A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945)

Lehr, Edgar; Lundberg, Mikael; Cusi, Juan Carlos; Sites, Jack W.; Torres, Claudia; Aguilar-Puntriano, César

doi:10.3390/taxonomy5020018

Open AccessArticle

A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945)^†

by

Edgar Lehr

^1,2,*

,

Mikael Lundberg

²

,

Juan Carlos Cusi

²

,

Jack W. Sites, Jr.

³,

Claudia Torres

² and

César Aguilar-Puntriano

²

¹

Department of Biology, Illinois Wesleyan University, Bloomington, IL 61701, USA

²

Departamento de Herpetología, Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Avenida.Arenales 1256, Jesús María, Lima 15072, Peru

³

Department of Biology, Brigham Young University, 4102 Life Sciences Building, Provo, UT 84602, USA

^*

Author to whom correspondence should be addressed.

^†

zoobank: urn:lsid:zoobank.org:pub:90E2E300-ED3A-495B-B0CF-E59A6125F63C.

Taxonomy 2025, 5(2), 18; https://doi.org/10.3390/taxonomy5020018

Submission received: 6 January 2025 / Revised: 17 March 2025 / Accepted: 28 March 2025 / Published: 2 April 2025

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Abstract

:

We describe a new species of snake of the genus Tachymenoides using molecular and morphological evidence. The description is based on 21 specimens (4 females, 17 males) obtained in the regions of Pasco, Junín, and Puno between 2190 and 3050 m elevation. A maximum likelihood phylogenetic tree based on two mitochondrial (12S and cyt-b) genes and one nuclear (c-mos) gene shows that the new species is the sister taxon of T. affinis and distinct from Galvarinus tarmensis, which we transfer back to Tachymenis. The new species has smooth dorsal scales without apical pits usually in 19/17/15 series, 1 preocular, 2 postoculars, 1 loreal undivided nasal scale, 8 supralabials (4th and 5th in contact with the eye), 9 infralabials, 1–2+2–3 temporals, 139–157 ventrals, 52–67 subcaudals, and a divided cloacal scale. The longest specimen, a male, had a total length of 559 mm. Two females contained six and five eggs with small embryos. In life, the dorsum and flanks are olive brown to pale grayish brown with scattered black and cream flecks and no longitudinal stripes. Ventral coloration is highly variable, nearly uniformly black, mottled gray and dark-gray, mottled pale gray and tan, or pale grayish tan. Usually, three irregularly shaped, narrow, longitudinal ventral stripes are present. The iris is brown with a distinct yellowish-tan ringlet.

Keywords:

Squamata; reptiles; snakes; Andes; Pasco region; Junín region; Puno region; Tachymenis tarmensis

1. Introduction

A recent comprehensive phylogenetic analysis of Tachymenini snakes using molecular and morphological characters by Trevine et al. [1] resulted in the description of four new genera (Apographon, Galvarinus, Tachymenoides, and Zonateres), three revalidated genera, and many new combinations of snakes formerly incorrectly assigned to Tachymenis or Thamnodynastes. This research demonstrates a typical example of cryptic diversity and convergent evolution in Andean snakes. Trevine et al. [1] used Tachymenis affinis Boulenger, 1896 [2], as the type species for the new genus Tachymenoides, and a new species, Tachymenoides harrisonfordi, was described by Lehr et al. [3] 127 years after the discovery of Ta. affinis.

Tachymenis was described by Wiegmann in 1835 [4] from Peru with Tachymenis peruviana as a type species. Tachymenis tarmensis Walker, 1945 [5], was known from a single adult female specimen collected in the Junín region, Tarma, in 1914 and described 31 years after its preservation. Without genetic material and limited morphological characteristics, Trevine et al. [1] questioned the generic affiliation with Tachymenis and tentatively moved T. tarmensis to Galvarinus. In their Appendix S3, Trevine et al. [1] identified two additional specimens (AMNH 117734, a pregnant female from Tarma, Junín region, Peru, and MUSM 31152, identified as Thamnodynastes sp. in the collection, a male from Chacos, Pasco region, Peru) as G. tarmensis based on “melanic coloration, smaller SVL size, and ventral and subcaudal counts”, and on hemipenial characters that are shared with the other members of Galvarinus, such as the enlarged first row of calyces [1]. Trevine et al. [1] described the hemipenis for G. tarmensis based on MUSM 31152.

Lehr et al. [3] commented on potentially unnoticed cryptic species diversity within Andean snakes cataloged in museum collections as Tachymenoides affinis and Tachymenis peruviana Wiegmann, 1835 [4]. Our revision of Tachimenini snakes at the Natural History Museum in Lima, Peru, including the holotype of Tachymenis tarmensis (FMNH 5698), and using molecular and morphological characters, resulted in the identification of two female specimens of Galvarinus tarmensis from Junín. Furthermore, we show that the male specimen (MUSM 31152) from the Pasco region, previously identified as G. tarmensis by Trevine et al. [1], represents a new species that is distributed in the regions of Pasco, Junín, and Puno. Twenty additional specimens of our new species were misidentified in the herpetological collection of the Natural History Museum of the San Marcos University (Lima, Peru) as Tachymenis affinis and T. peruviana.

In this paper, we correct the generic affiliation of Galvarinus tarmensis and describe a new species of Tachymenoides using molecular and morphological characters.

2. Materials and Methods

2.1. Molecular Genetics

Tissue samples (liver and muscle) from two specimens of the new species (MUSM 23470, 25422) and two specimens of Galvarinus tarmensis (MUSM 31533, 38469) were taken before preservation and stored in 96%-proof ethanol. The samples were sequenced for 12S rRNA and c-mos. 12S rRNA, cytb, and c-mos sequences from GenBank representing Tachymenini taxa were included in the phylogenetic analyses (see Appendix A, Table A1). Double-stranded DNA polymerase chain reactions (PCRs) amplified target regions using forward 12S rRNA primer L1091mod (sequence: 5′ CAA ACT AGG ATT AGA TAC CCT ACT AT 3′; modified from Kocher et al. [6]) and reverse H1557mod (5′ GTA CRC TTA CCW TGT TAC GAC TT 3′; modified from Knight and Mindell [7]; and for c-mos forward primer S77 (5′ CAT GGA CTG GGA TCA GTT ATG 3′) and reverse S78 (5′ CCT TGG GTG TGA TTT TCT CAC CT 3′)), both from Lawson et al. [8].

PCR products were visualized on 10% agarose gels to ensure the targeted product was clearly amplified, then purified using a MultiScreen PCR (l) 96 (Millipore Corp., Billerica, MA, USA), and directly sequenced using the BigDye Terminator v 3.1 Cycle Sequencing Ready Reaction (Applied Biosystems, Foster City, CA, USA). The cycle sequencing reactions were purified using Sephadex G-50 Fine (GE Healthcare) and MultiScreen HV plates (Millipore Corp.). Samples were then analyzed on an ABI3730xl DNA Analyzer in the DNA Sequencing Center (DNASC) at Brigham Young University.

We used Geneious Pro versio 2025.0.2 (https://www.geneious.com) for sequence assembly and edition. Sequence alignment was performed in the MUSCLE 5.1 ([9]) plugin in Geneious Pro and trimmed to a length of 475 bp for 12S, 735 bp for cytb, and 688 bp for c-mos. Maximum likelihood (ML) phylogenetic analysis of the concatenated regions (with and without cytb) and by partition was performed using RAXML ([10]) plugin in Geneious Pro with GTR Gamma for substitution model and 1000 Rapid Bootstrapping replications and search for best scoring ML tree algorithm. We estimated clade distances on a 12S ML tree using the Species Delimitation ([11]) plugin in Geneious Pro.

2.2. Morphology

The specimens (17 males, 4 females) were fixed in 10% formol and stored in 70% ethanol. Taxonomy follows Trevine et al. [1]. The format of the description and terminology of the morphological characters follow mostly Bailey et al. [12], Franco et al. [13], and Trevine et al. [1]. Terminology of head scales follows Dixon et al. [14]. We recorded (Appendix B, Table A2) the number of supralabials and noted which of those were in contact with the eye, number of loreals, pre- and postoculars, temporals (distinguished as primary temporals in contact with the rear of the postoculars, and secondary temporals in contact with the rear of the primaries; primary temporals were separated by a plus sign from the secondary temporals), and infralabials, noting how many are in contact with the primary and secondary genials. Dorsal scales were counted in three different portions of the body (separated by a slash): one head length before the head, at the middle of the body, and one head length before the cloaca. We followed Dowling [15] for counting ventrals and subcaudals (excluding the spine). A slash separates character counts taken on opposite sides of the head, with the left side indicated first. The number of maxillary and dentary teeth on the left and right jaws was counted. Sex was determined by the presence or absence of inverted hemipenes by making a small incision midventrally on the base of the tail following Dowling and Savage [16]. All measurements are in mm. Snout–vent length (SVL, distance from the snout tip to the posterior end of the cloaca) and tail length (TaL, distance from the posterior end of the cloaca to the tip of the tail) were taken to the nearest millimeter from the specimen straightened against a meter stick. Other measurements taken with digital calipers to the nearest 0.1 mm include: eye distance (straight line distance between pupils), eye to nostril distance (straight line distance between anterior corner of orbita and posterior margin of external nares), snout length (straight line distance from tip of snout to posterior end of interprefrontal suture), head length (distance from tip of snout to posterior margin of jaw), head width (distance between lateral margins of supraoculars), head height (at tallest point), distance between nostrils, eye height, eye width, and ovarian egg length (distance between anterior and posterior end). Greatest length and greatest width of the following head scales (left if applicable) were measured with a digital caliper to the nearest 0.1 mm: internasal, loreal, prefrontal, frontal, and parietal. Photographs of preserved specimens (except for hemipenis) were taken while they were temporarily immersed in ethanol to minimize reflections. Hemipenial morphology follows Dowling and Savage [16] and Zaher [17] and is based on two specimens (holotype MUSM 31152, paratype MUSM 18580) with (partially) everted hemipenes.

Notes on the coloration in life were taken from field notes and photographs. Comparative data were taken from original species descriptions and the specimens examined (Appendix C). Collection acronyms are: AMNH = American Museum of Natural History, New York, USA. CORBIDI = Centro de Ornitología y Biodiversidad, Lima, Peru. MTD = Museum für Tierkunde Dresden, Dresden, Germany. MUSM = Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Lima, Peru. NZCS = National Zoological Collection of Suriname, Paramaribo, Suriname. Field number codes are: IWU = Illinois Wesleyan University, and ML = Mikael Lundberg. Threat status was evaluated using the IUCN criteria [18]. The map was designed with ArcGIS (ESRI 2019. ArcGIS Desktop 10.8 version 10.7.0. Redlands, CA: Environmental Systems Research Institute).

3. Results

3.1. Molecular Genetics

Our phylogeny (Figure 1), including all gene regions, recovered a new species as sister taxon of Tachymenoides affinis with a bootstrap support of 0.79 at the node of their most recent common ancestor. The new species and Ta. affinis share a common ancestry with Ta. harrisonfordi with a strong bootstrap support of 0.96 at the node. Furthermore, Tachymenis tarmensis is recovered as the sister taxon of T. peruviana with a bootstrap support of 0.69 at the node of its most recent common ancestor. Both species share a common ancestry with T. ocellata with a bootstrap support of 0.69 at the node of their most recent common ancestor. All three species (T. ocellata Duméril, Bibron, Duméril, 1854 [19], T. peruviana, and T. tarmensis) share a common ancestry with T. trigonatus with a strong bootstrap support of 0.94 at the node.

Our phylogeny without cytb (Figure S1) also shows the new species as the sister taxon of Tachymenoides affinis with a bootstrap support of 0.80 at the node of its most recent common ancestor. The new species and Ta. affinis share a common ancestry with Ta. harrisonfordi with a strong bootstrap support of 0.86 at the node. Tachymenis tarmensis is recovered as the sister taxon of T. peruviana with low support (0.32) at the node of its most recent common ancestor. Both species share a common ancestry with T. trigonatus with a bootstrap support of 0.32 at the node of their most recent common ancestor. All three species (T. trigonatus, T. peruviana, and T. tarmensis) share a common ancestry with T. ocellata with a bootstrap support of 0.41 at the node.

12S ML tree interclade distances between new species and Ta. affinis are similar (0.020) to interclade distances between Ta. harrisonfordi and Ta. affinis (0.027). Interclade distances between the new species and Ta. harrisonfordi is (0.026). Intraclade distance between new species is 0.003.

3.2. Tachymenis tarmensis Walker, 1945 [5]

Our phylogeny reveals that the tentative placement of Tachymenis tarmensis in Galvarinus by Trevine et al. [1] is incorrect, and we transfer it back to Tachymenis following Walker [5]. One specimen, a female (MUSM 38469), was photographed (Figure 2). In life, the dorsolateral view shows a pale yellowish-olive–gray coloration with scattered dark-brown flecks; the venter is pale gray. The postocular stripe is pale brown, and the supralabials are pale yellowish tan and marbled gray without labial bars. The iris is pale bronze. For the distribution, see Figure 3.

3.3. Generic Assignment

We place the new species in the genus Tachymenoides of the family Dipsadidae following Trevine et al. [1] and our molecular phylogeny (Figure 1).

3.4. Systematics

ZooBank LSD: urn:lsid:zoobank.org:pub:90E2E300-ED3A-495B-B0CF-E59A6125F63C.
Family Dipsadidae Bonaparte, 1838 [20].
Tribe Tachymenini Bailey, 1967 [21].
Genus Tachymenoides Trevine et al., 2022 [1].
Tachymenoides goodallae sp. nov. Lehr, Lundberg, Cusi, Sites, Torres and Aguilar-Puntriano, 2025.
ZooBank LSD: urn:lsid:zoobank.org:act:9B56676E-70A8-46E1-BA7E-E80F1B057DA4 (Figure 4, Figure 5 and Figure 6).
Galvarinus tarmensis in Trevine et al. [1].
Thamnodynastes sp. in Lehr et al. [22].

Figure 4. Living holotype of Tachymenoides goodallae sp. nov. (MUSM 31152) in dorsolateral (A), dorsal (B), and ventral views (C). Total length of the snake is 524 mm. Photos by EL.

Figure 5. Preserved holotype of Tachymenoides goodallae sp. nov. (MUSM 31152) in dorsal (A) and ventral (B) views. Red arrows indicate the longitudinal midventral and ventrolateral stripes. Total length of the snake is 524 mm. Photos by EL.

Figure 6. Head of the living holotype of Tachymenoides goodallae sp. nov. (MUSM 31152) in lateral (A), dorsal (B), and ventral (C) views. The head length is 13.8 mm. Photos by EL.

Holotype (Figure 4, Figure 5 and Figure 6): MUSM 31152 (field number IWU 100), a male from the mountain ridge close to the radio tower at Chacos (10°39′05.7″ S, 75°17′39.3″ W; WGS84) in the buffer zone of the Yanachaga Chemillén National Park, 2836 m a.s.l., Distrito Oxapampa, Provincia Oxapampa, Pasco, Peru, collected on 15 February 2012 by E. Lehr and J. C. Cusi.

Paratypes: n = 20 (4 females, 16 males), all from Peru, all collected by M. Lundberg. From along the road between Auquimarca and Uchuhuerta (10°43′29.2″ S, 75°41′57.3″ W), 2700 m a.s.l., Distrito Paucartambo, Provincia Pasco, Pasco, collected on 09 February 2000 (MUSM 17813, ML 9, male), on 21 March 2000 (MUSM 17828, ML 15, male), and on 19 February 2001 (MUSM 18122, ML 122, male). From Auquimarca (10°44′54.2″ S, 75°42′18.4″ W), 2650 m a.s.l., Distrito Paucartambo, Provincia Pasco, Pasco, collected on 04 April 2000 (MUSM 17839, ML 22, female), collected on 21 March 2002 (MSUM 18580, ML 340, male), collected on 10 November 2002 (MUSM 19051, ML 456, male), and collected on 28 March 2004 (MUSM 23422, ML 680, female). From Huallamayo (10°45′53.2″ S, 75°43′25.1″ W), 2500 m a.s.l., Distrito Paucartambo, Provincia Pasco, Pasco, collected on 13 March 2001 (MUSM 18104, ML 160, male). From Puagmaray (10°39′02.5″ S, 75°46′35.5″ W), 2600 m a.s.l., Distrito Huachon, Provincia Pasco, Pasco, collected on 26 April 2003 (MUSM 19167, ML 496, male), collected on 28 March 2003 (MUSM 19177, ML 484, male), collected on 29 April 2003 (MUSM 19187, ML 498, male), and collected on 23 March 2004 (MUSM 23420, ML 676, female). From the mountain ridge close to the radio tower at Chacos (10°39′30.1″ S, 75°17′52.2″ W), 2780 m a.s.l., Distrito Oxapampa, Provincia Oxapampa, Pasco, collected on 06 November 2004 (MUSM 23485, ML 996, male; MUSM 23492, ML 1024, male), collected on 16 January 2005 (MUSM 24715, ML 1144, male; MUSM 24716, ML 1145, male), and collected on 24 March 2006 (MUSM 25403, ML 1400, male; MUSM 25422, ML 1399, male, GenBank accession number PV272666). From Ingenio (11°27′01.2″ S, 75°41′10.6″ W), 3050 m a.s.l., Distrito Tarma, Provincia Tarma, Junín, collected on 19 September 2004 (MUSM 23470, female [tissue sample MTD 2037], ML 917, GenBank accession numbers PV272665, PV255662). From Pacchani (14°04′42.7″ S, 69°41′55.7″ W), 2190 m, Distrito Limbani, Provincia Sandia, Puno, collected on 06 November 2004 (MUSM 23483, ML 987, male).

Diagnosis: Usually 1 preocular, 10 maxillary teeth; 9 dentary teeth; dorsal scale rows smooth in 19/17/15 series; dorsal scales lacking apical pits; 140–157 ventrals in males (n = 17), 139–142 ventrals in females (n = 4); 52–67 paired subcaudals in males (n = 17), 52–56 paired subcaudals in females (n = 4); hemipenes with pronounced and deep spinulate calyces on the base of capitulum and distal portion of hemipenial body, extending onto half of the body on the asulcate side with the first proximal row of calyces distinctly larger, containing 9 calyces; in real life, the dorsum and flanks are olive brown to pale grayish brown with scattered black and cream flecks and no longitudinal stripes; the flanks are olive brown, dark brown, or dark gray, and darker than the dorsum toward the posterior body end; the head is laterally pale grayish brown or cream with a dark-brown lateral stripe from the nose to postoculars and diagonally to corner of the mouth, and contrasting dark-brown flecks on the cream supralabials and infralabials; the iris is brown with a distinct yellowish-tan ringlet; the ventral coloration is highly variable: nearly uniformly black, gray, and mottled dark gray; pale gray and mottled tan; or pale grayish tan; and usually, three irregularly shaped, narrow, longitudinal ventral stripes are present (one midventral and one ventrolateral on each side).

Comparisons: see Appendix D, Table A3. Tachymenoides goodallae sp. nov. and Ta. affinis are similar in size (maximum SVL 433 mm vs. 436 mm in Ta. affinis, [1]) and have hemipenes with large calyces restricted to the hemipenial body on the asulcate side. However, both species can be distinguished as follows: Tachymenoides goodallae sp. nov. has dorsal scale rows mostly in 19/17/15 (61.9%, n = 13) series (17/17/15 in Ta. affinis, 81.3%, n = 13, [1]); wider hemipenial calyces (narrower, Appendix S S11 in [1] for CORBIDI 11791); cream supralabials with black blotches (black blotches absent); hemipenis with the first row of calyces distinctly larger (not distinctly larger); and a pale-gray venter with three continuous, longitudinal black stripes, one midventrally and one ventrolaterally on each side (two to five non-continuous longitudinal ventral stripes in a few specimens, forming a dark band in each ventral scale, with a general darkening tendency toward the tail; some specimens have an overall dark venter, with no discernible stripes [1]). Both Tachymenoides goodallae sp. nov. and Ta. harrisonfordi have dorsal scale rows in 19/17/15 series, supralabials with black blotches, but Ta. goodallae sp. nov. lacks longitudinal body folds (present in Ta. harrisonfordi), and the iris is brown (copper) with a distinct yellowish-tan ringlet (absent).

Tachymenoides goodallae sp. nov., Ta. affinis, and Ta. harrisonfordi lack apical pits ([1,3], in this paper) and are immediately separated from all 15 Dryophylax spp., which have apical pits ([1,23]). Tachymenis currently contains four species: T. ocellata, T. peruviana, T. tarmensis, and T. trigonata. Tachymenoides goodallae sp. nov. has dorsal scales usually in 19/17/15 series whereas Tachymenis spp. has dorsal scales usually in 19/19/15 series [1]. Tachymenoides goodallae sp. nov. has contrasting dark-brown flecks on the cream supralabials and infralabials, whereas Tachymenis spp. has labials with a homogeneous clear background, with solid bands or small blotches under the eye [1]. Furthermore, the head of Ta. goodallae sp. nov. is shorter than the head of T. tarmensis (Figure 7).

Galvarinus contains two species, G. chilensis (Schlegel, 1837 [24]) and G. attenuatus Walker, 1945 [5], which have dorsal scales in 19/19/15 series and apical pits (see Appendix S3 in [1]). The dorsal scales in Ta. goodallae sp. nov. are mostly in 19/17/15 rows (61.9%, n = 13), followed by 19/19/15 (19.0%, n = 4), 19/18/15 (14.3%, n = 3), and 19/17/14 (4.8%, n = 1); see Table A2. Trevine et al. [1] mention the dorsal scale row variation in Galvarinus as 19/19/15 (79%, n = 93), 19/19/17 (13%, n = 14), and one specimen in Tachymenis tarmensis as 19/19/13. Tachymenoides goodallae sp. nov. differs from G. attenuatus in having males with less subcaudals (52–67 [n = 17] vs. 69–75 in G. attenuatus, [1]) and females with less subcaudals (52–56 [n = 5] vs. 63–65 in G. attenuatus, [1]).

Description of holotype: Adult male (MUSM 31152) with partially everted left hemipenis (right one cut off and missing) (Figure 4 and Figure 5); SVL 416 mm; TL 108 mm. Smooth dorsal scales in 19/17/15 rows, without apical pits; 155 ventral scales; 57/56 paired subcaudals; a divided cloacal scale; head scales consisting of one rostral, two internasals, two prefrontals, one frontal, one supraocular on each side of the head, and two parietals (Figure 6); 8/8 supralabials, 4th+5th/4th+5th contacting the orbit; 9/9 infralabials, 1st to 5th/1st to 5th contacting the first pair of genials, 5th/5th contacting the second pair of genials; two pairs of chin shields; 2+3/1+2 temporals; undivided nasal; 1/1 preocular, 2/2 postoculars; maxilla with 7/8 pre-diastemal teeth and 2/2 grooved and enlarged post-diastemal teeth; and 9/9 dentary teeth. Head length: 13.8 mm; head width: 6.6 mm; head height: 7.2 mm; snout length: 3.3 mm; distance between nostrils: 3.1 mm; eye height: 2.6 mm; eye width: 2.9 mm; distance between eyes: 8.3 mm; eye–nostril distance: 2.4 mm; internasal scale (left) wider than long (length: 1.7 mm, width: 2.3 mm); loreal scales: 1/1, slightly wider than long (length: 1.0 mm, width: 1.2 mm); prefrontal scale slightly wider than long (length: 2.2 mm, width: 2.3 mm); frontal scale longer than wide (length: 5.3 mm, width: 2.6 mm); and parietal scale longer than wide (length: 5.4 mm, width: 3.6 mm).

Trevine et al. [1] provided in their Appendix S3 the following description of the right hemipenis of MUSM 31152 but did not provide a photograph: “The hemipenis is faintly bilobed with a short capitulum covered in papillated calyces on the lobes and pronounced spinulate calyces on the base of capitulum and distal portion of hemipenial body, extending onto half of the body on the asulcate side. Five rows of body spines that progressively get larger towards the base of the body, and first row with 9 spines encircling the body. Sulcus spermaticus divided at the base of capitulum; subtle constriction between body and base of the hemipenis, base of the organ covered with spines (smaller than those on the body)”. The left hemipenis, not fully everted, is pictured in Figure 8 as a reference.

Coloration of holotype in life (Figure 4 and Figure 6): the dorsum and flanks are olive brown with scattered black and cream flecks. Flanks are slightly darker than the dorsum, especially toward the posterior body end. No longitudinal dorsal stripes are discernible. The head has a dark-brown dorsal stripe on each parietal, bifurcating slightly before the posterior end of the parietals, with one end running diagonally over the temporals to the corner of the mouth and the other end (3–4 scales wide) running straight dorsally and slightly passing the level of the jaw angles. Laterally (Figure 6A), the head has an irregularly shaped dark-brown canthal stripe covering the nasal, upper halves of loreal, and preocular; a dark-brown postocular stripe extends between the suture of postoculars, diagonally across the lower half of anterior temporal, upper half of the seventh supralabial, and eight supralabial to the corner of the mouth. Supraliabials are cream with irregular dark-brown flecks, the largest one on the fifth supralabial. The throat is yellowish cream with a grayish-brown streak along midventral suture of first and second genials, and on both the outer sutures of the first and second genials (Figure 6C). The infralabials are yellowish tan with grayish-brown streaks (Figure 6A,C). Other ventral surfaces are pale gray and mottled tan with three dark-gray, irregularly shaped, narrow longitudinal stripes: one midventral stripe from posterior end of the throat and growing wider toward the anal plate, and one ventrolateral stripe on each side running from the first ventral scale toward the anal plate. The tail is gray and mottled tan. The posterior half of the ventral body is slightly darker colored than the anterior body half. The iris is brown, with a distinct yellowish-tan ringlet around the elliptical pupil.

Coloration of holotype after 12 years in alcohol (Figure 5) is nearly identical as described for the life holotype, except that the coloration is slightly paler. The ventral surfaces are pale gray and mottled dark gray, with the three narrow longitudinal stripes discernible but slightly faded out. The iris is gray with a white ringlet.

Variation (Figure 9 and Figure 10): For variations in scutelation and measurements, see Appendix B, Table A2. Males are larger (total length: 348–559 mm, n = 17 vs. 291–322 mm, n = 4 in females), have more ventrals (140–157, n = 17 vs. 139–142 in females, n = 4), and more subcaudals (52–67, n = 17 vs. 52–56 in females, n = 4). Most specimens (n = 13) have the dorsal scales in 19/17/15 (61.9%, n = 13) series, one specimen has 19/17/14 (4.8%, n = 1), three specimens have 19/18/15 (14.3%, n = 3), and four specimens have 19/19/15 (19.0%, n = 4). Most specimens (95.2%, n = 20) have eight supralabials with the fourth and fifth contacting the orbit, one specimen (4.8%, n = 1, MUSM 19167) has, on the right side, nine supralabials with the fifth and sixth contacting the orbit. Most specimens (66.7%, n = 14) have 9/9 infralabials, three (14.3%) have 9/10, two (9.5%) have 8/8, one (4.8%) has 10/9, and one (4.8%) has 9/8. Most specimens (66.7%, n = 14) have 1–5 infralabials in contact with the first genial, but seven specimens (33.3%) have 1–4 infralabials in contact with the first genial. Most specimens (81.0%, n = 17) have the fifth infralabial in contact with the second genial, but three specimens (14.3%) have the fourth and fifth infralabials in contact with the second genial, and one specimen (4.8%) has the fourth infralabial in contact with the second genial. Most of the specimens (71.4%, n = 15) have 1/1 preoculars, five specimens (23.8%) have 2/2 preoculars, and one specimen (4.8%) has 1/2 preoculars. All specimens have 2/2 postoculars. Great variation can be observed in the number of anterior and posterior temporals: 12 specimens (57.1%) have 2+3/2+3, two specimens (9.5%) have 2+2/2+2, two specimens (9.5%) have 1+2/1+2, and five specimens (23.8%) have the left and right sides of the head in variable numbers (2+3/1+2, 2+3/2+2, 2+2/1+2, 1+2/1+3, 2+2/2+3). Male paratype MUSM 18580 has partially everted hemipenes and characteristics, as described for the holotype.

Dorsal and lateral coloration variations in life specimens are documented in Figure 9. Overall, the ground coloration is as described for the holotype, except for two specimens that have a pale grayish-brown dorsum (Figure 9B,D). The flanks are darker than the dorsum and are dark brown (Figure 9A,C,E), grayish brown (Figure 9B,D), and dark gray (Figure 9F). All specimens have heads with a dark-brown postocular stripe (not distinct in Figure 7E) and contrasting dark-brown flecks on the supralabials (not distinct in Figure 9C,E). Most contrasting head patterns of pale olive gray with brown stripes and flecks is shown in female MUSM 23470 (Figure 9B). All specimens have a brown iris with a distinct yellowish-tan ringlet (difficult to see in the small images in Figure 9).

Coloration and pattern variations in ventral surfaces (excluding head) can be observed in preserved specimens as follows (Figure 10): Five specimens (MUSM 17839, 19051, 19167, 19187, 18580) have nearly uniformly black ventral surfaces but ventrolateral longitudinal stripes (one on each side) that are still discernable (Figure 10A); four specimens (MSUM 18104, 24715, 24716, 25403) have gray and mottled dark-gray ventral surfaces with three distinct black longitudinal stripes (one midventrally, one ventrolaterally on each side) (Figure 10B); two specimens (MUSM 18122, 23429) have grayish-tan ventral surfaces with a broad, dark-gray, washed out midventral longitudinal band (Figure 10C); one specimen (MUSM 25422) has gray and mottled dark-gray ventral surfaces but the three black longitudinal stripes are not as discernable, except for the midventral stripe (Figure 10D; and six specimens (MUSM 17813, 17828, 19177, 23422, 23483, 23485) have, like the holotype, pale grayish-tan ventral surfaces, Figure 10E, and one specimen (MUSM 23420) has grayish-tan ventral surfaces with five narrow longitudinal stripes: one tan midventral stripe flanked by black stripes and one black ventrolateral stripe on each side (Figure 10F).

Etymology: We dedicate this species to ethologist and conservationist Dr. Jane Goodall for her scientific accomplishments and worldwide conservation activism, which inspire and encourage people to protect biodiversity. The English Common Name is Goodall’s Slender Snake.

Distribution, natural history, and threat status: The type locality, Chacos (Figure 3 and Figure 11), is within the buffer zone of the Yanachaga Chemillén National Park and is located south of the town of Oxapampa on a mountain ridge close to the radio tower. The vegetation consists of sclerophyllous forests, bushes, mosses, lichens, and many epiphytic bromeliads, orchids, and ferns, surrounded by primary montane forests at slightly lower elevations. Several small scattered lagunas (Figure 11B). Sympatric reptiles include two gymnophthalmid lizards, Dendrosauridion yanesha (MUSM 23524, 25345) and Proctoporus sp. (MUSM 23545–23547), and two colubrid snakes, Atractus nigricaudus (MUSM 23500, 25429, 25379) and Erythrolamprus epinephelus (MUSM 24764, 24765, 25420). Sympatric amphibians include two strabomantid frogs, Pristimantis aniptopalmatus (MUSM 31151) and P. bromelicaceus (MUSM 46309–46311), and the tree frog Scinax oreites (MUSM 46490–46493). The holotype was found in the afternoon sun, basking on brown and white lichens (Figure 11D), and was slightly covered by them. Two females (MUSM 23420, 23422) contained, respectively, six and five yolky eggs with small embryos. The longest egg measured 25.6 mm (Figure 12B). Although viviparous reproduction is known for Tachymenini [1], the early pregnancy status of the two observed females of Ta. goodallae sp. nov. does not allow us to conclude viviparity. Tachymenoides goodallae sp. nov. is ophistoglyphous and venomous but harmless to humans. Based on the large distributional area (Figure 3), we suggest classifying Ta. goodallae sp. nov. as “Least Concern” according to the IUCN red list criteria ([18]).

4. Discussion

Since the description of Tachymenoides by Trevine et al. [1] based on Ta. affinis, two additional species (Ta. goodallae, Ta. harrisonfordi) were added to this genus. Consequently, the diagnosis of Tachymenoides needs to be revised as follows: smooth dorsal scale rows are usually in 17/17/15 rows (Ta. affinis [1]) or 19/17/15 rows (Ta. goodallae sp. nov. and Ta. harrisonfordi [3]), see Appendix D, Table A3. However, additional research and specimens are needed for Ta. harrisonfordi to describe the hemipenial morphology and pholidosis range of males and females. The distributional area of Tachymenoides goodallae sp. nov. is large (see Figure 3), with a gap between the northern and southernmost collecting sites. A similar distributional gap is known for Dendrosauridion yanesha, which has the same type locality as Ta. goodallae sp. nov. and is known from a second locality in Cusco [22]. In Junín, Tachymenoides goodallae sp. nov. is sympatric or even syntopic with Tachymenis tarmensis. We follow an integrative taxonomic approach and consider our species description justified. However, the unbalanced sex ratio in the type series (17 males, 4 females) and conclusions regarding the larger size of males (total length: 348–559 mm, n = 17) compared to females (total length: 291–322 mm, n = 4) could be a potential sampling artifact. Our phylogeny, which includes tissues of Tachymenis tarmensis, confirmed its generic affiliation as proposed by Walker [5] and helped revalidate its status. Although we did not include cytb sequences of this taxon, and our phylogeny without this marker shows low support for the generic position of T. tarmensis, we predict that the addition of cytb sequences will corroborate our phylogenetic hypothesis. Future research and fieldwork, especially in southeastern Andean Peru, will help to fill the current distribution gaps for Ta. goodallae sp. nov. and T. tarmensis and better understand their ecological preferences.

Naming a new species after a person, a so-called eponym or wildlife eponym, is a common practice in taxonomy. According to Wulf [25], 100 animals and nearly 300 plants have been named after Alexander von Humboldt. By dedicating a new species to a person, they are honored for their scientific accomplishments and contributions to species discoveries or society. However, this practice has received some backlash in recent years, especially when species dedicated to people from the past were no longer considered worthy based on their misconduct, racism, or misogyny, resulting in changes in popular North American bird species [26]. However, eponyms are a “good attention-getting device” [27] and are useful for raising awareness about undiscovered biodiversity and the need for conservation, especially during the current biodiversity crisis. With our eponym and 2025 being the Chinese New Year of the Wood Snake, we hope that our dedication to Dr. Jane Goodall helps to create awareness for the protection of snakes, many of which are killed out of fear.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/taxonomy5020018/s1, Figure S1: Maximum likelihood (ML) tree showing relationships of the new species and other Tachymenini genera. Phylogenetic analysis was based on concatenated 12S and c-mos gene regions. Bootstrap values are shown on nodes.

Author Contributions

Conceptualization: E.L.; Funding acquisition: E.L. and J.W.S.J.; Investigation: E.L., C.A.-P., C.T., J.C.C., J.W.S.J. and M.L.; Methodology: E.L., C.A.-P., C.T., J.C.C., J.W.S.J. and M.L.; Project administration: E.L.; Writing—original draft: E.L.; Writing—review and editing: E.L., C.A.-P., C.T., J.C.C., J.W.S.J. and M.L. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Illinois Wesleyan University, including an ASD grant awarded to E.L. and by a grant from UNMSM (project code: B24101931) to C.A.-P.

Institutional Review Board Statement

Ethical review and approval (protocol 11-003) were granted by the IACUC committee of Illinois Wesleyan University.

Data Availability Statement

All DNA sequences used in this study are in GenBank https://www.ncbi.nlm.nih.gov (accessed on 28 October 2024 and 19 February 2025).

Acknowledgments

The IACUC Committee (institutional animal care and use committee) of Illinois Wesleyan University approved this study (Protocol 11-003) in 2011, and the Ministerio del Ambiente, Lima, Peru, granted research (No -060-2003-INRENA-IFFS-DCB, No -408-2009-AG-DGFFS-DGEFFS, No -008-2011-SERNANP-DGANP-JEF) and export permits. We thank A. Resetar (FMNH) for granting the loan of the holotype of Tachymenis tarmensis, D. Kizirian (AMNH) and L. Vonnahme (AMNH) for providing access to T. tarmensis (AMNH 117734) and for providing photos of this specimen. We are grateful to P. Bowles (IUCN), N. Cox (IUCN), C. Schmidt, and F. Schmidt for their suggestion on how to reach out to the Jane Goodall Institution. M. Lewis from the Jane Goodall Institution kindly helped us to communicate with J. Goodall.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

Appendix A

Table A1. GenBank accession numbers for the taxa and genes sampled in this study.

Terminals/Organism	Voucher	12S	cytb	c-mos
Calamodontophis paucidens	MCP 5737	OP024352	OP036772	OP036907
Dryophylax chaquensis	UFMT 1387	OP024305	OP036793
Dryophylax chimanta	no number	OP024324	OP036819
Dryophylax gambotensis	MHUA-R 14364	OP024321	OP036805
Dryophylax hypoconia	UFRGS 4980	OP024306	OP036794
Dryophylax paraguanae	G.Rivas no number	OP024323	OP036807	OP036939
Galvarinus chilensis	PUC Chile PC114/RC159	OP024298	OP036831
Gomesophis brasiliensis	IBSP 72604	OP024287	OP036773	OP036961
Mesotes rutilus	no number	OP024325	OP036811	OP036917
Mesotes strigatus	IBSP 88213	OP024327	OP036818
Ptychophis flavovirgatus	MZUSP 17229	OP024290	OP036774	OP036909
Tachymenis trigonatus	FML 16130	OP024329	OP036833	OP036908
Tachymenis peruviana	FML 16131	OP024294	OP036834
Tachymenis ocellata	MACN 45865	OP024288	OP036835	OP036911
Tachymenis tarmensis	CAP 1051	PV253003		PV255660
Tachymenis tarmensis	CTG 1585	PV253004		PV255661
Tachymenoides harrisonfordi	MUSM40925	OQ883945	OQ870342
Tachymenoides affinis	MUSA 027	OP024304	OP036775	OP036916
Tachymenoides goodallae sp. nov.	MTD2037Junin	PV272665
Tachymenoides goodallae sp. nov.	ML1399Pasco	PV272666		PV255662
Thamnodynastes sertanejo	MTR (MTJ) 0503	OP024291	OP036829	OP036943
Tomodon dorsatus	MZUSP 13969	OP024300	OP036808

Appendix B

Table A2. Measurements (in mm) and pholidosis characters of the type series of Tachymenoides goodallae sp. nov. TaL is tail length, ToL is total length, * indicates an incomplete (broken and healed) tail, m = male, and f = female. For other abbreviations and symbols see Section 2.

Characters	MUSM 31152	MUSM 24715	MUSM 25422	MUSM 19051	MUSM 23485	MUSM 18580	MUSM 17813
Sex	m	m	m	m	m	m	m
SVL	416	433	422	410	418	392	396
TaL	108	126	124	132	116	127	115
ToL	524	559	546	542	534	519	511
Dorsal scales	19/17/15	19/17/15	19/17/14	19/17/15	19/19/15	19/18/15	19/17/15
Ventral scales	155	157	155	148	155	150	144
Subcaudals	57/56	65/63	64/63	67/65	63/62	58/58	59/59
Supralabials [Contacting orbit]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]
Infralabials	9/9	9/9	9/9	9/9	9/10	9/9	9/9
Left infralabials with 1 genial	1–5	1–4	1–5	1–4	1–5	1–5	1–5
Left infralabials with 2 genials	5	4–5	5	5	5	5	5
Loreals	1/1	1/1	1/1	1/1	2/1	1/1	1/1
Preoculars	1/1	1/1	1/1	1/1	1/1	1/1	2/2
Postoculars	2/2	2/2	2/2	2/2	2/2	2/2	2/2
Temporals	2+3/1+2	1+2/1+2	2+3/2+2	2+2/1+2	2+3/2+3	2+3/2+3	2+3/2+3
Characters	MUSM 23429	MUSM 19167	MUSM 18112	MUSM 23483	MUSM 24716	MUSM 19187	MUSM 17828
Sex	m	m	m	m	m	m	m
SVL	375	405	352	328	324	304	295
TaL	117	6.1 *	104	83	84	80	82
ToL	492	466	456	411	408	384	377
Dorsal scales	19/17/15	19/19/15	19/17/15	19/17/15	19/18/15	19/17/15	19/17/15
Ventral scales	149	143	144	145	140	150	140
Subcaudals	66/66	29/29 *	58/58	54/54	59/59	58/58	56/56
Supralabials [Contacting orbit]	8/8 [4+5]	8/9 [4+5]/[5+6]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]
Infralabials	9/9	9/10	9/9	10/9	9/9	8/8	9/10
Left infralabials with 1 genial	1–5	1–5	1–5	1–4	1–5	1–4	1–4
Left infralabials with 2 genials	5	5	5	4–5	5	4	5
Loreals	1/1	1/1	1/1	1/0	1/1	1/1	2/2
Preoculars	1/1	2/2	1/2	1/1	1/1	2/2	1/1
Postoculars	2/2	2/2	2/2	2/2	2/2	2/2	2/2
Temporals	1+2/1+3	2+2/2+3	2+3/2+2	2+3/2+3	2+3/2+3	2+3/2+2	2+3/2+3
Characters	MUSM 19177	MUSM 18104	MUSM 25403	MUSM 23420	MUSM 23422	MUSM 17839	MUSM 23470
Sex	m	m	m	f	f	f	f
SVL	296	293	272	314	294	291	322
TaL	77	58 *	76	100	76	77	92
ToL	373	351	348	414	370	368	414
Dorsal scales	19/19/15	19/17/15	19/17/15	19/18/15	19/17/15	19/17/15	19/19/15
Ventral scales	149	143	143	139	141	142	142
Subcaudals	60/60	36/36 *	56/55	53/52	56/55	56/54	52/52
Supralabials [Contacting orbit]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]	8/8 [4+5]
Infralabials	9/9	8/8	9/8	9/9	9/9	9/9	9/9
Left infralabials with 1 genial	1–4	1–4	1–5	1–5	1–5	1–5	1–5
Left infralabials with 2 genials	5	4–5	5	5	5	5	5
Loreals	1/1	1/1	1/1	1/1	1/1	1/1	1/1
Preoculars	2/2	1/1	1/1	1/1	2/2	1/1	1/1
Postoculars	2/2	2/2	2/2	2/2	2/2	2/2	2/2
Temporals	2+3/2+3	1+2/1+2	2+3/2+3	2+2/2+2	2+2/2+2	2+3/2+3	2+3/2+3

Appendix C

In addition to the type material of the new species, the following specimens were examined: Tachymenoides affinis: Peru: Huánuco: Ambo: Conchamarca, 2800 m: MUSM 38744, Huánuco: Acomayo, 3000 m: MUSM 2723–25, Huánuco: Conchamarca (Santa Rosa), ca. 2900 m: SMF 80049; Tachymenoides harrisonfordi: Peru: Junín: Otishi National Park, 3248 m: MUSM 40925; Tachymenis peruviana: Peru: Ayacucho: Parinachocas Chumpi, Breapampa: MUSM 26386, Apurimac: Cotabambas, Callhuahuacho, 4144 m; Tachymenis tarmensis: Peru: Junín, Tarma: FMNH 5698 (holotype), Peru: Junín, Concepción, Aco, 3507 m: MUSM 38469, Peru: Junín, Pichjapuquio: MUSM 31533, Peru: Junín, 3 km from Acobamba, 23 km from Tarma: AMNH 117734 (analyzed by digital images); Thamnodynastes pallidus: Suriname, unknown location: NZCS R129; Indira Gandhiweg km 14: NZCS R283.

Appendix D

Table A3. Comparative diagnostic characters of Tachymenoides goodallae sp. nov., Ta. affinis, and Ta. harrisonfordi. Data for Ta. affinis taken from [1]; for T. harrisonfordi taken from [3]. Females for Ta. harrisonfordi are unknown. Coloration is described from real life; measurements are in mm.

Characters	Ta. goodallae sp. nov.		Ta. affinis		Ta. harrisonfordi
Sex	males	females	males	females	males
Sample size	17	4	9	7	1
SVL	272–433	291–322	183–436	162–400	315
TaL	77–126	76–100	44–129	47.5–107	92
Apical pits	absent		absent		absent
Dorsal scales	19/17/15 (n = 13)		17/17/15 (n = 13)		19/17/15
	19/18/15 (n = 3)		19/17/15 (n = 2)
	19/17/14 (n = 1)		17/17/13 (n = 1)
Ventral scales	140–157	139–142	141–153	141–149	139
Subcaudals	52–67	52–56	48–64	52–65	54
Cloacal shield	divided		divided		divided
Hemipenis	hemipenis with large calyces restricted to the capitulum on the asulcate side		hemipenis with large calyces restricted to the capitulum on the asulcate side		unknown
Dorsal skin folds	absent		absent		present
Dorsal coloration	the dorsum is olive brown to pale grayish brown with scattered black and cream flecks, no longitudinal stripes		uniform, with sparse scales bordered in black along the body		pale yellowish brown with scattered black blotches that form a paravertebral stripe (one scale wide) on each side at the posterior half of the body
Ventral coloration	nearly uniformly black, mottled gray and dark gray, mottled pale gray and tan, or pale grayish tan; usually, three irregularly shaped, narrow longitudinal ventral stripes		two to five non-continuous longitudinal ventral stripes in a few specimens, forming a dark band in each ventral scale, with a general darkening tendency towards the tail; some specimens have an overall dark venter, with no discernible stripes		throat and first 60 mm of the ventral body are pale yellowish tan with black flecks, and a ventrolateral black stripe on each side of the same length, the remaining venter and tail entirely black
Iris coloration	brown		unknown		copper

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Figure 1. Maximum likelihood (ML) tree showing relationships of the new species and other Tachymenini genera. Phylogenetic analysis is based on concatenated 12S, cytb, and c-mos gene regions. Bootstrap values are shown on nodes. Species of interest are highlighted in bold and the new species is in red.

Figure 2. Live Tachymenis tarmensis (MUSM 38468) from Tarma, Junín, Peru. Photo by CT.

Figure 3. Map of Peru with the distribution of Tachymenoides goodallae sp. nov. and Tachymenis tarmensis. The pink star is the type locality (Chacos) of Ta. goodallae sp. nov.; the white star is the type locality (Tarma) of T. tarmensis. Pink circles (1–7) are collecting sites of paratypes of Ta. goodallae sp. nov.: 1 = Chacos, type locality (Pasco), 2 = Puagmaray (Pasco), 3 = road between Auquimarca and Uchuhuerta (Pasco), 4 = Auquimarca (Pasco), 5 = Huallamayo (Pasco), 6 = Ingenio (Junín), and 7 = Pacchani (Puno). The distance (airline) from the northernmost (2) to the southernmost (7) collecting site is 782.27 km. White triangles (1–2) are our collecting sites of T. tarmensis: 1: Aco (Junín) and 2: Pichjapuquio (Junín). The type locality (Tarma, white star) of T. tarmensis and the collecting site of Ta. goodallae sp. nov. at Ingenio (pink circle 6) overlap; both localities are separated by 3.5 km airline. Green stars associated with numbers (1, 6; 1, 2) indicate collecting sites for specimens whose tissues were used in the phylogeny (Figure 1).

Figure 7. Comparisons of heads in lateral, dorsal, and ventral views of the holotype of Tachymenis tarmensis (FMNH 5698, (A–C), head length: 17.0 mm), T. tarmensis (MUSM 38469 from Junín, (D–F), head length: 16.7 mm), and paratype Tachymenoides goodallae sp. nov. (MUSM 23470 from Junín, (G–I), head length: 13.6 mm). Photos by CA.

Figure 8. Left hemipenes of the holotype of Tachymenoides goodallae sp. nov (MUSM 31152) in the sulcal (A), left side (B), right side (C), and asulcal side (D). Red arrows point to sulcus spermaticus, blue arrows to the bulge, and green arrow to the first row of nine enlarged calyces. Photos by EL.

Figure 9. Life specimens of Tachymenoides goodallae sp. nov. (A): male (MUSM 23485, total length 534 mm) from Chacos (Pasco); (B): female (MUSM 23470) from Ingenio (Junín); (C): male (MUSM 17813, total length 511 mm) between Auquimarca and Uchuerta (Pasco); (D): female (uncollected, total length 670 mm) from Auquimarca (Pasco); (E): female (uncollected, total length 430 mm) from Auquimarca (Pasco); (F): male (ML 446, total length 410 mm) from Puagmaray (Pasco). Photos by ML.

Figure 10. Variation in ventral coloration of preserved paratypes of Tachymenoides goodallae sp. nov. (A): nearly uniformly black (MUSM 17839), (B): gray and mottled dark gray with three distinct black longitudinal stripes (MUSM 25403), (C): grayish tan with a broad dark-gray, washed out midventral longitudinal band (MUSM 18112), (D): gray and mottled dark gray but the three black longitudinal stripes are not as discernable, except for the midventral stripe (MUSM 25422), (E): pale grayish tan (MUSM 17828), and (F): grayish tan with five narrow longitudinal stripes (MUSM 23420).

Figure 11. The type locality of Tachymenoides goodallae sp. nov. at Chacos. Mountain ridge with the radio antenna (A), forests and scattered lagunas at lower elevations (B), close-up view of the vegetation (C), and basking location of the holotype (D). Photos by EL taken on 15 February 2011.

Figure 12. Female Tachymenoides goodalle sp. nov. (MUSM 23420) in dorsal (A) and ventral views (B) with body opened to expose the six eggs (egg length in mm in the head-to-tail order: 20.6, 18.5, 19.3, 20.1, 21.6, 25.6). The upper scale row is in centimeters. Photos by EL.

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Lehr, E.; Lundberg, M.; Cusi, J.C.; Sites, J.W., Jr.; Torres, C.; Aguilar-Puntriano, C. A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945). Taxonomy 2025, 5, 18. https://doi.org/10.3390/taxonomy5020018

AMA Style

Lehr E, Lundberg M, Cusi JC, Sites JW Jr., Torres C, Aguilar-Puntriano C. A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945). Taxonomy. 2025; 5(2):18. https://doi.org/10.3390/taxonomy5020018

Chicago/Turabian Style

Lehr, Edgar, Mikael Lundberg, Juan Carlos Cusi, Jack W. Sites, Jr., Claudia Torres, and César Aguilar-Puntriano. 2025. "A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945)" Taxonomy 5, no. 2: 18. https://doi.org/10.3390/taxonomy5020018

APA Style

Lehr, E., Lundberg, M., Cusi, J. C., Sites, J. W., Jr., Torres, C., & Aguilar-Puntriano, C. (2025). A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945). Taxonomy, 5(2), 18. https://doi.org/10.3390/taxonomy5020018

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A New Species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from Peru with Comments on the Taxonomic Status of Galvarinus tarmensis (Walker, 1945)^†

Abstract

1. Introduction