Research Article |
Corresponding author: Christoph I. Grünwald ( cgruenwald@switaki.com ) Academic editor: Ben Wielstra
© 2023 Christoph I. Grünwald, Carlos Montaño-Ruvalcaba, Jason M. Jones, Iván Ahumada-Carrillo, André J. Grünwald, Jiacheng Zheng, Jason L. Strickland, Jacobo Reyes-Velasco.
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:
Grünwald CI, Montaño-Ruvalcaba C, Jones JM, Ahumada-Carrillo I, Grünwald AJ, Zheng J, Strickland JL, Reyes-Velasco J (2023) A novel species of piping frog Eleutherodactylus (Anura, Eleutherodactylidae) from southern Mexico. Herpetozoa 36: 95-111. https://doi.org/10.3897/herpetozoa.36.e104707
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We describe a new species of Eleutherodactylus (subgenus Syrrhophus) from Guerrero, Mexico, based on morphological and molecular data, as well as advertisement call analysis. Eleutherodactylus franzi sp. nov. has unique features including widely expanded fingertips, indistinct, but visible lumbo-inguinal glands, an immaculate white venter and dark reticulations on a cream dorsal background colouration. The new species belongs to the Eleutherodactylus nitidus species group. Eleutherodactylus franzi sp. nov. is micro-endemic, restricted to a small range in karstic hillsides on the southern extreme of the Mexican Transverse Ranges. We discuss conservation needs of this species, which we provisionally classify as Critically Endangered.
Describimos una especie nueva de Eleutherodactylus (subgénero Syrrhophus) de Guerrero, México basado en datos morfológicos y moleculares, así como análisis de llamadas de apareamiento. Eleutherodactylus franzi sp. nov. tiene características únicas, incluyendo puntas de los dedos ampliamente expandidas, glándulas lumbo-inguinales indistintas pero visibles, un vientre blanco inmaculado y una coloración dorsal de reticulaciones oscuras sobre un fondo crema. La especie nueva pertenece al grupo de especies de Eleutherodactylus nitidus. Eleutherodactylu. franzi sp. nov. es micro-endémica, restringida a un pequeño rango en laderas kársticas en el extremo sur del Eje Neovolcánico. Presentamos mapas de distribución y gráficos de llamadas de apareamiento de los machos de la nueva especie y sus parientes más cercanos, así como recomendaciones de conservación.
Wir beschreiben eine neue Art von Eleutherodactylus (Untergattung Syrrhophus) aus Südmexiko, auf der Grundlage morphologischer und molekularer Daten sowie einer Analyse des Anzeigerufs. Eleutherodactylus franzi sp. nov. ist einzigartig in der Gattung durch eine Kombination von Merkmalen, darunter weit verbreiterte Fingerspitzen, undeutliche, aber sichtbare lumbo-inguinale Drüsen, ein makellos weißer Bauch und eine einzigartige Rückenfärbung aus dunklen Netzen auf einem cremefarbenen Hintergrund.. Die neue Art gehört zur Eleutherodactylus nitidus Artengruppe. Eleutherodactylus franzi sp. nov. ist mikroendemisch und auf ein kleines Verbreitungsgebiet in Karsthängen am südlichen Ende der Mexikanischen Querketten beschränkt. Wir präsentieren eine Verbreitungskarte der neuen Art, analysieren ihren Anzeigeruf und geben Empfehlungen für ihren Schutz.
amphibians, Anura, conservation, Guerrero, Mexican Transverse Ranges, taxonomy
anfibios, Anura, conservación, Eje Neovólcanico, Guerrero, taxonomía
Amphibien, Anura, Eje Neovólcanico, Guerrero, Naturschutz, Taxonomie
Frogs of the genus Eleutherodactylus Duméril & Bibron, 1841 from continental North America are amongst the least understood and most taxonomically challenging groups of New World anurans (
Recent studies have expanded the diversity within the E. nitidus species group.
The present study provides evidence, based on molecular, morphological and mating call data, for the occurrence of an additional undescribed species of Syrrhophus in the State of Guerrero, in southern Mexico. Our molecular phylogenetic analysis indicates that the species belongs to the Eleutherodactylus nitidus species group and is most closely related to E. humboldti. We analyse the male advertisement call of the new species, compare it to closely-related species and discuss its limited distribution. Finally, we make conservation recommendations.
We examined specimens of all currently recognised species of the subgenus Syrrhophus (Frost, 2020) and measured specimens of all species, except for the enigmatic E. verruculatus (Peters, 1870), whose existence has been questioned by several authors (
We photographed all specimens used in this study alive, including dorsal, lateral and ventral profiles, as well as photographs of each showing colours of flanks and flash colours on the groin and thigh. We then euthanised the frogs with 10% ethanol or with topical benzocaine and took tissue samples from the thigh muscle or liver upon death and preserved them in 96% ethanol. We preserved specimens in 10% formalin and transferred them to 70% ethanol for storage. We measured additional specimens of the subgenus Syrrhophus in the Museo de Zoología, Facultad de Ciencias (MZFC) of the Universidad Nacional Autónoma de México (
We did not measure type specimens of some previously-described taxa so we used the measurements of the type specimens provided in their original descriptions and published literature. Measurements of Eleutherodactylus dilatus (Davis & Dixon, 1955), E. maurus (Davis & Dixon, 1955) (= E. fuscus), E. albolabris (Taylor, 1943) are given in their original descriptions and in
The material collected was deposited at the Instituto de Investigaciones sobre los Recursos Naturales (INIRENA), which is now officially known as Colección Herpetológica de la Universidad Michoacana (CHUM) of the Universidad Michoacana de San Nicolás de Hidalgo in Morelia, Michoacán, Mexico; at the Museo de Zoología, Facultad de Ciencias (MZFC) of the Universidad Nacional Autónoma de México (
The characters and terminology we use herein follow those of
Measurements were made with Truper (Mexico) brand digital calipers and rounded to the nearest 0.1 mm. The sex of adult specimens was determined by presence of vocal slits.
DNA extraction and PCR amplification
A detailed description of the DNA extraction and PCR amplification protocols can be found in
Sequence alignment and phylogenetic analysis
We included additional sequences of multiple members of the subgenus Syrrhophus obtained from GenBank to infer the phylogenetic relationships of the new individuals sequenced in this study. We have included all sequences used in this study with their accession numbers in GenBank in Appendix
We removed regions with poor-quality base calls by manually trimming the 5’ and 3’ ends of all sequences using the programme Geneious v.6.1.6 (Biomatters Ltd., Auckland, NZ). We then aligned all sequences in Muscle (
We performed Bayesian Inference of phylogeny (BI) in MrBayes v.3.2.2 (
We recorded vocalisations of several individuals of the new species described here, as well as all other members of the Eleutherodactylus (Syrrhophus) nitidus species group (sensu
We isolated the individual calls from other calls and background noise using Adobe Audition CC, using default settings in the application. We then analysed the calls using the software Raven Pro 1.5 (The Cornell Lab of Ornithology 2014). Spectrograms were constructed using a Blackman-type window with a size of 5 ms, 80% overlap and DFT of 512 samples. Temporal parameters were measured from the oscillogram in ms. Spectrogram and oscillogram graphics were generated using Seewave v.1.6 (Sueur et al. 2008) in RStudio v.1.1.423 (RStudio Team 2016). Values in the call descriptions are given as mean ± standard deviation. 2D spectrograms were visualised using a sliding window analysis of short-term Fourier transform calculations.
Our phylogenetic results (Fig.
Bayesian phylogenetic inference of members of the Eleutherodactylus subgenus Syrrhophus, with a focus on the E. nitidus species group, based on the mitochondrial locus 16S rRNA. Black circles represent nodes with a posterior support of 1. All nodes with support of less than 0.5 are collapsed.
Holotype. INIRENA 2900 (CIG 01725). Adult male (Fig.
Some of the paratypes of Eleutherodactylus franzi sp. nov., in life. A–C. INIRENA 2902 (CIG 01717) El Cucharillo, Municipio de Ixcateopan de Cuauhtémoc, Guerrero, Mexico; D–F. INIRENA 2910 (CIG 01726) El Cucharillo, Municipio de Ixcateopan de Cuauhtémoc, Guerrero, Mexico; G–I. INIRENA 2895 (CIG 01727), 3 km E of Ixcateopan de Cuauhtémoc, Municipio de Ixcateopan de Cuauhtémoc, Guerrero, Mexico; J–L. INIRENA 2896 (CIG 01728), 3 km E of Ixcateopan de Cuauhtémoc, Municipio de Ixcateopan de Cuauhtémoc, Guerrero, Mexico; M–O. INIRENA 2898 (CIG 01731), between El Cucharillo and Chichila, Municipio de Taxco de Alarcón, Guerrero, Mexico.
Based on our phylogenetic analysis, this is a member of the genus Eleutherodactylus, subgenus Syrrhophus, as defined by
Eleutherodactylus franzi can be distinguished from all species in the Eleutherodactylus (Syrrhophus) longipes species series by: possessing a small, indistinct tympanum with no tympanic annulus visible and with a diameter less than 50% of the diameter of the eye; by possessing a ventral epidermis which is semi-translucent and combined with a visceral peritoneum which is not white, an abdominal vein on the venter is not clearly evident against a white background in life; by possessing indistinct, but visible raised lumbo-inguinal gland above the inguinal region.
Eleutherodactylus franzi can be distinguished from most species of the Eleutherodactylus (Syrrhophus) modestus species group by the combination of possessing a compact, protruding lumbo-inguinal gland above the inguinal region, digital tips which are expanded more than 1.8 times the width of the narrowest part the finger on the third and fourth fingers and the lack of a distinct interorbital bar a colour distinct from the dorsal ground colouration. It can further be distinguished from the superficially similar E. grunwaldi by its smaller body size, 25.6–29.5 mm (vs. 28.4–32.4 mm), less expanded fingertips, 1.8–2.5 times the width of the narrowest part of the finger on fingers three and four (vs. 2.8–3.2) and the presence of a visible raised lumbo-inguinal gland. From the superficially very similar E. saxatilis, it can be distinguished by snout shape, eye size and head colouration. These two frogs, although not closely related, are very similar in appearance, but come from two widely-separated mountain ranges in central Mexico and are genetically distinct. Eleutherodactylus franzi has an angular canthus rostralis, with a shorter snout that is distinctly truncated from a lateral profile. It has a larger eye, with a larger ETD and, generally, there is a pale interorbital region that lacks dark markings. In E. saxatilis, the canthus rostralis is noticeably rounded and the snout is acuminate from a lateral profile. The eyes are smaller and located closer to the tympanum and there is no noticeable lack of dark dorsal markings in the interorbital area.
Within its own species group, E. franzi can be distinguished from most species by possessing a compact inguinal gland that is indistinct, but visible in live specimens. This character may or may not be visible in preserved specimens depending on how they were preserved. This species differs from E. pipilans, E. erythrochomus and E. nebulosus, which lack visible compact lumbo-inguinal glands altogether. All other known species in the E. (Syrrhophus) nitidus species group have readily visible compact lumbo-inguinal glands above the inguinal region, except E. maculabialis, which has similarly visible, but indistinct lumbo-inguinal glands. Eleutherodactylus franzi can be further distinguished from E. pipilans and E. nebulosus by possessing digital tips which are expanded more than 1.8 times the width of the narrowest part of the finger and from E. erythrochomus by possessing digital tips which are more than 1.5 times, but less than 3.0 times, the width of the narrowest part of the finger. It is distinguished from E. albolabris, E. nitidus, E. petersi, E. jamesdixoni and E. orarius by the combination of larger size, smoother skin, longer limbs and tips of digits which are expanded more than 1.8 times the narrowest part of the finger on the third and fourth fingers. This species is distinguished from E. dilatus, E. humboldti, E. maurus and E. sentinelus by its larger size, smoother skin, pale dorsal colouration with dark reticulations and lack of a pale interorbital bar which is paler than the pale dorsal colouration. Furthermore, all these species, except E. humboldti, present multi-note whistles, while E. franzi has a call that consists of a single note, low-pitched pipe. Eleutherodactylus franzi may be distinguished from E. maculabialis and E. syristes by its larger size, more expanded fingertips, lack of inguinal flash colouration, as well as those species’ unique advertisement call which consists of a trill rather than a short pipe. In Guerrero, two other species of saxicolous Eleutherodactylus (E. pipilans and E. erythrochomus) have similar colouration, similar smooth skin and widely expanded digital pads. Both are readily distinguishable from E. franzi by possessing an indistinct lumbo-inguinal gland which is barely visible in life and by a male advertisement call which is a peep instead of a pipe. Furthermore, E. franzi can be distinguished from E. pipilans by its more expanded digital tips on the third and fourth fingers, 1.8–2.5 times the width of the narrowest part of the finger (vs. 1.3–1.9) and distinct lack of dark markings in the interorbital region (vs. no difference from rest of dorsum). It can further be distinguished from E. erythrochomus by the conspicuous dark pattern on a pale dorsal colouration, lesser expanded digital tips no more than 2.5 times the width of the narrowest part of the digit on the third and fourth fingers (vs. 2.3–3.8) and a less distinct tympanum. General characteristics for the Eleutherodactylus nitidus species group are given in Table
Key Comparative Characters of the Eleutherodactylus nitidus species group.
E. albolabris | E. dilatus | E. erythrochomus | E. franzi sp. nov. | E. humboldti | E. jamesdixoni | E. maculabialis | E. maurus | E. nebulosus | E. nitidus | E. orarius | E. petersi | E. pipilans | E. rubrimaculatus | E. sentinelus | E. syristes | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Size | Medium | Medium | Large | Large | Medium | Medium | Small | Small | Medium | Medium | Medium | Medium | Large | Small | Medium | Small |
SVL adult males (range) in mm | 23.0–26.8 | 23.8–25.7 | 24.9–30.0 | 25.6–29.5 | 23.4–25.7 | 24.3–26.3 | 17.9–24.7 | 20.7–24.3 | 22.9–28.3 | 24.3–26.3 | 24.6–28.0 | 23.9–26.3 | 25.5–29.6 | 18.2–23.5 | 23.3–25.3 | 21.7–24.6 |
Condition of Interorbital Bar | Indistinct, Pale | Present, Pale | Absent | Absent | Usually Present, Pale |
Indistinct, Pale | Present, Pale | Present, Pale | Absent | Indistinct, Pale | Indistinct, Pale | Indistinct, Pale | Absent | Absent | Present, Pale | Present, Pale |
Pale Mid-dorsal Blotch | Sometime Present | Absent | Absent | Absent | Present | Present | Absent | Absent | Absent | Present | Sometimes Present | Present | Absent | Absent | Present | Absent |
Colouration of Lip | White | Dark with pale flecking | As head | Grey, with flecking of dorsal colouration | Pale, with white flecking, | Mottled | Dark with pale spots | Dark with pale flecking | Dark with pale spots | Mottled | White, mottled with dark | Mottled | As head | Dark with white or pale spots | Dark with pale flecking | Dark, variedly spotted |
Inguinal Flash Colouration | Orange | Yellow | Absent | Absent | Yellow or orange | Faint, yellow-orange | Varied, Yellow / Orange | Varied, usually Absent | Absent | Faint, yellow-orange | Faint, yellow-orange | Faint, yellow-orange | Absent | Absent | Yellow | Varied, Yellow / Orange |
Ventral Colouration | White with black spots | Grey with white and black | Transparent | White or pinkish | Grey, with white and black | White with dark mottling | Grey with white and black | Grey with white and black | Transparent | White with dark mottling | White with dark mottling | White with dark mottling | Transparent | Pale grey with white spotting | Transparent with white and black | Transparent and White, with black spots |
Dorsal Skin Texture | Smooth | Not Smooth | Smooth | Smooth | Not Smooth | Not Smooth | Smooth | Not Smooth | Smooth | Not Smooth | Not Smooth | Not Smooth | Smooth | Smooth | Smooth | Smooth |
Ventral Skin Texture | Slightly Rugose | Rugose | Smooth | Smooth | Rugose | Slightly Rugose | Smooth | Rugose | Smooth | Slightly Rugose | Slightly Rugose | Slightly Rugose | Smooth | Smooth | Smooth | Smooth |
Condition of Lumbo-Inguinal Gland | Very distinct | Very distinct | Indistinct | Distinct | Very distinct | Very distinct | Distinct | Very distinct | Indistinct | Very distinct | Very distinct | Very distinct | Indistinct | Indistinct | Very distinct | Distinct |
3FPW/3FW | 1.3–1.9 | 1.5–1.7 | 2.3–3.8 | 1.8–2.5 | 1.9–2.4 | 1.1–1.5 | 1.4–2.1 | 1.3–1.6 | 1.1–1.5 | 1.1–1.5 | 1.2–1.4 | 1.1–1.7 | 1.5–1.9 | 1.5–1.9 | 1.7–2.3 | 1.1–1.9 |
4FPW/4FW | 1.3–1.9 | 1.5–1.8 | 2.3–3.5– | 1.8–2.5 | 1.3–2.1 | 1.1–1.5 | 1.4–2.1 | 1.3–1.7 | 1.1–1.5 | 1.1–1.5 | 1.2–1.4 | 1.1–1.7 | 1.5–2.0 | 1.5–1.9 | 1.5–2.3 | 1.2–1.8 |
TW/ED | 0.27–0.32 | 0.25–0.35 | 0.33–0.51 | 0.40–0.61 | 0.37–0.52 | 0.25–0.29 | 0.25–0.28 | 0.28–0.31 | 0.35–0.38 | 0.25–0.29 | 0.25–0.29 | 0.25–0.29 | 0.30–0.36 | 0.25–0.36 | 0.26–0.28 | 0.25–0.29 |
Call | Whistle | Peep | Peep | Pipe | Pipe | Whistle | Trill | Pipe | Peep | Whistle | Whistle | Whistle | Peep | Peep | Peep | Trill |
Adult male, relatively large (26.2 mm SVL); head as wide (9.5 mm) as long (9.5 mm), head wider than body; snout rounded from a dorsal view and rounded to slightly truncate from a lateral profile; tympanum indistinct, rounded with no supra-tympanic fold present; tympanum small, oval, greatest width of tympanum 1.4 mm; greatest diameter of eye 2.8 mm; tympanum width to eye-diameter 0.51; eyelid width 1.6 mm, approximately 38% of the IOD; first finger shorter than second finger; finger lengths from shortest to longest I-II-IV-III; digital pads on fingers two, three and four expanded, 2.1 times the narrowest point of the digit on fingers three and four; expanded finger pads widely expanded, truncate, three palmar tubercles; inner palmar tubercle 70% of middle palmar tubercle and outer palmar tubercles about 60% as large as middle palmar tubercle, (Fig.
A. Ventral aspect of the hand of the holotype of Eleutherodactylus franzi sp. nov., INIRENA 2900 (CIG 01725) from El Cucharillo, Municipio de Ixcateopan de Cuauhtémoc, Guerrero, Mexico; B. Ventral aspect of the hand of a Eleutherodactylus humboldti, INIRENA 2911 (CIG 01703) from 9.4 km N of Valle de Bravo junction on Valle de Bravo – Toluca toll road, near San Bartolo, Municipio de Amanalco, Estado de México, Mexico.
In life, the holotype had a yellowish-tan dorsal colouration on the back, with darker brown blotches on the back and flanks. Head yellowish-tan with some dark brown speckling, no pale interorbital bar; however, the interorbital region mostly lacks darker brown speckling giving it the resemblance of a pale interorbital bar. Labial region pale grey with some tan and some white speckling. Three and four white-tipped tubercles present at the rictal. A dark brown stripe present from the tip of the snout posteriorly through loreal region, eye and tympanum to right above rictal tubercles. Forearms, thighs, femur and tarsus tan with indistinct pale brown banding. The upper arms were unmarked, tan to slightly orange. No inguinal flash colouration was present on groin or thighs. Ventral colouration was lavender with some sparse white and on sides, ventral colouration on throat grey. Ventral skin was slightly translucent and visceral peritoneum clear, so no visible red abdominal vein and viscera were visible in life.
Colouration in preservative is pale tan on dorsum, with darker brown reticulations. Pale tan interorbital bar present. Dark canthal bar is dark brown. Unmarked upper arms are cream to white. Limbs cream with dark brown cross-banding. The dorsal surfaces of the legs are light brown and the groin and posterior surfaces of the thighs are brown. Ventral surfaces yellowish-cream, unmarked, slightly darker brownish pigmentation on throat and chin. Ventral surfaces of hands and feet brown, with dark brown spots. (Fig.
IND 2.2, IOD 4.2, END 2.5, ETD 0.9, UpL 6.4, FoL 7.5, PaL 2.3, HaL 6.7, F1L 2.3, F1PW 0.5, F1W 0.4, F2L 2.8, F2PW 0.8, F2W 0.4, F3L 4.7, F3PW 1.1, F3W 0.5, F4L 3.5, F4PW 1.1, F4W 0.5, IPTL 0.7, MPTL 1.0, OPTL 0.6, FeL 10.0, TL 11.0, TaL 7.1, TotFL 11.4, T1L 2.4, T1PW 0.62, T1W 0.5, T2L 3.6, T2PW 0.7, T2W 0.5, T3L 4.4, T3PW 0.7, T3W 0.5, T4L 6.0, T4PW 0.7, T4W 0.5, T5L 2.5, T5PW 0.6, T5W 0.4, IMTL 1.1, OMTL 0.6, FeL/SVL 38%, TL/SVL 42%, HaL/SVL 26%, TotFL/SVL 56%, HL/SVL 35%, HW/SVL 36%.
SVL from 25.6–29.5 mm (27.15 ± 1.77). Expanded finger pads vary from 1.8–2.5 times the narrowest part of the digit on the third finger and from 1.8–2.6 times the narrowest part of the digit on the fourth finger, with average 2.0 ± 0.21 on the third finger and average 2.2 ± 0.21 on the fourth finger. Dorsal ground colouration cream or tan, but varied with some greenish, reddish or yellowish tinge, always with darker brown blotches or reticulations. The extent of the darker brown blotches or reticulation varied greatly. The condition of the interorbital area ranged from unmarked and same colour as ground colouration to heavily marked by dark speckling same colour as dark dorsal blotches or reticulation. Venter always lavender, but with varying amounts of white spots. Morphological variation of E. franzi is presented in Table
Eleutherodactylus franzi sp. nov. Morphological Measurements (in millimetres).
INIRENA 2901 | INIRENA 2902 | INIRENA 2903 | INIRENA 2904 | INIRENA 2905 | INIRENA 2906 | INIRENA 2907 | INIRENA 2908 | INIRENA 2909 | INIRENA 2900 | INIRENA 2910 | INIRENA 2895 | INIRENA 2896 | INIRENA 2897 | INIRENA 2898 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SVL | 27.72 | 25.58 | 27.72 | 26.86 | 26.4 | 26.04 | 28.62 | 26.68 | 25.55 | 26.24 | 27.97 | 28.7 | 27.12 | 29.51 | 26.56 |
HL | 9.08 | 8.76 | 9.35 | 9.1 | 8.61 | 8.33 | 8.76 | 8.73 | 8.68 | 9.12 | 9.38 | 9.06 | 8.79 | 9.04 | 8.61 |
HW | 9.78 | 9.03 | 9.81 | 9.78 | 9.82 | 9.07 | 9.4 | 9.41 | 9.42 | 9.5 | 9.99 | 9.68 | 9.67 | 9.66 | 9.61 |
TW | 1.53 | 1.36 | 1.52 | 1.88 | 1.74 | 1.45 | 1.38 | 1.47 | 1.32 | 1.4 | 1.39 | 1.12 | 1.39 | 1.34 | 1.42 |
ED | 3.74 | 2.62 | 2.79 | 3.08 | 2.9 | 2.63 | 2.74 | 2.78 | 2.81 | 2.76 | 2.77 | 2.78 | 2.38 | 2.8 | 2.94 |
ElW | 1.13 | 1.39 | 1.52 | 1.68 | 1.19 | 1.45 | 1.49 | 1.67 | 1.69 | 1.59 | 1.35 | 1.78 | 1.51 | 1.62 | 1.9 |
IOD | 5.1 | 5.02 | 5.19 | 5.09 | 5.04 | 4.32 | 5.06 | 5.09 | 5.05 | 4.24 | 4.4 | 5.09 | 5.14 | 5.08 | 5.37 |
IND | 2.25 | 2.29 | 2.42 | 2.31 | 2.28 | 2.16 | 2.32 | 2.38 | 2.12 | 2.17 | 2.53 | 2.6 | 2.37 | 2.21 | 2.3 |
END | 3.15 | 2.61 | 2.51 | 2.53 | 2.7 | 2.77 | 2.78 | 2.96 | 2.79 | 2.47 | 2.61 | 2.69 | 2.33 | 2.55 | 2.37 |
ETD | 0.96 | 0.92 | 0.98 | 1 | 0.86 | 1.02 | 0.82 | 0.85 | 0.89 | 0.89 | 0.76 | 0.84 | 0.76 | 0.95 | 0.98 |
UpL | 7.1 | 6.19 | 6.31 | 6.02 | 6.5 | 6 | 6.08 | 6.62 | 6.31 | 6.43 | 6.39 | 6.31 | 6.39 | 6.88 | 6.58 |
Fol | 7.86 | 7.39 | 7.78 | 7.49 | 7.28 | 7.28 | 7.32 | 7.58 | 7.56 | 7.51 | 7.82 | 7.51 | 7.58 | 7.92 | 7.62 |
HaL | 7.57 | 6.82 | 7.3 | 6.41 | 6.23 | 6.6 | 6.72 | 6.28 | 6.46 | 6.86 | 6.93 | 6.7 | 6.58 | 7.24 | 7.53 |
F3PW/F3W | 1.78 | 1.85 | 1.94 | 2.33 | 2.23 | 1.77 | 1.97 | 2.13 | 2.08 | 2.09 | 1.75 | 2.43 | 1.87 | 2.12 | 2.53 |
F4PW/F4W | 1.77 | 2.34 | 2.15 | 2.38 | 2.10 | 1.84 | 1.94 | 2.12 | 2.17 | 2.13 | 2.42 | 2.44 | 2.46 | 1.97 | 2.64 |
FeL | 10 | 9.4 | 9.11 | 9.24 | 9.19 | 9.87 | 9.57 | 9 | 9.32 | 9.95 | 9.81 | 9.82 | 9.34 | 9.22 | 9.9 |
TL | 12.78 | 11.79 | 11.88 | 11.82 | 11.46 | 11.5 | 11.42 | 12.41 | 12.69 | 11.05 | 11.81 | 11.47 | 11.21 | 12.39 | 11.34 |
TaL | 7.41 | 7.47 | 7.36 | 7.51 | 7.31 | 7.18 | 7.93 | 6.84 | 7.92 | 7.14 | 7.42 | 7.59 | 7.42 | 7 | 7.83 |
FL | 4.49 | 4.33 | 4.97 | 4.31 | 3.9 | 4.22 | 4.49 | 4.12 | 4.21 | 4.58 | 4.82 | 4.35 | 4.25 | 4.58 | 4.75 |
3TL | 3.72 | 3.74 | 3.64 | 3.59 | 3.18 | 3.16 | 3.44 | 3.18 | 3.17 | 3.53 | 3.42 | 3.34 | 3.48 | 3.38 | 3.71 |
TotFL | 11.9 | 11.37 | 11.08 | 11.41 | 11.15 | 10.82 | 12.44 | 11.16 | 11.44 | 11.42 | 11.47 | 11.31 | 11.42 | 11.96 | 12.46 |
IPT | 0.68 | 0.89 | 0.86 | 0.88 | 0.86 | 0.64 | 0.92 | 0.91 | 0.86 | 0.68 | 0.62 | 0.94 | 0.8 | 0.8 | 0.78 |
MPT | 1.09 | 1.19 | 1.1 | 1.12 | 1.16 | 1.18 | 1.17 | 1.06 | 1.14 | 1 | 1.06 | 1.02 | 1.08 | 1.18 | 1.26 |
OPT | 0.52 | 0.56 | 0.58 | 0.54 | 0.62 | 0.58 | 0.76 | 0.66 | 0.68 | 0.58 | 0.52 | 0.6 | 0.71 | 0.76 | 0.64 |
IMTL | 1.02 | 1.2 | 1.36 | 1.2 | 1.14 | 1.01 | 1.12 | 1.26 | 1.18 | 1.1 | 1.16 | 1.15 | 1.16 | 1.13 | 1.1 |
OMTL | 0.72 | 0.77 | 0.6 | 0.78 | 0.74 | 0.92 | 0.75 | 0.68 | 0.74 | 0.62 | 0.78 | 0.65 | 0.91 | 0.81 | 0.98 |
TW/ED | 0.41 | 0.52 | 0.54 | 0.61 | 0.60 | 0.55 | 0.50 | 0.53 | 0.47 | 0.51 | 0.50 | 0.40 | 0.58 | 0.48 | 0.48 |
F3PW | 1.14 | 1.26 | 1.28 | 1.26 | 1.36 | 1.12 | 1.42 | 1.32 | 1.25 | 1.13 | 1.1 | 1.36 | 1.12 | 1.4 | 1.62 |
F3W | 0.64 | 0.68 | 0.66 | 0.54 | 0.61 | 0.67 | 0.72 | 0.62 | 0.6 | 0.54 | 0.63 | 0.56 | 0.6 | 0.66 | 0.64 |
F4PW | 1.1 | 1.36 | 1.33 | 1.31 | 1.3 | 1.14 | 1.4 | 1.44 | 1.41 | 1.13 | 1.26 | 1.34 | 1.23 | 1.38 | 1.48 |
F4W | 0.62 | 0.58 | 0.62 | 0.55 | 0.62 | 0.62 | 0.72 | 0.68 | 0.65 | 0.53 | 0.52 | 0.55 | 0.5 | 0.7 | 0.56 |
The advertisement call of the males of this species consists of a single, short, low-pitched pipe that lasts about 106 ms and has a dominant frequency of 2612.7 ± 40.6 Hz (Fig.
Advertisement call data of Eleutherodactylus franzi sp. nov. and the sympatric Eleutherodactylus humboldti.
Eleutherodactylus franzi sp. nov. | Eleutherodactylus humboldti | |
---|---|---|
Individuals | 4 | 3 |
Call type | Pipe | Pipe |
Dominant frequency (kHz) | 2.61 ± 0.04 | 2.89 ± 0.23 |
Call length (ms) | 106.0 ± 5.2 | 256.8 ± 12.8 |
Call rate (/m) | 6.88 ± 0 | 2.16 ± 1.73 |
Call rise time (ms) | 63.64 ± 4.06 | 90.82 ± 45.37 |
Pulse Rate | – | 2.05 ± 0.62 |
Call Interval | 13.03 ± 3.35 | 39.75 ± 14.77 |
Eleutherodactylus franzi appears to be endemic to the Sierra de Taxco Region of northern Guerrero (Fig.
Map showing the type localities and distribution of Eleutherodactylus which are either closely related or superficially similar to the species described herein from southern Mexico. White diamond represents the type locality of E. franzi sp. nov. and white circle represents additional locality.
This species is named after Héctor Franz-Chávez, Mexican herpetologist and avid field collector who collected the type material and who helped collect an extensive sampling of the Eleutherodactylus specimens to be used in our succession of studies.
In recent years, the subgenus Syrrhophus has received increasing attention from researchers. With the description of the new species here, the number of the taxa in the subgenus Syrrhophus increases to 44, with two in Cuba and 42 in mainland North America. Guerrero remains the most diverse State for the subgenus with 11 species occurring within its borders. The assignation of the novel species to the E. nitidus species group raises the number of species in that group to 16 (
One of the most pressing questions regarding the taxonomy of these frogs is whether the subgenus Syrrhophus should be considered a genus. The genus Eleutherodactylus currently includes over 200 species distributed throughout North and Central America, including the Caribbean (
The species described herein has a very limited range, known only from near the type locality and restricted to one biogeographical formation as defined by Grünwald et al. (2015), namely the Central Mexican Transverse Range Pine-Oak Woodland (#44 on p. 409 in Grünwald et al. (2015).
Eleutherodactylus franzi is known from a karstic mountain range that has about 75 km2 of habitat at elevations sufficiently high enough to support populations. It has not been collected anywhere else and may be restricted to the immediate vicinity of the type locality (Fig.
The frog subgenus Syrrhophus is the fastest-growing group of frogs in Mexico, in terms of the number of new taxa being described. Despite the recently augmented interest in the group, we believe that the current diversity may still be underestimated and might continue to increase in the coming years. Apart from collecting in regions currently under-sampled, like the mountains of Guerrero and Oaxaca, we suggest that it is important to re-examine specimens already housed in herpetological collections, as there are multiple errors in the identification of specimens of Eleutherodactylus and un-identified new taxa may be hiding in collections.
Additionally, we believe that it is of great importance that future molecular studies include additional markers, as this allows for a much better understanding of the relationships within this group of frogs. Finally, our data suggest that it is time to revise the status of the subgenus Syrrhophus and to determine if it should be elevated to generic level.
The authors have declared that no competing interests exist.
First and foremost, we thank our field crew for their consistent and impressive efforts in the field, even under adverse conditions. We are extremely indebted to Raquel Austria-Hernández for her extensive help with the call analysis in this paper. We are equally indebted to María de los Ángeles Palma-Irizarry of the Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT) for providing collecting permits (SGPA/DGVS/002288/18). We thank Dr. Ireri Suazo-Ortuño and Jonatan Torres-Pérez-Coeto from the Colección Herpetológica de la Universidad Michoacana for receiving and cataloguing the type material. Biencom Real Estate, Biodiversa A.C. and Herp.mx A.C. provided important funding for the fieldwork. We want to thank the community of Ixcateopan de Cuauhtémoc, who allowed us to collect frogs on their communal lands despite having shut down the entrances to their town due to the Covid-19 pandemic. We thank our children Ámbar Lanomy, Ximena and Emilio for lending their parents time to conduct field research. Finally, we thank Dr. Ben Wielstra and two anonymous reviewers for their valuable revisions of this manuscript.
Specimens examined
Eleutherodactylus albolabris (n = 20): MEXICO: Guerrero: MZFC 33025–33030 (CIG 00327–00332), 33082–33085 (CIG 00390– 00393), MZFC 33108–33109 (CIG 00441–00442), MZFC 33230 (CIG 00668), MZFC 33300–33301 (CIG 00903–00904), MZFC 33323 (CIG 00953), MZFC 33325–33326 (CIG 00955–00956), JAC 25586, 25642.
Eleutherodactylus. angustidigitorum (n = 20): MEXICO: Jalisco: MZFC 33127–33130 (CIG 00476–00479), MZFC 33224–33225 (CIG 00662–00663), MZFC 33386–33388 (CIG 00991–00993), JAC 24912; Michoacán: MZFC 33015–33017 (CIG 00316– 00318), MZFC 33065–33070 (CIG 00373–00378), JAC 26977.
Eleutherodactylus campi (n = 13): MEXICO: Nuevo León: MZFC 33195–33198 (CIG 00606–00609); UNITED STATES: Texas: JHM 1390–1394.
Eleutherodactylus colimotl (n = 20): MEXICO: Colima: MZFC 29282 (CIG 00468), MZFC 33115–33120 (CIG 00462–00467), MZFC 33237–33239 (CIG 00682–00684), MZFC 33299 (CIG 00901), MZFC 33329–3330 (CIG 00960–00961), JAC 30498– 30499, 30631; Michoacán: MZFC 33036 (CIG 00340), JAC 23999–24001.
Eleutherodactylus cystignathoides (n = 6): MEXICO: Veracruz: MZFC 33351–33353 (CIG 01163–01165), MZFC 33354 (CIG 01170), JAC 30000–30001.
Eleutherodactylus dennisi
(n = 13): MEXICO: Tamaulipas: MZFC 33255–33261 (CIG 00822–00828),
Eleutherodactylus dilatus
(n = 19): MEXICO: Guerrero: MZFC 33089–33094 (CIG 00405–00410), MZFC 33097 (CIG 00428), MZFC 33231 (CIG 00669),
Eleutherodactylus erendirae (n = 25): MEXICO: Jalisco: MZFC 33000–33008 (CIG 00300–00309), MZFC 33226–33229 (CIG 00664–00667), MZFC 33232 (CIG 00673), MZFC 33234–33235 (CIG 00679–00681); Michoacán: MZFC 29274, 33019– 33024 (CIG 00319–00325).
Eleutherodactylus erythrochomus (n = 2): MEXICO: Guerrero: INIRENA 2923–24 (CIG 01922–01923).
Eleutherodactylus franzi sp. nov. (n = 15): MEXICO: Guerrero: INIRENA 2895–98, 2900–2910 (CIG 01716–01729, 01731).
Eleutherodactylus floresvillelai (n = 12): MEXICO: Michoacán: MZFC 33053–33064 (CIG 00361–00372).
Eleutherodactylus grandis
(n = 1): MEXICO: Ciudad de Mexico:
Eleutherodactylus grunwaldi (n = 12): MEXICO: Colima: MZFC 27467–27475, MZFC 27484, MZFC 33298 (CIG 00898); JRV 00230.
Eleutherodactylus guttilatus (n = 10): MEXICO: Guanajuato: MZFC 33367–33369 (CIG 01248–01250); San Luis Potosí: MZFC 33200–33206 (CIG 00619–00625).
Eleutherodactylus humboldti (n = 17): MEXICO: Estado de México: INIRENA 2911–2916 (CIG 01703, 01702, 01704–01710); Guerrero: INIRENA 2899, 2920–2922 (, 01730, CIG 01713–01715); CIG 00962–00965.
Eleutherodactylus interorbitalis (n = 7): MEXICO: Sinaloa: MZFC 33186–33187 (CIG 00584–00585), MZFC 33190–33194 (CIG 00600–00604).
Eleutherodactylus jaliscoensis (n = 15): MEXICO: Jalisco: MZFC 33131–33141 (CIG 00480–00490), MZFC 33274–33276 (CIG 00861–00863), MZFC 33280 (CIG 00876).
Eleutherodactylus jamesdixoni (n = 14): MEXICO: Jalisco: MZFC 33010–33014 (CIG 00310–00314), MZFC 33034–33035 (CIG 00336–00337), MZFC 33110 (CIG 00457), MZFC 33273 (CIG 00860), JAC 28612; Nayarit: MZFC 33211 (CIG 00649), MZFC 33240–33242 (CIG 00685–00687).
Eleutherodactylus leprus (n = 7): MEXICO: Veracruz: MZFC 33345–33350 (CIG 01139–01144), CIG 01270.
Eleutherodactylus longipes
(n = 3): MEXICO: Nuevo León: MZFC 33199 (CIG 00611); Querétaro:
Eleutherodactylus maculabialis (n = 27): MEXICO: Guerrero: MZFC 33307–33319 (CIG 00916–00923, 00940–00941, 00945–00947), MZFC 33321 (CIG 00949), MZFC 33323 (CIG 00953), CIG 01484–01485, 01501, JAC 25643–25646.
Eleutherodactylus marnocki (n = 3): USA: Texas: JHM 1427–1429.
Eleutherodactylus manantlanensis (n = 14): MEXICO: Colima: MZFC 33372–33377 (CIG 00530–00535), MZFC 33379–33381 (CIG 00646–00648), MZFC 33292–33296 (CIG 00892–00896).
Eleutherodactylus maurus (n = 19): MEXICO: Estado de México: MZFC 33071–33076 (CIG 00379–00384), MZFC 33355 (CIG 01174); Morelos: MZFC 33077–33080 (CIG 00385–00388), INIRENA 2925–30 (CIG 01733–01737, 01742).
Eleutherodactylus modestus (n = 34): MEXICO: Colima: MZFC 26888–26889, MZFC 33263–33270 (CIG 00850–00857), MZFC 33291 (CIG 00891), MZFC 33297 (CIG 00897); Jalisco: MZFC 33144–33149 (CIG 00493–00498), MZFC 33150–33154 (CIG 00505–00509), MZFC 33161 (CIG 00522), MZFC 33183–33185 (CIG 00570–00572), MZFC 33217–33223 (00655–00661).
Eleutherodactylus nebulosus (n = 6): MEXICO: Chiapas: MZFC 33361–33366 (CIG 01236–01241).
Eleutherodactylus nietoi (n = 13): MEXICO: Michoacán: MZFC 33121 (CIG 00299), MZFC 33042–33045 (CIG 00346–00349), MZFC 33050–33052 (CIG 00355–00357), MZFC 33336–33337 (CIG 00974–00975), MZFC 33342–33343 (CIG 00983– 00984), MZFC 33344 (CIG 00994).
Eleutherodactylus nitidus (n = 31): MEXICO: Estado de México: JAC 27237; Guerrero: MZFC 33096–33097 (CIG 00411–00412), MZFC 33104–33105 (CIG 00437–00438), JAC 25815; Morelos: MZFC 33081 (CIG 00389); Oaxaca: MZFC 33357–33358 (CIG 01211–01212); Puebla: MZFC 33356 (CIG 01181), JAC 27256–27276.
Eleutherodactylus orarius (n = 13): MEXICO: Colima: MZFC 26890, MZFC 33262 (CIG 00849); Michoacán: MZFC 33037 (CIG 00341), MZFC 33335 (CIG 00973), JAC 24020, 25526, 25563–25564, 29107, 30500–30501, 30517, 30625.
Eleutherodactylus pallidus (n = 13): MEXICO: Jalisco: MZFC 33271–33272 (CIG 00858–00859); Nayarit: MZFC 33189 (CIG 00588), MZFC 33212–33216 (CIG 00650–00654), MZFC 33243–33245 (CIG 00688–00690), MZFC 33018 (CIG 00995); Sinaloa: MZFC 33188 (CIG 00586).
Eleutherodactylus petersi (n = 11): MEXICO: Guerrero: MZFC 33034–33035 (CIG 00336–00337); JAC 25219, 25265–25266, 25299; Jalisco: MZFC 33010–33014 (CIG 00310–00314), MZFC 33034–33035 (CIG 00336–00337), MZFC 33110 (CIG 00457), MZFC 33273 (CIG 00860), JAC 28612; Michoacán: MZFC 33382–33385 (CIG 00675–00677), JAC 26947; Nayarit: MZFC 33211 (CIG 00649), MZFC 33240–33242 (CIG 00685–00687).
Eleutherodactylus. pipilans (n = 15): MEXICO: Guerrero: MZFC 33086–33088 (CIG 00396–00398), MZFC 33106–33107 (CIG 00439–00440), MZFC 33322 (CIG 00952), CIG 1465; Oaxaca: MZFC 33210 (CIG 00645), JAC 24283, 25809–25811.
Eleutherodactylus rubrimaculatus (n = 3): MEXICO: Chiapas: MZFC 33249–33251 (CIG 00753, 00755–00756),
Eleutherodactylus rufescens (n = 40): MEXICO: Jalisco: MZFC 33122–33126 (CIG 00471–00475), MZFC 33162–33164 (CIG 00527–00529), MZFC 33165–33174 (CIG 00544–00553), MZFC 33385 (CIG 00678); Michoacán: MZFC 33038–33041 (CIG 00342–00345), MZFC 33046–33049 (CIG 00350–00353), MZFC 33175–33182 (CIG 00559–00566), MZFC 33233 (CIG 00674), MZFC 33338 (CIG 00976), MZFC 33339–33341 (CIG 00980–00982).
Eleutherodactylus saxatilis (n = 4): MEXICO: Sinaloa: MZFC 26893, 26896, 26898–26899.
Eleutherodactylus sentinelus (n = 8): MEXICO: Guerrero: MZFC 33031–33033 (CIG 00333–00335), MZFC 33302– 33306 (CIG 00907–00913).
Eleutherodactylus syristes (n = 21): MEXICO: Guerrero: ANMO 2999; MZFC 33098–33103 (CIG 00431–00436), MZFC 33324 (CIG 00954), MZFC 33327–33328 (CIG 00957–00958) JAC 25701–25703; Oaxaca: MZFC 33207–33208 (CIG 00627–00628), MZFC 33209 (CIG 00644), 33378 (CIG00643), MZFC 33246–33247 (CIG 00713–00714), MZFC 33359–33360 (CIG 01232– 01233).
Eleutherodactylus teretistes (n = 5): MEXICO: Jalisco: MZFC 33142–33143 (CIG 00491–00492), MZFC 33277–33279 (CIG 00864–00866).
Eleutherodactylus verrucipes (n = 3): MEXICO: Tamaulipas: MZFC 33253–33254 (CIG 00813–00814); Querétaro: CIG 01273.
Eleutherodactylus wixarika (n = 3): MEXICO: Jalisco: MZFC 27477–27479.
Field number | Organism | Museum number | Locality | GenBank number |
---|---|---|---|---|
CIG-00953 | E. albolabris | MZFC-33323 | Mexico: Guerrero | MG856956 |
JAC-25642 | E. albolabris |
|
Mexico: Guerrero | MT872448 |
CIG-00392 | E. albolabris | MZFC-33084 | Mexico: Guerrero | MT872468 |
CIG-00441 | E. albolabris | MZFC-33108 | Mexico: Guerrero | MT872476 |
CIG-00331 | E. albolabris | MZFC-33029 | Mexico: Guerrero | MT872482 |
CIG-00332 | E. albolabris | MZFC-33030 | Mexico: Guerrero | MT872483 |
CIG-00477 | E. angustidigitorum | – | Mexico: Jalisco | MG856963 |
CIG-00479 | E. angustidigitorum | – | Mexico: Jalisco | MG856964 |
CIG-00407 | E. dilatus | MZFC33091 | Mexico: Guerrero | MG856973 |
CIG-00408 | E. dilatus | MZFC33092 | Mexico: Guerrero | MG856974 |
CIG-00070 | E. dilatus | – | Mexico: Guerrero | OP895113 |
EU186711 | E. erythrochomus | MZFC 16254 | Mexico: Guerrero | EU186711 |
RPA-0183 | E. erythrochomus | – | Mexico: Guerrero | MZ203201 |
RPA-0185 | E. erythrochomus | – | Mexico: Guerrero | MZ203202 |
CIG-01716 | E. franzi sp. nov. | INIRENA-2901 | Mexico: Guerrero | OP888987 |
CIG-01717 | E. franzi sp. nov. | INIRENA-2902 | Mexico: Guerrero | OP888988 |
CIG-01729 | E. franzi sp. nov. | INIRENA-2897 | Mexico: Guerrero | OP888989 |
CIG-01248 | E. guttilatus | – | Mexico: Guajuato | OP895114 |
CIG-01249 | E. guttilatus | – | Mexico: Guajuato | OP895115 |
CIG-01702 | E. humboldti. | INIRENA-2912 | Mexico: Mexico | OP888990 |
CIG-01703 | E. humboldti. | INIRENA-2913 | Mexico: Mexico | OP888991 |
CIG-01713 | E. humboldti | INIRENA-2920 | Mexico: Guerrero | OP888992 |
CIG-01730 | E. humboldti | INIRENA-2899 | Mexico: Guerrero | OP888993 |
GP-2422 | E. humboldti | – | Mexico: Mexico | OP888994 |
GP-2423 | E. humboldti | – | Mexico: Mexico | OP888995 |
RHA-0006 | E. humboldti. | – | Mexico: Mexico | OP888996 |
RHA-0007 | E. humboldti | – | Mexico: Mexico | OP888997 |
CIG-00649 | E. jamesdixoni | MZFC-33211 | Mexico: Nayarit | MT872469 |
CIG-00687 | E. jamesdixoni | – | Mexico: Nayarit | MG857035 |
CIG-00686 | E. jamesdixoni | – | Mexico: Nayarit | OP895111 |
CIG-00860 | E. jamesdixoni | – | Mexico: Jalisco | OP895112 |
CIG-00921 | E. maculabialis | MZFC-33312 | Mexico: Guerrero | MT872460 |
CIG-00923 | E. maculabialis | MZFC-33314 | Mexico: Guerrero | MT872461 |
CIG-00940 | E. maculabialis | MZFC-33315 | Mexico: Guerrero | MT872462 |
CIG-00941 | E. maculabialis | MZFC-33316 | Mexico: Guerrero | MT872463 |
CIG-00946 | E. maculabialis | MZFC-33318 | Mexico: Guerrero | MT872464 |
CIG-00947 | E. maculabialis | MZFC-33319 | Mexico: Guerrero | MT872465 |
CIG-00893 | E. manantlanensis | MZFC-33293 | Mexico: Colima | MG857007 |
CIG-00388 | E. maurus | MZFC-33080 | Mexico: Morelos | MG857010 |
CIG-00380 | E. maurus | MZFC-33072 | Mexico: Mexico | MG857011 |
CIG-00382 | E. maurus | MZFC-33074 | Mexico: Mexico | MT872478 |
CIG-00385 | E. maurus | MZFC-33077 | Mexico: Morelos | MT872479 |
CIG-0-387 | E. maurus | MZFC-33079 | Mexico: Morelos | MT872480 |
CIG-01733 | E. maurus | INIRENA-2925 | Mexico: Morelos | OP888998 |
CIG-01734 | E. maurus | INIRENA-2926 | Mexico: Morelos | OP888999 |
CIG-00857 | E. modestus | MZFC33270 | Mexico: Colima | MG857021 |
CIG-00891 | E. modestus | – | Mexico: Colima | MG857012 |
CIG-00753 | E. nebulosus | MZFC-33249 | Mexico: Chiapas | MG857056 |
CIG-00755 | E. nebulosus | MZFC-33250 | Mexico: Chiapas | MG857057 |
CIG-01237 | E. nebulosus | MZFC-33362 | Mexico: Chiapas | MT872429 |
CIG-01238 | E. nebulosus | MZFC-33363 | Mexico: Chiapas | MT872430 |
CIG-01240 | E. nebulosus | MZFC-33365 | Mexico: Chiapas | MT872431 |
CIG-01241 | E. nebulosus | MZFC-33366 | Mexico: Chiapas | MT872432 |
EU186712 | E. nitidus | AMCC-118239 | Mexico: Puebla | EU186712 |
CIG-00715 | E. nitidus | MZFC-33248 | Mexico: Oaxaca | MG857030 |
CIG-00412 | E. nitidus | MZFC-33096 | Mexico: Guerrero | MG857031 |
CIG-00336 | E. nitidus | MZFC-33034 | Mexico: Guerrero | MG857032 |
CIG-00311 | E. nitidus | MZFC-33011 | Mexico: Jalisco | MG857033 |
JAC-25815 | E. nitidus |
|
Mexico: Guerrero | MT872459 |
CIG-00341 | E. orarius | MZFC-33037 | Mexico: Michoacán | MG857041 |
CIG-00460 | E. orarius | MZFC-33113 | Mexico: Colima | MG857042 |
JAC-24020 | E. orarius |
|
Mexico: Michoacán | MT872434 |
JAC-25343 | E. orarius |
|
Mexico: Michoacán | MT872442 |
JAC-25344 | E. orarius |
|
Mexico: Michoacán | MT872443 |
CIG-00458 | E. orarius | MZFC-33111 | Mexico: Colima | MT872477 |
JAC-25392 | E. orarius | – | Mexico: Michoacan | OP895116 |
CIG-00337 | E. petersi | MZFC-33035 | Mexico: Guerrero | MT872473 |
CIG-00310 | E. petersi | MZFC-33010 | Mexico: Jalisco | MT872487 |
CIG-00396 | E. pipilans | MZFC-33086 | Mexico: Guerrero | MG857054 |
CIG-00398 | E. pipilans | MZFC-33088 | Mexico: Guerrero | MG857055 |
CIG-00440 | E. pipilans | MZFC-33107 | Mexico: Guerrero | MT872475 |
JAC-31539 | E. pipilans | – | Mexico: Oaxaca | OP895117 |
JAC-31540 | E. pipilans | – | Mexico: Oaxaca | OP895118 |
CIG-00334 | E. sentinelus | MZFC-33032 | Mexico: Guerrero | MT872485 |
CIG-00335 | E. sentinelus | MZFC-33033 | Mexico: Guerrero | MT872486 |
CIG-00333 | E. sentinelus | MZFC-33031 | Mexico: Guerrero | MT872484 |
CIG-00954 | E. syristes | MZFC-33324 | Mexico: Guerrero | MG857070 |
CIG-00714 | E. syristes | MZFC-33247 | Mexico: Oaxaca | MG857072 |
CIG-00628 | E. syristes | MZFC-33208 | Mexico: Oaxaca | MG857073 |
CIG-00627 | E. syristes | MZFC-33207 | Mexico: Oaxaca | MT872467 |
CIG-00431 | E. syristes | MZFC-33098 | Mexico: Guerrero | MT872471 |
CIG-00434 | E. syristes | MZFC-33101 | Mexico: Guerrero | MT872472 |