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Where is the enigmatic Telmatobius halli Noble 1938? Rediscovery and clarification of a frog species not seen for 80 years

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Telmatobius halli was the first endemic Telmatobius species described in Chile, reported by Noble in 1938 near the locality of Ollagüe, in the high Andean zone of the Antofagasta region. To this date, there are no specimens assignable to this species other than the type series; although many expeditions have tried to search for T. halli, they have been unsuccessful, but they have found and described new species around this area. In order to clarify the origin of the enigmatic T. halli, we reviewed the itinerary of the expedition carried out by F. G. Hall in the Chilean Altiplano, to place a putative type locality. We contrast the morphology of the holotype, with that of recently collected specimens from the new putative type locality, to confirm the population’s identity; and finally, we perform phylogenetic analyses in order to clarify the systematic position of this taxon. The historical review of the expedition that collected these frogs shows that it is likely that Telmatobius halli had been collected near Collahuasi, about 50 km northwest of Ollagüe, site that we have assigned as a putative type locality for T. halli. The morphological analyses support this hypothesis, while phylogenetic results show that the specimens assigned to this species form a monophyletic group, and is a sister clade of T. chusmisensis. Thus, we propose that the type locality of T. halli be changed from “around Ollagüe” to the area of the Copaquire ravine, so its distribution would be restricted to this system and Choja-Chijlla ravine, both in the high Andean zone of the Región de Tarapacá, Chile.
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Accepted by D. Baldo: 10 Oct. 2018; published: 4 Dec. 2018
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https://doi.org/10.11646/zootaxa.4527.1.5
http://zoobank.org/urn:lsid:zoobank.org:pub:7C707F6E-E5E0-4940-9CCD-97769446A8ED
Where is the enigmatic Telmatobius halli Noble 1938? Rediscovery and
clarification of a frog species not seen for 80 years
PABLO FIBLA
1
, HUGO SALINAS
2
, GABRIEL LOBOS
2,3
, TALÍA DEL POZO
4
, ALEJANDRA FABRES
1
&
MARCO A. MÉNDEZ
1,5
1
Laboratorio de Genética y Evolución, Facultad de Ciencias, Universidad de Chile. Santiago, Chile
2
Ecodiversidad Consultores. Santiago, Chile.
3
Centro de Gestión Ambiental y Biodiversidad, Facultad de Ciencias Veterinarias y Pecuarias, Universidad de Chile. Santiago, Chile.
4
Laboratorio de propagación y conservación vegetal (Ceproveg), Facultad de Ciencias.
Universidad Mayor, Camino La Pirámide 5750. Huechuraba. Santiago, Chile.
5
Corresponding author. E-mail: mmendez@uchile.cl
Abstract
Telmatobius halli was the first endemic Telmatobius species described in Chile, reported by Noble in 1938 near the locality
of Ollagüe, in the high Andean zone of the Antofagasta region. To this date, there are no specimens assignable to this spe-
cies other than the type series; although many expeditions have tried to search for T. halli, they have been unsuccessful,
but they have found and described new species around this area. In order to clarify the origin of the enigmatic T. halli, we
reviewed the itinerary of the expedition carried out by F. G. Hall in the Chilean Altiplano, to place a putative type locality.
We contrast the morphology of the holotype, with that of recently collected specimens from the new putative type locality,
to confirm the population’s identity; and finally, we perform phylogenetic analyses in order to clarify the systematic po-
sition of this taxon. The historical review of the expedition that collected these frogs shows that it is likely that Telmatobius
halli had been collected near Collahuasi, about 50 km northwest of Ollagüe, site that we have assigned as a putative type
locality for T. halli. The morphological analyses support this hypothesis, while phylogenetic results show that the speci-
mens assigned to this species form a monophyletic group, and is a sister clade of T. chusmisensis. Thus, we propose that
the type locality of T. halli be changed from “around Ollagüe” to the area of the Copaquire ravine, so its distribution would
be restricted to this system and Choja-Chijlla ravine, both in the high Andean zone of the Región de Tarapacá, Chile.
Key words: Andean frogs, taxonomy, type locality
Introduction
The high Andean environments are the habitat of one of the most diverse groups of frogs from South-America, the
genus Telmatobius Wiegmann, 1834. The species of this genus live in aquatic ecosystems between 5° and 29° S
latitude (from Ecuador to northern Chile and Argentina); its altitudinal range is from 1000 to 5200 m (De la Riva &
Harvey 2003; Seimon et al. 2007). To date 62 species have been described in this genus, nine of which inhabit
Chile (Correa et al. 2017, Frost et al. 2017). The taxonomy and systematics of Telmatobius has been considered
complex, mainly because of the high degree of intra- and interspecific morphological variation, which precludes a
clear delimitation among the species of the genus (e.g. Trueb, 1979; Wiens, 1993; De la Riva et al., 2010). On the
other hand, there are species in this group whose taxonomic status is not clear, since they have only been reported
in their original description and have not been collected since then (see Correa et al. 2017, Fibla et al. 2017), which
has not allowed them to be integrated adequately in any of the recent systematic studies of the group (e.g. Sáez et
al. 2014; Barrionuevo 2016).
An enigma that has persisted for a long time in Chilean herpetology is the case of Telmatobius halli. This
species, described by Noble in 1938, was the first endemic species of Telmatobius described for Chile, based on six
females and six larvae collected “near Ollagüe” in northern Chile, on June 25, 1935 (Noble 1938). Ollagüe is a
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locality at about 3700 m a.s.l in the Antofagasta region. For a long time, other Telmatobius that were found in the
Loa River and Vilama creek, at about 2100 and 2500 m a.s.l respectively, were recognized in different studies as T.
halli (e.g. Cei 1962, 1986, Northland et al. 1990, Veloso et al. 1982). However, Formas et al. (1999, 2003)
recognized these Telmatobius as new species (Telmatobius dankoi and T. vilamensis), which means that T. halli has
never been recorded since the capture of the type series, even though a number of field expeditions have searched
for it, specially near its type locality (e.g. Formas et al. 2003), and two additional species have been described near
Ollague (T. fronteriensis and T. philippii).
There is an important gap in the original description of T. halli , which has probably prevented populations of
this species from being discovered in the field. The main doubt about the type locality is that Noble (1938)
indicates that the type locality of T. halli was “a warm spring near Ollagüe, Chile, 10000 ft. altitude”. The phrase
“…near Ollagüe”, is ambiguous, it does not clearly indicate the locality where the species were first found;
additionally, Ollagüe is at about 3700 m a.s.l. and not at 3000 m a.s.l, as indicated by Noble (1938) for the type
locality. The description did not include geographic coordinates, which was normal for species descriptions at this
time. However, for T. halli this seems to be the main impediment for obtaining a correct identification of the
species and new reports.
In order to clarify the origin of the enigmatic Telmatobius halli, first we reviewed the itinerary of the
expedition carried out by F. G. Hall in the Chilean Altiplano, then we assigned a putative type locality for T. h alli,
and finally we contrasted the morphology of the holotype with that of recently collected specimens from the new
putative type locality, to confirm the population’s identity. In addition, we performed phylogenetic analyses to
assess the systematic position of this taxon.
Material and methods
Historical record: Itinerary of the “International High Altitude Expedition in Chile”. With the objective of
clarify the itinerary of the expedition in which T. halli was collected, we consulted three bibliographic sources that
have not been reviewed before. The first was chapter 7 of the book “High life: A history of high altitude physiology
and medicine” (West 1998), which describes the International High Altitude Expedition of Chile in 1935 (hereafter
‘the expedition’), which was led by David Bruce Dill, in which Frank Gregory Hall participated as the zoologist
who collected specimens used to describe species. The second source was the paper “Individual variations in
ability to acclimatize to high altitude” (Keys et al. 1938), which indicates the time and date of all activities of ‘the
expedition’ in Chile. The third was the paper “Case History of a Physiologist: F. G. Hall” (Dill 1979), which
reviews the career of F. G. Hall, and provides supplementary information about the expedition.
TABLE 1. Dates and altitudes of the places visited by the International High Altitude Expedition of Chile (Modified
from Keys et al. 1938).
The primary information on the expedition in which Telmatobius halli was collected had not been previously
consulted, remaining in a “herpetological anonymity”. The expedition was made from April 8
th
to July 18
th
, 1935 by
train, visiting localities between Chuquicamata (22°17’26’’S 68°54’07’’W) and Collahuasi (20°59’13.0’’S
68°43’02.0’’W). In this trajectory, the expedition stopped in different localities to perform their studies (Keys et
al.1938; West 1998) (Table 1). It helps to understand that the objective of the expedition was to study the
physiological adaptations to altitude in persons and animals (West 1998). Frank G. Hall studied the adaptations to
altitude of fauna (wild and domestic animals) (Hall et al. 1936), thus the capture of amphibians was not the central
objective of this expedition.
Train station Period Altitude (m)
Chuquicamata April 8 to June 4 2.810
Ollagüe June 5–13; June 25 to July 18 3.660
Collahuasi (Montt) June13– 25 4.700
Aucanquilcha June 26 to July15 5.340
Punta de Cerro June 29 to July14 6.140
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REDISCOVERY OF TELMATOBIUS HALLI
FIGURE 1. Map of the study area showing Telmatobius populations distributed in the zone. Localities visited by the
expedition carried out by F.G. Hall are indicated by stars.
According to D.B. Dill (1979), Hall was in Ollagüe for 10 days, after which he travelled by train to the
Collahuasi mine, where Montt station is located at 4998 m a.s.l. The expedition remained there for 10 days,
performing studies in adaptation to altitude. Near the end of their stay in Collahuasi, on a “free” Sunday, the
scientists visited a recreation area for the miners, located about 915 m a.s.l lower than the mining area (at about
4000 m a.s.l.). Dill (1979), indicates that it was here where F.G. Hall collected the amphibians that he later sent to
the museum. The original description indicated, “…a concrete swimming pool filled with spring water was the
major attraction. Greg (as Frank Gregory Hall was called) searched the area for animal life and captured a frog that
he preserved and eventually sent to the National Museum where it proved to be a new species; appropriately it was
named Telmatobius halli”. Considering this information from Dill (1979), the individuals were most likely
collected on Sunday, June 23
rd
, 1935 and not on the 25
th
as indicated in the original description, which was most
likely the date in which the specimens were sent. According to Keys et al. (1938), on June 25
th
, 1935 the expedition
moved from Montt station in the Collahuasi mine to Ollagüe (50 km northwest) after a 12 day stay at the mine.
In the year of the expedition, mining operations were concentrated in La Ponderosa, from Collahuasi shaft
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(Millán 2006). Figure 1, shows that the nearest populations of Telmatobius are those from the stream system of the
Choja–Chijlla ravines, less than 5 km away from La Ponderosa. Another system is the wetland called Vega Paco–
Paco, which begins less than 2 km away from La Ponderosa and connects with Huinquintipa ravine; both are
affluents of the Quebrada de Copaquire. Considering the difficulty of movement at this time, it is most likely that
the members of the expedition visited the streams located in the same watershed of La Ponderosa, near the camp.
For this reason, we have focused on populations of Telmatobius that have previously been reported in the area of
Choja–Chijlla (21°5’10.33’’S, 68°51’57.40’’W) and Copaquire (20°55’45.37’’S, 68°53’6.53’’W), whose
distribution fits well with the evidence presented here.
Biological samples. To obtain biological samples for phylogenetic analyses from putative T. h alli populations,
animals were anesthetized using 0.2% tricaine methanesulfonate (MS-222; modified from Mitchell 2009), and a
small piece of interdigital membrane was cut from adults and caudal membrane from larvae. After the animals
recovered, they were released on site. We collected three specimens in Copaquire (a female, a male and a juvenile),
and seven specimens in Choja (two females, three males, a juvenile and a larvae). Samples were stored in 2 ml
microtubes with 100% ethanol. Two specimens (one male and one female) per locality (except for Chijlla) were
deposited in the Herpetological Collection of the Departamento de Biología Celular y Genética of the Universidad
de Chile (DBGUCH). Total DNA was isolated using the salting-out method (modified from Jowett 1986). Partial
sequences from the mitochondrial marker Cytochrome b (Cytb) were obtained using primers and PCR conditions
described in Sáez et al. (2014). Sequences were aligned and edited in BioEdit v.7.2.0 (Hall 1999) using the option
ClustalW, and later by visual inspection.
Morphological analyses. In order to establish the degree of morphological differentiation between putative T.
halli populations and other Telmatobius species that were previously confused with this species, we performed linear
morphometric analyses using a dataset composed of morphometric measurements from preserved specimens
deposited in the DBGUCH (including collected specimens), plus morphometric variables that were measured by
Formas et al. (2003) on the holotype (AMNH A–44753) and a paratype (AMNH A–44754) of T. halli. The collection
codes and sex of the measured specimens are detailed in Appendix 1 (n=52).
Six external variables were measured on each specimen: snout-vent length (SVL), tibia length (TL), head width
(HW), head length (HL), inter-orbital distance (IOD) and inter-nostril distance (IND). All bilateral traits were
measured on the right side. Measurements were taken with a digital Vernier (precision = 0.05 mm). The pattern of the
morphological variation was evaluated using Principal Components Analysis (PCA) on the covariance matrix of size-
corrected variables. Size correction was done by applying the Burnaby’s allometric method for morphometric
variables. All morphometric analyses were performed in the PAST 3.0 software (Hammer et al. 2001).
In addition to morphometric analyses, we made a brief morphological description of the specimens collected
from the putative T. halli populations to contrast their morphology with that of the type series (photos) and with the
morphological descriptions of the T. halli holotype made by Noble (1938) and Formas et al. (2003).
Phylogenetic analyses. To establish the phylogenetic position of the samples collected from the putative
Telmatobius halli populations, we conducted analyses using Maximum Parsimony (MP), Maximum Likelihood
(ML), and Bayesian Inference (BI). Reconstructions were based on partial sequences of the cytochrome b gene.
The dataset included Telmatobius sequences which were used in Fibla et al. (2017) and Sáez et al. (2014) (T.
marmoratus, T. dankoi, T. vilamensis, T. chumisensis, T. fronteriensis, T. philippii, T. pefauri, T. huayra, T. hintoni,
and Telmatobius sp. from the salt pans of Carcote and Ascotán, Región de Antofagasta, Chile), plus sequences
obtained from Telmatobius samples collected in this study (GenBank Accesion Numbers MH646652-MH646661).
Phylogenetic reconstructions were rooted using the T. bolivianus group (sensu De la Riva et al. 2010) as outgroup.
The MP analysis was performed in PAUP v.4.0 program (Swofford 2002), using a heuristic search with the TBR
(Tree Bisection-Reconnection) branch swapping algorithm. The ML reconstruction was performed in RAxML
GUI (Stamatakis 2014). The statistical support in the MP and ML analyses was estimated by non-parametric
bootstrap with 1000 pseudoreplicates. The BI analysis was performed in Mr. Bayes v.3.2.3 (Ronquist &
Huelsenbeck 2003). Two independent MCMC analyses of 10 million generations each were performed with four
Markov chains, sampling every 1000 generations and discarding the first 25% of trees and parameter estimations as
the burn-in period. Convergence was assessed checking ESS (Effective Sample Size) and PSRF (Potential Scale
Reduction Factor) values. Both the ML and BI analyses used the GTR (General Time Reversible) model plus the
gamma parameter (GTR+G), which was selected using JmodelTest 2.0 (Darriba et al. 2012). BI analysis was
performed in the CIPRES Science Gateway V. 3.3 (Miller et al. 2010).
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Results
Morphological analyses. The PCA generated six components; the first two explain about 83% of the
morphometric variation, 63% for PC1 and 20% for PC2. We found no correlation between SVL and PC1
(r=0.0363; p=0.7982) suggesting that the size correction was effective. The confidence interval of the eigenvalues
generated by 1000 bootstrap replicates indicated that only PC1 was significant (not shown). The factor loadings
showed that PC1 was most related to the distance between nares (IND), the tibia length and the inter-orbit distance
(IOD); the former two positives and the latter negative (Table 3). Therefore, taxa with higher PC1 values had
relatively longer hindlimbs, more separated nostrils, shorter inter-orbital distances, as well as a narrower head (e.g.
T. dankoi or T. vilamensis). The graph of PC1 vs. PC2 (Figure 2) shows that the Telmatobius specimens from
Copaquire and Choja were located in the center of the morphometric space, not differentiated from T. halli and T.
chusmisensis, while the other species were separated to the left (T. fronteriensis, T. philippii, and Telmatobius sp.
from Ascotán and Carcote) and right (T. dankoi and T. vilame n sis) in the morphological space. This pattern was
mainly due to PC1; the taxa were not differentiated by PC2.
FIGURE 2. Results of the Principal Component Analysis using morphometric variables of Telmatobius specimens. Percentage
of variance explained by each axis is showed between parentheses.
TABLE 2. Morphological measurements of the Telmatobius specimens collected from Choja and Copaquire. The
abbreviations are showed in the text (see Morphological analyses).
Phylogenetic analyses. The parsimony analysis retained 100 trees (Tree length= 153, Consistency index=
0.6863, Homoplasy index=0.3137, Retention index= 0.9474, Rescaled consistency index= 0.6502). The ML
analysis obtained a final ML Optimization Likelihood of -2395.834. In the BI analysis, the PSRF converged to 1
and the ESS average was 32516.59, suggesting that the analysis reached convergence.
Locality Sex DBGUCH code SVL HW HLt TbL IND IOD
Choja female 1710093 40.58 14.58 9.53 17.85 3.05 4.77
Choja male 1710094 47.97 16.67 10.79 22.07 3.48 5.87
Copaquire female 1710090 47.49 16.91 12.51 20.32 3.22 5.67
Copaquire male 1710091 42.38 14.93 11.44 17.14 2.88 4.81
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TABLE 3. Variables Loadings, eigenvalues and variance explained for the first two Principal Components.
The topologies found by the three different phylogenetic analyses were congruent, and show that the
Telmatobius samples collected from the Choja and Copaquire ravines form a well-supported monophyletic group
along with the sequence from Chijlla; this group is a sister clade of T. chusmisensis (Figure 7).
Description of the specimen DBGUCH-1710090, female collected at Quebrada Copaquire
(20°55’45.37’’S, 68°53’6.53’’W, 3542 m elevation), Comuna de Pozo Almonte, Región Tarapacá, Chile, on
October 26
th
, 2017 by H. Salinas, G. Lobos and P. Fibla. Description (Figure 3; Table 2): moderate sized female,
SVL 47.49 mm (Figures 3A and 3B). Large depressed head (HL/SVL=0.26, wider than long (HL/HW=0.74), with
a rounded dorsal profile of the snout. The lateral profile is rounded, with flared lips (Figure 3C). Small nostrils,
barely protuberant, with flanges, located slightly closer to the tip of the snout than to the eyes. Internarial region
flat. Loreal zone nearly horizontal. Cantus rostralis poorly defined. Tympanic ring not evident. Supratympanic fold
poorly developed. Large eyes (diameter = 4.71 mm), positioned in the front of the head with frontal orientation.
Premaxillary and maxillary teeth totally embedded into the labial mucosa. Premaxillary teeth protrude only when
lifting the upper lip (Figures 4A, 4C and 4E). Dentigerous processes of the prevomers absent. Small subcircular
choanae, widely separated in the middle. Thick circular tongue with distal border free and unnotched, anteriorly
attached 73% of its length. Forelimbs thin, without a dermic fold on wrist, relative finger length I>II<III>IV
(Figure 3E). Palmar webbing absent and tip of fingers slightly expanded in spherical pads. Lateral margins of digits
have differentiated fringes distinguishable on the inner margin of fingers II and III. Inner palmar tubercles long,
elliptic. Outer palmar tubercles prominent and triangular. One large rounded subarticular tubercle present on each
of the first two fingers; two smaller rounded subarticular tubercles on fingers III and IV. Supernumerary palmar
tubercles present at the base of each finger. Hindlimbs and toes long and thin. Relative toe length I<II<III<IV>V.
Webbing of toes concave, emarginated, almost reaching the tip of the toes, decreasing distally to shape wide fringes
along lateral margins of toes (Figure 3D). Outer border of finger V widely fringed almost to the tip. Tips of toes
expanded and spherical. Inner metatarsal tubercles elliptic, prominent and long. Outer metatarsal tubercles rounded
and smaller than inner tubercle. Subarticular tubercles present, but small; distribution of subarticular tubercles
(rounded) in toes I(1)–II(1)–III(2)–IV(3)–V(2), with some indistinct tubercles on toes III and IV. Tiny tubercles
present along ventral surface of tarsus. Postfemoral fold poorly developed. Tarsal fold reduced to a fringe along the
tarsus and decreasing distally towards fringe of internal margin of first toe. “Bagginess” absent, but ventral surface
of the thighs loose, folded. Dorsal skin smooth. A profusion of very small tubercles on cloacal region, ventral
surface of forearms, knees, external surfaces of tibias and tarsi, and posteroventral surfaces of thighs. Cloacal
aperture directed towards posterior at the dorsal level of thighs, ornamented below with folded skin and small
tubercles.
Coloration: in life dorsum reddish-brown with yellow-golden spots; venter cream yellow color with
translucent ventral surfaces of arms and legs (Figure 5A), in preservative dorsum brown with minute pale spots;
venter predominantly pale cream with translucent ventral surfaces of arms and legs. Presence of pale spots around
the vent and ventral surfaces of thighs.
Morphological variable PC1 PC2
Snout to vent lenght 0.0878 0.0241
Head width -0.2166 -0.0513
Head lenght -0.2843 0.8041
Tibia lenght 0.4962 0.0544
Internarial distance 0.5745 -0.1926
Anterior interorbital distance -0.5369 -0.5569
Eigenvalue 0.0031 0.0010
% variance 63.181 20.338
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FIGURE 3. Specimen of Telmatobius halli DBGUCH–1710090 fixed in formalin. A–dorsal view of the body; B–ventral view
of the body; C–lateral profile of the head; D–Plantar view of the foot; E–palmar view of the arm.
Va ri at io n : the most notorious differences in the specimens collected from the different localities are in body
color, both the dorsal and ventral pattern. In live individuals from Choja (Figure 5B) had an olive brown dorsal
surface with darker spots, and a cream color venter. Those from Chijlla (Figure 5C) were similar to those from
Choja. In the case of Copaquire, we found different coloration patterns, occasionally with darker spots. Males with
profusion of minute dark spines on the chest and nuptial pad on the base of the thumb. Female SVL 42–57.06 mm;
male SVL 38.02–47.97 mm.
Taxonomic assignation: in the base of the following characters we assign the studied populations to
Telmatobius halli: premaxillary and maxillary teeth completely embedded into labial mucosa, vomerine teeth
absent, toes extensively webbed, outer border of finger V widely fringed almost to the tip, hindlimbs long, presence
of supernumerary tubercles along palmar and plantar surfaces, nostrils slightly nearer to the tip of the snout than to
the eye, pale venter and dorsum brown color occasionally with darker spots (in preservative; Figure 6A and 6B).
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FIGURE 4. Premaxillary and maxillary teeth in Telmatobius halli (C and E; DBGUCH–1710090) and T. chusmisensis (D and
F; DBGUCH–0812028). In A and B the premaxillary zone of T. halli specimens (DBGUCH–1710090 and holotype
respectively) is showed.
Discussion
The evidence presented here suggests that Telmatobius halli would have been collected around Collahuasi (“La
Ponderosa”), specifically in drainages associated with the ravines of Choja-Chijlla and Copaquire. Considering that
the closest ravine to “La Ponderosa” is that of Huinquintipa-Copaquire and that the morphology of these individuals is
similar to that of the type specimens, we propose this area as the type locality of T. halli (following recommendations
76A.1.2 and 76A.2 of the International Code of Zoological Nomenclature), and that its distribution is restricted to the
ravines of Choja–Chijlla and Copaquire. Although we did not find any swimming pool on the zone, water temperature
at the collection site in Copaquire was 19°C, which is a similar a conditions to spring systems found in the Chilean
Altiplano, different from Choja’s creek which records 13.3°C (Márquez-García et al. 2009).
According to the information we provide, Ollagüe appears to be the locality from which the holotype of T. halli
was sent to the American Museum of Natural History, instead of being the site where the holotype was captured. To
understand this, we must consider that Ollagüe was one of the main border crossings at the beginning of the XX
century and that its train station was one of the most important in northern Chile. These arguments would explain why
T. halli has never been seen again around Ollagüe, as well as its morphological differences with species with which it
has previously been confused with (T. dankoi and T. vila m ensis), and with those described later from this area (T.
fronteriensis and T. philippii), which involve dentition, foot webbing and morphometric variables. Lobos et al. (2018)
recently assigned the Telmatobius population from the Ascotan salt pan (which has been confused with T. ha l l i) to T.
philippii, which fits well with the results obtained here, and confirms the absence of T. halli around Ollagüe.
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FIGURE 5. Geographic variation in body color pattern of live specimens of T. halli from Copaquire (A), Choja (B) and Chijlla
(C).Ventral surface is shown at the right for each case.
Although we were not able to differentiate Telmatobius halli from T. chusmisensis with morphometrics, the
phylogenetic analyses show that the localities assigned to T. halli form a well-supported monophyletic group which
is a sister clade to T. chu s misens i s . Morphological differences between T. halli and T. chusm i s ensis are also present,
for example, the absence of vomerine teeth in T. halli, versus T. chusmisensis, as well as the position of the nostrils
(nearer to the eyes in T. chusmisensis). We found that the maxillary and premaxillary teeth are very similar in both
species (Figure 4C-F), although previous descriptions suggest the opposite. Noble (1938) points out that: “In the
present species (i.e. T. halli) the maxillary teeth are mere rudiments a half or a third this size (in contrast to
Telmatobius peruvianus teeth). In all of the paratypes the teeth do not penetrate the mucosa of the jaws but may be
found with the help of a needle on the more anterior sections of the maxillae”. He did not mention anything about
premaxillary teeth on the text, but he states that teeth may be found on the anterior sections of the maxillae, which
could be referring to premaxillary teeth. These antecedents fit well with teeth morphology of the specimens
collected from Copaquire and Choja, but not with the description of Telmatobius halli made by Formas et al.
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(2003). Noble (1938) and this paper agree with Formas et al. (2003), in state that maxillary teeth in T. halli are
rudimentary, however, the diagnosis provided by Formas et al. (2003) specifies the lack of premaxillary teeth in
this species. The discordance between those descriptions could be a fact of the preservation state of the T. halli’s
holotype, which mouthparts are deteriorated, especially in the premaxillary region (Figure 4B). Either way, Formas
et al. (2003) did not make any reference about the presence or absence of premaxillary teeth in the description of
the T. ha l l i holotype (as Noble), so the lack of premaxillary teeth as a diagnostic character, could also have been the
result of following Noble’s description.
FIGURE 6. Dorsal and ventral view of the holotype AMNH–A44753 (A) and paratype AMNH–A44758 (B) of T. halli.
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FIGURE 7. Phylogenetic relationships between Chilean and Bolivian–Altiplanic Telmatobius species including populations
assigned to Telmatobius halli. Majority–rule consensus Bayesian tree. The lineage associated to T. halli is indicated with a
black bar at the right. Statistical support (bootstrap and posterior probability) is given below or above the corresponding node
(MP/ML/IB).
As in the re-discovery of Telmatobius pefauri (Fibla et al. 2017), and the refutation of the validity of T. laevis
as a species (Correa et al. 2017), this study shows the complexity of the taxonomy of this genus; their morphology
is notably variable both intra- and interspecifically, and it has a number of species that have only been seen once,
whose taxonomic status is unclear, at least in this part of their distribution. Added to this, is the difficulty related to
species that have probably gone extinct (e.g. Merino-Viteri et al., 2005; Barrionuevo & Mangione 2006), and the
lack of herpetological exploration along their distributional range (Sáez et al. 2014). However, comparing the
different sources of evidence that provide small clues about the existence of these “lost” taxa could be useful in this
group, as shown by the studies mentioned above.
Although there were no collections for 80 years after the description of T. h alli, this species has been listed as
Critically Endangered by the Reglamento de Clasificación de Especies de Chile (2011) and as Data Deficient by
the UICN (IUCN SSC Amphibian Specialist Group, 2015). The area occupied by these amphibians is very
restricted, and most of the water systems or wetlands mentioned here are seriously affected by mining activities—
water extraction, stream pollution and habitat destruction—placing this species in an area under high extinction
risk.
Acknowledgements
We thank FONDECYT 1140540 (MM). We are deeply grateful to Lauren Vonnahme and David Kizirian
(A.M.N.H.) for providing photos of the type material, and Vinko Malinarich (Servicio Agrícola Ganadero, Chile)
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for his help during field campaign. P. Fibla thanks CONICYT PCHA/Doctorado Nacional/2014-21140355. This
study was authorized by the Servicio Agrícola y Ganadero Resolución Exenta # 6300/2017.
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APPENDIX 1. Specimens measured for the morphometric analyses. DBGUCH: Herpetological Collection of the
Departamento de Biología Celular y Genética of the Universidad de Chile.
Telmatobius chusmisensis: CHILE: Región de Tarapacá: Chusmiza, DBGUCH 805022 (female), 812025 (male), 812028
(female); Laonzana, DBGUCH 1111012–3 (female and male).
Telmatobius dankoi: CHILE: Región de Antofagasta: Calama, DBGUCH 744 (female), 766–8 (females), 803–4 (females), 808
(female), 1304025 (female).
Telmatobius fronteriensis: CHILE: Región de Antofagasta: Puquios, DBGUCH 1309009–12 (females).
Telmatobius halli: CHILE: Región de Tarapacá: Quebrada Choja, DBGUCH 1111001–2 (juveniles), 1710093–4 (female and
male); Quebrada Copaquire, DBGUCH 703004 (male), 809058 (male), 1109005 (male), 1710090–2 (one female, two
males).
Telmatobius philippii: CHILE: Región de Antofagasta: Quebrada Amincha, DBGUCH 1110054–5 (females), 1211037
(female), 1309015–7 (one male, two females).
Telmatobius sp.: CHILE: Región de Antofagasta: Salar de Ascotán, DBGUCH 604016–7 (female), 809054–5 (females),
1211024 (female), 1211034 (female), 1304001 (female), 1501026 (female).
Telmatobius vilamensis: CHILE: Región de Antofagasta: San Pedro de Atacama: Arroyo Vilama, DBGUCH 812036–8
(females), 1108020 (female), 1309024–6 (females), 1501028 (female).
... Recently, Fibla et al. (2018) and Cuevas et al. (2020) independently claimed to have rediscovered T. halli. Bibliographic sources describing the IHAEC's activities were used in both studies, but each focused on different known populations of Telmatobius. ...
... Bibliographic sources describing the IHAEC's activities were used in both studies, but each focused on different known populations of Telmatobius. Thus, Fibla et al. (2018) assigned the southernmost populations of T. chusmisensis Formas, Cuevas & Nuñez, 2006(sensu Sáez et al. 2014 to T. halli (Copaquire, Quebrada Choja, Quebrada Chiclla), while Cuevas et al. (2020) did the same with a population from the Carcote Salt Flat (specifically, from the hot spring Aguas Calientes) (Fig. 1A, B). Previously, a Carcote population (coordinates not specified) was considered as Telmatobius sp. by Sáez et al. (2014) or T. cf. ...
... The diary also confirms that the date of the departure of the IHAEC from Collahuasi (railway station Montt) (Fig. 1B) back to Ollagüe was Tuesday, 25 June 1935. As stated in Fibla et al. (2018), this means that the original collection date was June 23 and not June 25, as specified by Noble (1938). Regarding McFarland's video material (Suppl. ...
Article
Full-text available
Telmatobius halli was the first representative of its genus to be described exclusively for Chile, yet for 85 years no new individuals could be located due to the vagueness with which its type locality was described. The type series was collected by one of the members of the International High Altitude Expedition to Chile (IHAEC) of 1935. Recently, three studies successively claimed to have located the type locality in different places. The third study proved, according to the chronicles of the IHAEC, that the actual locality is Miño, at the origin of the Loa River, where currently there are no published records of Telmatobius . In this study, additional documentary antecedents and graphic material are provided that corroborate that Miño is indeed the type locality of T. halli. Additionally, the recently rediscovered Telmatobius population from Miño and the environment it inhabits are described. The external characteristics of the frogs are consistent with the description of T. halli . Furthermore, molecular phylogenetic analyses were performed that showed that T. halli , T. dankoi , and T. vilamensis , all known only from their type localities in Chile, comprise a clade without internal resolution. A detailed comparison of the diagnoses of the three species revealed that the few phenotypic differences between these taxa were based on characteristics that vary widely within and between populations of the genus, hence their conspecificity is proposed. The implications of this synonymy for the taxonomy, biogeography, and conservation of the Telmatobius from the extreme south of its distribution in Chile are discussed.
... living in Chile and Bolivia. We propose a hypothesis about the exact loca-tion of the type locality of T. halli and discuss it with that recently proposed by Fibla et al. (2018). We comment the taxonomic status of T. halli, in relation with T. fronteriensis, T. huayra, T. philippi, and two Chilean Telmatobius sp. ...
... Consequently, they propose to move the type locality from "near Ollagüe" (Noble 1938) to a presumed area around Copaquire ravine, near Collahuasi, located 73 km north-west of Ollagüe. The frogs collected in Copaquire and Choja-Chijlla, identified as T. halli by Fibla et al. (2018), is valuable information that could extend the distribution of this species, but it needs verification. Copaquire, Choja-Chijlla, or neighboring areas, do not meet the necessary conditions as described in Noble (1938) and additional literature (Dill 1979;1980;Keys 1936aKeys , 1936b to be proposed as a type locality of T. halli. ...
... The chronicles of the activities of the IHAEC include different types of information; some subject to interpretation and others comparable with field observations. Here, taking into account the verifiable information and our field observations, we propose a working hypothesis different from that proposed by Fibla et al. (2018). Our hypothesis that Aguas Calientes is the type locality of T. halli is constructed based on different sources: the description of Noble (1938), the itinerary of the IHAEC indicating the collection site as located 3,000 feet down of Collahuasi near Ollagüe, the collection of larvae and adults, the information provided by the local inhabitants and comparison of the new animals with reference material. ...
Article
We report the rediscovery of Telmatobius halli (Hall’s water frog), which had not been found since its description (over 80 years) since its type locality was not clearly established. “Aguas Calientes” near Ollagüe is hypothesized as the original type locality where Frank Gregory Hall collected the type material in 1935. The tadpole is re-described, and new data on the external and internal morphology of adults is provided. These new morphological data are compared with Telmatobius spp. inhabiting geographically close to T. halli in Chile and Bolivia. In addition, comments on its ecology, conservation, and taxonomic status in relation with other Telmatobius spp. inhabiting nearby areas in Ascotán and Carcote salt pans are provided. No evidence of Batrachochytrium dendrobatidis and Ranavirus infection was found in T. halli and a sympatric amphibian species. Our work supports the validity of T. halli and suggests this species should be considered as Data Deficient in the IUCN Red List assessment until taxonomic issues are resolved.
... Rediscovery of species in Chile is a common phenomenon thanks to systematic collections across many ecosystems (see examples in Vera 2017, Vianna et al. 2017, Araya and Bitner 2018, Fibla et al. 2018) and intensive sampling efforts in unexplored ecosystems, filling current knowledge gaps on biodiversity. The localities where we rediscovered L. pulla belong to the Valdivian evergreen forest, which still maintains large patches of ancient forest. ...
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Lasia pulla has not recorded since their description in 1865 by Philippi. New specimen records and an updated taxonomical description is provided here. This hummingbird fly species is endemic from the Los Ríos Region, Chile. In this area, many ecosystems are still unexplored, but anthropic activities are currently fragmenting the evergreen forests. A IUCN Red List assessment is suggested.
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The natural history of most species of Telmatobius that live in the Altiplano of the Andes is unknown due to the difficulty of performing long-term studies and the logistics of working in these remote areas with extreme environmental conditions. One of these anurans inhabits the springs of the Ascotán salt flat of Chile. Here, we provide for the first time information on its distribution, habitat, microhabitat, density, diet and reproductive activity. Suitable habitat for the specie is restricted to a few springs that drain into the salt flat. Amphibian density was variable among springs and seasons and diet was composed mainly of bottom-dwelling invertebrates. We also found evidence of selection for some benthonic prey. Reproductive activity occurred mainly at night. Larvae were found during the whole year, which suggests a long larval developmental period. The salt flat is under strong anthropic pressure, although so far this anuran has managed to persist in spite of habitat perturbations. Our biological and ecological data might increase our ability to act and protect this high Andean anuran species.
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The taxonomy and systematics of Andean frogs of the genus Telmatobius have been considered complex, due mainly to the high levels of inter and intraspecific variation in morphological characters. Recently, Cuevas (2013) revalidated the species T. laevis Philippi 1902, which was originally described from two syntypes (one currently lost) collected in the late nineteenth century, based on photographs of the only preserved specimen. He also used bibliographic material showing that the original type material constituted two different taxa and that its type locality, previously not located, is Potrero Grande in the Andes Range of central Chile (33°28’S). Biogeographically, this implies a geographic distribution extension for the genus of more than 450 km on the other side of Andes, and that T. laevis inhabits within the distribution range of the phenotypically similar Alsodes montanus. Here I critically review the arguments of Cuevas (2013) and show that his main evidence for revalidating T. laevis, the location of its type locality, is based on an erroneous interpretation of the literature. Moreover, I point out several deficiencies and inconsistencies of the description and redescriptions of this taxon that were not addressed by Cuevas (2013). Reanalysing the literature and photographs of the only known specimen, and incorporating new geographic data, I suggest instead that the only known specimen of T. laevis belongs to T. marmoratus, its original designation, and came from an undetermined place within the traditional known range of the genus in Chile. However, this proposal is problematic due to the high degree of morphological variation exhibited by T. marmoratus, the uncertain taxonomic status of its Chilean populations and the unclear origin of the specimen. Therefore, I consider T. laevis as a species inquirenda until these issues are clarified or new biological material is obtained. Furthermore, I provide photographic and geographic data of frogs from Potrero Grande belonging to the genus Alsodes.
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On the basis of molecular and morphological evidence, we evaluated the taxonomic identity of two species of Andean frogs of the genus Telmatobius: Telmatobius pefauri and T. zapahuirensis, present in the western Andean slopes at the northern extreme of Chile. We also investigated the taxonomic assignment of five populations of Telmatobius recently discovered around the type localities of these two species. The results indicate that T. pefauri inhabits, not only Murmuntani its type locality, but also the montane localities of Belén, Copaquilla, Lupica, Saxamar and Socoroma. Our study also shows that T. pefauri and T. zapahuirensis are the same taxon. Therefore, Telmatobius zapahuirensis Veloso, Sallaberry, Navarro, Iturra, Valencia, Penna & Díaz, 1982 would be a subjective junior synonym of Telmatobius pefauri Veloso & Trueb, 1976.
Chapter
Molecular biology studies usually involve molecular cloning, characterization, and analysis of gene expression. These procedures rely heavily on nucleic acid hybridization, such as Southern blots, Northern blots, and dot blots. One of the most important aspects of these protocols is the use of either radioactively or nonradioactively labeled nucleic acids (deoxyribonucleic acid [DNA] or ribonucleic acid [RNA]) as probes that specifically hybridize with their complementary DNA or RNA strands. The quality of the probe plays an essential role in detecting specific DNA or RNA sequences of interest. Therefore, the preparation of a probe with high specific activity is critical in nucleic acid hybridization. ¹ The present chapter describes in detail reliable methods for the labeling of DNA and RNA. ²⁻⁵ These methods are well established and have been routinely used in our laboratory.
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A phylogenetic hypothesis for the frogs of the genus Telmatobius that includes a comprehensive sample of the morphological and geographical variation is lacking. Obtaining such a hypothesis constitutes the main focus of this contribution. A phylogenetic matrix was generated based on 97 phenotypic characters and 56 terminals. A parsimony analysis of this matrix was performed with TNT. Telmatobius is found to be monophyletic and well supported by 11 synapomorphies. Although the consensus tree shows several polytomies, four main groups have been recovered. The well-supported T. verrucosus Group includes forest and sub-paramo species from Bolivia and Peru, and is the sister group of the remaining species. The T. bolivianus Group includes forest and inter-Andean valley species from Argentina and Bolivia but it is poorly supported. Two supported high-altitude groups have been recovered, the T. macrostomus Group from the Central Andes of Peru, and the T. marmoratus Group from the Altiplano- Puna Plateau of Argentina, Bolivia, Peru and Chile and its adjacent Pacific and Northern slopes. The synapomorphies proposed for Telmatobius are discussed as well as the evolution of some of these synapomorphies and other characters within the genus.
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The nuclear DNA content (DNA/N), of 12 anuran species belonging to three tribes of lower Leptodactylids is shown. The amount of DNA/N determined by cytophotometric measurements in erythrocyte interphase nuclei ranged between 14.39 pg/N in Caudiverbera caudiverbera (2n = 26) to 5.81 pg/N in Telmatobius pefauri (2n = 26). Intergeneric DNA/N variability is higher than intrageneric variability except in Batrachyla. The DNA/N amount in these species is not clearly associated with diploid chromosome number variability except in the Telmatobiini, where high diploid number species (2n = 28, 30), have a larger genome size.
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Three species of Telmatobius occur above 2,450 m in the Ecuadorian Andes. Telmatobius niger (including T. cinereus, new synonym) is the most widespread, whereas T. vellardi and T. cirrhacelis, new species, have restricted distributions in the cordilleras of southern Ecuador. Included in the species accounts are redescriptions of the holotypes of T. niger and T. vellardi, descriptions of the larvae of these species, and a discussion of morphological variation in T. niger. On the bases of observations reported here, certain emendations to the definition of the genus Telmatobius are suggested.
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.