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15
New endangered species of Pristimantis from Peru
Open access at hp://www.salamandra-journal.com
© 2021 Deutsche Gesellscha für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany
15 February 2021 ISSN 0036–3375
SALAMANDRA 57(1): 15–26 SALAMANDRA
German Journal of Herpetology
A new, critically endangered species of Pristimantis
(Amphibia: Anura: Strabomantidae) from a
mining area in the Cordillera Occidental
of northern Peru (Región Cajamarca)
E L,, S L A C
1) Department of Biology, Illinois Wesleyan University, P.O. Box 2900, Bloomington, IL 61701, USA
2) Departamento de Herpetología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos,
Av. Arenales 1256, Jesús María, Lima 15072, Peru
3) School of Biological Sciences, Georgia Institute of Technology, Atlanta, GA 30332, USA
4) Florida International University, Department of Biological Sciences, 11200 SW 8th Street, Miami, FL 33199, USA
Corresponding author: E L, e-mail: elehr@iwu.edu
Manuscript received: 4 September 2020
Accepted: 26 November 2020 by J K
Abstract. We describe a new species of Pristimantis from high Andean grasslands (jalca) at m above sea level in
northern Peru (Región Cajamarca) based on morphological and molecular characters. e new species is known from
four males and ve females, which were found sheltering in the rosettes of Puya fastuosa (Bromeliaceae). e phylogenetic
analysis of a fragment of the mitochondrial S rRNA gene suggests that the new species is a sister taxon of Pristimantis
simonsii. e new species diers from its congeners by having a black dorsum speckled with white ecks and a dark brown
groin with white spots. Furthermore, adult males have a snout–vent length of .–. mm (n = ), and adult females of
.–. mm (n = ). Intensive mining activities apparently have extirpated the new species at its type locality and it is
therefore considered critically endangered. We discuss the impact of mining on biodiversity and biological surveys in Peru.
Key words. Extirpation, gold mining, Hualgayoc, jalca, morphology, phylogeny, systematics, Pristimantis astralos new spe-
cies, Peru.
Resumen. Describimos una nueva especie de Pristimantis de pastizales altoandinos (jalca) a m s.n.m. en el norte de
Perú (Región Cajamarca) en base a caracteres morfológicos y moleculares. La nueva especie se conoce de cuatro machos
y cinco hembras que encontramos escondidos en rosetas de Puya fastuosa (Bromeliaceae). El análisis logenético de un
fragmento del gen mitocondrial S rRNA sugiere que la nueva especie es el taxón hermano de Pristimantis simonsii. La
nueva especie se distingue de sus congéneres por tener un dorso negro con manchas blancas rociadas y una ingle de color
marrón oscuro con manchas blancas. Además, los machos adultos tienen una longitud hocico-cloaca de .–. mm (n
= ), y las hembras adultas de .–. mm (n = ). Debido a las intensas actividades mineras, la nueva especie ha sido ex-
tirpada en su localidad tipo. Discutimos el impacto de la minería sobre la biodiversidad y los estudios biológicos en Perú.
Introduction
Peru’s complex oreographic structure provides a wide va-
riety of microclimates and ecological zones along its altitu-
dinal gradients, harbouring exceptionally rich biodiversity
characterized by high endemism in both plants and ani-
mals, placing Peru amongst the top “megadiverse” coun-
tries in the world (MN et al. , M et
al. , M et al. ). Conservation International
designated the Tropical Andes as one of the global bio-
diversity “hotspots” because of their high species richness
and endemism (M et al. ). e National Service
of Natural Protected Areas (SERNANP) in Peru is aware
of its responsibility and preserves . of its national ter-
ritory with national, regional, and private natural
areas (SERNANP ). Among them are een nation-
al parks, which are the areas with the highest protection
status. However, fauna and ora outside of protected ar-
eas suer from diverse anthropogenic inuences such as
agriculture, deforestation, pollution, and mining. Peru has
some of the world’s richest mineral deposits (B ),
one of which is the Hualgayoc district (Regíon Cajamar-
ca), ca. km north of the city Cajamarca. e Spanish
discovered the silver deposits of the Hualgayoc district in
(O’P G ), and they have been mined
since colonial times (B S ). Today, other
16
E L et al.
minerals such as zinc, lead, copper, and gold are extracted
through conventional open-pit mining as well (C
). Mines are required to provide reports of biolog-
ical surveys conducted by consultants (e.g., biologists)
to facilitate evaluation of the environmental impacts of
proposed mining operations by the Ministerio de Ener-
gia y Minas (MINEM) and Servicio Nacional Forestal y de
Fauna Silvestre (SERFOR) (Gold Fields , ).
A review of the collection of Pristimantis specimens
at the Museo de Historia Natural Universidad Nacional
Mayor de San Marcos in made the senior author
discover unidentiable specimens (Figs , ) that had
been collected during a biological survey in the Hual-
gayoc district. Herein, we describe these as a new species
based on molecular and morphological data, discuss the
threat status of the new species, and the impact of mining
on the conservation of the new species.
Materials and methods
Morphology and voucher specimens
e format for the description follows L D
(), except that the term “dentigerous processes of vom-
ers” is used instead of “vomerine odontophores” (D-
et al. ), and diagnostic characters follow those of
D L (). Taxonomic classication fol-
lows H et al. () and H et al. (), ex-
cept that we follow P W () for family place-
ment, and P et al. () for names of species groups
in Pristimantis. Information on species for comparative
diagnoses was obtained from D L ()
and from original species descriptions. Specimens were
xed in formol and stored in ethanol. Specimens
were sexed externally by the presence or absence of vocal
slits and internally by the condition of the gonads. Meas-
urements, which were taken with digital callipers under a
microscope by SL and rounded to the nearest . mm, are:
snout–vent length (SVL), tibia length (TL, distance from
the knee to the distal end of the tibia), foot length (FL, dis-
tance from the proximal margin of inner metatarsal tuber-
cle to tip of Toe IV), head length (HL, from angle of jaw to
tip of snout), head width (HW, at level of angle of jaw), hor-
izontal eye diameter (ED), horizontal tympanum diame-
ter (TY), interorbital distance (IOD), upper eyelid width
(EW), internarial distance (IND), eye–nostril distance (E-
N, straight line distance between anterior corner of orbit
and posterior margin of external nares), and egg diame-
ter. Fingers and toes are numbered preaxially to postaxially
from I–IV and I–V, respectively. We compared the lengths
of toes III and V by adpressing both toes against Toe IV;
lengths of ngers I and II were compared by adpressing the
ngers against each other. All drawings were made by SL
using a stereomicroscope with a drawing tube attachment.
Photographs of preserved specimens where taken by SL.
Photographs of life specimens taken by Carlos Diaz were
used for descriptions of skin texture and coloration in life.
Specimens examined are listed in Appendix I; collec-
tion acronyms are: AMNH = American Museum of Nat-
ural History, New York, USA; MCZ = Museum of Com-
parative Zoology, Cambridge, USA; MUSM = Museo de
Historia Natural Universidad Nacional Mayor de San Mar-
cos, Lima, Peru; NMPV = National Museum Prague,
Prague, Czech Republic. reat status was assessed using
the IUCN criteria (). e book “Composition of Scien-
tic Words” by B () was used to form the name
for the new species.
Molecular genetics
We used phylogenetic analyses to conrm the generic
placement of the new species within Pristimantis. We used
a fragment of the mitochondrial S rRNA fragment, be-
cause this gene is the most frequently sequenced gene for
species of Pristimantis (F et al. , H et
al. , V et al. ). For our focal specimens, we
used liver tissues from MUSM and , and ex-
tracted DNA with a commercial extraction kit (IBI Scien-
tic, Peosta, USA). We also obtained DNA sequences from
morphologically similar, or putatively related, species (on
the basis of similarity from BLAST search) of Pristiman-
tis from GenBank (Appendix ). We followed H et
Figure . Map of Peru indicating the type locality (red star) of
Pristimantis astralos sp. n. Map by E. L.
17
New endangered species of Pristimantis from Peru
al. () for DNA amplication and sequencing, with the
Sar (forward) and Sbr (reverse) primers. e thermo-
cycling conditions during the polymerase chain reaction
(PCR) using a Proex thermal cycler (Applied Biosys-
tems) were: one cycle at °C/ min; cycles at °C/s,
°C/ s, °C/. min; and one cycle at °C/ min. We
puried PCR products with Exosap-IT (ermoFisher),
and shipped puried samples to MCLAB (South San Fran-
cisco, CA, USA) for sequencing.
We used Geneious, version .. (Biomatters, http://
www.geneious.com/) to align sequences with the MAFFT
v. alignment program (K S ), aer
manually removing hyper-variable regions, and trimmed
sequences to a length of bp. Our analysis included
terminals. We used MEGA v. (K et al. ) to de-
termine the best evolutionary model. We employed a Max-
imum Likelihood (ML) approach to infer a molecular phy-
logeny using RAxML v. .. (S ) based
on the GTR+G+I model. We assessed node support using
, bootstrap replicates. We also estimated pairwise,
uncorrected genetic distances (p-distances) for S rRNA
between the new species and other species of Pristimantis.
Nomenclatural acts
e electronic edition of this article conforms to the re-
quirements of the amended International Code of Zoo-
logical Nomenclature, and hence the new names contained
herein are available under that Code from the electronic
edition of this article. is published work and the nomen-
clatural acts 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:DD-D-F-BE-CCEBDE.
e electronic edition of this work was published in a jour-
nal with an ISSN, has been archived, and is available from
the following digital repositories: www.salamandra-jour-
nal.com and www.zenodo.org.
Results
According to our maximum likelihood phylogeny (Fig.),
the new species is most closely related to Pristimantis
simonsii. ese two sister species are part of a polytomy
that includes P. rhodoplichus, as well as a group of spe-
cies consisting of P. simonbolivari and species related to
P. croceoinguinis and P. platydactylus. However, there is
little support for several of these nodes (Fig. ), as could
be expected given the known issues with the alignment of
S fragments, reduced fragment length, and small sam-
ple size. Future analyses should expand the dataset to in-
clude other mitochondrial genes. Uncorrected genetic
distances for S and BLAST searches conrm that the
most similar species is P. simonsii (p-distance .–.,
Figure 2. Satellite imagery from GoogleEarth showing the Cerro Corona-Gold Fields Cima Mine (Regíon Cajamarca, Hualgayoc
district) and the type locality (yellow star) of Pristimantis astralos sp. n. is satellite image was taken prior to the destruction of the
type locality at the end of 2014. Figure by E. L.
18
E L et al.
.–. pairwise identity from BLAST search). Fur-
thermore, genetic distances and BLAST searches returned
only three additional specimens with values ~. ese
specimens are a paratype of P. rhodoplichus, KU
(p-distance ., . pairwise identity), a paratype of
P. simon bolivari, KU (p-distance ., . pair-
wise identity), and a member of the P. platydactylus Group
(p-distance ., . pairwise identity; GenBank ac-
cession code EU). All other species of Pristimantis
available on GenBank had uncorrected p-distances >
compared to the focal specimens.
Pristimantis astralos sp. n.
(Figs 4–8)
urn:lsid:zoobank.org:act:DF-FB-BA-C-
F
Holotype: MUSM , adult male (Figs –), from
the Cerro Corona mining concession (°’.’’ S,
°’.’’W), m a.s.l., Comunidad Tingo, Distri-
to Hualgayoc, Provincia Bambamarca, Regíon Cajamarca,
Peru, collected on January by C. D.
Paratypes: ve adult females (MUSM , , ,
, , Fig. ), four adult males (MUSM , ,
, , Fig. ), all collected at the type locality along
with the holotype by C. D.
Generic placement: We assign this species to Pristimantis
based on our molecular data (Fig. ) and its overall mor-
phological similarity to other members of this genus.
Diagnosis: () Skin on dorsum tuberculate, skin on venter
coarsely areolate; discoidal and thoracic folds present, dor-
solateral folds present; () tympanic membrane and tym-
panic annulus present, distinct, visible externally; () snout
broadly rounded to truncate in dorsal view, round in lateral
view; () upper eyelid lacking conical tubercles; EW small-
er than IOD; cranial crest absent; () dentigerous process-
es of vomers oblique; () males with vocal slits, subgular
vocal sac, and with nuptial pads; () Finger I shorter than
Finger II; discs of digits broadly expanded, round, bear-
ing circumferential grooves; () ngers with lateral fring-
es; ()ulnar and tarsal tubercles present; () heel lacking
conical tubercles; inner tarsal fold present; () inner meta-
tarsal tubercle ovoid, – times larger than outer one, outer
metatarsal tubercle small, ovoid; numerous supernumer-
Figure . Maximum Likelihood (ML) phylogeny (best tree) based on a sequence of 503 bp of 16S ribosomal RNA inferred by using
the GTR + G + I model of nucleotide substitution in RAxML. ML bootstrap values > 50% are indicated at each node.
19
New endangered species of Pristimantis from Peru
ary tubercles; () toes with lateral fringes; basal toe web-
bing present; Toe V longer than Toe III; toe discs about as
large as those on ngers, bearing circumferential grooves;
() in life, dorsum ranges from black to dark brown with
or without white ecks; anterior and posterior face of
thighs black to dark brown with white spots; anks black
to dark tan, with or without white or cream ecks; groin
black to dark brown with or without white spots; venter
dark to pale grey with black dots; iris dark copper-colored
with ne black vermiculations; () SVL in adult females
.–. mm (n = ), in adult males .–. mm (n = ).
Comparisons: Pristimantis astralos is distinguished from
its congeners in Peru ( species, AmphibiaWeb ) by
the following combination of characters: skin on dorsum
tuberculate, tympanum and tympanic annulus distinct,
weakly dened dorsolateral folds, uniform black to dark
brown dorsal ground coloration with or without irregular
white ecks, groin black to dark brown with white spots,
and anterior and posterior faces of thighs dark brown with
white spots. Pristimantis astralos can be distinguished from
other Andean species of Peruvian Pristimantis (P. atten-
boroughi, P. ardalonychus, P. atrabracus, P. coronatus, and
P.vilcabambae) that have a dark dorsum as follows (char-
acters of P. astralos in parenthesis):
Pristimantis attenboroughi L M, from al-
titudes of – m a.s.l. in central Peru (Regíon
Junín) shares with P. astralos the presence of dorsolateral
folds, and the usually dark grey dorsal coloration. How-
ever, P. atten boroughi lacks a visible tympanic membrane
(present), a visible tympanic annulus (present), circum-
ferential grooves (present), vocal slits (present), and nup-
tial pads (present). Pristimantis ardalonychus (D
P, ) from altitudes of – m a.s.l. in
northern Peru (Regíon San Martín) shares with P. astralos
a dark dorsal coloration, a tympanic membrane, tympanic
annulus, a reticulated pattern on venter and lateral fring-
es. However, P. ardalonychus has the dorsum smooth with
scattered low tubercles (tuberculate), and smaller females
with SVL . and . mm (.–. mm, n = ; D-
P ). Pristimantis atrabracus (D-
P, ) from altitudes of – m
a.s.l. in northern Peru (Regíon Amazonas) and P. astralos
both have a tympanic membrane, tympanic annulus, vo-
cal slits, lateral fringes on ngers and toes, and a reticu-
lated pattern on their venters. However, P. atrabracus has
the skin on dorsum shagreen (tuberculate), dentigerous
processes of vomers absent (present), and no nuptial pads
(present). Pristimantis coronatus L D,
from m a.s.l. in northern Peru (Regíon Piura) shares
with P.astralos a blackish brown dorsum with white ecks.
However, P. coronatus has the groin marked with orange-
red blotches (white), lacks a tympanic membrane (present),
lacks a tympanic annulus (present), and lacks dorsolater-
al folds (present). Pristimantis vilcabambae L,
from m a.s.l in southern Peru (Regíon Cusco) shares
with P. astralos a dark dorsal coloration and white spots
on the groin. However, P. vilcabambae lacks dorsolateral
folds (present), a tympanic membrane (present), tympanic
annulus (present), vocal slits (present), and nuptial pads
(present).
Pristimantis chimu L, from – m a.s.l.
of the Pacic versants of northern Peru (Regíon Cajamar-
ca) occurs syntopically with P. astralos. Pristimantis chimu
shares with P. astralos a tympanic membrane and annulus,
rounded snout, dentigerous processes of vomers, and lat-
eral fringes on toes and ngers. However, P. chimu lacks
nuptial pads (present), vocal slits (present), and has a white
groin with brown blotches (black with white dots). Fur-
thermore, P. chimu has smaller males (SVL .–. mm
[n = ] vs. SVL .–. mm [n = ] in P. astralos) and
smaller females (SVL .–. mm [n = ] vs. SVL .–
. mm [n = ] in P. astralos; Lehr ).
Pristimantis pinguis (D P, ) from
altitudes of – m a.s.l. (D P
, D L ) occurs syntopically with
P. astralos. Pristimantis pinguis has smaller females (SVL
.–. mm, n = vs. .–. mm, n = , in P. astra-
los), lacks dorsolateral folds (present), and has the venter
dull yellow with greyish brown reticulations (dark to pale
grey with black dots).
Pristimantis mariaelenae V D,
from m a.s.l. in the humid puna (Regíon Lambayeque)
in northeastern Peru shares with P. astralos a visible tym-
panic membrane and tympanic annulus, rounded snout,
dentigerous processes of vomers, vocal slits, nuptial pads,
and white blotches in the groin area. However, P. mariae-
lenae lacks lateral fringes on ngers and toes (present),
and its dorsal coloration is reddish brown (black to dark
brown). Furthermore, P. mariaelenae has smaller males
(SVL .–. mm [n = ] vs. SVL .–. mm [n= ]
in P. astralos) and smaller females (SVL .–.mm [n=
] vs SVL .–. mm [n = ] in P. astralos; V
D ). Pristimantis stipa V D-
, from m a.s.l. in the humid puna (Regíon
Lambayeque) in northeastern Peru shares with P. astralos
a similar SVL, dorsolateral folds, and a visible tympanic
membrane and tympanic annulus. However, P. stipa has
digits without circumferential grooves (present), and ulnar
tubercles that are coalesced into a low fold (absent; V-
D ).
Genetically, P. astralos is most closely related (.–.
S genetic distance, Fig. ) to P. simonsii (B,
). Pristimantis simonsii inhabits the humid puna at al-
titudes of – m a.s.l. in northern Peru (Regíon Ca-
jamarca) (D L ) and occurs syntopi-
cally with P. astralos. Both species are of similar size (male
SVL .–. mm in P. simonsii vs. .–. mm [n = ]
in P. astralos; female SVL .–. mm vs. .–. mm
[n = ] in P. astralos; D L ), share the
presence of dorsolateral folds, and nuptial pads. Howev-
er, P. simonsii lacks a tympanic membrane and a tympan-
ic annulus (both present), has males that lack vocal slits
(present), lacks ngers with lateral fringes (present), lacks
circumferential groves (present), lacks dentigerous proc-
esses of vomers (present), and has in life the dorsum dull
20
E L et al.
brown to pinkish tan with dark brown spots (black to dark
brown with irregular scattered white ecks).
Description of the holotype: Head narrower than the body,
slightly longer than wide; head width . of SVL; head
length . of SVL; cranial crest absent; snout bluntly
rounded in dorsal and lateral views (Figs A, B), eye–nos-
tril distance . of eye diameter; nostrils protuberant,
directed dorsolaterally; canthus rostralis straight in dorsal
view, rounded in prole; loreal region slightly concave; lips
rounded; upper eyelid without enlarged tubercles; upper
eyelid width . of IOD; supratympanic fold distinct,
narrow, extending diagonally from posterior margin of up-
per eyelid towards insertion of arm, covering upper mar-
gin of the tympanum (Fig. A); tympanic membrane and
tympanic annulus distinct, externally visible, tympanum
. of ED; two conical postrictal tubercles present bilat-
erally (Fig. A). Choanae small, teardrop-shaped; dentiger-
ous processes of vomers oblique and small; tongue width
is half of the tongue length, notched posteriorly, posterior
third free; vocal slits slightly curved, located in posterior
half of mouth oor between tongue and margin of jaw;
subgular vocal sac distinct (Fig. C).
Skin on dorsum tuberculate, dorsolateral fold weakly
dened and in weak contrast to the tuberculate skin tex-
ture (Fig. A); skin on anks tuberculate with tubercles
coalescing into short ridges; skin on throat and chest sha-
green, belly coarsely areolate; discoidal and thoracic weak-
ly dened (Fig. C); cloacal sheath short.
Outer surface of ulnar with minute tubercles; outer pal-
mar tubercle bid (Fig. A); distinct supernumerary tu-
bercles, ovoid, approximately half the size of subarticular
tubercles; subarticular tubercles well dened, most prom-
inent on ngers, round in ventral view, conical in lateral
view; ngers with narrow lateral fringes; Finger I shorter
than Finger II (Fig. A); discs on digits of ngers broadly
expanded, rounded, bearing circumferential grooves.
Hind limbs short, slender, tibia length . of SVL;
foot length . of SVL; upper surface of hind limbs tu-
berculate with few scattered tubercles; inner surface of
thighs smooth, posterior and ventral surfaces of the thighs
areolate; heels without conical tubercles; outer surface
of tarsus with minute tubercles; inner tarsal fold short
(Fig.B); inner metatarsal tubercle ovoid, two times the
size of ovoid outer metatarsal tubercle; subarticular tuber-
cles well dened, round in ventral view, conical in lateral
view; plantar supernumerary tubercles distinct, about one-
fourth the size of subarticular tubercles; toes with narrow
lateral fringes, fringe of Toe V undulated; basal webbing
present, most prominent between toes IV and V; discs ex-
panded, slightly truncated, approximately the same size as
discs on ngers, bearing circumferential grooves; relative
length of toes: <<<< (Fig. B).
Measurements (in mm) of the holotype: SVL .;
TL.; FL .; HL .; HW .; ED .; TY .; IOD .;
EW .; IND .; E-N ..
Coloration in preservative (Fig. ): e dorsal ground
coloration is dark charcoal with irregular creamy white
Figure . Holotype of Pristimantis astralos sp. n. (MUSM 32752) in alcohol in dorsal (A) and ventral (B) views. SVL = 27.2 mm.
Photos by S. L.
21
New endangered species of Pristimantis from Peru
ecks; anks charcoal with relatively larger, irregular,
cream, rounded dorsolateral white ecks; groin and ante-
rior faces of thighs brown with a cream tint; chest and belly
cream and dark brown, with reticulations forming an ir-
regular dark brown stripe along the midline of venter; ven-
tral faces of thighs cream, throat pale cream and mottled
with pale brown; palmar and plantar faces, and ngers and
toes cream; iris pale bluish grey.
Coloration in life (Fig. ): e dorsal ground coloration
is black with white ecks and spots; anks black, with rela-
tively large, irregular, rounded white spots; groin and an-
terior faces of thighs dark charcoal with white spots; chest
and belly pale grey and dark brown, with an irregularly
reticulated dark brown stripe along the midline of ven-
ter; posterior faces of thighs black with white spots; throat
greyish purple with a pale grey tint; palmar and plantar
faces, and ngers and toes brown; iris dark copper with
ne black vermiculations.
Variation: All paratypes (Figs , ) are similar to the holo-
type regarding morphology and proportions. Besides dif-
ferences in size, morphological variation is noticeable in
the dorsolateral folds, which may be prominent (MUSM
, , Figs G, H) or weakly dened (MUSM ,
, Figs A, B; MUSM , Figs D, E; MUSM ,
, Figs A, B; MUSM , Figs D, E). Skin texture
on the anks is also variable, with tubercles either fused
into prominent, continuous ridges (MUSM , Fig. G)
or weakly dened ridges (MUSM , , Figs A, B;
MUSM , , Figs D, E; MUSM , , Figs
A, B; MUSM , Figs D, E). Most specimens have bi-
laterally two distinct conical postrictal tubercles, where-
as some specimens have the postrictal tubercles fused
into short ridges (MUSM , Fig. B), being elongated
(MUSM , Fig. E), or weakly dened (MUSM ).
e dorsal coloration in life ranges from black (MUSM
, Fig. B; MUSM , , Fig. D; MUSM ,
Figure . Holotype of Pristimantis astralos sp. n. (MUSM 32752)
in life, in lateral (A), dorsal, (B), and ventral (C) views. Photos
by C. D.
Figure . Ventral views of hand (A) and foot (B) of the holotype
of Pristimantis astralos sp. n. (MUSM 8796). Drawings by S. L.
22
E L et al.
Fig. A; MUSM , Fig. D) to dark brown (MUSM
, , Fig. D; MUSM , Fig. G). Some speci-
mens have white, irregularly shaped ecks scattered on the
dorsum (MUSM , , Fig. A). While the groin is
dark grey with creamy white patches in all specimens, the
number of patches ranges from one (MUSM , ,
Fig. H) to four (MUSM ). e venter may be dark
grey (MUSM , , Fig. F), pale grey (MUSM
, , Fig. C) or creamy white (MUSM , ,
Fig. C; MUSM , Fig. F; MUSM , Fig. I).
Some specimens have the reticulated dark brown ventral
blotches fused into a thin midline (MUSM , Fig. C;
MUSM , Fig. I), or dark brown dots (MUSM ,
, , Fig. F; MUSM , Fig. C; MUSM ,
Fig. F). Only one female has a reddish brown interorbital
bar (MUSM , Fig. G).
Etymology: e species epithet astralos is a Greek adjec-
tive meaning “spotted with stars” or “speckled”. e name
refers to the dorsal white spots on black background of the
species that is reminiscent of stars in the night sky.
Distribution, natural history, and threat status: Hualgayoc
is situated in the Cajamarca Regíon of northern Peru, be-
tween and m a.s.l., about km north of the
city Cajamarca, and east of the continental divide of the
Cordillera Occidental (C , Fig. ). Specimens
were obtained on January in the concession of the
Mina Cerro Corona-Gold Fields La Cima, Comunidad
Tingo, in the high Andean grasslands (jalca) consisting of
Peruvian feather grass and Puya fastuosa M, (Bro-
meliaceae). Specimens of P. astralos were found sheltering
in P. fastuosa plants. Up to specimens (juveniles, males,
females) were found in the same plant, and P. astralos ap-
parently prefers hiding at the bases of larger plants under
dead leaves where humidity levels are higher (C. D,
pers. comm.). Syntopic frogs at the time of the nd includ-
ed Pristimantis chimu, P. pinguis, P. simonsii, and a syn-
topic lizard was Stenocercus stigmosus C, , all of
which use Puya fastuosa plants as refuges (C. D, pers.
comm.).
Pristimantis astralos is so far only known from its type
locality at m a.s.l. in the Cordillera Occidental. Min-
ing operations, and especially large-scale dumping of
mining debris at the type locality (pers. comm. C.D)
by the end of , have destroyed the type locality and
caused the extirpation of the population of P. astralos at
the type locality (Figs , ). We are unaware of similar
habitats with Puya fastuosa in the proximity of the type
locality where P.astralos could be still in existence. Con-
Table 1. Morphometrics (in mm) of the individual type specimens of Pristimantis astralos sp. n.
Characters MUSM
32755
MUSM
32759
MUSM
32753
MUSM
32760
MUSM
32758
MUSM
32752
MUSM
32757
MUSM
32754
MUSM
2756
Sex F F F F F M M M M
SVL 32.8 31.5 31.0 27.8 25.6 27.2 27.0 24.9 23.6
TL 12.1 11.5 12.3 12.0 11.0 10.0 10.0 9.3 8.5
FL 12.2 12.7 11.9 12.2 11.5 9.6 11.0 9.6 9.4
HL 11.6 10.6 12.0 9.1 8.8 10.5 9.9 8.8 8.8
HW 12.6 11.2 12.0 9.8 8.2 9.5 10.3 9.4 8.9
ED 3.1 3.2 3.2 3.2 3.2 3.0 3.2 3.1 2.9
TY 1.8 1.8 1.6 1.8 1.6 1.4 1.6 1.4 1.3
IOD 3.3 3.5 3.9 3.5 3.2 3.0 3.2 2.9 3.0
EW 2.7 2.5 2.1 2.3 2.2 2.1 2.3 2.0 2.3
IND 2.5 2.2 2.3 3.7 2.0 2.0 2.3 1.9 2.5
E–N 3.2 2.7 3.2 3.7 2.4 2.3 2.1 2.3 2.0
Table 2. Morphometrics (in mm) and proportions of the type
series of Pristimantis astralos sp. n.; ranges followed by means
and one standard deviation in parentheses.
Characters Males (n = 4) Females (n = 5)
SVL 23.6–27.2 (25.7 ± 1.5) 25.6–32.6 (29.7 ± 2.6)
TL 8.5–10.0 (9.5 ± 0.6) 11.0–12.3 (11.8 ± 0.5)
FL 9.4–11.0 (9.9 ± 0.6) 11.5–12.7 (12.1 ± 0.4)
HL 8.8–10.5 (9.5 ± 0.7) 8.8–12.0 (10.4 ± 1.3)
HW 8.9–10.3 (9.5 ± 0.5) 8.2–12.6 (10.8 ± 1.6)
ED 2.9–3.2 (3.1 ± 0.1) 3.1–3.2 (3.2 ± 0.0)
TY 1.3–1.6 (1.4 ± 0.1) 1.6–1.8 (1.7 ± 0.1)
IOD 2.9–3.2 (3.0 ± 0.1) 3.2–3.9 (3.5 ± 0.2)
EW 2.0–2.3 (2.2 ± 0.1) 2.1–2.7 (2.4 ± 0.2)
IND 1.9–2.5 (2.2 ± 0.2) 2.0–3.7 (2.5 ± 0.6)
E–N 2.0–2.3 (2.2 ± 0.1) 2.4–3.7 (3.0 ± 0.4)
TL/SVL 0.36–0.37 0.37–0.43
FL/SVL 0.35–0.41 0.37–0.45
HL/SVL 0.35–0.39 0.33–0.39
HW/SVL 0.35–0.38 0.32–0.39
HW/HL 0.9–1.0 0.9–1.1
E–N/ED 0.66–0.77 0.75–1.16
EW/IOD 0.69–0.77 0.54–0.82
TY/ED 0.45–0.50 0.50–0.58
23
New endangered species of Pristimantis from Peru
sequently, we suggest the threat status for P. astralos to
be ‘critically endangered’ (CR). e current conserva-
tion status of Puya fastuo sa is ‘endangered’ (EN, T
Z ).
Discussion
Peru is the leading gold producer in South America, and
with metric tons per year the sixth largest in the world
Figure 7. Coloration variation of male Pristimantis astralos sp. n.: Le column MUSM 32754, SVL 24.9 mm; right column MUSM
32756, SVL 23.6 mm in dorsal (A, D), lateral (B, E), and ventral (C, F) views. Photos by C. D.
24
E L et al.
in (World Atlas ). Metals have been mined for
thousands of years in Peru and used by various pre-Co-
lumbian cultures for artworks and tools long before the
Spanish colonization. Alexander von Humboldt visited the
Andean silver mines of Hualgayoc in and commented
critically on the destructive techniques employed and the
inhumane treatment of the workers (O’P G
). e widespread habitat destruction has continued
until today (Fig. ). e Cerro Corona – Gold Fields La
Cima Mine in Peru produces copper and gold by conven-
tional open-pit mining methods. With a gold content of ca.
g per ton, and the expectation to mine . MOZ (= .
metric tons) of gold over a -year period (Gold Fields
), the accumulated debris is immense.
In areas with intensive mining, habitat destruction and
pollution can be drastic and threaten endemic species. Ac-
cording to Peruvian law, companies that have an expected
impact on nature are obliged to hire consultants (e.g., bio-
logists, engineers) to conduct environmental impact as-
sessments. ese assessments include lists of species found
before a company starts their activities. e nal technical
reports are then sent for review to the Ministry of the En-
vironment.
e Tropical Andes are recognized as a biodiversity
hotspot (M et al. ) for animals and plants, many
of which have small-scale distributions that render them
highly sensitive to environmental impacts. Mining com-
panies are aware that the presence of potentially new or
endangered species can interfere with their activities. As
a result, new species may not be highlighted in survey re-
ports by environmental consulting rms under pressure
from mining operators. Instead, potential new species may
be identied only to genus level (e.g., as Pristimantis sp.),
or tentatively assigned to the most similar species (e.g., as
Pristimantis cf. simonsii, as was done for the new species
described herein). In order to better understand the threat
caused by mining activities in the Peruvian Andes, A-
et al. () surveyed amphibians in mining conces-
sions, and noted a decline in species richness in two con-
cessions where two previously recorded species (Atelopus
peruensis G C, ; Nannophryne copho-
tis (B, )) are now absent. For the latter two
Figure 8. Coloration variation of female Pristimantis astralos sp. n.: Le column MUSM 32758, SVL = 25.6 mm; middle column
MUSM 32759, SVL = 31.5 mm; right column MUSM 32760, SVL = 27.8 mm in dorsal (A, D, G), lateral (B, E, H), and ventral views
(C, F, I). Photos by C. D.
25
New endangered species of Pristimantis from Peru
critically endangered species (IUCN SSC Amphibian Spe-
cialist Group a, b), additional factors apart from habi-
tat destruction/pollution such as diseases (e.g., Batracho-
chytrium dendrobatidis, also see C ) may
have contributed to their local disappearance.
B et al. () analysed the land-use conicts be-
tween vertebrate biodiversity conservation and extractive
industries in the forested eastern Peruvian Andes from
to m a.s.l., and found that of endemic-rich are-
as overlapped with mining concessions. Furthermore, B
et al. () reported that of all endemic species have
geographical distributions that overlap by more than
with concession areas, and that less than of all endemic
mammal, bird, amphibian and reptile species are protect-
ed within reserves (B F ). We assume
that conservation gaps of similar severity exist for endemic
species in the western Andes of Peru. Pristimantis astra-
los exemplies how quickly habitat destruction in a mining
concession can extirpate species. In their report, Gold
Fields mentioned that Lupinus peruvianus (ora), amphib-
ians, and reptiles had been relocated in (Gold Fields
: ), but the success of these relocations remains un-
evaluated and unknown. e naming of new species is the
rst important step towards conservation, and we hope
that MINEM and SERFOR will initiate surveys at Hual-
gayoc and its surroundings to assess the current popula-
tion status of P. astralos.
Acknowledgements
We thank J. K for his suggestions and comments, which
helped to improve our manuscript. For loan of material we thank
J. C and C. A (MUSM), J. R (MCZ), D. K-
(AMNH), and J. M (NMP). We thank C. D for
making available his photos. We thank C. A, J. B, P.
C, and J. P for their comments and information on
mining in Peru.
References
A, C., R. G, C. R, J. S, C. T
K. S-T (): Anbios andinos y estudios de impacto
ambiental en concesiones mineras de Perú. – Alytes, : –
.
AmphibiaWeb (): AmphibiaWeb. Information on amphi-
bian biology and conservation. – http://amphibiaweb.org/, ac-
cessed July .
B, V. W. F (): Conservation gaps and priori-
ties in the Tropical Andes biodiversity hotspot: implications
for the expansion of protected areas. – Journal of Environ-
mental Management, : –.
B, V., W. F A. D (): Land-use con-
icts between biodiversity conservation and extractive indus-
tries in the Peruvian Andes. – Journal of Environmental Man-
agement, : –.
B S, E. (): e Bishop’s Utopia: Envisioning
Improvement in Colonial Peru. – University of Pennsylvania
Press, pp.
B, R. W. (): Composition of Scientic Words. – Smith-
sonian Institution Press, Washington and London, pp.
B, J. (): Mining mountains: neoliberalism, land tenure,
livelihoods, and the new Peruvian mining industry in Ca-
jamarca. – Environment and Planning A : –.
C, S. (): Stratabound ore deposits of Hualgayoc, Ca-
jamarca, Peru. – pp. – in: F, L., G. C. A,
M. C, E. C J. F (eds): Stratabound Ore
Deposits in the Andes. – Special Publication No. of the Soci-
ety for Geology Applied to Mineral Deposits, vol. . Springer,
Berlin, Heidelberg, pp.
C, A. (): State of the world’s amphibians. – Annual
Review of Environment and Resources, : –.
D, W. E. E. L (): Terrestrial-breeding Frogs
(Strabomantidae) in Peru. – Natur und Tier-Verlag, Münster,
Germany, pp.
D, W. E. J. B. P (): Frogs of the genus Eleu-
therodactylus (Anura: Leptodactylidae) in the Andes of north-
ern Peru. Scientic Papers. – Natural History Museum, Uni-
versity of Kansas, : –.
D, W. E., E. L P. V () Two new species
of Eleutherodactylus (Anura: Leptodactylidae) from northern
Peru. – Zootaxa, : –.
F, A., M. V, M.-D. S, A. M, C. M-
, M. B A. G (): Revealing cryptic diversity
using molecular phylogenetics and phylogeography in frogs of
the Scinax ruber and Rhinella margaritifera species groups. –
Molecular Phylogenetics and Evolution, : –.
Gold Fields (): Reporte integrado. – Gold Fields en Perú, pp.
. Accessible at: https://www.goldelds.com.pe/
Gold Fields (): Reporte integrado. – Gold Fields en Perú, pp.
. Accessible at: https://www.goldelds.com.pe/
H, S. B., W. E. D M. P. H (): New
world direct-developing frogs (Anura: Terrarana): molecular
phylogeny, classication, biogeography, and conservation. –
Zootaxa, : –.
H, M. P., W. E. D, L. T, D. B. M, R. D.
MC S. B. H (): A new frog family (An-
ura: Terrarana) from South America and an expanded direct-
developing clade revealed by molecular phylogeny. – Zootaxa,
: –.
IUCN SSC Amphibian Specialist Group (a): Atelopus pe-
ruensis. – e IUCN Red List of reatened Species :
e.TA,downloaded on June .
IUCN SSC Amphibian Specialist Group (b): Nannophryne
cophotis. – e IUCN Red List of reatened Species :
e.TA,downloaded on June .
IUCN Standards and Petitions Subcommittee (): Guidelines
for using the IUCN Red List categories and criteria. Version
. – Prepared by the Standards and Petitions Subcommittee.
Downloadable from http://www.iucnredlist.org/documents/
RedListGuidelines.pdf.
K, K. D. M. S (): MAFFT multiple sequence
alignment soware version : improvements in performance
and usability. – Molecular Biology and Evolution, : –
.
K, S., G. S K. T (): MEGA: Molecular
Evolutionary Genetics Analysis version . for bigger datasets.
– Molecular Biology and Evolution, : –.
26
E L et al.
L, E. (): New eleutherodactyline frogs (Leptodactylidae:
Pristimantis, Phrynopus) from Peru. – Bulletin of the Muse-
um of Comparative Zoology. Cambridge, Massachusetts, :
–.
L, E. R. M (): A new species of terrestrial-breed-
ing frog (Amphibia, Craugastoridae,Pristimantis) from high
elevations of the Pui Pui Protected Forest in central Peru. –
ZooKeys, : –.
L, J. D. W. E. D (): Frogs of the genus Eleu-
therodactylus in western Ecuador: systematics, ecology, and
biogeography. – Special Publication Natural History Museum
University of Kansas, : –.
MN, J. A., K. R. M, W. V. R, R. A. M
T. B. W (): Conserving the World’s Biological Di-
versity. – IUCN, World Resources Institute, Conservation In-
ternational, WWFUS and the World Bank: Washington, DC.
M, N., R. M, C. M, G. F
J. K (): Biodiversity hotspots for conservation priori-
ties. – Nature, : –.
O’P G, S. (): Chapter : A German mineralo-
gist visits Peru. – pp. – in: E, R., M. A. F
B. S (coord.): Alexander von Humboldt. From the
Americas to the Cosmos. – Bildner Center for Western Hemi-
sphere Studies, e Graduate Center, e City University of
New York, pp. .
P, J. M., T. G D. R. F (): Molecular sys-
tematics of terraranas (Anura: Brachycephaloidea) with an as-
sessment of the eects of alignment and optimality criteria.
– Zootaxa, : –.
P, R. A. J. J. W (): A large-scale phylogeny of Am-
phibia including over species, and a revised classication
of extant frogs, salamanders, and caecilians. – Molecular Phy-
logenetics and Evolution, : –.
SERNANP ): Guía Ocial De Áreas Naturales Protegidas
Del Perú. – Lima, Peru, pp. .
S, A. : RAxML version : a tool for phylogenetic
analysis and post-analysis of large phylogenies. – Bioinformat-
ics, : –.
T Z, I. (): Puya fastuosa. – e IUCN Red
List of reatened Species : e.TA,
downloaded on June .
V, P. J. W. E. D (): Two syntopic new spe-
cies of the Pristimantis orestes Group (Anura: Strabomantidae)
from northwestern Peru. – Zootaxa, : –.
World Atlas (): Available online: https://www.worldatlas.
com/articles/top--gold-producers-in-the-world.html, ac-
cessed on July .
Appendix 1
Comparative material examined.
Pristimantis aaptus: Colombia: Amazonas: Puerto Nariño: MCZ
A-. Pristimantis attenboroughi: Peru: Junín: Pui Pui Protect-
ed Forest: Hatunpata, m: NMPV , Antuyo Bajo,
m a.s.l.: NMPV . Pristimantis chimu: Peru: Cajamarca:
–km NW El Pargo (Llama-Huambos Rd.), – m a.s.l.:
MCZ –. Pristimantis mariaelenae: Peru: Lambayeque:
Cañaris, – m a.s.l.: MUSM . Pristimantis puipui:
Peru: Junín: Pui Pui Protected Forest, Laguna Sinchon, m
a.s.l.: MUSM (holotype), MUSM , , NMPV
–, –, all paratypes. Pristimantis seorsus: Peru: Cus-
co: Cordillera Vilcabamba, m: AMNH A–. Pristi-
mantis simonsii: Peru: Cajamarca: . km NE Encanada, m
a.s.l.: MUSM –. Pristimantis stipa: Peru: Lambayeque:
Cañaris, – m a.s.l.: MUSM –. Pristimantis vilca-
bambae: Peru: Cusco: Cordillera Vilcabamba, m: AMNH
A–.
Appendix 2
GenBank accession codes for the taxa used in the molecular ge-
netic analysis in this study.
Species Voucher 16S rRNA
Pristimantis altamazonicus CORBIDI 16778 MG820143
Pristimantis aradalonychus KU 212301 EU186664
Pristimantis astralos sp. n. MUSM 32753 MT968733
Pristimantis astralos sp. n. MUSM 32755 MT968732
Pristimantis attenboroughi NMP6V 75524 KY594754
Pristimantis attenboroughi NMP6V 75525 KY594755
Pristimantis bounides NMP6V 75097 KY962797
Pristimantis bounides MUSM 31198 KY962794
Pristimantis buccinator MUSM 33269 KY652650
Pristimantis cf. carvalhoi CORBIDI 16294 KY652651
Pristimantis croceoinguinis MC 11557 DQ195455
Pristimantis croceoinguinis QCAZ 18231 MH516171
Pristimantis croceoinguinis QCAZ 25444 MH516173
Pristimantis croceoinguinis QCAZ 25788 MH516176
Pristimantis cf. croceoinguinis MUSM 31154 KY594759
Pristimantis fenestratus CORBIDI 16222 MT968731
Pristimantis humboldti MUSM 31194 KY962798
Pristimantis humboldti NMP6V 75538 KY962799
Pristimantis lindae MUSM 27902 KY652653
Pristimantis ockendeni RvM 5.12 KY652654
Pristimantis phoxocephalus KU 218025 EF493349
Pristimantis platydactylus MVZ 272359 KY652656
Pristimantis platydactylus MNCN-DNA
4138 EU712671
Pristimantis puipui NMP6V 75542 KY962800
Pristimantis rhodoplichus KU 219788 EF493674
Pristimantis ridens AJC 1778 KR863320
Pristimantis simonbolivari KU 218254 EF493671
Pristimantis simonsii N/A AM039641
Pristimantis simonsii KU 212350 EU186665
Pristimantis salaputium MUSM 27916 KY652658
