Research Article |
Corresponding author: Abel Batista ( abelbatista@hotmail.com ) Academic editor: Johannes Penner
© 2022 Konrad Mebert, Macario González-Pinzón, Madian Miranda, Edgardo Griffith, Milan Vesely, P. Lennart Schmid, Abel Batista.
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:
Mebert K, González-Pinzón M, Miranda M, Griffith E, Vesely M, Schmid PL, Batista A (2022) A new rainfrog of the genus Pristimantis (Anura, Brachycephaloidea) from central and eastern Panama. ZooKeys 1081: 1-34. https://doi.org/10.3897/zookeys.1081.63009
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Substantial molecular and morphological character differences lead us to the description of a new species of the genus Pristimantis from the cloud forest of Cerro Chucantí, Maje Mountains, Darien Province, as well as from several other mountain ranges in eastern and central Panama. Pristimantis gretathunbergae sp. nov. is a sister species to the allopatric P. erythropleura-penelopus group from northern Colombia with a mtDNA sequence divergence of > 4.4% at 16S and > 14.6% at COI. Its closest congener in sympatry is P. cruentus that differs by a large sequence divergence of > 9.6% in 16S mtDNA and 19.0% at COI, and from which it differs also by ventral and groin coloration, unusually prominent black eyes, a contrasting light upper lip, commonly a single conical to spine-like tubercle on the upper eyelid, and a larger head. While the habitat continuity at most sites in eastern Panama is moderate, habitats in central Panama are severely fragmented. Cerro Chucantí and the surrounding Maje Mountains are highly threatened by rapid deforestation and replaced by plantations and cattle pastures. Thus, investigations on the ecology of the new species and its population status, especially at the type locality, are highly recommended. As a flagship species, this new frog can help to preserve the Chucantí cloud forest including several recently described species known only from this isolated area in eastern Panama.
Chucantí, Craugastoridae, Greta Thunberg’s Rainfrog, Maje, Pristimantis gretathunbergae sp. nov., Strabomantidae, Terraranae
Tropical regions are extraordinarily rich in biodiversity which is caused by the combination of historical, climatic, and geographic characteristics that favor high speciation rates, as for example in anurans (
Currently, there are 13 species of Pristimantis frogs known to occur in Panama (
Our study focuses on Darién Province with the principal material originating from Cerro Chucantí. This mountain supports a remarkable diversity of organisms with 13 recently described new species, including plants (
Herein we describe a new species of Pristimantis based on molecular and morphological characters of specimens from Cerro Chucantí, Maje Mountains, and other mountain ranges in eastern and central Panama. Additionally, we present sequences and photographic vouchers (photo panels in online Suppl. material
The primary study site is Cerro Chucantí that includes the highest peak (1439 m a.s.l. at 8.8046°N, 78.4595°W; Fig.
Additional material for molecular and/or morphological analysis was obtained from specimens collected in eastern Panama (location names in bold as used in article): 1) Maje Ambroya, Maje Mountains, Panama Province; 2) Cerro Chucantí, Province Darién; 3) Río Tuquesa, Cerro Pechito Parao at Bajo Pequeño, Darien Mountains, Embera-Wounaan Comarca (= indigenous autonomous region). Third party material was collected in Central Panama from: 4) Cerro Brewster, Piedras-Pacora Mountains of Chagres National Park, Province Panama; 5) only photographic vouchers from Cerro Bruja, Chagres National Park, Province Colon; 6) a single DNA sequence from El Cope National Park at Rio Blanco, Penonomé Mountains, Province Cocle; and 7) Altos del Maria, vicinity of Gaita Hills, Province Panama Oeste (Fig.
For molecular analyses of Panamanian samples, DNA was extracted from fresh liver tissue. The 16S mtDNA extraction and sequencing follow previously described routines (
A Maximum Likelihood analysis (MA) was conducted for both genetic markers using IQ-TREE (
Collecting permits for 2009 (SC/A-8-09, SC/A-28-09), 2011 (SC/A-37-11), 2012 (SC/A-33-12), 2016 (SE/A-60-16) and 2018 (SE/A-33-18) as well as export permits for 2012 (SC/A-33-12) and 2013 (SEX/A-7-13) were provided by UNARGEN-Ministerio de Ambiente, Panama. Finally, we applied the traditional Environmental Vulnerability Score (EVS) methodology by
Specimens were collected by hand, photographed alive, euthanized with the Solution Tanax T-61, fixed with a preservative solution of 5 mL formalin (36%) in 1 L ethanol (94%), and subsequently stored in ethanol (70%) following the protocol of
Sex of specimens was determined by morphometric characters and presence of eggs in Panamanian samples. Measurements were taken to the nearest 0.1 mm, using a stereomicroscope and precision digital calipers. Following variables were measured according to
SVL Snout-Vent Length
HW Head Width, measured between posterior end of jaws
HL Head Length, measured between posterior end of jaws and tip of snout
InD Internarial Distance as shortest line between inner edges of narial openings
IoD Interorbital Distance as shortest distance between visible eyes, reflecting size of braincase
EW Eyelid Width, perpendicular distance to the outer edge of the upper eyelid
ED Eye Diameter as length of exposed eye
EN Eye-Nostril distance as shortest distance between anterior corner of eye and posterior margin of nostril
TY Tympanum Diameter
TL Tibial Length from knee to distal end of tibia
FL Foot Length between proximal edge of inner metatarsal tubercle to tip of fourth toe
FAL Forearm Length between elbow and hand
HAL Hand Length between proximal edge of palmar tubercle to tip of third finger
BW Body Width as largest width on trunk
AGD Axilla-Groin Distance as length between hind and front limbs along the trunk
TrL Trunk Length as SVL minus HL
3FW Width of 3rd Finger at penultimate phalanx just anterior to disc
3FD Disk Width of 3rd Finger
3TW Width of 3rd Toe at penultimate phalanx just anterior to disc
3TD Disk Width of 3rd Toe
4TW Width of 4th Toe at penultimate phalanx just anterior to the disc
4TD Disk Width of 4th Toe
Interspecific differences among Pristimantis spp. and related species are known to be relative lengths of heads, hind limbs, and feet (
We conducted a Multiple Correspondence Analysis (MCA) in R (Version 4.0.3), using the FactoMineR package, on categorical variables to compare the presence/absence of certain color pattern and tubercle characters between sympatric Pristimantis gretathunbergae sp. nov. and P. cruentus. We assessed the character state of six variables from photographic material of 26 P. gretathunbergae sp. nov. and 17 P. cruentus, whereby several specimens were collected and inspected by us, and their taxonomic allocation confirmed through molecular means. The value 0 was assigned to the morphological state typical for P. gretathunbergae sp. nov., whereas the value 2 is typical for P. cruentus, and the value 1 represents an intermediate expression. Follow do the descriptive states for these six variables: A) iris coloration: 0 = blackish to very dark red; 2 = whitish, golden, or light reddish, B) iris reticulation: 0 = no pattern, some with golden red sparkling; 1 = some dark red, small patches; 2 = reticulation, C) upper eyelid tubercle: 0 = single conical to spine-like; 1 = short singular, but spine-like, higher than other subtriangular humps; 2 = not singular or none at all, D) upper lip coloration: 0 = light colored and sharply demarcated to darker snout coloration; 1 = light color with some dark patches ingressing vertically from the snout, but upper border of light colored parts on the lip still contrastingly sharp bordered; 2 = no light color or very diffuse, no upper dark border, E) groin coloration: 0 = colors are relatively uniform white, yellow, light olive, or red, but some show a flecking pattern of these colors; 2 = dark brown to black flecks or patches on a light ground, F) ventral coloration: 0 = unicolored or fine spotting on white, yellow or orange: 2 = heavily dark mottled.
In the following, we present information on both genes (16S and COI) separately, as well as their combined results. However, we focus more on 16S for the presentation on closely related taxa, as 16S is more widely used and thus comparable with many Neotropical anurans (
The ABGD analysis generated ten groups of species by initial partition with prior maximal distance P = 1.45e-02 (Distance K80 Kimura MinSlope = 0.5) and a relative width of barcoding gap of 0.05 X-value (Fig.
Estimates of net evolutionary divergence (mean %) between groups (G-numbers from the ABGD analysis) of sequences of two mtDNA genes, 16S and COI. For every group, the estimated average evolutionary divergence over sequence pairs within groups is shown in parenthesis, with n: number of samples included in each group, followed by origin of country: CO (Colombia), CR (Costa Rica), EC (Ecuador), HO (Honduras), PA (Panama).
Species | 16S and COI evolutionary divergence between groups | ||||||||
---|---|---|---|---|---|---|---|---|---|
G1 | G2 | G3 | G4 | G5 | G6 | G7 | G8 | G9 | |
G1 P. gretathunbergae sp. nov. (5%; n: 10, PA) | – | – | – | – | – | – | – | – | – |
G2 P. penelopus (3%; n: 3, CO) | 9.6 | – | – | – | – | – | – | – | – |
G3 P. erythropleura (1%; n: 2 CO) | 11.0 | 8.1 | – | – | – | – | – | – | – |
G4 P. viejas (n.a.: n: 1, CO) | 13.7 | 12.7 | 14.6 | – | – | – | – | – | – |
G5 P. cruentus (12%; n: 22, PA) | 14.9 | 13.4 | 13.1 | 17.0 | – | – | – | – | – |
G6 P. cerasinus (6%; n: 2, CR, PA) | 14.6 | 12.8 | 14.3 | 13.2 | 17.5 | – | – | – | – |
G7 P. calcaratus (0%; %; n: 2, EC) | 15.1 | 12.2 | 13.2 | 13.2 | 16.5 | 15.9 | – | – | – |
G8 P. museosus (n.a; n:1, PA) | 16.4 | 12.7 | 14.2 | 15.7 | 18.0 | 16.0 | 18.3 | – | – |
G9 P. ridens (11%; n: 3, CR, HO) | 18.6 | 15.4 | 15.7 | 17.3 | 19.2 | 17.9 | 16.7 | 21.1 | – |
G10 P. taeniatus (13%; n: 3, CO) | 22.0 | 20.6 | 20.5 | 22.4 | 22.7 | 22.0 | 20.6 | 22.3 | 21.7 |
Phylogenetic tree of Pristimantis spp. based on mtDNA genes 16S and COI performed by a Shimodaira-Hasegawa approximate likelihood ratio test (SH-aLRT test). Numbers on nodes indicate estimated SH-aLRT support/bootstrap support with SH-aLRT values ≥ 80% are considered reliable for a clade (
On a wider perspective, the phylogenetic inference based on combined 16S and COI sequences placed the Pristimantis erythropleura-penelopus clade, P. cruentus, P. cisnerosi (data of cisnerosi available only for 16S, see also Suppl. material
Pristimantis gretathunbergae sp. nov. reveals a genetic variation of < 3% for 16S between our focus populations from Maje Mountains (Cerro Chucantí and Maje Ambroya) to related populations from Rio Tuquesa (divergence of 1.5–2.8%) and Cerro Brewster (1.9–2.9%), but also to a single sequence from farther west at El Cope, Central Panama (2.3%). This corresponds to the suggested and applied minimum sequence divergence of 3% between Neotropical frog species (
The shortest genetic distance for 16S mtDNA between the new Pristimantis species and any other Pristimantis sample was 4.4% and pertains to two specimens of allopatric Colombian relatives, one P. erythropleura (minimum of 14.6% at COI to the new species) and one P. penelopus (min. of 15.5% at COI to the new species). The mean difference at 16S in these Colombian subsamples to P. gretathunbergae sp. nov. is 4.8%, that increased with the addition of a few samples from other sites in the same general region to 5.9% (P. erythropleura), respectively 6% (P. penelopus, see also Suppl. material
Results of morphometric measurements of adult specimens of Pristimantis gretathunbergae sp. nov. are presented in Table
Morphometric characters of adult Pristimantis gretathunbergae sp. nov. with mean ± SD (range, followed by n); all values are in mm and separated by sex. Abbreviations of raw variables: Snout-Vent Length (SVL), Head Width (HW), Head Length (HL), Internarial Distance (InD), Interorbital Distance (IoD), Eyelid Width (EW) Eye Diameter (ED), Eye-Nostril Distance (EN), Tympanum Diameter (TY), Tibial Length (TL), Foot Length (FL), Forearm Length (FAL), Hand Length (HAL), Body Width (BW), Axilla-Groin Distance (AGD), 3rd Finger Width (3FW), 3rd Finger Disk Width (3FD), 3rd Toe Width (3TW), 3rd Toe Disk Width (3TD), 4th Toe Width (4TW), and 4th Toe Disk Width (4TD); see methods for definitions.
Measurement | Females | Males |
---|---|---|
SVL | 42.66±3.71(38.15–46.3; 4) | 31.24±3.52(26.9–36.7; 8) |
HW | 19.17±0.99(17.84–20; 4) | 12.39±1.55(10.7–15.9; 8) |
HL | 18.14±1.49(16.46–19.9; 4) | 12.44±1.69(10–14.7; 7) |
InD | 3.05±0.21(2.9–3.2; 2) | 2.37±0.55(1.67–3.3; 7) |
IoD | 5.1±0.71(4.6–5.6; 2) | 3.11±0.45(2.6–3.9; 7) |
EW | 6.3±0.14(6.2–6.4; 2) | 4.79±0.43(4.3–5.4; 7) |
TL | 23.15±2.05(21.7–24.6; 2) | 14.61±5.07(3.5–18.8; 7) |
FL | 22.1±0.14(22–22.2; 2) | 14.45±1.99(11.7–17.8; 7) |
TY | 2.85±0.92(2.2–3.5; 2) | 1.34±0.49(0.65–2; 7) |
ED | 5.45±0.21(5.3–5.6; 2) | 4.11±0.61(3.7–5.3; 7) |
EN | 5.65±0.49(5.3–6; 2) | 3.55±0.67(3.02–5; 7) |
FAL | 11.2±1.13(10.4–12; 2) | 7.73±0.85(6.5–8.86; 7) |
HAL | 9.15±6.72(4.4–13.9; 2) | 9.22±1.27(7.5–11.4; 7) |
3FW | 1.45±0.21(1.3–1.6; 2) | 0.72±0.28(0.41–1.1; 7) |
3FD | 3.15±0.07(3.1–3.2; 2) | 1.66±0.26(1.2–2; 7) |
3TW | 1.25±0.07(1.2–1.3; 2) | 0.59±0.28(0.19–0.94; 7) |
3TD | 2.35±0.07(2.3–2.4; 2) | 1.24±0.35(0.66–1.7; 7) |
4TW | 1.4±0(1.4–1.4; 2) | 0.62±0.16(0.39–0.83; 7) |
4TD | 2.5±0.14(2.4–2.6; 2) | 1.39±0.11(1.3–1.6; 7) |
BW | 16.75±1.34(15.8–17.7; 2) | 8.46±1.2(7.14–10.8; 7) |
AGD | 21.4±0.85(20.8–22; 2) | 12.85±1.73(10.5–14.8; 5) |
A PCA revealed the following relative variables to contribute mostly to the principal components: TrL/HL, 3TD/3TW, 3FD/3FW, 4TD/4TW with strong loadings and IoD/HL, ED/HL, EW/HL with medium loadings (Suppl. material
A subsequent Linear Discriminant Analysis LDA correctly separated P. gretathunbergae sp. nov. (n = 9) from P. cruentus from eastern Panama (n = 27) and western Panama (n = 37). Pristimantis cruentus had to be split into two separate geographic groups based on LDA-conditions (LDA graph in Suppl. material
A final univariate analysis corroborates that in four morphometric variables used in the LDA P. gretathunbergae sp. nov. differ significantly from P. cruentus, for which Eastern and Western populations were combined (unlike in the LDA): P. cruentus exhibits a relatively larger eye (mean ED/HL = 0.414; mean P. gretathunbergae sp. nov. = 0.322; Welch Two Sample t-test, t = 6.297, df = 25.65, p < 0.001), and eyelid width (mean sp. nov. EW/HL = 4.50, mean P. gretathunbergae sp. nov. = 0.376; Welch Two Sample t-test, t = 4.667, df = 25.97, p < 0.001); a longer trunk (mean TrL/HL = 2.109; mean P. gretathunbergae sp. nov. = 1.529; Mann‑Whitney‑U‑Test, W = 548, p < 0.001), and a wider head (mean IoD/HL = 0.368; mean P. gretathunbergae sp. nov. = 0.259; Welch Two Sample t-test, t = 7.591, df = 18.8, p < 0.001). These results indicate that the head morphology relates primarily to the separation of P. gretathunbergae sp. nov. and all P. cruentus. Toe characters, important in the LDA, are only significantly different between P. cruentus from eastern Panama and western Panama (4TD/4TW: Mann‑Whitney‑U‑Test, W = 686, p = 0.011), whereas they are marginal to not different between P. gretathunbergae sp. nov. and P. cruentus from eastern Panama (4TD/4TW: Mann‑Whitney‑U‑Test, W = 73, p = 0.079) or from western Panama (4TD/4TW: Mann‑Whitney‑U‑Test, W =151, p = 0.683), respectively.
A Multiple Correspondence Analysis (MCA) of six categorical variables of color pattern and tubercle properties results in a clear distinction between the new Pristimantis gretathunbergae sp. nov. and its closest relative in sympatry, P. cruentus. The most frequent and/or typical expression of these variables in Pristimantis gretathunbergae sp. nov. (with the comparative expression of P. cruentus in parenthesis) are: 1) blackish eyes or iris (light colored iris in P. cruentus), 2) no iris reticulation (reticulated), 3) a single conical tubercle on the upper eyelid (rarely so, generally more variable from subtriangular to spine-like, and from none at all to several ones), 4) light upper lip contrastingly bordered to dark coloration on snout above (none, diffusely colored lips, or light, but not demarcated), 5) coloration of groin, as well as 6) venter is unicolored whitish, yellow or reddish, sometimes with fine spotting (heavily black and white to dark and light mottled, see methods for a more detailed and expanded species-specific variable definition and quality scoring). A photographic example of a Pristimantis gretathunbergae sp. nov. and a P. cruentus in a face-off position is depicted in Fig.
The first and second dimension of the MCA describe 73.89% of the total variance, allowing a conclusive two-dimensional display of the scores (Fig.
Map of the Multiple Correspondence Analysis (MCA) of P. gretathunbergae sp. nov. (red dots) and P. cruentus (black dots): Number labels for individual frog with lines pointing to specimen location on the map. Following correlation ratio (Dimension 1/Dimension 2) resulted from the MCA: iris coloration 0.937/0.001; iris reticulation 0.933/0.637; upper eyelid tubercle 0.751/0.331; upper lip coloration 0.735/0.326; groin coloration 0.852/0; ventral coloration 0.810/0.001. The qualitative scoring of the variables and its species-specific expression is explained in the methods.
Two distinct clusters appear in the MCA that clearly represent the two sympatric species P. gretathunbergae sp. nov. and P. cruentus (Fig.
U‑Test, W = 133, p < 0.001, and Dimension 2: Mann-Whitney U-Test, W = 321, p = 0.026). These results strongly separate the two Pristimantis species on qualitative morphological characters, with the distinctive eye color and pattern being a particular easy and obviously useful character to separate P. gretathunbergae sp. nov. from P. cruentus (see Suppl. material
Based on molecular divergence and morphological consensus, we assign the undescribed Pristimantis sp. with the type material from Cerro Chucantí, Maje Mountains as a new species to science. It belongs to the Pristimantis ridens species group (sensu
MHCH 3082 (original field number AB 1059), an adult male (Figs
Coloration in life of specimens of Pristimantis gretathunbergae sp. nov. and P. cruentus from eastern Panama A holotype male (MHCH 3082), Cerro Chucantí B paratype female (SMF97520), Cerro Chucantí C left, paratype female (MHCH 3081), right P. cruentus female (MHCH3034) D female from Cerro Chucantí, not collected E female (MHCH3115) La Javillosa F female, Cerro La Javillosa, Ambroya, Maje Mountain Range (SMF97517) G female (MHCH3079), Rio Tuquesa. Colored lines point to some diagnostic characters as follow: red: blackish iris; yellow: single spine-like tubercle; turquoise: light-colored upper lip; pink: cream, yellow to red groin (red groin also shown in Suppl. material
Seven males, three females. Male and female
Pristimantis gretathunbergae sp. nov., a member of the Pristimantis ridens species group (sensu
Pristimantis gretathunbergae sp. nov. differs markedly from all other Pristimantis species in central and eastern Panama by its very dark to black, non-reticulated iris, respectively entire eyes (iris pale and/or with heavy pale flecking in other species). Some fine golden to dark red speckling or flecking might be visible in some P. gretathunbergae sp. nov. In sympatry, the new species is most similar to the equally large and bulky P. cruentus (Fig.
This comparison includes only members of the Pristimantis ridens species group sensu
Two additional rainfrog taxa inhabit northwestern Colombia that are closely related to P. gretathunbergae sp. nov. First, Pristimantis penelopus, sister species to P. erythropleura, was originally known to inhabit montane areas higher than 1000 m a.s.l. in northwestern Colombia (
Both, P. penelopus and P. sanguineus (examples in Suppl. material
Further detailed comparisons to similar rainfrog species, e.g., P. viejas, P. latidiscus, P. laticlavius, P. cisnerosi, and P. paisa is provided in the Suppl. material
(Figs
SVL 34.6, HW 12.8, HL 14.2, InD 2.4, IoD 4.1, EW 5.4, ED 4.6, EN 3.6, TY 1.9, TL 17.7, FL 16.2, FAL 8.8, HAL 10.2, BW 8.7, 3FW 0.8, 3FD 1.5, 3TW 0.9, 3TD 1.4, 4TW 0.7, 4TD 1.3.
(MHCH 3082; Fig.
(Fig.
(Fig.
The specific name is a noun in the genitive case and is a patronym in honor for Greta Thunberg, a Swedish student, and her global climate activism. Greta initiated a “School Strike for Climate Action” outside the Swedish parliament to demand a radical response to the threat by the ongoing climate change. Then sixteen-year-old Thunberg’s example has inspired students worldwide to carry out similar strikes called Fridays For Future that started in August 2018. In December 2018 she addressed world leaders at the COP24 climate talks in Katowice, Poland, with sharp and unmasked words, and equally impressed a global audience in January 2020 with her unpolitical, direct speech down to the point on “Averting a Climate Apocalypse” at the WEF (World Economic Forum) in Davos, Switzerland. Just recently, she publicly slammed the world leaders at the 26th UN Climate Change Conference of the Parties (COP26) in Glasgow, November 2021, for not doing enough to meet the demands of the climate emergency. Greta Thunberg represents the authentic voice that exposes the motivations behind the diplomatic curtain of politicians and business stakeholders. Her voice is essential if we want to revert to and maintain a healthy environment on the planet we all share, and not least, learn to respect its magnificent mega-diversity of life that took millions of years to evolve.
Pristimantis gretathunbergae sp. nov. is endemic to Panama, but it could occur on near mountains along the border in Colombia. Its currently known distribution covers eastern Panama with records from the Darien Mountains within Embera Comarca and the Maje Mountains within Darien and Panama Provinces, including the type locality at Cerro Chucantí. The distribution continues west into Central Panama, including records from Piedras-Pacora Mountains, Panama Province, and Cerro Bruja, Colon Province, both within Chagres National Park. Farther west across the Panama Canal, P. gretathunbergae sp. nov. is present at Altos del Maria, region of Gaita Hills, Panama Oeste Province, and in the region of El Cope, Omar Torrijos National Park, Coclé Province.
Color pattern of specimens from Cerro Brewster, not included in the LDA (DFA) analysis, are consistent with the specimens from Maje Mountains in having a cream dorsum coloration, the margin of the upper lip in females yellow, an iris nearly black with pale dots or speckles, venter dirty white, and general stocky body and head. Due to the unique combination of characters of P. gretathunbergae sp. nov., in particular the blackish non-reticulated iris and light, unpatterned upper lip, that differs from any other related rainfrog in Panama and Colombia, we confidently allocate specimens available only as photo vouchers from Cerro Bruja, Colon Province, and Altos del Maria, Gaita Hills, Panama Oeste Province to the same species. The latter two localities substantially reduce the gap to El Cope, Cocle Province, the origin of the most western specimen of our Group 1. So far, we have not received photographic vouchers for the specimen from El Cope, but the low 16S-divergence of 2.3% clearly links it to the undescribed species from the Maje Mountains (see above).
Pristimantis gretathunbergae sp. nov. has been recorded at altitudes between 718–1439 m a.s.l. and occupies most frequently montane forest, a cloud forest consisting predominantly of trees covered with moss and a large variety of understory and midstory bromeliads (
Habitat, mating, and parental care in females of Pristimantis gretathunbergae sp. nov. from Cerro Chucantí A Habitat on Cerro Chucantí at ca. 1300 m a.s.l. B understory bromeliad with a P. gretathunbergae sp. nov. in situ (blue line) and zoomed in on inset (MHCH 3115) C amplectant pair on axillary part of bromeliad leaf (not collected) D same female after amplexus guarding eggs E female of P. gretathunbergae taking care of its eggs with a male P. cruentus species holding on the female in reverse position (not collected) F female with eggs about to hatch, note the transparency of the egg membrane (not collected).
Habitats occupied by P. gretathunbergae sp. nov. are under latent threat. For example, anthropogenic pressure around Cerro Chucantí and the Maje Mountains most likely will lead to declines of populations through habitat destruction (
1 | Dorsal ground color uniform blackish or grayish, with white or orange blotches on groin, if not, the color is gray to pink, some species have an orange or yellow dorsolateral stripe, continuous or interrupted | 2 |
– | Dorsal ground color, cream, reddish, brown tones, green or olive, uniform or darker blotches or reticulations, without white or orange blotches on groin | 5 |
2 | White or orange blotches on groin | 3 |
– | Groin uniform | 4 |
3 | Well defined white blotches on groin | Pristimantis pardalis |
– | Well defined orange blotches on groin | Pristimantis altae |
4 | Dorsal ground color uniform gray to pink without dorsolateral stripes | Pristimantis pirrensis |
– | Orange or yellow dorsolateral stripes, continuous or interrupted, some specimens lack stripes, but dorsal color is blackish, never gray or pink | Pristimantis gaigei |
5 | Dorsal ground color green or olive green, uniform or with darker blotches or reticulations | 6 |
– | Dorsal ground color, cream, reddish, brown, dark brown, or olive, uniform or with darker blotches or reticulations | 7 |
6 | Dorsal ground color uniform green, with or without a reddish brown transverse interorbital band, dorsal skin smooth | Pristimantis moro |
– | Dorsal color green or olive, with irregular blotches or reticulations, brown, olive or reddish color, dorsal skin tuberculated | Pristimantis museosus |
7 | Heel smooth or with one to several similar small sized tubercles scattered over upper surface of hind limb; enlarged tubercle on upper eyelid present or not | 8 |
– | Well-developed pointed calcars, usually enlarged tubercle on upper eyelid | 10 |
8 | Presence of a dorsolateral granular folds, dorsal pattern with chevrons | Pristimantis achatinus |
– | Dorsolateral region smooth, dorsal pattern uniform, never with chevron pattern | 9 |
9 | Anterior and posterior surfaces of thighs, calves, and feet red | Pristimantis ridens |
– | Uniform posterior surface of thigh | Pristimantis taeniatus |
10 | Posterior thighs uniform | 11 |
– | Posterior thighs dark brown with red-orange dots | Pristimantis adnus |
11 | Dorsal skin granulate or tuberculate, rarely smooth, head about as broad as long; snout rounded | 12 |
– | Dorsal skin smooth, long and pointed snout | Pristimantis caryophyllaceus |
12 | No W-shape on dorsum, iris variable in color, usually highly reticulated or blackish | 13 |
– | W-shape ridge that extent from the back of the head to the shoulder region, groin, anterior, and posterior thigh red, iris usually pale golden without reticulation, eyes usually with an orange perimeter | Pristimantis cerasinus |
13 | Iris variably light colored, cream, yellow or reddish and strongly reticulated, venter heavily mottled with dark pigment to almost uniform black, upper surfaces gray, brown, brownish black; tympanic annulus partially evident in females, upper lips with dark patches, with light colored lips or parts of it little or not dark-bordered above | Pristimantis cruentus |
– | Iris black, some very dark red, without reticulation, venter white, dirty white, yellow or red, upper surfaces reddish brown or yellow, white to yellow upper lips, contrastingly dark-bordered above, some with dark patches, tympanic annulus not visible in females | Pristimantis gretathunbergae sp. nov. |
The genus Pristimantis is one of the most species rich genera of amphibians in the Neotropics (Lehr and Duellman 2009). It is primarily distributed in South America with a few species reaching Central America. Based on a combination of molecular data (small sequence divergence of mtDNA 16S and COI) and a consensus of conspicuous morphological characters (e.g., unusually dark to black eyes, spine-like single tubercle on upper eyelid, sharply dark-bordered upper light lip, large body size), we could identify seven locations in Darien and Central Panama that relate to a new species, Greta Thunberg’s Rainfrog Pristimantis gretathunbergae sp. nov. This is the 14th or 15th known Pristimantis species in Panama, depending on the author (see Introduction). Initially, P. gretathunbergae sp. nov. thought to be related to P. latidiscus (
Within Panama, Pristimantis gretathunbergae sp. nov. is most closely related to P. cruentus, a rainfrog species with a large variation in morphology and genetics (
Whereas all interpopulation divergence between the type series with any of the other sites remains below 3%, other population comparisons can vary and increase up to 5.4% between single individuals from Cerro Brewster and El Cope, both central Panama. Similar minimum genetic differentiation of > 3–4% of the 16S rRNA gene have been found to associate to CCS and UCS (not yet described Confirmed and Unconfirmed Candidate Species) of frogs in Madagascar, Africa, and Amazon Basin, South America (
Cloud forests in general and isolated mountain tops in particular are highly vulnerable to climate change due they low range of mobility and high habitat specialization of its denizen (
We thank Luis DeLeon, Jesús Pérez, Juan Zarzavilla, Yorlis Cáceres, Hugo Martínez, and Gilberto Torres for field assistance, and Guido Berguido for his support during our stay at the Chucantí private reserve. We also thank to Sarah Farinelli, Juan Daza, Mauricio Rivera-Correa, and an anonymous reviewer for their valuable comments during the review process, which greatly improved the manuscript. To Marcos Ponce and Angel Sosa a thank you for providing photos of a specimen from Cerro Brewster. This work was financially supported by Asociación ADOPTA el Bosque Panama, Sistema Nacional de Investigación (SNI) of the Secretaría Nacional de Ciencia, Tecnología e Innovación (SENACYT, Panamá), and the Rainforest Trust (US). We thank Sebastian Lotzkat, for his assistance in the field and on the manuscript. Many colleagues have provided photographs for the species panels in the Suppl. material
Tables S1–S5
Data type: docx file
Explanation note: A new rainfrog of the genus Pristimantis (Anura, Brachycephaloidea) from central and eastern Panama
Figures S1–S16
Data type: docx file
Explanation note: A new rainfrog of the genus Pristimantis (Anura, Brachycephaloidea) from central and eastern Panama