New Mannophryne from Sierra de San Luis, Venezuela

ANARTIA

Publicación del Museo de Biología de la Universidad del Zulia

ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)

https://doi.org/10.5281/zenodo.10429600 / Anartia, 36 (junio 2023): 63-74

http://zoobank.org/urn:lsid:zoobank.org:pub:4E4F7C71-F68D-4BC0-B6DD-83A5CFCDB6E6

Some new species can be foretold: An endemic collared frog

(Aromobatidae: Mannophryne La Marca, 1992) is discovered

in a still herpetologically unexplored mountain range in

northern Venezuela

Algunas especies nuevas se pueden predecir: una rana endémica con collar

(Aromobatidae: Mannophryne La Marca, 1992) es descubierta en una serranía

herpetológicamente inexplorada del norte de Venezuela

Enrique La Marca

1,2

, Abraham Mijares-Urrutia

, Luis A. Saavedra

& Carlos Gottberg

Escuela de Geografía, Facultad de Ciencias Forestales y Ambientales, Universidad de Los Andes, Mérida 5101, Venezuela

REVA (Rescue of Endangered Venezuelan Amphibians) Conservation Center, and Fundación BIOGEOS para el estudio

de la diversidad biológica; Mérida 5101, Venezuela

Conservation and Ecosystem Management Teaching Section, Technical and Further Education Institute, TAFE Digital Campus,

Sydney, NSW 2135, Australia

Departamento de Biología, Facultad de Ciencias, Universidad de Los Andes, Mérida 5101, Venezuela

Correspondencia: email: enrique.lamarca@gmail.com

(Received: 29-06-2023 / Accepted: 30-11-2023 / On line: 24-12-2023)

ABSTRACT

A morphological study of small body sized Mannophryne frog specimens from the Sierra de San Luis, Venezuela, revealed

that they belong to an undescribed species. By having a narrow collar, the new species dierentiates from all Mannophryne

having a wide collar (a character present in almost half of the species included in this genus). From the rest of narrow-col-

lared Mannophryne it is easily dierentiated by a combination of dierent foot-web formula and pattern of coloration. e

geographically closest species is M. caquetio Mijares-Urrutia & Arends, 1999, from which it diers by having a distinctive

pattern of coloration, dierent shape of the tip of the snout, size of tympanum, degree of detachment of tongue from oor

of mouth, by having lateral fringes along toes not forming ap folding on digits, and by having a more extensive foot-web.

e most closely resembling frog to the new species is M. lamarcai Mijares-Urrutia & Arends, 1999, from which it diers

by having a distinctive pattern of coloration, of which it stands out the uniformly dark dorsum without dorsolateral bands;

also, by having the tip-of-snout truncated, lateral aps along toes not folding onto digits, and less foot-webbing. e new

species comes from a Premontane humid forest that is completely isolated by surrounding dry forests and raises to three

the known members of the genus in Falcon State, a Venezuelan geopolitical division still lacking complete herpetological

inventories. It is the northernmost Venezuelan and continental South American Mannophryne, rising to 21 the known

species in the genus. In this paper we describe the new species, provide a diagnosis comparing it to all known members of

the genus, and provide data on its biogeography, ecology, and conservation.

Key words: Aromobatidae, biogeography, conservation, Dendrobatoidea, Falcon state.

E. La Marca, A. Mijares-Urrutia, L. A. Saavedra & C. Gottberg

RESUMEN

Un estudio morfológico de ejemplares de pequeñas ranas Mannophryne de la Sierra de San Luis, Venezuela, reveló que

pertenecen a un taxón previamente no descrito. Al tener un collar estrecho, la nueva especie se diferencia de todos los

Mannophryne que tienen un collar ancho (carácter presente en casi la mitad de las especies incluidas en este género). Del

resto de Mannophryne con collar estrecho se diferencia por la combinación de diferente extensión de membranas en las

patas y por el patrón de coloración. La especie geográcamente más cercana es M. caquetio Mijares-Urrutia & Arends,

1999, de la cual se diferencia por tener distinto patrón de coloración, diferente forma de la punta del hocico, tamaño del

tímpano, grado de desprendimiento de la lengua desde el piso de la boca, por tener rebordes laterales a lo largo de los dedos

de los pies que no forman solapas plegables sobre los dedos y por tener una membrana podal más extensa. La rana que más

se parece a la nueva especie es M. lamarcai Mijares-Urrutia & Arends, 1999, de la cual se diferencia por tener un patrón de

coloración distinto, del cual destaca el dorso uniformemente oscuro y sin bandas dorsolaterales; también por tener la punta

del hocico truncada, solapas laterales a lo largo de los dedos de los pies que no se doblan sobre los dedos y por poseer menor

extensión de las membranas en los pies. La nueva especie proviene de un bosque húmedo premontano aislado y comple-

tamente rodeado por bosques secos, y eleva a tres los miembros conocidos del género en el estado Falcón, una división

geopolítica venezolana aún carente de inventarios herpetológicos completos. En este artículo describimos la nueva especie,

proporcionamos un diagnóstico donde se compara con todos los miembros conocidos del género y aportamos datos sobre

su biogeografía, ecología y conservación.

Palabras clave: Aromobatidae, biogeografía, conservación, Dendrobatoidea, estado Falcón.

MATERIALS AND METHODS

Specimens examined are listed in the Appendix 1. Ter-

minology and methods follow La Marca et al. (2004), Var-

gas Galarce & La Marca (2007); and La Marca (2009). Sex

was determined by gonadal inspection. Measurements (in

mm) were taken on the le side of animals under a Wild

stereomicroscope (TYP 181300, Heerbrugg, Switzer-

land), using a Surtek® digital caliper (model 122202) with

a precision of 0.01 mm.

Measurements taken for post-metamorphic specimens

were snout-to-vent length (SVL); head length: distance

from tip-of-snout to posterior corner of mouth (HL);

head width: maximum straight distance between angle of

jaws (HW); eye-to-naris distance: distance from anterior

corner of eye to center of naris (EN); internarial distance:

maximum straight length between centers of nares (IN);

nares to tip-of-snout distance (NTS); eye length: distance

from anterior to posterior corner of eye (EYE); horizon-

tal length of tympanum: distance between anterior and

posterior level of tympanum (T); hand length: distance

from proximal edge of palmar tubercle to tip of nger III

(HAND); tibia length: distance from outer edge of exed

knee to heel (TL); foot length: distance from proximal

edge of outer metatarsal tubercle to tip of toe IV (FOOT);

interorbital distance: distance between borders of upper

eyelids (IOD); upper eyelid width: distance between bor-

der of eye and base of eyelid (UEW); distance from an-

terior border of eye to tip-of-snout (ETS); distance from

anterior border of tympanum to posterior border of eye

(TE); length of shank from knee or tibio-tarsal joint to

INTRODUCTION

Mannophryne La Marca, 1992, is a genus of Neotropi-

cal frogs found almost exclusively in Venezuela (Rojas-

Runjaic et al. 2018). A total of 20 species (Frost 2023)

are known, distributed mainly in the mountain systems to

the north and west of this country, being more numerous

in the Cordillera de Mérida, followed by those located in

the mountains to the east of the country. is genus is also

represented in the Lara-Falcón Mountain System (Yústiz

1991, MijaresUrrutia & Arends 1999a, b, Mijares-Urrutia

2000), which is a kind of wedge between the Venezuelan

Andean and coastal systems, but the species inventory

there is still far from complete.

Mannophryne species are found mostly in humid forests,

specically those characterized as “Premontane moist for-

est” and “Premontane wet forest” in the Holdridge’s Life

Zone system (Ewel et al. 1976). is close association of

species in the genus and this Life Zone served to postulate

a hypothesis (La Marca 1994, 1995) which predicts the ex-

istence of representatives of the genus in these plant forma-

tions where these frogs have not been previously reported,

especially if they are naturally isolated from each other.

Studying frogs from the Sierra de San Luis, in the Fal-

cón state, we found some specimens belonging to the ge-

nus Mannophryne and coming from a Premontane humid

forest that is completely isolated by surrounding dry for-

ests. e morphological study of these animals revealed

that they belong to an undescribed taxon whose presence

was expected in this type of forest, as predicted. e de-

scription of this species is the main objective of this work.

New Mannophryne from Sierra de San Luis, Venezuela

heel (TARSUS); length of dermal fold on tarsus (LTF);

width of pad on third nger (WP3F); width of phalanx

adjacent to pad on third nger (WPAP).

A denition and a diagnosis are presented separately for

the new taxon before its description. ese concepts are, as

Simpson (1961:138) pointed out “importantly dierent,

but in fact taxonomists seldom use the terms consistently

and commonly assume that they are synonymous”. Gloyd

& Conant (1990:13) stated that “A denition explains

what a taxon is, whereas a diagnosis tells what it is not, es-

pecially if a comparison is made with one or more other

taxa”. Both terms, denition and diagnosis, have been con-

sistently used by the rst author in all his publications re-

garding the genus Mannophryne.

Mannophryne phylidros sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:F882F1AB-D8FB-4A32-8067-

53DB93C84DB9

(Figs. 1, 2)

Mannophryne sp. 2. Mijares-Urrutia & Arends 2000: 6.

Holotype. CHCG (Colección Herpetológica Carlos

Gottberg) 428; adult female (Fig. 1), collected by Carlos

Gottberg in Cataratas de Hueque (11° 11’N, 69° 34’W),

610 m asl, Sierra de San Luis, municipio Petit, estado Fal-

cón, Venezuela.

Paratypes. Topotypes (Fig. 2), same data as holo-

type. CHCG 425-427, and 429, four adult females; and

CHCG 430-431, two adult males.

Remark: the CHCG is housed as a separate collection

within the Colección de Vertebrados de la Universidad de

Los Andes (CVULA), Mérida, Venezuela.

Etymology. e specic name derives from the Greek

word “phylidros”, meaning “water-loving” (Brown 1978:

845), in loose relation to the association of the type speci-

mens with the waterfalls at the type locality.

Denition. A medium sized Mannophryne, mean SVL

males: 23.4 mm (range 23.2-23.7 mm), females: 26.4 mm

(range 25.3-27.6 mm), distinguished from other Man-

nophryne by the following combination of characters: (1)

skin of dorsum smooth, (2) lower part of tympanum mod-

erately conspicuous, (3) tip-of-snout truncated, (4) can-

thus rostralis ill-dened, sinuous, (5) rst nger equal in

length to second, (6) pad on third nger about twice wider

than adjacent phalanx, (7) ngers bearing lateral fringes,

(8) short cloacal fold with crenulate border, (9) tarsal fold

strong, not forming a ap, (10) foot web formula: I (1.5-

2.0)-0.5II 1.5-1 III 1.5-1IV (0.5-1)-1V, (11) toes with later-

al aps, not folding onto digits, (12) pale dorsolateral band

absent, dorsolateral dark band absent, (13) width of pad on

third toe slightly less than half the width of the adjacent

phalanx, (14) heel reaching to eye when leg is adpressed

forward, (15) short pale oblique inguinal band, (16) collar

dark-brown, relatively-narrow and with small cream-col-

ored blotches, (17) undened pale ventrolateral blotches,

(18) venter cream or dirty cream in females; black to gray

(especially darker on anterior part) in males.

Diagnosis. As other members in the genus, Man-

nophryne phylidros sp. nov. has a dark band (“collar”) on

chest. e new species has a relatively narrow collar, which

allows to dierentiate it from other members of the ge-

nus having a wide collar [M. collaris (Boulenger, 1912);

M. cordilleriana La Marca, 1994; M. herminae (Boettger,

1893); M. larandina (Yústiz, 1991); M. molinai Rojas-

Figure 1. Dorsal and ventral view of preserved holotype of Mannophryne phylidros sp. nov.

E. La Marca, A. Mijares-Urrutia, L. A. Saavedra & C. Gottberg

Figure 2. Dorsal and ventral views of preserved paratypes of Mannophryne phylidros sp. nov.

New Mannophryne from Sierra de San Luis, Venezuela

Runjaic et al. 2018; M. oblitterata (Rivero, 1984), M. riv-

eroi (Donoso-Barros, 1964); M. speeri La Marca, 2009;

M. trujillensis Vargas Gallarce & La Marca, 2007; and M.

yustizi (La Marca, 1989)]. Of those Mannophryne having

a narrow collar, the new species diers as follows (char-

acters of compared species are within parentheses): from

M. leonardoi Manzanilla et al. 2007, by lacking a dark

canthal band surrounding snout and by having a more ex-

tensive foot web (dark canthal band surrounds snout, and

feet basally webbed, I1-0.5II1-0.5III1-1IV0.5-1V); from

M. neblina (Test, 1956) by having a spotted dorsum and

a brown upper lip (uniformly brown dorsum; uniform

cream lip); from M. olmonae (Hardy, 1983) and M. trini-

tatis (Garman, 1887), by having a crenulate free cloacal-

sheath border (smooth free border); from M. orellana Bar-

rio-Amorós, Santos & Molina, 2010, by having a less-well

developed foot web (I1-2II1-3III2-3.5IV3-2V); from M.

urticans Barrio-Amorós, Santos & Molina, 2010, by being

smaller (females 30.8 mm, males 26.9 mm) and less-well

developed foot web (I1.5

-2.5

II2-3III2.5

-4IV4-2.5

V);

from M. vulcano Barrio-Amorós, Santos & Molina, 2010,

and M.venezuelensis Manzanilla, Jowers, La Marca & Gar-

cía-París, 2007, by having more developed foot web (only a

basal web between toes III and IV in M. vulcano, and basal

webbing between toes, absent between III and IV, in M.

venezuelensis). Only two other Mannophryne with narrow

collars are le to compare. One of them (M. lamarcai) is

the most closely resembling, while the other (M. caquetio)

is the geographically closest to the new taxon. ese two

species are compared as follows.

e new species diers from M. caquetio (diagnostic

characters of the later given within parentheses) by hav-

ing a uniformly dark colored dorsum without dorsolateral

bands and without pale bands on shoulders (very dark

conspicuous dorsolateral bands reaching to inguinal re-

gion; short pale bands on shoulders); a narrow dark su-

pratympanic fringe, extending from behind the eye to

arm insertion (a not-conspicuous narrow dark fringe goes

beyond and onto the ank in adults; juveniles with same

pattern as the new species); dark stippling on lateral side

of head from tip-of-snout to level of shoulder (reticulated

pattern); dark canthal band bordering tip-of-snout but

not very dierentiated from coloration of top of the head

(canthal band not very much dierent in coloration from

other lateral parts of the head, but well dierentiated from

top of the head); loreal region dark, with narrow medial

pale line (loreal region dark with little pale spots, with-

out pale line); dark upper lip with heavy dark stippling

and inconspicuous little pale spots (upper lip darker, with

large pale spots coalescent or not); tympanum bi-colored,

upper part dark and lower part pale brown (tympanum

uniformly dark colored, with pale portions on inferior

border); anks dark brown, as dorsum, with a pale oblique

inguinal band extending to half the ank (anks stained

with brown, and bearing a disordered oblique band, spot-

ted and inconspicuous, extending onto posterior 1/3 of

ank); chin and lateral parts of throat, dark (chin not or in-

conspicuously dark-colored, inconspicuously darker along

borders of throat); dark collar with pale stippling which

is more blurred towards the sides of collar (collar broader

and more pale-stippled); extremities not conspicuously

banded (well-dened dark bands on extremities). Tip-of-

snout truncated (tip-of-snout semicircular); disk on third

nger covers 1/3 of tympanum (disk covers ½ of the tym-

panum); posterior end of tongue detached ½ of its exten-

sion from the oor of mouth (tongue almost completely

detached, about ¾ of its length); cloacal ap crenulate,

with dark border (cloacal ap slightly crenulate, bearing

small blunt papillae, with brown border, undierentiated;

lateral fringes along toes, not forming a ap folding onto

the digits (lateral fringes on toes, folding on digits); foot-

web I(1.5-2.0)-(0.5-1)II1.5-1III(1.0-1.5)-(1.0-1)IV(0.5-

1.0)-(0.5-1)V (foot-web less-extensive, I1.5-0.5II1.5-

1.0III0.5-1.5IV0.5-1.5V).

e new species diers from M. lamarcai (characters of

the later given within parentheses) by having a uniformly

dark dorsum without dorsolateral bands (dark dorsolat-

eral bands present and tending to blur backwards); loreal

region, dark, with a narrow medial pale line bordering

tip-of-snout (loreal region dark, with a medial pale line

twice as wide as that of the new species); dark canthal

band bordering tip-of-snout but not very dierentiated

from coloration of top of the head (dark canthal band

very well dierentiated from top of the head); dark up-

per lip with heavy dark stippling and inconspicuous little

pale spots (upper lip cream to pale brown); a narrow dark

supratympanic fringe (dark supratympanic band, ill-de-

ned); tympanum bi-colored, upper part dark and lower

part pale brown (tympanum bicolored, inferior 2/3 pale

brown, superior 1/3 dark); extremities without conspicu-

ous bands (extremities bearing conspicuous dark bands);

chin and lateral parts of throat, dark (chin somehow

dark-colored, without conspicuous marks on lateral sides

of throat); dark and well-dened collar with conspicuous

and well-spaced little pale spots (collar diuse, with not-

conspicuous little pale spots). Tip-of-snout truncated

(tip-of-snout from almost truncate to semicircular); pos-

terior end of tongue detached ½ of its extension from the

oor of mouth (tongue extensively, but not completely

detached; although less detached than in M. caquetio);

tarsal fold conspicuous (tarsal fold not very conspicu-

ous); cloacal ap crenulate, with dark border (cloacal ap

E. La Marca, A. Mijares-Urrutia, L. A. Saavedra & C. Gottberg

almost smooth to crenulate, with brown border, undier-

entiated); lateral aps along toes, not folding onto digits

(well-developed lateral fringes on toes, folding on digits);

foot-web I(1.5-2.0)-(0.5-1)II1.5-1III(1.0-1.5)-(1.0-1

)

IV(0.5-1.0)-(0.5-1

)V (foot-web a little more extensive,

I(1.0-2.0)-(0.5-1)II(1.5-2.0)-(1.0-1.5)III(1.5-2.0)-(1.0-

1.5)IV(0.5-1.0)-(1.0-2.0)V).

Description of holotype. Adult female with deep-

ly convoluted oviducts and mature ova (largest ovum,

2.0mm). Head slightly wider than long; interorbital re-

gion smooth; interorbital distance 1.5 times greater than

upper eyelid width; canthus rostralis ill-dened, sinuous;

nares slightly elevated, directed slightly posterolaterally;

loreal region slightly convex; snout truncated in dor-

sal view; tip-of-snout truncated, protruding not much

beyond lower jaw; length of eye about 1.5 times eye-to-

nostril distance; internarial distance about 1.7 times eye-

to-nostril distance; tympanum with elevated anterior

and inferior parts, and upper 1/3 concealed by a thick

supratympanic ridge; tympanum separated from eye by

about half its horizontal length; a single, rounded, rather

large tubercle behind angle of jaws and below tympanum;

tongue elongate, entire, posterior 1/3 not adherent to

oor of mouth; lingual papillae absent; choanae rounded,

almost completely concealed by palatal shelf of maxillary

arch; maxilla and premaxilla toothed; teeth minute.

Dorsum smooth, with low-elevated tubercles on low-

er back, although conspicuous on cloacal fold; anks

shagreened, with low tubercles towards groin; venter

shagreened; palmar tubercle single, rounded, three times

larger than thenar; thenar with lacrimal shape, about twice

longer than wide; supernumerary tubercles present; sub-

articular tubercles rounded, elevated; ngers with small-

sized pads; largest pad on third nger, covering about half

the size of tympanum when placed on it; pads as long as

wide; pad on third nger about twice wider than adjacent

phalanx; ngers free, with lateral fringes along ngers II,

III, and along internal border of I and external border of

IV; rst nger equal in length to second.

Cloacal opening above midlevel of thighs, directed ven-

trally, covered by a medium-sized cloacal ap with crenu-

late border; border of cloacal ap dark, dierentiated; su-

pra-cloacal ap thick, bearing few blunt tubercles; thighs

and shanks without conspicuous tubercles; strong tarsal

fold, not forming a ap, from base of rst toe to a point

about 3/5 of distance from tibiotarsal articulation to base

of foot; no conspicuous tubercles on tarsal fold or behind

it; outer metatarsal tubercle elevated, rounded in outline,

subconical in lateral prole; inner metatarsal tubercle

elongate, about 3 times longer than wide, about twice the

size of outer; no supernumerary tubercles; subarticular

tubercles moderate-sized, rounded to oval, attened; toes

with very little foot webbing; foot-web formula (le foot)

I2.0-0.5II1.5-1.0III1.5-1IV0.5-1V (right foot similar

webbed, except that between toes IV and V the foot web

is a little bit more extensive, starting at rst tubercle on

toe IV); toes with well-developed lateral aps; thick ap

along border of h toe, from base of pad to almost falling

short of outer metatarsal tubercle; pads wider than long;

largest pad on third toe, slightly wider than pad on fourth

toe; pad on third toe slightly less than half the width of

adjacent phalanx; heels do not overlap when thighs are

held at right angles to body axis, reaching to eye when leg

is adpressed forward.

Measurements (in mm) of the holotype. For abbre-

viations, see the section on Materials and Methods. SVL

26.7; TL 12.1; HW 9.3; HL 8.7; IOD 2.8; UEW 1.9;

NTS 1.1; EN 2.1; EYE 3.2; IN 3.5; T 1.7; ETS 4.2; TE

1.0; HAND 6.9; FOOT 10.8; TARSUS 67.5; LTF 4.7;

WP3F 0.9; WPAP 0.5.

Coloration in preservative (ethanol 70%, aer 10%

formaldehyde xation) of the holotype. Dorsum choc-

olate brown, uniform; under magnication there appear

small pale blotches, especially evident towards anterior

part of dorsum, with larger pale blotches between upper

eyelids; a diuse wide dark-brown band between upper

eyelids; a short whitish line between anterior part of up-

per eyelids; dorsum of head, in anterior part, as dark as

dorsum; upper eyelids darker than dorsum; loreal region

brown, as dark as dorsum of head; a pale line between an-

terior part of eye to naris or slightly beyond, but not bor-

dering tip of snout; upper lip brown, heavily stippled with

dark brown; border of eye membrane with a narrow band

heavily stippled with dark brown; tympanum bicolored,

upper half dark brown, lower half cream but with heav-

ily dark stippling; a narrow dark band below tympanum

to posterior border of eye, connecting to dark band on

middle anterior part of arm; short bracelets on upper part

of forearms; ulnar tubercles showing pale tips with dark

brown base; upper part of anterior extremities same as

dorsum; no bands evident on hands nor on ngers, except

for inconspicuous band on fourth nger; anks slightly

darker than dorsum with a short oblique inguinal band

extending from groin to about 2/5 of distance from groin

to arm insertion; undened pale ventrolateral blotches on

inferior part of anks.

Lower extremities, in general, with same background col-

oration as dorsum, except on posterior part of thighs which

are darker, with pale blotches forming an irregular band at

mid-level of thighs; a cream irregular blot on ventrolateral

part of thighs; the mid-level cream band tends to connect to

inguinal band through an irregular band passing alongside

New Mannophryne from Sierra de San Luis, Venezuela

cloaca and insertion of thighs; two dark brown transversal

bands on dorsal and anterior part of thighs, connecting to

longitudinal dark band from groin to knee, becoming dif-

fuse as irregular blotches on shanks; under parts of lower

and upper extremities cream-colored, with dense min-

ute stippling; transversal dark bands on dorsal surfaces of

shanks, tarsi and toes; palms and soles dark brown.

Lower part of throat cream, almost immaculate; ante-

rior part of throat and chin dark brown (densely stippled);

collar dark brown with small blotches, cream-colored; a

few small cream blotches on collar, towards right part of

body; venter dark, darker towards anterior part; posterior

part of chest cream, constituting a kind of band that con-

nects ventral cream parts of upper extremities. ere are

small pale blotches, that seem the product of loosening

skin, on dorsum, posterior extremities, and anks. Venter

dark, darker towards anterior part.

Variation in paratypes. Color variation of paratypes,

as compared to holotype, as follows: CHCG 425: In-

conspicuous pale spots on anterior part of head and be-

tween upper eyelids; inconspicuous narrow dark bands on

ngers; inguinal band ⅔ longer on le side than on right

side, extending to just ½ the ank on the later; palms and

soles slightly darkened. CHCG426: Fingers bearing dark

bands; dark spots on dorsum; cream inguinal band from

groin to anterior upper part of thigh, not connecting with

pale inguinal band; external half of ventral portion of

forearms dark brown with pale dots, as in most paratypes;

pale venter. CHCG 427: Upper lip with less dark stip-

pling than holotype; inconspicuous dark band below tym-

panum, ngers banded; cream band besides cloaca, not

connecting to inguinal band. CHCG429: Inconspicuous

pale spots on anterior part of head or between upper eye-

lids; dark bands on ngers; pale inguinal band extending

along ⅔ of distance from groin to insertion of arm; pale

venter. CHCG 430: Anterior part of venter, chest and

throat black, no indication of collar; inguinal stripe very

short, ill-dened; upper parts of legs blackened; ⅓ lower

part of tympanum pale-colored. CHCG 431: Anterior

part of venter, and throat, gray; a faint indication of collar;

inguinal stripe short, ill-dened; upper part of legs faintly

banded; lower half of tympanum pale-colored. Variation

in measurements of paratypes are indicated in Table 1.

Foot-web variation in paratypes, as compared to ho-

lotype, as follows: there is no appreciable variation in the

type series. ree specimens (CHGC 425, 429, 430) are

slightly less webbed between rst and second toe (I1.5-

0.5II); and specimen CHGC 427 has a slight dierence

between third and fourth toe (II1.0-0.5IV).

Biogeography. Mannophryne frogs are mostly endemic

species with very restricted distributions and are known

to inhabit the Holdridge’s Life Zone systems (Ewel et al.

1976) of moist and wet Premontane forests (La Marca

1992a). e type locality of Mannophryne phylidros sp.

nov. lies within a Premontane moist forest (bosque húm-

edo Premontano, according to Ewel et al. 1976). e dis-

junct distribution of the humid forests in the northwestern

part of the country, being separated by semiarid regions,

makes this locality a “biogeographical island”. As such, it

is separated from similar other regions to the south, espe-

cially those occupied by Mannophryne caquetio and M. la-

marcai (Fig. 3). Rivas et al. (2021) indicated that the area

surrounding the Sierra de San Luis is extremely dry.

Mannophryne phylidros sp. nov. is the northernmost

Venezuelan and continental South American species in

the genus (Fig. 3). is addition raises to 21 the known

Mannophryne frogs (see Frost 2023). Other 16 amphib-

ians, 15 of them anurans (see La Marca 1992b, Mijares-

Urrutia & Arends 2000, 2001, Barrio-Amorós et al. 2019,

Rivas et al. 2021, Frost 2023), were previously known

from the Sierra de San Luis: Allobates pittieri (La Marca et

al. 2004), Boana platanera (La Marca et al. 2021), Boana

sp., Bolitoglossa cf. borburata, Dendropsophus luteocellatus

(Roux, 1927), D. microcephalus (Cope, 1886), D. minu-

Table 1. Measurements (in mm) of specimens in the type series of Mannophryne phylidros sp. nov. Holotype indicated by

an asterisk. Abbreviations as explained in methods; additionally, F: Female, M: Male.

Museum number SVL HW HL T Eye EN IN Hand Foot Sex

CHCG 0425 26.4 9.4 8.3 1.8 3.8 2.2 3.7 6.6 12.5 F

CHCG0426 26.2 9.1 8.0 2.3 3.4 2.6 4.1 6.4 11.4 F

CHCG0427 25.3 9.4 8.2 1.8 3.4 2.1 3.4 6.7 12.3 F

CHCG0428* 26.7 9.3 8.7 1.7 2.8 2.1 3.5 6.6 12.4 F

CHCG0429 27.6 9.3 8.3 1,9 3.9 2,4 3.6 6.3 12.7 F

CHCG0430 23.7 8.1 7.1 1.7 3.3 2.3 3.3 6.1 10.5 M

CHCG0431 23.2 8.4 7.1 2.0 3.3 2.0 3,4 6.2 11.3 M

E. La Marca, A. Mijares-Urrutia, L. A. Saavedra & C. Gottberg

tus (Peters, 1872), Flectonotus pygmaeus (Boettger, 1893),

Hyalinobatrachium sp., Phyllomedusa trinitatis (Mertens,

1926), Pristimantis sp., Rhinella horribilis (Wiegmann,

1833), R. sternosignata (Günther, 1858), R. sclerocephala

(Mijares-Urrutia & Arends-R, 2001) and Scinax rostra-

tus (Peters, 1863). Only two (Bolitoglossa cf. borburata

and Mannophryne phylidros sp. nov.) out of the total 16

amphibians, are endemics of the Sierra de San Luis. is

11% of amphibian endemics is higher than the degree of

endemicity of the ora of this mountain range (estimated

in 6%, Steyermark 1975), although amphibian inventories

are far from complete.

ere are few mentioning of Mannophryne frogs re-

ported previously from the Sierra de San Luis. Mijares-

Urrutia & Arends (2000) mentioned a “Mannophryne

sp. 2” from middle to highest elevations of Sierra de San

Luis, coming from the localities of Cerro Galicia, Curima-

gua, Cataratas de Hueque (Municipio Petit), and near La

Chapa (Municipio Miranda). ey are considered here

as Mannophryne phylidros new species. ere is a single

collared frog specimen reported as Mannophryne a. ca-

quetio by Barrio-Amorós et al. (2010) from Sierra de San

Luis (at Cabure, San José). e later is probably the same

individual depicted in Barrio-Amorós et al. (2019), but

the lack of museum numbers or more precise information

impedes to better ascertain this. is (or these) animal(s)

may be conspecic with M. phylidros sp. nov., but proper

comparisons should await until more specimens or precise

data become available. ere is a listing of a “Mannophryne

sp. 1” coming from Sierra de San Luis in Rivas et al. (2021,

Supporting Information S4 Table). No specic locality

nor museum number(s) were given for this mentioning,

Figure 3. Map of northwestern Venezuela depicting the type locality of Mannophryne phylidros sp. nov. (red star), and localities of the

geographical closest M. lamarcai (red circles), M. caquetio (red square) and M. molinai (red triangle). Records are based on La Marca

(1996), Mijares & Arends (1999a, b), Morán et al. (2016) and Rojas-Runjaic et al. (2018).

New Mannophryne from Sierra de San Luis, Venezuela

Figure 4. Cataratas de Hueque, type locality of Mannophryne phylidros sp. nov. Photo by José R. Ochoa.

making it dicult to ascertain the taxonomic identity of

this collared frog.

Ecology. e type series of Mannophryne phylidros sp.

nov. was collected by day, during the morning hours, along

the banks of a mountain stream around the waterfalls

known as “Cataratas de Hueque”. Some males were singing

among leaf litter in very humid places (C. Gottberg, eld

notes). e leaf litter on the oor is, according to Ewel et

al. (1976) the product of some deciduous trees which lose

the leaves during the short dry season but is even percep-

tible during the rainy season.

e type locality (Fig. 4) has a sub-humid climate

type, characterized by having a mean annual precipita-

tion between 800 and 1,500 mm, with two dry seasons

(usually lasting from 4 to 6 months, mainly between De-

cember and April) and one humid season (usually lasting

from 6 to 8 months, mainly between July and Septem-

ber). e maximum mean annual temperature is around

25 °C (Pla et al. 1978, Diaz Zavala 2009). ese climate

parameters corroborate that the type locality lies within

the Premontane moist forest (bosque húmedo premon-

tano) of Ewel et al. (1976), who also indicated that the

median annual temperature of this life zone in Venezuela

is 18 to 24 °C.

Conservation. Mannophryne frogs are currently threat-

ened by a combination of factors, among which the an-

thropogenic activities seem to play the most important

role. In northwestern Venezuela, where the new species

was discovered, there are only two other collared frogs

known so far: M. caquetio and M. lamarcai. Both are en-

dangered species that are suering from habitat destruc-

tion (Mijares-Urrutia et al. 2008a, b; Barrio-Amorós et al.

2010) and both having populations with abnormalities

in their feet, attributed to pollution or some other cause

(Mijares-Urrutia et al. 2008a,b).

Populations of the Mannophryne phylidros sp. nov. are

currently favored by their occurrence in a protected area

(Juan Crisóstomo Falcón National Park), an area of roughly

20,000 hectares that was decreed in 1972. is protected

area houses 2,000 ha of evergreen forests at elevations be-

tween 200 and 1,500 m asl (Ataro 2001). In spite of this

protection, the evergreen forests in the Petit municipality,

where the type locality is situated, were considered under

the risk category of “Vulnerable” in the Red Book of Ter-

restrial Ecosystems of Venezuela (Zager & Carrasquel 2010:

285). e degree of intervention of these forests is medium-

to-high, and by its degree of threat they are considered as

endangered (Oliveira-Miranda et al. 2010: 132, Fig. 1C).

E. La Marca, A. Mijares-Urrutia, L. A. Saavedra & C. Gottberg

Rodríguez Olarte et al. (2018) evaluated the state of

conservation of the Hueque River basin, based on satellite

images, shape les of Venezuelan protected areas (sigot.

geoportalsb.gob.ve), data from the Venezuelan Red Book

of Ecosystems (Rodríguez et al. 2010), and the map of

degree of intervention of Venezuelan plant formations

(Madi et al. 2011). ey concluded that the Hueque River

basin was in the class 3 (out of 4 classes ranging from good

to very-poor conservation), meaning that its state of con-

servation is poor.

Cataratas de Hueque is a bathing resort, which may

pose some level of threat to the local population through

pollution, accidental killings or extraction of specimens.

We have second-hand information indicating that the gen-

eral area of the type locality is currently aected by nega-

tive practices such as illegal logging to make bonres, use

of soap by the bathers, use of detergents to wash vehicles,

garbage accumulation, disposal of organic and inorganic

waste, and decomposing plant and animal matter product

of Santeria rituals that have been in practice since a couple

of decades ago.

Although there is paucity of information regarding the

population status of the new species, given its reduced dis-

tribution, the degree of pollution at the type locality, and

the high amount of habitat fragmentation in the region,

we recommend applying the IUCN category of Vulner-

able (VU). We encourage to do more studies to better un-

derstand the biogeography and ecology of the new taxon,

to apply a more precise conservation category.

ACKNOWLEDGMENTS

We thank two anonymous reviewers for their useful

comments that improved the original manuscript. CG

is grateful to María José Praderio for eld assistance. Mi-

chelle Castellanos helped taking some data on museum

specimens. anks to José Rafael Ochoa who provided a

photo of the type locality (Fig. 4). Pascual Soriano grace-

fully provided room to host the original “teaching her-

petological collection” of Carlos Gottberg, which is now

an independent scientic collection (under the acronym

CHCG) within the larger CVULA collection. To all of

them our deepest gratitude. e Biogeos Foundation to

the study of biological diversity, and the REVA (Rescue

of Endangered Venezuelan Amphibians) conservation

center, provided working space, lab facilities and logistic

support. is research beneted secondarily from grants

provided by the Chessington Conservation Fund and the

Dendrobatidae Netherland to the REVA conservation

center in charge of ELM.

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APPENDIX 1

SPECIMENS EXAMINED

Mannophryne lamarcai. ULABG 4078-4087, Paratypes,

ESTADO FALCÓN: Municipio Mauroa, Cerro Socopo,

1,250 m asl.

Mannophryne larandina. ULABG 4800, Holotype, ES-

TADO LARA: Municipio Morán, Parque Nacional Dini-

ra, Hato Arriba, 1,800 m asl.

Mannophryne molinai. ULABG 7821, Paratype, ESTA-

DO YARACUY: Municipio Sucre, Sierra de Aroa, Que-

brada La Rondona, 10°18.7’N y 68°50,9’’W, 950 m asl.

Mannophryne phylidros new species. Holotype and paraty-

pes. Specimens’ numbers and data given herein.

Mannophryne trujillensis. ULABG 1160, Holotype, ESTA-

DO TRUJILLO: Quebrada Los Cedros, Paseo Los Ilus-

tres, 9°21’46.0’’N, 70°26’41.8’’W, 840 m asl.

e following specimens all come from Venezuela,

South America. Museum acronyms as follows: CVULA:

Colección de Vertebrados de la Universidad de Los An-

des, Mérida, Venezuela; ULABG: Colección de anbios y

reptiles del Laboratorio de Biogeografía, Escuela de Geo-

grafía, Universidad de Los Andes, Mérida, Venezuela.

Mannophryne caquetio. CVULA 8546-8548, CVULA

8549-8550, ESTADO FALCÓN: El Macano, Mapararí,

10 km from Churuguara, 900 m asl.

Mannophryne cf. caquetio. ULABG 5579-5583, ESTA-

DO LARA: Aguada Grande, Serranía de Parupano, 1,000

m asl.

Mannophryne cf. herminae. ULABG 4506-4508, ES-

TADO CARABOBO: La Entrada, near Valencia,

10°17’53’’N y 68°2’45’’W, 530 m asl.