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Zootaxa 4441 (3): 447–466 ISSN 1175-5326 (print edition)

http://www.mapress.com/j/zt/
Copyright © 2018 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4441.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:E8756978-D474-433C-8BE8-A3A0C0294AD0

An updated phylogeny and morphological study of the


Phymaturus vociferator clade (Iguania: Liolaemidae)

JAIME TRONCOSO-PALACIOS1,7, FRANCISCO FERRI-YÁÑEZ2,3,


ALEJANDRO LASPIUR4 & CÉSAR AGUILAR5, 6
1
Programa de Fisiología y Biofísica, Facultad de Medicina, Universidad de Chile, Independencia 1027, Casilla 70005, Santiago,
Chile. E-mail: jtroncosopalacios@gmail.com
2
Departamento de Biogeografía y Cambio Global, Museo Nacional de Ciencias Naturales, CSIC & Laboratorio Internacional en
Cambio Global CSIC-PUC (LINCGlobal), Calle José Gutiérrez Abascal, 2, 28006, Madrid, Spain.
E-mail: auroch.f@gmail.com
3
Centro Iberoamericano de Biodiversidad CIBIO, Universidad de Alicante, Campus Universitario San Vicente del Raspeig s/n, 03690
Alicante.
4
Departamento de Biología, Facultad de Ciencias Exactas, Físicas y Naturales (FCEFyN), Universidad Nacional Universidad Nacio-
nal de San Juan—Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). Av. José I. de la Roza 590 (0). J5402DCS,
Rivadavia, San Juan, Argentina. E-mail: laspiursaurus@gmail.com
5
Departamento de Herpetología, Museo de Historia Natural de San Marcos (MUSM), Av. Arenales 1256, Jesús María, Lima, Perú.
E-mail: caguilarp@unmsm.edu.pe
6
Instituto de Ciencias Biológicas Antonio Raimondi, Facultad de Ciencias Biológicas, Universidad Nacional Mayor de San Marcos,
Lima, Perú.
7
Corresponding author. E-mail: jtroncosopalacios@gmail.com.

Abstract

Species delimitation in Phymaturus has been a difficult task due to the highly conserved morphological and ecological
features present in this genus. Almost all species of Phymaturus have been described without DNA data or lacking statis-
tical analyses which makes even more difficult to compare species. Although two molecular phylogenetic studies have
been recently published, here we provide the first multilocus phylogenetic reconstruction including all Chilean species,
with samples from all type localities and some previously unsampled populations. We also estimate pairwise distances
among the Chilean species of Phymaturus (P. vociferator and P. mallimaccii clades) and compare our results with the P.
payuniae clade, where previous studies have used multiple lines of evidence. Additionally, we performed univariate and
multivariate morphological analyses and skeletal comparisons (clavicle) for the species of the P. vociferator clade. As a
result of this integrative approach, we describe a new species.

Key words: lizard, Phymaturus palluma, Chile, Andes, multilocus, new species

Introduction

The liolaemid genus Phymaturus is a peculiar group of viviparous and herbivorous South American lizards (Lobo
& Quinteros 2005), characterized by a broad flattened body, spiny caudal scales in regular whorls and lateral
nuchal skin folds with fat-filled pouches (Etheridge 1995). Diversity within this genus has been underestimated,
with one species and two subspecies in 1966 (Donoso-Barros 1966), 10 species in 1995 (Etheridge 1995) and
currently comprises 43 species (Abdala & Quinteros 2014), plus more than 20 candidate species (sensu Morando et
al. 2013; Lobo et al. 2016). All species inhabit the rocky highlands of Chile and Argentina, and the tablelands and
steppe environments of Patagonia in Argentina (Cei 1986; Lobo & Quinteros 2005; Lobo et al. 2012a; Lobo &
Nenda 2015).
Etheridge (1995) subdivided Phymaturus into two groups “palluma” and “patagonicus”, an arrangement that
has been supported by phylogenetic hypotheses based on morphological (Lobo & Quinteros 2005; Lobo et al.

Accepted by L. Avila: 18 Apr. 2018; published: 28 Jun. 2018 447


2012a) and molecular data (Espinoza et al. 2004; Morando et al. 2013; Lobo et al. 2016). Species of the P. palluma
group occur in Chile and Argentina, whereas the species of the P. patagonicus group are endemic to Argentina
(Lobo et al. 2012c). Additionally, Morando et al. (2013) proposed the subdivision of each group in several clades,
in particular, splitting the P. palluma group into four clades: mallimaccii, roigorum, verdugo and vociferator. Later,
Lobo et al. (2016) proposed a similar split of the P. palluma group, but with the verdugo clade nested inside the
roigorum clade. However, morphology and ecological traits of the Phymaturus species show low variability and
are highly conserved (Cruz et al. 2009; Lobo et al. 2012a), which has made difficult the delimitation among
species, especially when most taxa of this genus have been described and diagnosed based on overall
morphological features and distributional information (González-Marín et al. 2016a,b). Moreover, species limit
hypotheses have lacked molecular data and in several cases statistical analyses.
For many years, all populations of Phymaturs were assigned to P. palluma (Molina 1782) (see Donoso-Barros
1966; Núñez & Jaksic 1992), until Pincheira-Donoso (2004) described the southernmost populations of
Phymaturus as P. vociferator, based mainly in the color pattern of the head and shape of the gular scales. Later, P.
palluma was considered an endemic species, present only in Argentina (Scolaro 2010; Lobo & Etheridge 2013),
and subsequently several populations from Chile were described, based on some skeletal, color pattern and scale
features, but lacking any molecular data or morphological statistical analyses: P. alicahuense Núñez, Veloso,
Espejo, Veloso, Cortés & Araya 2010; P. darwini Núñez, Veloso, Espejo, Veloso, Cortés & Araya 2010; P.
maulense Núñez, Veloso, Espejo, Veloso, Cortés & Araya 2010; P. paihuanense Núñez, Veloso, Espejo, Veloso,
Cortés & Araya 2010; P. damasense Troncoso-Palacios & Lobo 2012; and P. aguedae Troncoso-Palacios &
Esquerré 2014. In addition, P. bibronii (Guichenot 1848) was resurrected and P. paihuanense was proposed as a
junior synonym of P. bibronii based on color pattern, scale features and the proximity between type localities
(Troncoso-Palacios et al. 2013). To date, no molecular phylogenetic study has included all Chilean Phymaturus
species, and several aspects of its taxonomy remain uncertain. Here, we provide the first multilocus molecular
phylogeny with samples of all described species from Chile, including type localities (with the exception of P.
alicahuense, collected at 2 km from the type locality) plus some previously unsampled populations, with a special
focus in the P. vociferator clade. We also examined some skeletal and morphological features, and propose a new
species in this clade, which we describe herein.

Materials and methods

Lizards were collected with a noose and euthanized with sodium thiopental. Specimens were fixed in 99% ethanol
and preserved in 70% ethanol, with the exception of the specimens SSUC Re 410–11, which were fixed in
formalin. All specimens were deposited in Colección de Flora y Fauna, Profesor Patricio Sánchez Reyes of the
Pontificia Universidad Católica de Chile (SSUC).
Laboratory procedures and phylogenetic reconstruction. DNA was extracted using the Qiagen® DNeasy®
96 Tissue Kit (Qiagen, Valencia, CA, USA) from frozen or ethanol preserved muscle tissue samples. Two
mitochondrial and two protein-coding nuclear genes were amplified and sequenced for seven specimens of the P.
vociferator clade and six of the P. mallimaccii clade. Mitochondrial 12S rRNA (12S: 790 bp) and cytochrome b
(cyt-b: 620 bp), serine/threonine-protein kinase mos gene (C-MOS: 400 bp) and pinin desmosome associated
protein (PNN: 960 bp) were amplified via Polymerase Chain Reactions (PCR) following Ferri-Yáñez (2016,
Appendix I). PCRs were performed as follows: an initial denaturation at 94ºC for 2 min followed by 39 cycles with
denaturation at 94 ºC for 30 sec, annealing at 51ºC for 30 sec, extension at 72ºC for 2 min; and after the cycles, a
final extension of 5 min at 72ºC. PCR products were visualized in a Dark ReaderTM transilluminator (Clare
Chemical Research, Inc.) after loaded onto 1% agarose gels and stained with GelStarTM gel stain (Cambrex Bio
Science, Inc.). Products were purified using 2 μL, from a 1:4 dilution of ExoSAP-IT (Amersham Biosciences,
Uppsala, Sweden) per 2 μL of PCR product prior to cycle sequencing. A 10 μL sequencing reaction included 2 μL
of template, 2.5 μL of sequencing buffer, 0.8 μL of 10 pmol primer, 0.4 μL of BigDye® Terminator version 3.1
Sequencing Standard (Applied Biosystems, Foster City, CA-USA) and 4.2 μL of water. The sequence reaction was
35 cycles of 10 s at 96° C, 10 s at 50° C and 4 min at 60° C. Cycle sequencing products were purified by a pipetting
robot. Sequence data collection and visualization were performed on an ABI 3730xl automated sequencer (Applied
Biosystems, Foster City, CA-USA).

448 · Zootaxa 4441 (3) © 2018 Magnolia Press TRONCOSO-PALACIOS ET AL.


The accession numbers of these sequences and those obtained from GenBank are shown in Appendix II. We
include several described species, plus nine candidate species (P. sp. 2, P. sp.3, P. sp. 4, P. sp. 5, P. sp. 6, P. sp. 7, P.
sp. 8, P. sp. 9 and P. sp. 10) from Morando et al. (2013) and two candidate species (P. sp. Uspallata and P. sp. Sierra
de Fiambalá) from Lobo et al. (2016). Liolaemus monticola Müller & Hellmich 1932 was selected as the outgroup.
Sequences of fifty-one specimens were aligned using MUSCLE (Edgar 2004) and concatenated using Mesquite
v3.31 (Maddison & Maddison 2017). We used JModelTest v2.1.7 (Guindon & Gascuel 2003; Darriba et al. 2012)
to select an appropriate substitution model using the Bayesian Information Criteria (BIC): GTR + G + I for cyt-b
and 12S, HKY for CMOS and GTR + G for PNN. We performed two Bayesian inference (BI) analyses with
MrBayes v3.2.6 (Ronquist et al. 2012), one for mitochondrial data (cyt-b + 12S) and another for the mitochondrial
and nuclear markers. Each analysis consisted of two groups of four chains that run independently for 10.0 x 106
generations and at a sample frequency of 1000. The first 25% of samples were discarded as burnin and
convergence diagnostics were assessed by examining the values of average standard deviation of the Potential
Scale Reduction Factor (PSRF) for all parameters. We consider the posterior probabilities (PP) ˃ 95% as strong
support for the nodes (Wilcox et al. 2002). Additionally, we performed Maximum Likelihood (ML) and Maximum
Parsimony (MP) analyses for the mitochondrial data (cyt-b + 12S) and another for the concatenated mitochondrial
and nuclear markers, using MEGA v6.06 (Tamura et al. 2013). Bootstrap values were calculated for 1000
replicates. ML bootstrap values ˃ 70 were considered as strong support for the nodes (Hillis & Bull 1993).
Non-redundant cyt-b haplotypes of the terminal taxa were identified using DnaSP v5.10 (Librado & Rozas
2009; Appendix III), using only the sites without missing data. We used the cyt-b marker because is the locus with
more available data in GenBank, has the largest number of variable sites between the loci studied in Phymaturus
(Morando et al. 2013), and has been also used for calculate pairwise differences in several clades of the
Phymaturus sister genus Liolaemus (Morando et al. 2007; Avila et al. 2009, 2010; Esquerré et al. 2014; Medina et
al. 2014). Uncorrected average pairwise differences were calculated using MEGA v6.06 (Tamura et al. 2013) and
standard errors were estimated using the Bootstrap method with 1000 replications. We made the calculations for
the two clades that occur in Chile (P. vociferator and P. mallimaccii clades) and used the species of the P. payuniae
clade for comparisons, because it is the only group of Phymaturus with both, high support values in a molecular
phylogeny (Morando et al. 2013) and species diagnosed using multiple lines of evidence (González-Marín et al.
2016b).
Morphological comparisons. Morphological comparisons were performed only in species of the P.
vociferator clade. We examined 55 specimens of five species from Chile and Argentina: P. damasense (n=10), P.
dorsimaculatus (n=9), P. sp. 2 (n=10), P. maulense (n=10) and P. vociferator (n=14). Details on number of males,
females and juveniles for each species and the vouchers numbers are shown on Appendix IV. Data for P.
dorsimaculatus Lobo & Quinteros 2005, were provided by F. Lobo and S. Valdecantos. Description of scalation,
color pattern and external morphology characters follows Lobo et al. (2010, 2012a). Because it has been pointed
out as a key diagnostic characteristic in other Phymaturus species (Núñez et al. 2010; Lobo et al. 2012a), we
examined the clavicle shape for all species, except in P. dorsimaculatus. The clavicle was examined in seven
specimens: P. damasense (n=1), P. loboi (n=3), P. maulense (n=2) and P. vociferator (n=1). The description of color
in life was based on photographs of recently collected specimens.
Body measurements were taken with a Vernier digital calliper (to the nearest ±0.02 mm). Scales were observed
using lenses, and scalation and measurements were recorded on the right side of the specimen unless otherwise
indicated. Data are presented as mean ± standard deviation (SD). Morphological analyses were performed only in
adult specimens, and all specimens were sexed through internal examination of testes/oocytes. We performed
nonparametric Kruskal-Wallis test on the meristic variables using Dunn test for post hoc comparisons (Dunn 1964).
The chosen meristic variables follow González-Marín et al. (2016b) with few modifications: scales around the
midbody (SAMB), dorsal scales (DS), ventral scales (VS), scales around the interparietal (SAI), subocular
fragmentation (SF), fourth finger lamellae (FL), fourth toe lamellae (TL), scales around the mental (SAM), scales
between the second pair of chin shields (SBS), infralabial scales (ILS), supralabial scales (SLS), scales around the
nasal (ANAS) and the scales between the last subocular scale and the supralabials (in vertical count, SLSS).
Moreover, we included the following meristic variables used by other authors in Phymaturus descriptions: scales
between the nasal and the rostral (SNR, Cei 1980), number of rostral scales (RS, single or fragmented, Lobo &
Abdala, 2007), number of precloacal pores in males and females (PCPM, PCPF, Lobo & Nenda 2015), number of
upper ciliare scales (UCS, Lobo et al. 2012c) and number of scales between the interparietal and the rostral scale

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE Zootaxa 4441 (3) © 2018 Magnolia Press · 449
(SIR, Scolaro & Ibargüengoytía 2008). Additionally, we performed a Principal Component Analysis (PCA) to
visualize and discriminate species in the morphological space. For this purpose we used as data the residuals of a
regression between the SVL and the following variables: head length (HL), head width (HW), head height (HH),
snout-vent length (SVL), axilla-groin distance (AGD), distance from the tip of the third finger to the elbow (TFE),
distance from the tip of the fourth toe to the heel (FTH), tibia length (TBL), distance between the interparietal and
the rostral scale (IR), distance between the naris and the eye (NE) and the distance between the nares (DBN). PCA
was performed with FactoMineR, Geomorph, Rgl and Ape packages in RStudio and missing data were previously
imputed with missMDA (RStudio Team 2015).

FIGURE 1. Bayesian mitochondrial consensus tree (cyt-b and 12S), showing phylogenetic relationships of Phymaturus
palluma group and P. payuniae clade. Left values on nodes indicate the posterior probabilities (in percentage). Bootstrap values
of the congruent clades found in the ML (middle) and maximum parsimony (right) are also shown on nodes. A = P. vociferator
clade, B = P. roigorum-verdugo clade and C = P. mallimaccii clade. Outgroup (Liolaemus monticola) not shown.

Results

Phylogenetic reconstruction. The Bayesian mitochondrial tree (mtDNA; Fig. 1) show strong support for the
Phymaturus vociferator, P. roigorum-verdugo and P. payuniae clades but not for the P. mallimaccii clade. In the P.
vociferator clade (Fig. 1, clade A), we found two strongly supported groups ((P. dorsimaculatus + P. vociferator +
P. sp. 3 + P. sp. 4) and (P. sp. 2 + P. sp. 6)). Inside the P. mallimaccii clade (Fig. 1, clade C), P. bibronii forms an
unsupported clade and a polytomy with P. extrilidus, P. punae and P. williamsi. A similar topology was found in the

450 · Zootaxa 4441 (3) © 2018 Magnolia Press TRONCOSO-PALACIOS ET AL.


Bayesian concatenated tree, but P. mallimaccii clade has strong support and individuals of P. bibronii form a well-
supported clade (Fig. 2; clade C). The topology found in the ML and MP mtDNA analyses was the same in the
Bayesian analysis for all deep nodes, but some differences were found at shallow nodes. In fact, in the ML mtDNA
tree P. maulense is not monophyletic because the sample for which we have only 12S data (MZUC 35960) is basal
to all species of the P. vociferator clade, but it is monophyletic in the MP analysis. Moreover, in the MP analysis P.
sp. 5 is basal to the entire P. vociferator clade. Finally, the clade formed by P. sp. 2 + P. sp. 6 has strong support in
both analyses. Moreover, same results were found in the all genes tree. Support values are shown in the Figures 1
and 2.

FIGURE 2. Bayesian inference of all genes consensus tree (cyt-b, 12S, CMOS and PNN), showing phylogenetic relationships of
Phymaturus palluma group and P. payuniae clade. Left values on nodes indicate the posterior probabilities (in percentage).
Bootstrap values of the congruent clades found in the ML (middle) and maximum parsimony (right) are also shown on nodes. A =
P. vociferator clade, B = P. roigorum-verdugo clade and C = P. mallimaccii clade. Outgroup (Liolaemus monticola) not shown.

Uncorrected average pairwise differences are provided in the Tables 1–3. Interspecific distances in the P.
payuniae clade show a higher variation (0.9–4.6 %; Table 1) than in the P. vociferator clade (0.5–1.6 %; Table 2).
In this clade, interdistance values lower than 0.9% were found in P. sp. 2 vs P. sp. 6 (0.7 %), P. sp. 3 vs P. sp. 5 (0.8
%), P. sp. 3 vs P. dorsimaculatus (0.8 %), P. sp. 3 vs P. vociferator (0.6 %), P. sp. 5 vs P. maulense (0.5 %), P. sp. 5
vs P. vociferator (0.8 %) and P. damasense vs P. maulense (0.8 %). The P. mallamacii clade shows variation
between 0–5.1 % (Table 3), with values below 0.9 % for P. denotatus vs P. laurenti (0 %), P. antofagastensis vs P.
laurenti (0.5 %) and P. antofagastensis vs P. denotatus (0.5 %).

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TABLE 1. Percentages of average cyt-b pairwise differences for non-redundant haplotypes (below diagonal) and
standard errors estimated with bootstrap (above diagonal) between taxa of the Phymaturus payuniae clade.
Phymaturus payuniae clade
delheyi nevadoi payuniae rahuensis sitesi zapalensis
delheyi (n=4) — 0.04 0.04 0.07 0.03 0.07
nevadoi (n=1) 1.5 — 0.04 0.07 0.03 0.07
payuniae (n=1) 1.7 0.9 — 0.07 0.04 0.07
rahuensis (n=1) 4.2 4.1 4.6 — 0.06 0.05
sitesi (n=6) 1.2 0.9 1.2 3.7 — 0.06
zapalensis (n=3) 3.9 3.8 4.4 1.9 3.8 —

TABLE 2. Percentages of average cyt-b pairwise differences for non-redundant haplotypes (below diagonal) and
standard errors estimated with bootstrap (above diagonal) between taxa of the Phymaturus vociferator clade. Values
lower than the minimum distance found within P. payuniae clade (0.9) are shown in bold.
Phymaturus vociferator clade
sp. 2 sp. 3 sp. 4 sp. 5 sp. 6 damasense dorsimaculatus maulense vociferator
sp. 2 (n=2) — 0.04 0.04 0.03 0.03 0.04 0.04 0.04 0.04
sp. 3 (n=1) 1.1 — 0.04 0.03 0.04 0.04 0.04 0.04 0.03
sp. 4 (n=2) 1.6 1.2 — 0.04 0.05 0.04 0.05 0.04 0.04
sp. 5 (n=1) 0.9 0.8 1.1 — 0.04 0.03 0.04 0.03 0.04
sp. 6 (n=4) 0.7 1.4 1.8 1.2 — 0.05 0.05 0.04 0.04
damasense (n=4) 1.3 1.5 1.4 1.0 1.6 — 0.04 0.03 0.04
dorsimaculatus (n=1) 1.1 0.8 1.5 1.0 1.5 1.6 — 0.04 0.04
maulense (n=1) 1.1 1.0 1.2 0.5 1.4 0.8 1.1 — 0.04
vociferator (n=1) 1.1 0.6 1.2 0.8 1.4 1.5 0.8 1.0 —

TABLE 3. Percentages of average cyt-b pairwise differences for non-redundant haplotypes (below diagonal) and
standard errors estimated with bootstrap (above diagonal) between taxa of the Phymaturus mallimaccii clade. Values
lower than the minimum distance found within P. payuniae clade (0.9) are shown in bold.
Phymaturus mallimaccii clade
sp. Fiambalá aguedae alicahuense antofagastensis bibronii darwini
sp. Fiambalá (n=1) — 0.09 0.08 0.04 0.05 0.08
aguedae (n=3) 5.3 — 0.07 0.08 0.08 0.05
alicahuense (n=1) 4.8 3.4 — 0.08 0.08 0.06
antofagastensis (n=1) 1.5 4.7 4.2 — 0.05 0.08
bibronii (n=2) 2.3 5.0 4.0 2.1 — 0.08
darwini (n=1) 4.8 1.7 2.9 4.2 4.2 —
denotatus (n=1) 1.3 5.2 4.4 0.5 2.1 4.7
extrilidus (n=1) 1.9 4.6 3.9 1.6 1.1 3.6
laurenti (n=1) 1.3 5.2 4.4 0.5 2.1 4.7
malliiccii (n=2) 2.0 4.8 4.1 1.7 2.5 4.8
punae (n=3) 2.3 4.5 4.0 2.0 1.5 4.0
williamsi (n=1) 1.8 4.8 4.0 1.8 1.5 4.7

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continued.
Phymaturus mallimaccii clade
denotatus extrilidus laurenti mallicciii punae williamsi
sp. Fiambalá (n=1) 0.04 0.05 0.04 0.06 0.05 0.05
aguedae (n=3) 0.09 0.08 0.09 0.08 0.08 0.08
alicahuense (n=1) 0.08 0.08 0.08 0.08 0.08 0.08
antofagastensis (n=1) 0.03 0.05 0.03 0.05 0.05 0.05
bibronii (n=2) 0.05 0.03 0.05 0.06 0.04 0.04
darwini (n=1) 0.08 0.07 0.08 0.08 0.08 0.08
denotatus (n=1) — 0.05 0 0.05 0.05 0.05
extrilidus (n=1) 1.8 — 0.05 0.06 0.04 0.04
laurenti (n=1) 0 1.8 — 0.05 0.05 0.05
malliiccii (n=2) 1.5 2.3 1.5 — 0.06 0.05
punae (n=3) 2.2 1.1 2.2 2.6 — 0.04
williamsi (n=1) 1.6 1.1 1.6 1.9 1.3 —

Morphological analyses. Morphological univariate analysis revealed significant differences in five meristic
variables among species of the P. vociferator clade: SAMB, DS, VS, SAI and SF. The only species that differs
statistically from all other species in at least one variable is Phymaturus sp. 2 (Table 4).

TABLE 4. Results for the Kruskal-Wallis test (H) with degrees of freedom (df) among selected species of the P.
vociferator clade and the post hoc comparison (Dunn’s test). Only significant results are provided. See Material and
Methods for abbreviations.
Kruskal-Wallis test Dunn’s test
H P df
SAMB 11.445 ˂ 0.05 4 P. dorsimaculatus ˃ P. damasense
DS 14.353 ˂ 0.01 4 P. sp. 2 ˃ P. dorsimaculatus
VS 13.795 ˂ 0.01 4 P. sp. 2 ˃ P. damasense; P. sp. 2 ˃ P. maulense
SAI 15.898 ˂ 0.01 4 P. sp. 2 ˃ P. dorsimaculatus; P. sp. 2 ˃ P. vociferator
SF 13.605 ˂ 0.01 4 P. damasense ˃ P. sp. 2

The PCA results (Table 5) show that the first three principal components (PCs) explain 73.52% of the
cumulative variance. PC1 accounts for 43.69% of the variance and explained mainly by HL, HW and NE variables
(Table 5). PC2 accounts for 17.19% of the variance and explained mainly by TBL, IR and AGD variables (Table
5). PC3 accounts for 12.63% of the variance and explained mainly by AGD, FTH and TBL variables (Table 5). In
the PC1vs PC2 plot, P. dorsimaculatus shows almost no overlap with others species (ellipses with 95% confidence
interval, Fig. 3A); Phymaturus sp. 2 overlaps with P. damasense and P. maulense and includes all individuals of P.
vociferator. In the PC1vs PC3 plot (Fig. 3B), all species overlap but only marginally between Phymaturus sp. 2 and
P. vociferator.
We found three different clavicle shapes (Fig. 4): 1) with a marked middle process and an additional process
near the interclavicle area (in P. damasense and P. maulense); 2) with an inconspicuous middle process (in
Phymaturus sp. 2); and 3) with a very developed middle process (in P. vociferator). No data for P. dorsimaculatus
was available.

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TABLE 5. Results of the PCA for the morphological measures among selected species of the P. vociferator clade. See
material and methods for abbreviations.
PCA
Eigenvalues PC1 PC2 PC3
Eigenvalue per Component 4.37 1.72 1.26
% of Variance 43.69 17.19 12.63
Cumulative % of Variance 43.69 60.89 73.52
Eigenvectors PC1 PC2 PC3
HL 0.874 -0.074 -0.197
HH 0.766 -0.099 -0.273
HW 0.814 -0.324 -0.206
AGD -0.426 0.472 0.583
TFE 0.720 0.315 0.197
FTH 0.538 -0.307 0.576
TBL 0.398 -0.697 0.437
IR 0.554 0.652 -0.246
NE 0.782 0.292 0.154
DBN 0.538 0.432 0.349

Taxonomy remarks in the P. vociferator clade. Although samples of Phymaturus damasense and P. maulense
form two reciprocally monophyletic groups (Figs. 1 and 2), they have low cyt-b pairwise difference (Table 2). In
fact, distances are lower than the minimum cyt-b pairwise distance found among species of the P. payuniae clade
(used here for comparisons; see Materials and Methods). The description of P. damasense stated three meristic
characters (PCPF, SF and SLSS) and four qualitative characters (scales on the auditory meatus of P. damasense,
enlarged scales on the chest of the females of P. damasense, different tail color of the males and different dorsal
pattern of the females) as diagnostic between both species (Troncoso-Palacios & Lobo 2012). However, there are
no statistical differences in our univariate analysis, which include these and all the meristic variables proposed as
diagnostic traits in Phymaturus species descriptions (Table 4). We confirm one marked qualitative difference
(enlarged scales on the chest of P. damasense females) but both differences referred to the color pattern are minor.
In regards to the projected scales on the auditory meatus, these are not always found in P. damasense from
Planchón, then it should not be taken as a diagnostic trait. Moreover, both species shared the same clavicle shape.
Although they do not overlap or show almost no overlap in our morphometric PC analysis (Fig. 3). We interprets
these minimum genetic distances and phenotypic differences as within species variation or between population
differences of the same species, but it also could be interpreted as evidence of incipient species formation (see
Wake 1997). In order to keep a conservative interpretation of our results, we suggest consider to P. damasense as a
junior synonym of P. maulense.
Our results also reveal that Phymaturus sp. 2 and Phymaturus sp. 6 do not form two reciprocally monophyletic
clades (Figs. 1 and 2) and the cyt-b pairwise distance between these two candidate species is extremely low (Table
2). Additionally, in our DnaSP analysis we found that the same cyt-b haplotype is shared by these four individuals
LJAMM-CNP 3440, SSUC Re 693 (Phymaturus sp. 2), LJAMM-CNP 5259 and 5260 (Phymaturus sp. 6). Based
on these evidences, we propose that both Phymaturus sp. 2 and Phymaturus sp. 6 might be considered conspecific.
Although PCA results show overlap between Phymaturus sp. 2 and other species of P. vociferator clade,
Phymaturus sp. 2 has strong statistical differences in meristic traits with all described species of the P. vociferator
clade (Table 4) and has a conspicuous clavicle shape not shared with most species of this clade. In addition,
Phymaturus sp. 2 forms a distinct and strongly supported monophyletic group in our phylogenetic trees. Moreover,
pairwise distance between this taxon and most species of this clade is greater than 0.9 (the minimum pairwise
distance found for species of the P. payuniae clade; Tables 1 and 2). All these evidences show that the Phymaturus
sp. 2 (including Phymaturus sp. 6) requires recognition as full species and is described below.

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FIGURE 3. Principal Component Analysis (PCA) results. On the left, ellipses representing the 95% confidence interval around
the centroid for each species. On the right, contribution of each variable to PCs .

FIGURE 4. Comparison of clavicle process. Upper: clavicle with two process found in Phymaturus damasense and P.
maulense (here illustrated in P. damasense). Middle: Clavicle of P. sp. 2. Bottom: Clavicle of P. vociferator. A: Marked middle
process. B: Additional process near the interclavicle area. C: Inconspicuous middle process. Black bar = 4 mm.

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE Zootaxa 4441 (3) © 2018 Magnolia Press · 455
Phymaturus loboi sp. nov.
(Fig. 5)

1966 Phymaturus palluma palluma (in part), Donoso-Barros, Rept. Chile: 349.
2005 Phymaturus flagellifer (in part), Pincheira-Donoso & Núñez, Pub. Ocas. Mus. Nac. Hist. Nat. 59: 24.
2005 Phymaturus cf. palluma (in part), Lobo & Quinteros, Pap. Avul. Zool. 45: 155.
2007 Phymaturus cf. palluma (in part), Lobo & Abdala, Cuad. Herpetol. 21: 113.
2012 Phymaturus maulense (in part), Troncoso-Palacios & Lobo, Cuad. Herpetol. 26: 73.
2013 Phymaturus maulense, Esquerré, Núñez & Scolaro, Zootaxa 3619: 439.
2013 Phymaturus maulense (in part), Troncoso-Palacios, Lobo, Etheridge, Acosta & Laspiur, Zootaxa 3652: 561.
2013 Phymaturus cf. palluma, Lobo & Etheridge, South Am. J. Herpetol. 8: 100.
2013 Phymaturus maulense, Garin, Troncoso-Palacios, Ferri-Yáñez & Lobos. Cuad. Herpetol. 27: 67.
2013 Phymaturus sp. 2 Morando, Avila, Pérez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701.
2013 Phymaturus sp. 6 Morando, Avila, Pérez, Hawkins, Sites, Mol. Phyl. Evol. 66: 701.
2016 Phymaturus maulense Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 649.
2016 Phymaturus sp. 6 Lobo, Barrasso, Hibbard & Basso. Zool. J. Linn. Soc. 176: 653.

Holotype. SSUC Re 691. Adult male. Collected at National Route CH115, 9 km NW of Maule Lagoon (35˚57' S,
70˚34' W), 1900 m asl, Maule Region, Chile by J. Troncoso-Palacios, A. Laspiur and R. Gómez. January 25, 2013
(Fig. 5a,b).
Paratypes. SSUC Re 692, male and 693, 694 females (Fig. 5). Same data as the holotype. SSUC Re 410–11
(male and juvenile male). Collected road to Maule Lagoon, Maule Region, Chile, by F. Ferri-Yáñez. November 11,
2011
Other specimens referred. MNHNCL 2353, 2460–61, males. Baños del Campanario, 1500 m asl, San
Clemente, Maule Region, Chile. J.C. Torres-Mura coll. February 2-3, 1992.
Etymology. The species is named after our colleague, the herpetologist Fernando Lobo, a prominent specialist
in the Phymaturus genus, who has participated in the description of 16 species in this genus (Lobo and Quinteros
2005, Lobo and Abdala 2007, Lobo and Nenda 2015, Lobo et al. 2010, 2012b, 2012c, 2013, Troncoso-Palacios and
Lobo 2012) and the identification of the true type locality and redescription of P. palluma (Lobo and Etheridge
2013). The suggested English common name for this species is "Lobo’s Rocky Lizard" and in Spanish, "Matuasto
de Lobo”.
Diagnosis. Phymaturus loboi belongs to the palluma group (Etheridge 1995) because it has short, non-
imbricate superciliary scales; fragmented subocular scale; and strongly spiny tail scales. Within the palluma group,
P. loboi belongs to the P. vociferator clade (Figs. 1 and 2) and differs from the species within this clade in the
following characteristics.
Phymaturus loboi differs from P. dorsimaculatus, in having higher number of dorsal scales (161.0±7.5 vs.
135.0±4.0, Tables 4 and 6), and a higher number of scales around the interparietal (9.1±1.2 vs. 6.2±2.0, Tables 4
and 6). Moreover, the males of P. loboi have green spotted shoulders and the area behind the shoulders, with an
inconspicuous scapular spot (Fig 5), whereas the males of P. dorsimaculatus have black color on the shoulder and
in the area behind the shoulder with a bold scapular spot.
Differs from P. vociferator, in having higher number of scales around the interparietal scale (9.1±1.2 vs
7.2±1.2, Tables 4 and 6). Males of P. loboi have a green spotted shoulder with an inconspicuous scapular spot (Fig.
5) whereas the males of P. vociferator have black color on the shoulders and in the area behind the shoulders, and a
marked scapular spot. Additionally, P. loboi does not vocalize.
Differs from P. maulense, in having higher number of ventral scales (184.0±7.6 vs 169.8.0±7.7, Tables 4 and
6). Males of P. loboi have an inconspicuous scapular spot and females have no precloacal pores, whereas the males
of P. maulense have a marked scapular spot and all females have precloacal pores.
Differs from Phymaturus damasense, in having higher number of ventral scales (184.0±7.6 vs 166.8±6.5,
Tables 4 and 6) and fewer subocular scales (2.0±0.9 vs 3.6±0.5, Tables 4 and 6). Additionally, females of P. loboi
never have precloacal pores, while 40% of the females of P. damasense have it.
Description of the holotype. SSUC Re 691. Adult male. Measurements in mm: snout–vent length (SVL):
102.5; trunk width: 41.7; head length: 20.7; head width: 21.1; head height: 10.9; interorbital distance: 8.4; distance
between the nares: 3.9; distance between the naris and the eye: 4.9; distance between rostral and interparietal scale:
12.7; axilla-groin distance: 51.7; arm length (from tip of toe of the 3rd toe to elbow): 29.7; right foot length: 24.1

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(from tip of toe of the 4th toe to heel); and tail length: 105.8 (with autotomized tip). Ten scales in contact with
interparietal scale; 16 scales between interparietal and rostral (excluding both). Parietal eye inconspicuous and
whitish. Supraorbital semicircles incomplete in both sides. Superciliary scales, 12/12 (left/right) juxtaposed and
flat. Subocular scales, two. Posterior subocular vertically separated from supralabials by 2 row of lorilabials.
Preocular scale smaller than canthal. Preocular and supralabials separated by 4 loreal scales. Canthal separated
from nasal by 2 scales. Nasal in contact with 8 scales and separated from rostral by 3 scales. Rostral scale
undivided. Nine rhomboidal or pentagonal, slightly keeled temporal scales. Three scales slightly projected on the
anterior border of auditory meatus. Eleven supralabial scales and 10 infralabial scales. Mental pentagonal, in
contact with 6 scales. Fifty six gular scales, counted transversally between mouth´s commissures. Secondary chin
shields separated by 2 scales. Well-developed antehumeral and neck fold. Conspicuous gular and antegular folds.
No enlarged scales on posterior margin of gular fold or on chest. Dorsal scales rounded, smooth and juxtaposed.
Dorsal scales (counted between the occiput and the level of the anterior border of the hindlimbs), 157. Middorsal
scales larger than those on flanks. Ventral scales larger than dorsals, quadrangular, smooth, and juxtaposed. Ventral
scales (counted from mental scale to the anterior margin of cloacal opening), 175. Scales around of midbody, 216.
Precloacal pores, 11. Supernumerary pores, 7 (1 above and 6 below precloacal pores). No enlarged pigal scales.
Scales of the dorsal surface of the femoral area smooth, rounded to pentagonal and juxtaposed or sub-imbricated.
Scales of the ventral surface of the femoral area smooth, pentagonal or hexagonal, and juxtaposed. Scales of the
dorsal surface of forearm rounded, smooth, and juxtaposed or sub-imbricated. Scales of the ventral surface of
forearm rounded, smooth, and juxtaposed. Subdigital lamellae of fingers: I: 9, II: 14, III: 19, IV: 21 and V: 13.
Subdigital lamellae of the toes : I: 9, II: 15, III: 20, VI: 24 and V: 15. Tarsal scales smooth or with 2/3 keels. Caudal
scales arranged in spiny whorls, slightly keeled with projected mucrons.

TABLE 6. Selected meristic variables in species of the Phymaturus vociferator group: scales around the midbody
(SAMB), dorsal scales (DS), ventral scales (VS), scales around the interparietal (SAI), subocular fragmentation (SF),
scales between the second pair of chin shields (SBS), scales around the mental (SAM), infralabial scales (ILS),
supralabial scales (SLS), scales around the nasal (ANAS) and the scales between the last subocular scale and the
supralabials (in vertical count, SLSS), fourth finger lamellae (FL), fourth toe lamellae (TL) and precloacal pores (PP). M
= males; F = females. Min–Max (Mean±SD).
P. damasense P. dorsimaculatus P. loboi sp. nov. P. maulense P. vociferator
N=8 N=5 N=9 N=10 N=10
(M = 3, F = 5) (M = 2, F = 3) (M = 6, F = 3) (M = 6, F = 4) (M = 5, F = 5)
SAMB 184–222 213–244 198–222 206–230 206–242
(205.4±13.6) (232.8±12.3) (211.6±7.5) (217.0±6.9) (217.8±11.4)
DS 139–151 131–139 150–168 143–160 135–157
(146.9±5.2) (135.0±4.0) (161.0±7.5) (150.0±6.8) (143.5±8.2)
VS 158–176 160–198 171–193 158–181 166–187
(166.8±6.5) (177.5±15.7) (184.0±7.6) (169.8.0±7.7) (173.8±9.7)
SAI 7–11 (8.1±1.2) 4–9 (6.2±2.0) 8–11 (9.1±1.2) 8–10 (8.6±0.7) 5–8 (7.2±1.2)
SF 3–4 (3.6±0.5) 2–4 (3.2±0.8) 1–3 (2.0±0.9) 2–3 (2.5±0.5) 2–4 (2.8±1.0)
SBS 2 0–2 (0.6±0.9) 0–3 (1.8±1.0) 0–3 (1.1±1.1) 0–4 (1.6±1.1)
SAM 6 3–6 (5.4±1.3) 5–6 (5.9±0.3) 5–7 (6.2±0.8) 5–8 (6.0±1.4)
SLS 9–11 (10.3±0.7) 9–12 (10.8±1.5) 10–11 (10.3±0.5) 9–11 (9.9±0.8) 10–11 (10.3±0.5)
ILS 8–10 (8.5±0.8) 7–11 (8.6±1.5) 6–10 (8.4±1.4) 8–9 (8.4±0.5) 8–9 (8.8±0.5)
ANAS 7–10 (8.8±1.2) 7–9 (7.8±0.8) 8–10 (8.8±0.7) 7–10 (8.3±1.1) 7–9 (8.0±0.8)
SLSS 1–2 (1.4±0.5) 1–2 (1.4±0.6) 1–3 (1.5±0.8) 1–2 (1.3±0.5) 1–2 (1.3±0.5)
FL 18–20 (19.1±0.6) 17–22 (20.0±2.2) 20–22 (20.8±0.8) 19–22 (20.4±1.0) 20–22 (21.0±1.0)
TL 22–24 (23.2±1.0) 22–27 (24.5±2.1) 24–26 (24.6±0.9) 23–25 (24.0±0.9) 24–25 (24.5±0.7)
PP M = 9–10; F=0–6 M = 10 F=0 M = 8–11 F=0 M = 8–10 F=4–10 M = 6–10 F=0

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE Zootaxa 4441 (3) © 2018 Magnolia Press · 457
FIGURE 5. Phymaturus loboi sp. nov. (A) and (B) Holotype SSUC Re 691 (SVL = 102.6 mm). (C) and (D) Paratype female
SSUC Re 694 (SVL = 93.1 mm). (E) Male not collected (picture by J. E. Domínguez). (F) Female not collected (picture by C.
Garin).

Color in life of the holotype. Head with black background color, spotted with green in nasal and temporal
areas, without spots on the cheeks. Dorsum green, with faint black reticulation, without ocelli. Neck green with
black reticulation to the level of the lower meatus (transversally), but is black with very few green spots below the
level of the lower meatus (transversally). Scapular spot inconspicuous. Limbs light green, with faint black
reticulation. Flanks yellowish green. Base of the tail green, the rest yellowish green (lighter than the body). Throat
and chest black, extending ventrally to the insertion of the forelimbs. Abdomen with a “∩” yellowish green area
(limited by the chest and flanks) with scattered black spots. Center of the belly light green with no appreciable
yellow shading on the cloaca. Ventral surface of the limbs light green with few black spots. Ventral surface of the
tail light green. Precloacal pores intense orange.
Variation. Measurements in mm. Adult males (n = 6): SVL, 102.1–102.8 (102.5 ± 0.3); head length, 20.3–21.8
(20.9 ± 0.8); head width, 20.3–21.5 (21.0 ± 0.4); head height, 10.9–13.2 (11.9 ± 0.9); axilla–groin distance, 50.9–
51.7 (51.4 ± 0.4); arm length, 29.7–33.0 (31.2 ± 1.7); foot length, 24.1–27.5 (25.7 ± 1.8); tibia length, 14.8–17.8
(16.3 ± 1.5); distance between the interparietal and the rostral, 11.9–12.7 (12.3 ± 0.4); distance between the nares
and the eye, 4.9–5.7 (5.2 ± 0.4); and distance between the nares, 3.2–3.9 (3.6 ± 0.3). Adult females (n = 3): SVL,
93.1–98.6 (96.6 ± 3.0); head length, 19.1–19.8 (19.4 ± 0.3); head width, 17.8–18.5 (18.2 ± 0.4); head height, 9.1–
11.0 (10.1 ± 0.1); axilla–groin distance, 45.4–50.7 (48.5 ± 2.8); arm length, 28.5–30.1 (29.0 ± 0.9); foot length,

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23.2–24.3 (23.8 ± 0.6); tibia length, 14.3–15.1 (14.7 ± 0.4); distance between the interparietal and the rostral, 12.3–
12.4 (12.4 ± 0.1); distance between the nares and the eye, 5.0–5.2 (5.0 ± 0.1); and distance between the nares, 3.5–
3.8 (3.6 ± 0.2). Variation in the meristic variables (scale count) is provided in the Table 6. Parietal eye
inconspicuous, whitish and opaque. No enlarged scales on posterior margin of gular fold or on chest. In males, 8–
11 (9.7 ± 1.5) precloacal pores, 0–6 supernumerary pores, with no enlarged postcloacal scales. Tarsal scales smooth
or bearing 2–3 keels. Males exhibit the same color pattern as holotype, with variation in color shades. One male has
the limbs of the same shade of the dorsum and less yellowish on the flanks than other males. Head brown in
females, slightly darker than the body. Occipital area with several dark brown spot, surrounding the interparietal
(one female), on the supraocular area (“O” shaped spot), the superciliaries, the snout and temporal area (except
postocular area). Scattered light brown scales on the head, between the occipital area and the snout, on the
postocular area, and between the subocular and the rictal. Dorsum light brown, with a thin dark brown reticulation
forming three short stripes on the neck, but not forming dark crossbars on the dorsum. A “ɔ” shaped dark brown
spot on the shoulder. Dorsolateral area is darker than the occipital area. The limbs, lateral neck and flanks are light
brown with scattered dark brown spots. Tail light brown with diffuse dark brown rings. Throat variegated, and one
specimen has dark brown color only to the level of the meatus. The belly and ventral surfaces of the limbs and the
tail are whitish with some scattered dark brown spot (especially on the chest). Flanks with oxide color. The juvenile
male (SVL = 80.4 mm) has a color pattern similar to the females, but the dorsal reticulation is thicker than in
females, the dark brown spot on the shoulder is “O” shaped, the head is noticeable more darker than in females, the
throat is black and the flanks are light brown.

FIGURE 6. Distribution map for Phymaturus loboi sp. nov. and all species and candidate species of the vociferator clade. Red
star: P. loboi (formerly known as P. sp. 2), Maule Lagoon, type locality. Light orange star: P. loboi (formerly known as P. sp. 6),
Near Domuyo Volcano. Green square: P. damasense (Termas del Flaco and Planchón). Black square: P. maulense (Lircay).
White diamond: P. vociferator (Los Barros). Green diamond: P. dorsimaculatus (Copahue). Black diamond: P. sp. 3 (Near
Loncopue). Blue diamond: P. sp. 4 (Near El Cholar). Black pentagon: P. sp. 5 (Near Epulafquen).

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE Zootaxa 4441 (3) © 2018 Magnolia Press · 459
Distribution and natural history. In Chile Phymaturus loboi is known from the type locality and adjacent
areas in the Region del Maule. This is an abundant lizard that inhabits rocky environments from the road to Maule
Lagoon (35˚57' S, 70˚34' W, 1900 m asl, Fig. 6) and the Termas del Campanario (35˚55' S, 70˚35' W, 1800 m), at 3
km NW from type locality. In the type locality Phymaturus loboi occurs syntopically with Liolaemus buergeri, L.
carlosgarini and L. flavipiceus and the vegetation is dominated by Hypochaeris tenuifolia and Festuca
acanthophylla. In Argentina, P. loboi is distributed near Domuyo Volcano (36˚39' S, 70˚32' W, 2200 m) at 68 km
SE from type locality.
At the date of capture (January), one female had one embryo and several small oocytes. Another female had
several oocytes. The intestines of one individual (SSUC Re 410) contained plant remnants and a large number of
nematodes of an unidentified species, probably of the genus Spauligodon or Parapharyngodon (C. Garin pers.
comm.).

Discussion

Delimiting species in Phymaturus is a difficult task since the morphological features are highly conserved among
species (Cruz et al. 2009) and our phylogenetic results show, in general, short branch lengths (low divergence)
among species of the same clade. In this sense, an assessment of the taxonomic status of the described and
candidate species using multiple lines of evidence is a more useful approach (Aguilar et al. 2013, 2017; González-
Marín et al. 2016a,b). PCA results show a marked morphological overlap among the species of the P. vociferator
clade, similar to the results reported for the species of the P. payuniae clade (González-Marín et al. 2016b).
However, compared with González-Marín et al. (2016b), the variation in the axes is explained by different main
variables: PC1 only coincide in HL as main explanatory variable, PC2 has no similarities and PC3 agrees in AGD
and FTH. Interestingly, we found significant differences for some meristic variables among the species of the P.
vociferator clade, two of these (SAMB, VS) coincide with González-Marín et al. (2016b) results for the P.
payuniae clade while others were found significant only for P. vociferator (DS) or P. payuniae clade (SLS, FL,
TL). All of these suggest the existence of different sets of meristic and morphometric traits useful for diagnosis
among main clades of Phymaturus.
Regarding the cyt-b pairwise distances, we found a variation between 0.9 and 4.6 % in the P. payuniae clade in
which all species have been delimited with different lines of evidence (González-Marín et al., 2016b). The
variation found between the species of the P. vociferator clade is lower (maximum of 1.7%) than those in the P.
payuniae clade. Although in Liolaemus, the sister genus of Phymaturus, a 3% in the pairwise differentiation for the
mDNA (cyt-b + 12S) between sister species has been proposed (Breitman et al. 2012), other authors recorded
much smaller differences for cyt-b pairwise differences between widely accepted species: L. riojanus vs L.
multimaculatus (1.23%; Avila et al. 2009), L. goestchi vs L. casamiquelai (0.62%; Avila et al. 2010) and L.
leopardinus vs L. ubaghsi (1.2%; Esquerré et al. 2014). The minimum cyt-b pairwise differences for the P.
vociferator clade among known species is 0.8% (e.g., P. vociferator vs P. dorsimaculatus) very similar to the 0.9%
found for the P. payuniae clade.
The species described here, Phymaturus loboi, is distributed in central Chile, in the surroundings of Maule
Lagoon (Fig. 6). It was previously assigned to P. maulense (with type locality in Lircay, Fig. 6) by Troncoso-
Palacios and Lobo (2012) due to geographical proximity between both populations (approximately 60 km), lack of
DNA data at the time of publication and because they studied very few specimens of P. loboi (one adult and one
juvenile males). This misidentification was later followed by other authors (Esquerré et al. 2013; Garin et al. 2013;
Troncoso-Palacios et al. 2013; Lobo et al. 2016).
In regards to the Chilean species of the P. mallimacci clade, Lobo et al. (2016) provided the cyt-b sequence of
P. aguedae SSUC Re 588 (KT203826). However, JTP sent to F. Lobo tissue samples only from P. aguedae SSUC
Re 595 and 596, therefore sequences of SSUC Re 588 by Lobo et al. (2016) corresponds to another P. aguedae
specimen. Nevertheless, our phylogenetic and cyt-b pairwise results support all Chilean species of the P.
mallimaccii clade, since all are monophyletic and show high genetic distances. However, more accurate
morphological analyses for this clade are still needed and it will be published elsewhere.

460 · Zootaxa 4441 (3) © 2018 Magnolia Press TRONCOSO-PALACIOS ET AL.


Acknowledgments

We thank M. Penna (Programa de Fisiología y Biofísica, Facultad de Medicina, Universidad de Chile) for his
support, and P. Zavala (Pontificia Universidad de Católica de Chile) for allowing us to deposit material in the
SSUC Re Collection. We are grateful to the following colleagues (and museums) for allowing us to study
specimens: H. Núñez (Museo Nacional de Historia Natural de Chile), F. Troncoso (Museo de Historia Natural de
Concepción), J. Artigas (Museo de Zoología, Universidad de Concepción). Special thanks to David Vieites (Museo
Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas) for his inestimable contribution
providing the laboratory equipment and in generating the DNA sequences. F. Lobo and S. Valdecantos for provide
data of P. dorsimaculatus. R. Gómez Alés, H. A. Díaz and F. Urra for their help in the field collections. D. Reyes
Arellano identified the plant species and C. Garin speculated as to the possible genus of the nematodes. C. Garin, F.
Cabello and J. Domínguez Bertelotto shared some pictures with us. C. Garin, H. Núñez, D. Esquerré and R.
MacCulloch for their comments to the early version of the manuscript. R. Buff for his help with the map. We thank
the Servicio Agrícola y Ganadero (SAG, Chile) for authorizing the capture of specimens by permission number
6967/2012. Financial support was partially provided to (AL) by Consejo de Investigaciones Científicas y
Tecnológicas and partially through the LINCGlobal CSIC-PUC and the NICHE project (CGL2011-26852) of the
Spanish Ministry of Economy and Competitiveness as well as the CSIC-JAE PhD studentship (2010 00735)..

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https://doi.org/10.1016/S1055-7903(02)00244-0

APPENDIX I. Primers used in this study were taken from Ferri-Yáñez (2016).
Locus Primer name Primer sequence
12S LIO_DRV_12S_F 5'-CATCCGCAYMCCAGTGAGAATGCC-3'
LIO_DRV_12S_R 5'-GAGGAGGGTGACGGGCGGTGTGTG-3'
CYTB LIO_DRV_CYTB_F 5'-CCCATCAAACATCTCTGCMTGATG-3'
LIO_DRV_CYTB_R 5'-TGGTGTRAAGTTTTCTGGGTCTCC-3'
C-MOS LIO_DRV_CMOS_F 5'-AGTAAGAATAGTTTGGCATCACGG-3'
LIO_DRV_CMOS_R 5'-TCAGGGGCACGATGAGTGTATGTC-3'
PNN LIO_DRV_PNN_F 5'-GTAGAAATGGAAGAGGCAGGGGAG-3'
LIO_DRV_PNN_R 5'-TCTGATATAGACCTCTTTCTGTCCGAC-3'

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE Zootaxa 4441 (3) © 2018 Magnolia Press · 463
$33(1',;,,*HQH%DQNDFFHVVLRQQXPEHUVRIWKHVSHFLPHQVXVHGLQSUHYLRXVVWXGLHVDQGJHQHUDWHGLQWKLVVWXG\

6SHFLHV 9RXFKHUV 6 &\WE &026 311 /RFDOLW\
Phymaturus VS /-$00±&13 -; -; ² -; /DJXQDGHO0DXOH ƒ¶6ƒ¶: 
P. VS.  668&5H 0+ 0+ 0+ 0+ /DJXQDGHO0DXOHÛ
6Û
:
P. VS.  /-$00±&13 -; -; ² -; /RQFRSXH ƒ¶6ƒ¶: 
P. VS.  /-$00±&13 -; -; -; -; (O&KRODU ƒ¶6ƒ¶: 
P. VS.  /-$00±&13 -; -; ² -; 0LQDV ƒ¶6ƒ¶: 
P. VS.  /-$00±&13 -; -; -; -; $JXDV&DOLHQWHV ƒ¶6ƒ¶: 
P. VS.  /-$00±&13 -; -; -; -; $XFD0DKXLGD ƒ¶6ƒ¶: 
P. VS.  /-$00±&13 -; -; -; -; 'RPX\R ƒ¶6ƒ¶ 
P. VS.  /-$00±&13 -; -; -; -; %XWDFR ƒ¶6ƒ¶ 
P. VS.  /-$00±&13 -; -; -; -; 3DVR3HKXHQFKH ƒ¶6ƒ¶: 
P. VS. 8VSDOODWD /-$00±&13 -; -; -; -; 9DOOHFLWRV ƒ¶6ƒ¶: 

464 · Zootaxa 4441 (3) © 2018 Magnolia Press


P. VS. )LDPEDOD 0&1 .7 .7 .7 ² 3XHVWROD/DJXQLWD XQNQRZQFRRUGLQDWHV 
P. aguedae 668&5H 0+ 0+ 0+ 0+ &HUUR&DUSD Ħ6Ħ: 
P. aguedae 668&5H 0+ 0+ 0+ 0+ &HUUR&DUSD Ħ6Ħ: 
P. aguedae 668&5H 0+ 0+ ² 0+ &HUUR&DUSD ƒ¶6ƒ¶: 
P. aguedae 668&5H 0+ 0+ 0+ ² &HUUR3URYLQFLDƒƍ6ƒƍ: 
P. aguedae 668&5H 0+ 0+ 0+ 0+ &HUUR3URYLQFLD ƒƍ6ƒƍ: 
P. alicahuense -7 0+ 0+ 0+ 0+ /DJXQD&KHSLFDO ƒƍ6ƒƍ: 
P. alicahuense -7 ² 0+ 0+ 0+ /DJXQD&KHSLFDO ƒƍ6ƒƍ: 
P. antofagastensis 0$&1 .7 .7 .7 ² 3DVR6DQ)UDQFLVFR XQNQRZQFRRUGLQDWHV 
P. bibronii 668&5H .7 .7 .7 ² /RV3LXTXHQHV ƒƍ6ƒƍ:
P. bibronii 668&5H .7 .7 .7 ² 3DVR/RV3DWRV ƒƍ6ƒƍ: 
P. damasense 668&5H 0+ 0+ 0+ 0+ 7HUPDVGHO)ODFR ƒƍ6ƒƍ: 
P. damasense 668&5H 0+ 0+ ² 0+ 7HUPDVGHO)ODFR ƒƍ6ƒƍ: 
P. damasense -7 0+ 0+ 0+ 0+ (O3ODQFKyQ ƒƍ6ƒƍ: 
P. damasense -7 0+ 0+ 0+ 0+ (O3ODQFKyQ ƒƍ6ƒƍ: 
P. darwini -7 0+ 0+ ² ² 5LHFLOORV ƒƍ6ƒƍ: 
P. darwini -7 0+ 0+ ² ² ,QFDODJRRQ ƒƍ6ƒƍ: 
     ……continued on the next page

TRONCOSO-PALACIOS ET AL.
     
$33(1',;,, &RQWLQXHG      
6SHFLHV 9RXFKHUV 6 &\WE &026 311 /RFDOLW\
P. delheyi /-$00&13 -; -; -; -; %XWDFR ƒƍ6ƒ¶: 
P. denotatus 0&1 .7 .7 .7 ² /DJXQD%ODQFD ƒƍ6ƒƍ: 
P. dorsimaculatus /-$00±&13 -) -) -; -; &RSDKXH ƒ¶6ƒ¶: 
P. extrilidus /-$00±&13 -; -; -; ² 'RQ&DUPHOR ƒ¶6ƒ: 
P. laurenti /-$00±&13 -; -; -; -; 4XHEUDGDGH5DQGROIR ƒ¶6ƒ¶: 
P. mallimaccii /-$00±&13 -; -; -; -; /D0HMLFDQD ƒ¶6ƒ¶: 
P. maulense 0=8& 0+ 0+ 0+ 0+ (QODGULOODGR ž¶6ž¶: 
P. maulense 0=8& 0+ ² 0+ 0+ (QODGULOODGR ž¶6ž¶: 
P. nevadoi /-$00&13 -; -; -; -; /D9HQWDQD Ħ6Ħ: 
P. palluma 0&1 .7 .7 .7 ² /DJXQD'LDPDQWH ƒƍƎ6ƒƍƎ: 
P. payuniae /-$00&13 -; -; -; -; 3D\XQ Ħ6Ħ: 
P. punae /-$00±&13 -; -; -; ² /D/DJXQLWD ƒ¶6ƒ¶: 
P. querque /-$00±&13 -; -; -; -; /DJXQD%ODQFD ƒ¶6ƒ¶: 

UPDATED PHYLOGENY OF THE PHYMATURUS VOCIFERATOR CLADE


P. rahuensis /-$00&13 -; -; -; -; 5DKXH Ħ6Ħ: 
P. roigorum /-$00±&13 -; -; -; ² 3D\~Q ƒ¶6ƒ¶: 
P. sitesi /-$00&13 -; -; -; -; $XFD0DKXLGD B6B: 
P. tromen /-$00±&13 -; -; -; -; 7URPHQ ƒ¶6ƒ¶: 
P. verdugo /-$00±&13 -; -; -; -; 7HUPDVGHO$]XIUH B6B: 
P. vociferator /-$00±&13 -; -; -; -; /DJXQDGHO/DMD ƒ¶6ƒ¶: 
P. williamsi 0&1 .7 .7 ² ² &DOLQJDVWD ƒƍ6ƒƍ: 
P. zapalensis /-$00&13 -; -; -; -; /DJXQD%ODQFD Ħ6Ħ: 
Liolaemus carlosgarini 668&5H 0+ 0+ ² ² /DJXQDGHO0DXOH ƒ¶6ƒ¶: 
L. monticola 668&5H 0+ 0+ ² ² )DUHOORQHV ƒ¶6ƒ¶: 


Zootaxa 4441 (3) © 2018 Magnolia Press ·


465
APPENDIX III. GeneBank accession numbers of the cyt-b sequences, used for calculation of average pairwise
differences. Only non-redundant haplotypes for each terminal taxon were used for the analysis.

PHYMATURUS MALLIMACII CLADE: P. sp. Fiambala: KT203831. P. aguedae: MH198588, MH198589, MH198590,
MH198591 and MH198592. P. alicahuense: MH198593 and MH198594. P. antofagastensis: KT203828. P. bibronii:
KT203829 and KT203832. P. darwini: MH198599 and MH198600. P. denotatus: KT203833. P. extrilidus: JX969019. P.
laurenti: JX969009. P. mallimaccii: JX948841 and JX969011. P. punae: JX948851, JX948852 and JX969013. P.
williamsi: KT203827. PHYMATURUS PAYUNIAE CLADE: P. delheyi: JX949085, JX949087, JX949088, JX949089,
JX949090 and JX969047. P. nevadoi: JX948955, JX949100, JX949101, JX949102, JX949103, JX949104, JX949105 and
JX969035. P. payuniae: JX948958, JX948959 and JX969037. P. rahuensis: JX948970 and JX969053. P. sitesi: JX948820,
JX948825, JX948826, JX948827, JX948828, JX948829, JX948830, JX948831, JX948834 and JX969049. P. zapalensis:
JX949124, JX949125, JX949126, JX949127, JX949128, JX949129, JX949130 and JX969042. PHYMATURUS
VOCIFERATOR CLADE: P. sp. 2: JX948887, JX969020 and MH198602. P. sp. 3: JX948971 and JX969021. P. sp. 4:
JX948968, JX948969 and JX969022. P. sp. 5: JX969023. P. sp. 6: JX948967, JX948988, JX948989, JX948990,
JX948991, JX948992, JX948993, JX948994, JX948997 and JX969024. P. damasense: MH198595, MH198596,
MH198597 and MH198598. P. dorsimaculatus: JX948838, JX948839, JX948880 and JX969010. P. maulense:
MH198601. P. vociferator: JX948886 and JX969016.

APPENDIX IV. Specimens examined. MCN, Museo de Ciencias Naturales, Universidad Nacional de Salta, Argentina.
MNHNCL, Museo Nacional de Historia Natural de Chile. MRC, Museo de Historia Natural de Concepción. MZUC,
Museo de Zoología, Universidad de Concepción. SSUC, Collection of the Pontificia Universidad Católica de Chile.

Phymaturus damasense (males = 3, females = 5, juveniles = 2). MNHNCL 4782. North shore of the Damas River,
approximately 1.5 Km to east from Termas del Flaco, Libertador Bernardo O´Higgins Region, Chile. Troncoso-Palacios J.
& F. Ferri colls. January 14, 2011. MNHNCL 4745–48. Termas del Flaco, Las Damas River, Libertador Bernardo
O´Higgins Region, Chile. Núñez H. & D. Esquerré colls. February 01, 2011. SSUC Re 413, 415, 417. North shore of the
Damas River, approximately 1.5 Km to east from Termas del Flaco, Libertador Bernardo O´Higgins Region, Chile.
Troncoso-Palacios J. & F. Ferri colls. January 14, 2011. SSUC Re 414, 416. South shore of the Damas River, Termas del
Flaco, Libertador Bernardo O´Higgins Region, Chile. Troncoso-Palacios J. & F. Ferri colls. January 14, 2011.
Phymaturus dorsimaculatus (males = 2, females = 3, juveniles = 6). MCN 1571–72, 1575–76, 1578. Copahue, Neuquén
Province, Argentina. Abdala C., Avila L., F. Lobo & M. Morando colls. January 13, 1999. MCN 1568–70. Termas de
Copahue, Neuquén Province, Argentina. January 13, 1999. MCN 1566–67. Copahue, Neuquén Province, Argentina. Pérez
D. coll. January, 1999. MCN 1577. Copahue, Neuquén Province, Argentina.
Phymaturus loboi (males = 6, females = 3, juveniles = 1). MNHNCL 2353, 2460, 2461. Baños del Campanario (1500 m),
Talca, San Clemente. J.C. Torres-Mura coll. February 2-3, 1992. SSUC Re 691–694, 737. National Route CH115, 9 km
NW of Maule Lagoon, Maule Region, Chile. Troncoso-Palacios J., A. Laspiur & R. Gómez colls. January 25, 2013. SSUC
Re 410–11. Road to Maule Lagoon, Maule Region, Chile. F. Ferri-Yáñez coll. November 28, 2011.
Phymaturus maulense (males = 6, females = 4, skeletal = 2). MNHNCL 3939–42, 3945–46, 4038–39, 3943–44 (skeletal).
Vilches Alto, El Enladrillado, Reserva Nacional Altos de Lircay. Veloso A., P. Espejo & S. Araya colls. April 16-17, 2005.
MZUC 35959-60. Enladrillado, Lircay, Provincia de Talca, Maule Region. Troncoso-Palacios J. coll. November 28, 2011.
Phymaturus vociferator (males = 5, females = 5, juveniles = 4). MNHNCL 3852. El Refugio, Laguna del Laja, Antuco, Bíobio
Region, Chile. Martín S. coll. December 01, 2000. MNHNCL 3854. El Refugio, Laguna del Laja, Antuco, Bíobio Region,
Chile. December 18, 2001. MNHNCL 3853, 3855. Antuco, Bíobio Region, Chile. MRC 20–23. Antuco, Chile.
Valdovinos-Zarges C. coll. (undated). MZUC 19510, 19513, 19517, 19518 (two specimens), 19540. Laja National Park.
Habit E. & P. Victoriano colls. March, 1987. SSUC Re 683. West shore of the Laja Lagoon. Troncoso-Palacios J., F. Urra
& H. A. Díaz colls. January 07, 2014.

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