Research Article |
Corresponding author: Raquel Alvarado-Larios ( raquel9.al@gmail.com ) Academic editor: Clara Stefen
© 2024 Raquel Alvarado-Larios, Pablo Teta, Pablo Cuello, J. Pablo Jayat, Andrea P. Tarquino-Carbonell, Guillermo D’Elía, Paula Cornejo, Agustina A. Ojeda.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Alvarado-Larios R, Teta P, Cuello P, Jayat JP, Tarquino-Carbonell AP, D’Elía G, Cornejo P, Ojeda AA (2024) A new living species of the genus Ctenomys (Rodentia: Ctenomyidae) from central-western Argentina. Vertebrate Zoology 74: 193-207. https://doi.org/10.3897/vz.74.e115242
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Abstract
The genus Ctenomys Blainville, 1826 includes 68 living species of small to medium-sized (100–1200 g) caviomorph rodents of subterranean habits. During the last decade, this genus has been the subject of numerous taxonomic studies, including the description of new species and the proposal of novel synonyms. Based on phylogenetic analysis of mitochondrial DNA sequences and qualitative and quantitative morphological traits, here we review the species boundaries of the tuco-tucos of the species group of C. mendocinus and describe a new species. The new species is morphologically distinct from other phylogenetically and geographically close species of Ctenomys (e.g., C. fochi, C. mendocinus), showing several differences in their craniodental traits (e.g., proportionally longer nasals and less globose tympanic bullae). The new species occurs in montane grasslands and shrublands of northwestern Mendoza (ca. 2710 m a.s.l.) and in lowlands (ca. 1000 m a.s.l.) of the Monte Desert ecoregion in an area highly impacted by accelerated processes associated with the wine industry.
Caviomorpha, Ctenomys fochi, Mendoza, Monte, Octodontoidea, Southern Andean steppe, tuco-tucos
The genus Ctenomys Blainville, 1826, with 68 living species, is one of the most speciose genera of the order Rodentia. It is restricted to southern South America, ranging from southern Peru to southernmost Argentina and Chile (
One of the groups that receive more attention (e.g.,
Here, based on sequences of the cytochrome b gene of the mitochondrial genome, and qualitative and quantitative morphological characters of skins and skulls, we show that the individual tuco-tucos from the “Arenales-Tupungato” lineage correspond in fact to an undescribed species for which no name is available. As such, we name and describe this taxon as a new species of the C. mendocinus group that is sister to C. fochi. In addition, we discuss and consider distinctiveness and geographic distribution of two other nominal forms, including C. fochi and C. johannis, of the C. mendocinus species group.
As an ontological (i.e., primary) species concept, we follow the General Lineage Species Concept; under this approach, species are considered as metapopulational lineages recognized by their emergent properties (e.g., monophyly, morphological diagnosability;
Phylogenetic analyses were based on 801 bp DNA sequences of the mitochondrial cytochrome b gene (cyt b). We expanded the large matrix of DNA sequences of Ctenomys used by
Sequences were aligned using CLUSTAL X (
We studied a total of 37 museum specimens of Ctenomys (see Appendix
Map of west-central Argentina depicting the studied samples. Green and red dots correspond to Ctenomys fochi and Ctenomys uco sp. nov., respectively. Numbers indicate the following localities: 1 Finca Zorro Pozo (Santiago del Estero); 2 Chumbicha (Catamarca); 3 El Chepical (San Juan); 4 Estancia La Providencia (Córdoba); 5 Finca Caicayén II (Mendoza); 6 Cajón de Arenales (Mendoza).
Anatomical terminology used to describe external and cranial traits follow
Morphological analyses were guided by the results of the phylogenetic analyses of molecular data (see below), geography, and current taxonomy (for a similar approach see
The phylogenetic analyses recovered a strongly supported monophyletic group of Ctenomys (BS = 100; PP = 1; Fig.
Phylogenetic analysis of mitochondrial DNA sequences. Maximum Likelihood tree (ln = –8860.2974) obtained in the analysis of 186 cytochrome b gene sequences of Ctenomys. Numbers indicate bootstrap (right of the diagonal) and posterior probability (left of the diagonal, found in a Bayesian analysis) values of adjacent nodes; a missing value indicates that the given node has less than 50% of posterior probability. Terminal labels indicate species name and GenBank accession numbers.
Percentage of average genetic variation (p distances), based on cytochrome b sequence data, observed within and between pairs of species and lineages of the Ctenomys mendocinus species group.
intraspecific | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | ||
1 | C. rionegrensis | 0.63 | |||||||||||
2 | C. flamarioni | 0.22 | 2.39 | ||||||||||
3 | C. coludo | 0.73 | 2.16 | 2.74 | |||||||||
4 | C. verzi | 0.00 | 2.22 | 2.60 | 1.92 | ||||||||
5 | C. emilianus | 0.47 | 2.56 | 2.62 | 2.27 | 2.40 | |||||||
6 | C. mendocinus | 1.17 | 2.44 | 2.33 | 2.36 | 2.33 | 2.71 | ||||||
7 | C. bergi | 0.13 | 1.45 | 2.26 | 1.67 | 1.64 | 1.98 | 2.08 | |||||
8 | Neuquén-La Pampa lineage | 0.69 | 2.70 | 2.74 | 2.95 | 2.45 | 2.97 | 2.11 | 2.68 | ||||
9 | C. fochi | 1.04 | 2.46 | 2.78 | 2.37 | 2.13 | 2.36 | 2.33 | 2.12 | 2.61 | |||
10 | C. uco sp. nov. | 1.14 | 2.52 | 2.90 | 2.62 | 2.26 | 2.68 | 2.89 | 2.03 | 3.06 | 1.85 | ||
11 | C. eileenae | 0.63 | 3.51 | 3.45 | 3.13 | 3.00 | 3.08 | 3.33 | 2.93 | 3.76 | 3.31 | 3.46 | |
12 | Quijadas lineage | 0.28 | 2.09 | 2.59 | 1.86 | 1.25 | 1.92 | 0.02 | 1.14 | 2.43 | 1.96 | 1.94 | 3.04 |
Qualitative differences between samples, including the external coloration and morphology of the skull, in particular those involving the “Arenales-Tupungato” lineage, are detailed below (see the Discussion). Descriptive statistics of cranial measurements (i.e., mean, standard deviation and range are summarized in Table
Summary statistics (N), mean, S.D., range (min.–max.) of cranial measurements (in mm) of adult samples (N = 35) of three species of the genus Ctenomys. See Materials and Methods for abbreviations.
C. fochi | C. mendocinus | C. uco sp. nov. | |||||||||||||||
N | Mean | S.D. | min. | max. | N | mean | S.D. | Min. | max. | N | Mean | S.D. | min. | max. | |||
TLS | 10 | 40.64 | 1.97 | 37.99 | 43.72 | 19 | 39.68 | 1.55 | 37.1 | 41.7 | 6 | 40.74 | 1.65 | 38.16 | 42.35 | ||
CIL | 10 | 39.17 | 1.82 | 36.68 | 41.99 | 19 | 38.37 | 1.35 | 36.11 | 40.51 | 6 | 37.26 | 0.97 | 35.75 | 38.15 | ||
NL | 10 | 14.32 | 0.94 | 12.61 | 15.61 | 19 | 13.21 | 0.91 | 11.74 | 14.92 | 6 | 14.81 | 0.58 | 14.25 | 15.58 | ||
NW | 10 | 6.33 | 0.37 | 5.98 | 6.99 | 19 | 5.83 | 0.37 | 5.15 | 6.48 | 6 | 5.85 | 0.23 | 5.55 | 6.20 | ||
RW | 10 | 9.25 | 0.63 | 8.67 | 10.63 | 19 | 8.77 | 0.46 | 7.91 | 9.49 | 6 | 9.17 | 0.34 | 8.80 | 9.60 | ||
FL | 10 | 11.41 | 0.53 | 10.47 | 12.15 | 19 | 12.34 | 0.89 | 10.76 | 13.95 | 6 | 11.64 | 0.50 | 11.05 | 12.35 | ||
IOC | 10 | 8.51 | 0.59 | 7.75 | 9.79 | 19 | 7.77 | 0.50 | 6.89 | 8.79 | 6 | 8.29 | 0.50 | 7.44 | 8.80 | ||
ZB | 10 | 24.2 | 1.01 | 22.86 | 26.04 | 19 | 23.37 | 1.12 | 21.11 | 25.49 | 6 | 23.36 | 0.95 | 22.26 | 24.9 | ||
BB | 10 | 16.73 | 0.74 | 15.82 | 18.35 | 19 | 16.45 | 0.70 | 15.16 | 17.72 | 6 | 16.68 | 0.75 | 15.65 | 17.55 | ||
BIB | 10 | 26.21 | 0.74 | 24.93 | 27.37 | 19 | 24.93 | 0.97 | 23.3 | 26.24 | 6 | 25.75 | 1.56 | 23.86 | 27.4 | ||
IFH | 10 | 7.63 | 0.53 | 6.87 | 8.51 | 19 | 7.25 | 0.43 | 6.41 | 7.96 | 6 | 7.63 | 0.23 | 7.30 | 8.02 | ||
DL | 10 | 10.23 | 0.61 | 9.51 | 11.39 | 19 | 10.03 | 0.70 | 8.94 | 11.16 | 6 | 10.38 | 0.32 | 10.03 | 10.9 | ||
PL | 10 | 16.94 | 1.13 | 15.47 | 18.97 | 19 | 16.63 | 0.82 | 15.2 | 17.81 | 6 | 17.73 | 1.03 | 16.11 | 18.9 | ||
UIW | 10 | 5.99 | 0.46 | 5.52 | 6.99 | 19 | 5.58 | 0.29 | 5.18 | 6.11 | 6 | 5.78 | 0.30 | 5.25 | 6.06 | ||
TRL | 10 | 8.50 | 0.39 | 7.96 | 9.11 | 19 | 8.43 | 0.46 | 7.61 | 9.19 | 6 | 9.00 | 0.38 | 8.40 | 9.40 |
The first 2 axes of the PCA accounted for 78.6% of the total variance (PC1 = 69.1%; PC2 = 9.5%; Fig.
Multivariate statistics for three species of Ctenomys from west-central Argentina. Individual scores of adult specimens of Ctenomys from west-central Argentina (N = 35) for (A) principal components 1 and 2; B canonical variates 1 and 2, extracted from three taxonomical group discriminant function analysis.
Results of principal components analysis performed on adult specimens of three species of Ctenomys (N = 35). See Materials and Methods for explanation of abbreviations.
PC1 | PC2 | CV1 | CV2 | |
TLS | 0.5590 | 0.0397 | 0.1514 | 0.0974 |
CIL | 0.4317 | –0.5297 | 0.0933 | –0.2549 |
NL | 0.2578 | 0.5718 | 0.1819 | 0.1766 |
NW | 0.0740 | –0.0400 | 0.0689 | –0.0384 |
RW | 0.1405 | 0.0988 | 0.0740 | 0.0271 |
FL | 0.0620 | –0.5135 | –0.1404 | –0.0393 |
IOC | 0.1296 | 0.1890 | 0.1121 | 0.0271 |
ZB | 0.3287 | –0.1622 | 0.1141 | –0.0724 |
BB | 0.1559 | –0.0073 | 0.0429 | 0.0141 |
BIB | 0.3305 | 0.1675 | 0.1931 | 0.0311 |
IFH | 0.1025 | 0.0585 | 0.0592 | 0.0327 |
DL | 0.1827 | 0.0448 | 0.0340 | 0.0425 |
PL | 0.2916 | 0.0891 | 0.0606 | 0.1591 |
UIW | 0.0907 | 0.0491 | 0.0605 | –0.0015 |
TRL | 0.1003 | 0.0991 | 0.0196 | 0.0906 |
The CVA correctly classified all individuals according to their taxonomic reference. Overall, this analysis was consistent with the PCA results (Fig.
Phylogenetic analysis of molecular markers (reciprocal monophyly) and genetic divergences of the cyt b gene suggest that the Arenales-Tupungato is an independent lineage of Ctenomys. The evaluation of integumental and craniodental morphological traits indicate this lineage is morphologically distinct and diagnosable, all of which provide strong evidence for its recognition as a new species of the genus, which is sister to C. fochi. As no name is available for it, here we describe it as a new species.
Chresonymy.
Ctenomys mendocinus
–
[Ctenomys] mendocinus 3 –
[Ctenomys] Arenales –
Ctenomys mendocinus
Tupungato –
[Ctenomys] “Arenales-Tupungato” lineage –
An adult male (CMI 7712), including skin, skeleton, and tissues, collected on 28 April 2023 by Raquel Alvarado-Larios, Pablo Cuello, Agustina Ojeda, Andrea P. Tarquino-Carbonell, and Paula Cornejo (original field number RCAL 91) (Figs
Argentina: Mendoza Province, Tunuyán Department, Cajón de Arenales, alongside Ruta Provincial N° 34, 2710 m a.s.l. (33.6213°S, 69.5112°W; Fig.
TOTL, 226; TAIL, 70; HFL, 31; EAR, 6.5; TLS, 39.66; CIL, 37.71; NL, 15.58; NW, 5.66; FL, 11.31; RW, 8.91; ZB, 22.26; IOB, 7.44; BB, 15.65; BIB, 23.90; IFH, 8.02; DL, 10.21; PL, 16.89; UIW, 6.06; TRL, 8.71. Weight, 110 g.
An adult male (CMI 7739), including skin, skeleton, and tissues, collected 28 April 2023 by Raquel Alvarado-Larios, Pablo Cuello, Agustina Ojeda, Andrea P. Tarquino-Carbonell, and Paula Cornejo (original field number RCAL 92). An 801 base-pair sequence of the cyt b gene has been deposited in GenBank with accession number (OR669673).
TOTL, 215; TAIL, 64; HFL, 31; EAR, 8; TLS, 38.16; CIL, 36.36; NL, 14.28; NW, 5.81; FL, 11.05; RW, 8.94; ZB, 22.77; IOB, 7.98; BB, 16.35; BIB, 23.86; IFH, 7.65; DL, 10.03; PL, 16.11; UIW, 6.00; TRL, 8.40. Weight, 109 g.
Three adult males (CMI 7735 [original field number RCAL 87], CMI 7737 [RCAL 89], CMI 7738 [RCAL 90]) and one adult female (CMI 7736 [RCAL 88]), including skin, skeleton, and tissues, collected 26 April 2023 by Raquel Alvarado-Larios, Pablo Cuello, Agustina Ojeda, Andrea P. Tarquino-Carbonell, and Paula Cornejo at the Finca Caicayén II (33.3925°S, 69.2007°W; Tupungato Department, Mendoza Province).
We named this species in reference to the region where the type locality lays, the Valle de Uco (Uco Valley), which includes the Tupungato, Tunuyán, and San Carlos Departments in Mendoza Province, central western Argentina; this valley is well known for its fine wines.
A small-sized tuco-tuco of the C. mendocinus species group (TOTL, 215–263 mm; TAIL, 64–79 mm; HFL, 31–36 mm: EAR, 6–8 mm; Weight, 109–138 g); dorsum Drab to Dusky Drab, which turns lighter on flanks; venter Light Brownish Drab to Light Drab, with line separating from dorsum scarcely defined; a patch above nose and forehead, blackish. Skull moderately robust, with rostrum and nasals proportionally long and narrow; premaxillary-frontal suture evident anterior to the naso-frontal suture; interorbital region with posteriorly divergent outer margins. Zygomatic arches thin, slightly divergent backwards in dorsal view; dorsal profile of cranium with a marked ventral inflection in the parietal-occipital region; incisive foramina short and narrow, recessed in a common fossa of posteriorly divergent outer borders; interpremaxillary foramen small or nearly absent; sphenopalatine vacuities nearly tear-shaped; auditory bullae moderately inflated and nearly oval, with salient auditory tubes.
Pelage dense, fine, and silky, about 12–15 mm long over back and rump (Fig.
Skull moderately robust with rostrum proportionally long and narrow, interorbital region with posteriorly divergent outer margins, and zygomatic arches thin and slightly divergent posteriad in dorsal view (Fig.
Mandible robust, markedly hystricognathous, with coronoid process long and pointed; condyloid process robust, bearing a poorly developed articulation flange. Postcondyloid process with small ventrolateral apophysis. Chin process small and moderately visible in lateral view; bottom of alveolar sheath of p4 weakly protruding (Fig.
Upper incisors large, robust, and orthodont to slightly ophistodont; frontal enamel surface Orange. Maxillary tooth rows slightly divergent posteriad. M3 reduced, with posterolingual face flat with an anterior protruding lobe (Fig.
One female from “Tupungato” has a 2n = 47, FN = 68 (
Ctenomys uco sp. nov. can be differentiated from C. coludo by its less inflated and proportionally smaller tympanic bullae, and its proportionally larger rostrum and narrower nasals. In addition, Ctenomys uco sp. nov. has large and wide incisive foramina, while in C. coludo these openings are short and wide.
Ctenomys uco sp. nov. differs from C. fochi (Figs
Cranial anatomy in four species of Ctenomys from west-central Argentina. Dorsal (A–D) and ventral (E–H) views of the skulls in four species of Ctenomys from west-central Argentina: A, E C. fochi (CFA-MA 12120); B, F C. uco sp. nov. (CMI 7712); C, G C. mendocinus (CMI 2497); D, H C. verzi. Scale = 5 mm.
Ctenomys uco sp. nov. can be differentiated from both C. mendocinus (Fig.
Ctenomys uco sp. nov. differs from C. verzi by its less robust appearance, narrower nasals and rostrum, and by the presence of an interparietal bone. In addition, Ctenomys uco sp. nov. has orthodont to slightly ophistodont upper incisors, while those of C. verzi are more proodont (Fig.
Pairwise genetic distances with other species of the Ctenomys mendocinus species group range from 1.85 to 3.76 % (Table
Ctenomys uco sp. nov. is known from only two localities, including its terra typica (see above) and Finca Caicayén II, near the small village of Sarmiento (33.3925°S, 69.2007°W; Tupungato Department, Mendoza Province) (Fig.
Mostly unknown. Field work indicates the species has solitary habits. The two known localities for this species are placed between 1000–2710 m a.s.l. The landscape in this area corresponds to an ecotone between the Low Monte and the Southern Andean Steppe (sensu Olsen et al. 2001), characterized by a mosaic of grasslands, shrublands, and vineyards (Fig.
Ctenomys uco sp. nov. apparently has a small geographic range (Fig.
Taxonomic work with Ctenomys is far from complete. Ongoing studies suggest that some nominal forms will likely fall into synonymy with others, while there are still many candidate species whose distinction needs to be tested (e.g., the “Quijadas” lineage; see
The case of C. fochi, the sister species to Ctenomys uco sp. nov., is an eloquent example of the limited ecological and systematic knowledge that characterizes most species of Ctenomys. During almost a century, this species was only known from four specimens, including its holotype, all coming from the species type locality (
Based on the results of the phylogenetic analysis conducted here, in which haplotypes of specimens assigned to C. johannis are nested within the clade corresponding to C. mendocinus and the morphological evidence provided by
We are grateful to A. Zarco, and the Terrazas de los Andes and Chandon Argentina Staff (D. Mezzatesta, M. Sonzogni, A. Sánchez, and C. Mamonte) for the logistical support in the field. We thank the Dirección de Recursos Naturales Renovables of Mendoza Province for the research permits (RES ANP EX-483484-2020). Also, we are grateful to curators and managers of the collections consulted: S. Bogan (CFA), B. Bender and R. Ojeda (CMI), and S. Lucero (MACN). This research was partially funded by PIP 03055 (PT) and PICT02019 (AO). Likewise, we are grateful for the revisions and comments made by Uliana Gorobeyko and Scott Gardner, which truly improved the manuscript.
List of specimens included in morphological comparisons. See “Materials and methods” for explanation of acronyms.
Catalog number | Species | Locality |
---|---|---|
CFA-MA 12120 | Ctenomys fochi t | Argentina, Catamarca, Capayán, Chumbicha |
CFA-MA 12118 | Ctenomys fochi t | Argentina, Catamarca, Capayán, Chumbicha |
CMI 7707 | Ctenomys fochi* | Argentina, Córdoba, Cruz del Eje, Estancia La Providencia |
CFA-MA 12028 | Ctenomys fochi | Argentina, San Juan, Jáchal, Valle del río Gualcamayo, El Chepical |
CFA-MA 11621 | Ctenomys fochi | Argentina, San Juan, Jáchal, Valle del río Gualcamayo, El Chepical |
CFA-MA 12031 | Ctenomys fochi | Argentina, San Juan, Jáchal, Valle del río Gualcamayo, El Chepical |
MACN-Ma 30872 | Ctenomys fochi | Argentina, Santiago del Estero, Giménez, Isca Yacu, finca Zorro Pozo |
MACN-Ma 30873 | Ctenomys fochi | Argentina, Santiago del Estero, Giménez, Isca Yacu, finca Zorro Pozo |
MACN-Ma 30874 | Ctenomys fochi | Argentina, Santiago del Estero, Giménez, Isca Yacu, finca Zorro Pozo |
MACN-Ma 30875 | Ctenomys fochi | Argentina, Santiago del Estero, Giménez, Isca Yacu, finca Zorro Pozo |
MACN-Ma 30876 | Ctenomys fochi | Argentina, Santiago del Estero, Giménez, Isca Yacu, finca Zorro Pozo |
CMI 7223 | Ctenomys mendocinus*t | Argentina, Mendoza, Capital, Cerro de la Gloria |
CMI 2806 | Ctenomys mendocinus t | Argentina, Mendoza, Capital, Cerro de la Gloria |
CMI 2650 | Ctenomys mendocinus | Argentina, Mendoza, Godoy Cruz, San Ignacio |
CMI 2664 | Ctenomys mendocinus | Argentina, Mendoza, Godoy Cruz, San Ignacio |
CMI 2658 | Ctenomys mendocinus | Argentina, Mendoza, Godoy Cruz, San Ignacio |
CMI 2804 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, Chacras de Coria |
CMI 2806 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, Chacras de Coria |
CMI 2519 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, La Puntilla |
CMI 2777 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, La Puntilla |
CMI 2812 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, La Puntilla |
CMI 2810 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, La Puntilla |
CMI 2786 | Ctenomys mendocinus | Argentina, Mendoza, Luján de Cuyo, La Puntilla |
CMI 2515 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2646 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2514 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2513 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2489 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2510 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2817 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 2512 | Ctenomys mendocinus | Argentina, Mendoza, Santa Rosa, Ñacuñán |
CMI 7712 | Ctenomys uco sp. nov.h | Argentina, Mendoza, Tunuyán, Cajón de Arenales |
CMI 7739 | Ctenomys uco sp. nov.p | Argentina, Mendoza, Tunuyán, Cajón de Arenales |
CMI7735 | Ctenomys uco sp. nov. | Argentina, Mendoza, Tupungato, Finca Caicayén II |
CMI7736 | Ctenomys uco sp. nov. | Argentina, Mendoza, Tupungato, Finca Caicayén II |
CMI7737 | Ctenomys uco sp. nov. | Argentina, Mendoza, Tupungato, Finca Caicayén II |
CMI7738 | Ctenomys uco sp. nov. | Argentina, Mendoza, Tupungato, Finca Caicayén II |
Table S1
Data type: .xlsx
Explanation notes: List of cytochrome b gene sequences of specimens of the Ctenomys mendocinus species group included in the genetic-based analyses. For each sequence, details of species/lineage (see text), GenBank accession numbers and geographic provenance are given. Sequences gathered here are indicated with an #. For these sequences catalog numbers are also provided (see collection acronyms in the main text). Species/lineage assignment follows the taxonomy discussed in the text, following