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A new, critically endangered species of Pristimantis (Amphibia: Anura: Strabomantidae) from a mining area in the Cordillera Occidental of northern Peru (Región Cajamarca)

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We describe a new species of Pristimantis from high Andean grasslands (jalca) at 3600 m above sea level in northern Peru (Región Cajamarca) based on morphological and molecular characters. The new species is known from four males and five females, which were found sheltering in the rosettes of Puya fastuosa (Bromeliaceae). The phylogenetic analysis of a fragment of the mitochondrial 16S rRNA gene suggests that the new species is a sister taxon of Pristimantis simonsii. The new species differs from its congeners by having a black dorsum speckled with white flecks and a dark brown groin with white spots. Furthermore, adult males have a snout-vent length of 23.6-27.2 mm (n = 4), and adult females of 25.6-32.8 mm (n = 5). 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. Resumen. Describimos una nueva especie de Pristimantis de pastizales altoandinos (jalca) a 3600 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 filogenético de un fragmento del gen mitocondrial 16S 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 23.6-27.2 mm (n = 4), y las hembras adultas de 25.6-32.8 mm (n = 5). 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.
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15
New endangered species of Pristimantis from Peru
Open access at hp://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 diers 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 (MN 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 suer from diverse anthropogenic inuences 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 unidentiable 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 classication 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; NMPV = National Museum Prague,
Prague, Czech Republic. reat status was assessed using
the IUCN criteria (). e book “Composition of Scien-
tic Wordsby B () was used to form the name
for the new species.
Molecular genetics
We used phylogenetic analyses to conrm 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-
tic, 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 amplication and sequencing, with the
Sar (forward) and Sbr (reverse) primers. e thermo-
cycling conditions during the polymerase chain reaction
(PCR) using a Proex 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
puried PCR products with Exosap-IT (ermoFisher),
and shipped puried 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 ), aer
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 Identier) for this publication is: urn:lsid:zoobank.
org:pub:DD-D-F-BE-CCEBDE.
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 conrm 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 dened 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 Pacic 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 prole; 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
dened 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 dened (Fig. C); cloacal sheath short.
Outer surface of ulnar with minute tubercles; outer pal-
mar tubercle bid (Fig. A); distinct supernumerary tu-
bercles, ovoid, approximately half the size of subarticular
tubercles; subarticular tubercles well dened, 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 dened, 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 dened (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 dened 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 dened (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 identied 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 conicts 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 exemplies 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.
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Appendix 1
Comparative material examined.
Pristimantis aaptus: Colombia: Amazonas: Puerto Nariño: MCZ
A-. Pristimantis attenboroughi: Peru: Junín: Pui Pui Protect-
ed Forest: Hatunpata,  m: NMPV , Antuyo Bajo, 
m a.s.l.: NMPV . 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 , , NMPV
–, –, 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
... The amphibian diversity in the tropical Andes is outstanding (Duellman, 1988;Myers et al., 2000;Hutter et al., 2013Hutter et al., , 2017. Each year, several species are described from montane forests of this biodiversity hotspot (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- Cisneros-Heredia and Gluesenkamp, 2010;Yánez-Muñoz et al., 2010a;Páez-Moscoso and Guayasamin, 2012;Almendáriz et al., 2014;Guayasamin et al., 2017aGuayasamin et al., , 2019Reyes-Puig et al., 2019c). ...
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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 relationships. The new species is differentiated from other species of Noblella by having discs of fingers 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. The 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.
... These cryptic species can be so similar that some species can only be distinguished by their advertisement call and genetic data [26,28,29]. The Andes of northern Peru, i.e., the Departments of Lambayeque, Piura, Cajamarca, La Libertad, Amazonas, and San Martín, harbor 52 species of Pristimantis [2,21,[30][31][32]. Sixty-seven percent of this species richness was described in the last three decades as a product of the herpetological exploration of remote places (i.e., Venegas and Duellman [32], Duellman and Wild [33], Lehr, et al. [34]) and systematics revisions (i.e., Duellman and Lehr [2], Duellman and Pramuk [21], Lehr [35]). ...
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We describe two new species of terrestrial-breeding frogs in the genus Pristimantis from the Andes of northeastern Peru, Amazonas Department. Both species share several characters with other congeners from northern Peru, such as the presence of prominent conical tubercles on their eyelids and heel, prominent conical tubercles along the outer edge of the tarsus, and discs on fingers and toes widely expanded. However, both species can be diagnosed from morphologically similar Pristimantis in the region. Pristimantis kiruhampatu has axillae, groins, and hidden surfaces of hindlimbs that are cherry with white minute flecks, tympanic membrane and tympanic annulus evident, conical tubercles along the edge of snout and outer edge of tibia, and \/ shaped folds in the scapular region. Pristimantis paulpittmani has yellow or dirty cream groins and hidden surfaces of hindlimbs, whitish cream irises with scattered dark brown reticulations, and a thin vertical dark brown streak at the middle of the eye, snout subacuminate with a conical tubercle at the tip, and lacks a tympanic annulus and membrane. Additionally, we provide a short description of the advertisement call of P. kiruhampatu.
... Indeed, major part of the Moquegua region territory is under mining concessions, however, only a small part of the area is currently being explored or mined (INGEMMET, 2019) (Fig. 19). The exploration phase, the vegetation removal and ground disturbance led to an immediate impact on the local biodiversity (Farrel & Kratzing, 1996), consequently the areas with intensive mining, habitat destruction and contamination can be drastic and threaten endemic species (Lehr et al., 2021). Considering that theraphosids have sedentary habits, limited dispersal mechanisms due their poor vagility (Ferretti et al., 2014) and very long reproductive periods because of their great longevity, habitat disturbance could contribute to the decreasing population densities of E. vanessae sp. ...
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A new species of Euathlus Ausserer, 1875 (Araneae: Theraphosidae: Theraphosinae), Euathlus vanessae sp. nov. is described and diagnosed. The genus is known from Chile and Argentina, and E. vanessae sp. nov. represents the first record for the genus in Peru, being the new northernmost limit of its known distribution. Distribution and natural history data are provided. Additionally, the conservation status of the new species is discussed and it is proposed as Endangered (EN) according to IUCN criteria.
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Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community. I present some of the most notable new features and extensions of RAxML, such as, a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX, and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date, 50 page user manual covering all new RAxML options is available. The code is available under GNU GPL at https://github.com/stamatak/standard-RAxML. Alexandros.Stamatakis@h-its.org.
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We report a major update of the MAFFT multiple sequence alignment program. This version has several new features, including options for adding unaligned sequences into an existing alignment, adjustment of direction in nucleotide alignment, constrained alignment and parallel processing, which were implemented after the previous major update. This report shows actual examples to explain how these features work, alone and in combination. Some examples incorrectly aligned by MAFFT are also shown to clarify its limitations. We discuss how to avoid misalignments, and our ongoing efforts to overcome such limitations.
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In December 1788, in the northern Peruvian city of Trujillo, fifty-one-year-old Spanish Bishop Baltasar Jaime Martínez Compañón stood surrounded by twenty-four large wooden crates, each numbered and marked with its final destination of Madrid. The crates contained carefully preserved zoological, botanical, and mineral specimens collected from Trujillo’s steamy rainforests, agricultural valleys, rocky sierra, and coastal desert. To accompany this collection, the Bishop had also commissioned from Indian artisans nine volumes of hand-painted images portraying the people, plants, and animals of Trujillo. He imagined that the collection and the watercolors not only would contribute to his quest to study the native cultures of Northern Peru but also would supply valuable information for his plans to transform Trujillo into an orderly, profitable slice of the Spanish Empire. Based on intensive archival research in Peru, Spain, and Colombia and the unique visual data of more than a thousand extraordinary watercolors, The Bishop’s Utopia recreates the intellectual, cultural, and political universe of the Spanish Atlantic world in the late eighteenth century. Emily Berquist Soule recounts the reform agenda of Martínez Compañón-including the construction of new towns, improvement of the mining industry, and promotion of indigenous education-and positions it within broader imperial debates; unlike many of his Enlightenment contemporaries, who elevated fellow Europeans above native peoples, Martínez Compañón saw Peruvian Indians as intelligent, productive subjects of the Spanish Crown. The Bishop’s Utopia seamlessly weaves cultural history, natural history, colonial politics, and art into a cinematic retelling of the Bishop’s life and work.
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Hualgayoc is one of the typical “complex mining districts” of the Central Andes, characterized by the influence of many superimposed geologic events and oreforming processes.
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The extant amphibians are one of the most diverse radiations of terrestrial vertebrates (>6800 species). Despite much recent focus on their conservation, diversification, and systematics, no previous phylogeny for the group has contained more than 522 species. However, numerous studies with limited taxon sampling have generated large amounts of partially overlapping sequence data for many species. Here, we combine these data and produce a novel estimate of extant amphibian phylogeny, containing 2871 species (∼40% of the known extant species) from 432 genera (∼85% of the ∼500 currently recognized extant genera). Each sampled species contains up to 12,712 bp from 12 genes (three mitochondrial, nine nuclear), with an average of 2563 bp per species. This data set provides strong support for many groups recognized in previous studies, but it also suggests non-monophyly for several currently recognized families, particularly in hyloid frogs (e.g., Ceratophryidae, Cycloramphidae, Leptodactylidae, Strabomantidae). To correct these and other problems, we provide a revised classification of extant amphibians for taxa traditionally delimited at the family and subfamily levels. This new taxonomy includes several families not recognized in current classifications (e.g., Alsodidae, Batrachylidae, Rhinodermatidae, Odontophrynidae, Telmatobiidae), but which are strongly supported and important for avoiding non-monophyly of current families. Finally, this study provides further evidence that the supermatrix approach provides an effective strategy for inferring large-scale phylogenies using the combined results of previous studies, despite many taxa having extensive missing data.
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Peru has been transformed over the course of the past decade into a neoliberal, mineral-based, export-oriented, country. The author evaluates the neoliberal transformation of the country in three parts. First he examines the economic and political transformation of the country in the early 1990s, particularly in terms of how the Peruvian state and economy were rapidly restructured according to neoliberal principles. He then illustrates how, both through privatization and through transnationalization, the mining sector has become a key element for future development opportunities. This discussion is then contextualised by an examination of how mining activities are affecting land-tenure patterns and livelihoods in the Cajamarca region of Peru. In particular, drawing upon case-study research, the author evaluates how Minera Yanacocha's transnational gold-mining operations are transforming land-tenure institutions, land values, and the spatial distribution of land-use patterns throughout the region. In addition, he evaluates how livelihoods are being transformed in response to Minera Yanacocha's activities through an examination of changing household access to natural, human, social, and economic resources.
  • A Catenazzi
Catenazzi, A. (2015): State of the world's amphibians. -Annual Review of Environment and Resources, 40: 91-119.
Guidelines for using the IUCN Red List categories and criteria. Version 12. -Prepared by the Standards and Petitions Subcommittee
  • Iucn Standards
  • Petitions Subcommittee
IUCN Standards and Petitions Subcommittee (2016): Guidelines for using the IUCN Red List categories and criteria. Version 12. -Prepared by the Standards and Petitions Subcommittee. Downloadable from http://www.iucnredlist.org/documents/ RedListGuidelines.pdf.
A German mineralogist visits Peru
  • S O'phelan Godoy
O'Phelan Godoy, S. (2004): Chapter 32: A German mineralogist visits Peru. -pp. 403-414 in: Erickson, R., M. A. Font & B. Schwartz (coord.): Alexander von Humboldt. From the Americas to the Cosmos. -Bildner Center for Western Hemisphere Studies, The Graduate Center, The City University of New York, pp. 537.