the largest impacts of the pipe-
line development. Eventually Mindo has transformed
into one of the most popular ecotourism destinations in
northwestern Ecuador, allowing several private protected
areas to preserve large tracks of mature forest (Welford
and Yarbrough, 2015). Unfortunately, twenty years later,
history repeats itself, now at Los Cedros reserve, but this
time with mining concessions, exploration and exploita-
tion (Roy et al., 2018). us, biodiversity conservation is
facing an uncertain future.
e increasing descriptions of new species within
the Ecuadorian territory have a practical application in
the conservation of biodiversity. By highlighting the
presence of new vertebrates with restricted distributions
in the Andes, the visualization of this unique biodiver-
sity is indisputable.
83
A new species of the genus Noblella from Ecuador
ACKNOWLEDGEMENTS
We conducted this research under research permits
and agreement for genetic resources access (MAE-DNB-
CM-2018-0106, 019-2018-IC-FAU-DNB/MAE) issued by
Ministerio del Ambiente del Ecuador. We carried out
this study following the guidelines for treatment and
management of live amphibians and reptiles in field and
laboratory investigations (Beaupre et al., 2004), recom-
mended by the American Society of Ichthyologists and
Herpetologists, the Herpetologists’ League and the Soci-
ety for the Study of Amphibians and Reptiles.
is study was developed as part of “Proyecto Des-
cubre Napo”, an initiative of Universidad San Francisco
de Quito in association with Wildlife Conservation Soci-
ety, and funded by the Gordon and Betty Moore Foun-
dation, as part of the project “WCS Consolidating Con-
servation of Critical Landscapes (mosaics) in the Andes”.
We express our gratitude to the following people for their
support: Ana Nicole Acosta-Vásconez, Susana Cárdenas,
Mariela Domínguez, Jonathan Guillemot, Andrés León-
Reyes, Emilia Peñaherrera, Carolina Proaño, Robert P.
Reynolds, Alejandra Robledo, Ana Sevilla, Rebecca Zug.
Work at Los Cedros Biological Reserve was developed as
part of project “Muestreo de Grupos Ecológicos Clave”,
research program “Evaluación biológica rápida del corre-
dor norte de la Reserva de Mashpi”, a joint initiative of
Fundación Futuro and Universidad San Francisco de
Quito USFQ (Instituto ECOLAP, Museo de Zoología,
Instituto iBIOTROP). is research was supported by
Universidad San Francisco de Quito USFQ through
research funds for the Museo de Zoología & Laboratorio
de Zoología Terrestre, Instituto de Diversidad Biológica y
Tropical iBIOTROP granted to DFCH; USFQ Collabora-
tion Grants and COCIBA Grants (project HUBI ID 34,
39, 48, 1057, 7703, 12268, 13524) granted to DFCH and
CRP; and Collaboration Grants and COCIBA Grants
(project HUBI ID 5521, 5467, 5447, 11164, 16871) grant-
ed by USFQ to JMG. Work by DFCH was supported by
Programa “Becas de Excelencia”, Secretaría de Educación
Superior, Ciencia, Tecnología e Innovación (SENES-
CYT), Ecuador; the Smithsonian Women’s Committee,
the 2002 Research Training Program, National Museum
of Natural History, Smithsonian Institution; María Elena
Heredia and Laura Heredia. We thank the Inédita Pro-
gram from the Ecuadorian Science Agency SENESCYT
(Respuestas a la Crisis de Biodiversidad: La Descripción
de Especies como Herramienta de Conservación; INED-
ITA PIC-20-INE-USFQ-001) that funded the molecular
component of this study. We are grateful for the com-
ments of Edgar Lehr and an anonymous reviewer to
improve the manuscript.
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APPENDIX I
Examined specimens.
Noblella lochites. Ecuador, Napo: ZSFQ 346, Archido-
na, Reserva Narupa, 1176 m; ZSFQ 347, Reserva Naru-
pa, 1152 m; ZSFQ 348, Reserva Narupa, 1167 m; Zamora
Chinchipe: ZSFQ 1119, Yantzaza, Concesión La Zarza,
1385 m; ZSFQ 1124, Concesión La Zarza, 1357 m; ZSFQ
1186, ZSFQ 1187, ZSFQ 1188, Yantzaza, Río Blanco, 1654
m; ZSFQ 1188, Río Blanco, 1830 m.
Noblella cf. lochites. Ecuador, Zamora Chinchipe:
ZSFQ 3262 – 326, Yantzaza, Estación Experimental El
Padmi UNL, 775 m. Noblella myrmecoides: Ecuador,
Napo: ZSFQ 670, Mera, Parque Nacional Llanganates,
1325 m; ZSFQ 671, Parque Nacional Llanganates, 1352
m; ZSFQ 672, Parque Nacional Llanganates, 1327 m.
Noblella cf. myrmecoides. Ecuador, Tungurahua:
ZSFQ 1341, Río Negro, Reserva Río Zuñag, 1269 m.
Noblella coloma. Ecuador: Pichincha: QCAZ 7277,
7412, 8701, 11614, 26307, 32702, Reserva Ecológica Río
Guajalito; 1800–2000 m.
Noblella heyeri. Ecuador, Loja: QCAZ 31470, 31471,
31473, Loja–Zamora road; 2385 m, QCAZ 22501, Zamo-
ra-Huaico; 2000 m.
Noblella worleyae. Ecuador, Imbabura: ZSFQ 345,
550, 551, 552, 2502, 2503, 2504, Reserva Manduriacu,
1184–1597 m.
