Stumpffia froschaueri, Crottini & Rosa & Penny & Cocca & Holderied & Rakotozafy & Andreone, 2020

Crottini, Angelica, Rosa, Goncalo M., Penny, Samuel G., Cocca, Walter, Holderied, Marc W., Rakotozafy, Lovasoa M. S. & Andreone, Franco, 2020, A new stump-toed frog from the transitional forests of NW Madagascar (Anura, Microhylidae, Cophylinae, Stumpffia), ZooKeys 933, pp. 139-164 : 139

Stumpffia froschaueri sp. nov. Figures 1 , 3

Notes.

This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The LSID (Life Science Identifier) for this publication is: urn:lsid:zoobank.org:pub:12D91167-C0F9-4DE2-924A-586A14C62E1D. The electronic edition of this work was published in a journal with an ISSN and has been archived in the following digital repository: https://zookeys.pensoft.net/

Remarks.

The species has been previously listed as Stumpffia sp. aff. pygmaea “Sahamalaza” in Penny et al. (2016) and Stumpffia sp. aff. pygmaea Ca “Sahamalaza” (UCS) in Penny et al. (2017), Stumpffia sp. 30 in Klages et al. (2013) and S. sp. Ca30 in Rakotoarison et al (2017). The latter two names only in the case of specimens DRV6457 and DRV6451, now considered conspecific with Stumpffia froschaueri sp. nov. Specimen DRV6487 should continue to be referred as S. sp. Ca30.

Type-locality.

Anketsakely (Anabohazo Forest, Sahamalaza Peninsula, north-western Madagascar), - 14.324712S, 47.910740E; ca 169 m a.s.l., fragment of dry littoral forest included in the buffer zone of the UNESCO Sahamalaza - Îles Radama Biosphere Reserve, G. M. Rosa and L. S. Rakotozafy leg.

Material examined.

Holotype. ZSM 169/2019 (ACZCV 0940) (Fig. 1A and Fig. 3 ), adult female collected on 30 January 2013 at Anketsakely (Anabohazo Forest, Sahamalaza Peninsula, north-western Madagascar), - 14.324712S, 47.910740E; ca 169 m a.s.l., fragment of dry littoral forest included in the buffer zone of the UNESCO Sahamalaza - Îles Radama Biosphere Reserve, G. M. Rosa and L. S. Rakotozafy leg.

Paratypes. ZSM 167/2019 (ACZCV 0968) (Fig. 1D ), subadult undetermined collected on 23 January 2013 at Ankarafa Forest (Sahamalaza Peninsula, north-western Madagascar), - 14.376367S, 47.761817E; ca 191 m a.s.l.; transitional forest, by G. M. Rosa and L. S. Rakotozafy; ZSM 168/2019 (ACZCV 0966) (Fig. 1C ), juvenile undetermined collected on 23 January 2013 at Ankarafa Forest (Sahamalaza Peninsula, north-western Madagascar), - 14.376441S, 47.761838E; ca 193 m a.s.l.; transitional forest, by G. M. Rosa and L. S. Rakotozafy; ZSM 166/2019 (ACZCV 0939) (Fig. 1B ), juvenile undetermined, collected on 24 January 2013 at Ankarafa Forest (Sahamalaza Peninsula, north-western Madagascar), - 14.376241S, 47.761224E; ca 211 m a.s.l.; transitional forest, by G. M. Rosa and L. S. Rakotozafy.

Diagnosis.

A species assigned to the small-sized/miniaturised species of Clade A (Clade A1 + A2) of the genus Stumpffia based on the small size, absence of digital reduction, absence of enlarged discs on fingers and toes, occurrence in the north-west of Madagascar. The species is placed in Clade A2, which contains four nominal species ( S. madagascariensis , S. pygmaea , S. yanniki Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. analamaina Klages, Glaw, Köhler, Müller, Hipsley & Vences, 2013), most similar to S. pygmaea and S. analamaina but strongly divergent in mitochondrial and nuclear DNA from these species (and see differential diagnosis below). Although we lack bioacoustic data for this taxon, we here suggest its status as new species due to the high genetic differentiation from all other species in Clade A (pairwise 16S distance ranging from 7.8% to 12.3%), a lack of haplotype sharing in the Rag-1 analysed fragment, and a combination of morphological characters: (1) miniature to small-sized species (SVL 8.9-12.8 mm); (2) manus with four fingers (not obviously reduced in length) and pes with five toes (first toe slightly reduced in length); (3) terminal phalanges of fingers and toes without enlarged discs; (4) relative hand and foot length, HAL/SVL 0.18-0.19, FOTL/SVL 0.59-0.69; (5) dorsum smooth or very slightly tubercular; (6) brownish colouration with indistinct pattern and without contrasted ventral colouration, red colour elements on ventral side, or sharp colour border between dorsum and flanks, presence of darker blotches in the lateral portion.

Distinguished from S. be Köhler, Vences, D’Cruze & Glaw, 2010, S. hara Köhler, Vences, D’Cruze & Glaw, 2010, S. megsoni , S. staffordi , S. meikeae Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. roseifemoralis Guibé, 1974, S. nigrorubra Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. achillei Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. diutissima Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. pardus Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. edmondsi Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. fusca Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. jeannoeli Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. analanjirofo Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. grandis Guibé, 1974 and S. kibomena Glaw, Vallan, Andreone, Edmonds, Dolch & Vences, 2015 by smaller body size (8.9-12.8 mm vs. 14.4-27.9 mm); from S. miery Ndriantsoa, Riemann, Vences, Klages, Raminosoa, Rödel & Glos, 2013, S. davidattenboroughi , S. tridactyla Guibé, 1975, S. contumelia Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. tetradactyla Vences & Glaw, 1991, S. makira Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. obscoena Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. betampona Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. dolchi Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. miovaova Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. spandei Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017 and S. garaffoi Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017 by a lower degree of digital reduction. Differs from most species in Clade A1 ( S. angeluci , S. gimmeli , S. huwei , S. iharana Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. larinki Rakotoarison, Scherz, Glaw, Köhler, Andreone, Franzen, Glos, Hawlitschek, Jono, Mori, Ndriantsoa, Raminosoa, Riemann, Rödel, Rosa, Vieites, Crottini & Vences, 2017, S. mamitika , S. maledicta and S. sorata ) by slightly smaller body size (8.9-12.8 mm vs. 11-18.1 mm).

Distinguished from S. psologlossa (the type species of the genus Stumpffia ) by manus with first finger not reduced in length (vs. slightly reduced), dorsum smooth (vs. tubercular), different colour pattern (absence of distinct dark brown patches on the back; absence of the brown bands along the flanks). Different from S. analamaina by manus with first finger not reduced in length (vs. slightly reduced) and smaller relative hand length (HAL/SVL 0.18-0.19 vs. HAL/SVL 0.20-0.24). Distinguished from S. gimmeli by smaller size (SVL 8.9-12.8 mm vs. adult male SVL 14.5 mm), manus with first finger not reduced in length (vs. slightly reduced), pes with first toe slightly reduced in length (vs. first toe almost not reduced in length), terminal phalanges of fingers and toes without enlarged discs, smaller relative hand length (HAL/SVL 0.18-0.19 vs. HAL/SVL 0.19-0.23), dorsum smooth (vs. tubercular), colour pattern (absence of yellow colour on the abdomen vs. presence). Differs from S. madagascariensis by manus with first finger not reduced in length (vs. slightly reduced), pes with first toe slightly reduced in length (vs. first toe strongly reduced in length), not enlarged terminal phalanges of toes (vs. slightly enlarged), larger relative hand length (HAL/SVL 0.18-0.19 vs. HAL/SVL 0.15-0.18), dorsum smooth (vs. tubercular), a different colour pattern (absence of sharp colour border between lighter dorsum and darker flanks vs. presence). Distinguished from S. pygmaea by the first finger not reduced in length (vs. slightly reduced) and a different colour pattern (presence of indistinct dorsal patter vs. absence; presence of darker blotches in the lateral portion vs. absence). Different from S. angeluci by smaller size (SVL 8.9-12.8 mm vs. SVL 13.7-16.1 mm), terminal phalanges of toes without enlarged discs (vs. slightly enlarged discs), smaller relative hand length (HAL/SVL 0.18-0.19 vs. HAL/SVL 0.20-0.25), dorsum smooth (vs. slightly tubercular), colour (dorsal brownish vs. apricot; ventrally absence of yellow colour on the abdomen vs. presence). Distinguished from S. huwei by smaller size (SVL 8.9-12.8 mm vs. SVL 12.5-14.8), terminal phalanges of toes without enlarged discs (vs. slightly enlarged discs), colour (dorsally brownish vs. greyish to reddish brown; ventrally cream vs. yellowish) and colour pattern (presence of darker blotches in the lateral portion vs. absence). Differs from S. iharana by smaller size (SVL 8.9-12.8 mm vs. SVL 14.0-15.5 mm), terminal phalanges of toes without enlarged discs (vs. slightly to moderately enlarged discs), dorsum smooth (vs. smooth with few scattered tubercles), colour (ventrally cream vs. yellowish) and colour pattern (presence of darker blotches in the lateral portion vs. absence). Distinguished from S. larinki by terminal phalanges of fingers and toes without enlarged discs (vs. slightly to moderately enlarged discs), smaller relative hand length (HAL/SVL 0.18-0.19 vs. HAL/SVL 0.22-0.24), colour (brownish vs. iridescent copper) and colour pattern (presence of darker blotches in the lateral portion vs. absence; ventrally uniform cream vs. presence of yellow blotches). Differs from S. maledicta by smaller size (SVL 8.9-12.8 mm vs. SVL up to 16.8 mm), manus with first finger not reduced in length (vs. weakly reduced), pes with first toe slightly reduced in length (vs. distinctly reduced), terminal phalanges of toes without enlarged discs (vs. slightly enlarged discs), dorsum smooth (dorsum slightly to moderately tubercular), colour (ventrally cream vs. translucent lemon yellow) and colour pattern (indistinct pattern vs. uniform colour; presence of darker blotches in the lateral portion vs. absence). Distinguished from S. mamitika by smaller size (SVL 8.9-12.8 mm vs. male SVL 12.7-15.0 mm), manus with first finger not reduced in length (vs. slightly reduced), terminal phalanges of toes without enlarged discs (vs. slightly enlarged discs), dorsum smooth (vs. smooth with few scattered tubercles or slightly tubercular), colour (dorsally brownish vs. russet). Distinguished from S. sorata by smaller size (SVL 8.9-12.8 mm vs. SVL 15.6-16 mm), manus with first finger not reduced in length (vs. slightly reduced), terminal phalanges of toes without enlarged discs (vs. slightly enlarged discs), dorsum smooth (vs. slightly to moderately granular), colour (dorsally brownish vs. taupe) and colour pattern (ventrally uniform cream vs. presence of yellow blotches). Different from S. yanniki by manus with first finger not reduced in length (vs. moderately reduced), pes with first toe slightly reduced in length (vs. distinctly reduced); colour pattern (indistinct pattern vs. well-contrasted central dark teddy bear-shaped middorsal marking; presence of darker blotches in the lateral portion vs. absence).

Description of the holotype

ZSM 169/2019, female (Figs 1A , 3 ). Specimen in good state of preservation, third, fourth and fifth toes of the left foot removed as a tissue sample for DNA extraction. Body roundish; head wider than long, narrower than body; snout rounded in dorsal view, slightly pointed in lateral view; nostrils directed laterally, not protuberant, nearer to tip of snout than to eye; canthus rostralis straight; loreal region straight and slightly oblique; tympanum distinct, about 58% of eye diameter; supratympanic fold slightly visible; tongue broadening posteriorly, ending slightly pointy, attached anteriorly, not notched; maxillary teeth and vomerine teeth absent; choanae round. Forelimbs slender; subarticular tubercles single, distinct; outer metacarpal tubercle distinct, single, oval; palmar tubercle distinct, single, oval, smaller in size to outer metacarpal tubercle; inner metacarpal tubercle, slightly smaller than the other carpal tubercles; fingers without webbing; no fingers reduced; relative length of fingers 1 <4 <2 <3; finger tips not expanded into discs. Hind limbs slightly slender; tibio-tarsal articulation reach tympanum when adpressed forward along the body, TIBL 38% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle distinct, small, and oval; outer metatarsal tubercle absent; no webbing between toes; toes not reduced; relative length of toes1 <2 <5 <3 <4; fifth toe distinctly shorter than third. Skin on dorsum smooth, without distinct dorsolateral folds; ventral skin smooth.

Colouration of the holotype

ZSM 169/2019, female (Fig. 3 ). After six years in 70% ethanol red-brownish colouration with indistinct darker markings. Two darker dots are visible over the anterior of the scapular region (above eye), forming the anterior ends of a faint X-like marking above the scapulae. Flanks with the same colour of the dorsum but with several small cream flecks. Four dark blotches are present in the lateral portion: first blotch on tympanum, two irregular blotches between arm and legs insertion, and a large, roundish blotch on the inguinal region. A darker blotch is present also in cloaca region. Nostril indistinctly surrounded by brown; lateral head same colour as dorsum. Abdomen and pectoral region cream, flecked with brownish spots, which become more abundant on chin and ventral surface of thigh; ventral shank uniform brown; sole of foot brown, lighter brown in correspondence to the subarticular tubercles; dorsal thigh brown as dorsum, with a defined darker brown crossband; dorsal shank brown, with a defined perpendicular darker brown crossband midway along its length, and in lateral view with several small cream flecks; posterodorsal surface of shank brown; dorsal foot brown with two slightly defined perpendicular darker brown crossbands dividing the foot in three segments of equal size; toes mottled brown. Arms dorsally light brown with darker (brownish) irregular flecks that become more abundant in the lower arm; hands speckled.

Colour in life

of the holotype ZSM 169/2019, female (Fig. 1 ). Dorsum burnt umber with undefined dark brown markings (Fig. 1A ). Slightly defined interocular bar, markings in suprascapular region forming a X-shaped marking, a weak anterior chevron from the inguinal region to the mid-back (Fig. 1A ). Flank with multiple cream flecks that become increasingly cream ventrally. Cream flecks present also in lateral head. Four large ebony patches: one less distinct patch runs from the posterior margin of the eye, curving toward the anterior insertion of the arm over the tympanum, two posteriorly to the arm insertion (Fig. 1A ) and one in the inguinal region (Fig. 1 ). One ebony spot in cloacal region (Fig. 1A ). Dorsal forelimbs dark orange, with irregular brown markings, forearm brown with two brown crossbands (Fig. 1A ). The fingers are mottled brown and cream (Fig. 1A ). The dorsal legs are as the back, with one crossband at the mid-thigh and one on the mid-shank (Fig. 1A ). Lower dorsal shank with several small cream flecks. The toes are mottled ebony and brown (Fig. 1A ). Ventral skin colouration in life unknown. The iris of the holotype is copper reticulated with black, becoming metallic red close to the anterior and posterior corner of the pupil.

Variation.

Dorsum can be light brown (Fig. 1C, D ) and have a dark cream vertebral line (Fig. 1B ). Dorsal markings can be more irregular (Fig. 1B, D ). Lateral cream flecks can be present also on tympanum (Fig. 1B ). The lateral ebony spots can be less defined and be partially fused (cf. Fig. 1B-D ). Toes are mottled ebony and brown and the fourth toe can have a white annulus before the terminal phalange (Fig. 1B ).

For variation in measurements among specimens, see Table 1 . ZSM 168/2019 (ACZCV 0966) and ZSM 166/2019 (ACZCV 0939), too small to be measured for all measurements. All examined specimens agree strongly with the holotype (although they are distinctly smaller in size) in hand and feet morphology, in having a smooth or very slightly tubercular dorsum and on the presence of an ebony spot over the tympanum, in the inguinal region and in cloacal region. Colour and colour pattern is variable. The degree of visibility of hindlimb crossbands varies strongly, but they are present to some degree in all specimens.

ZSM 167/2019 (ACZCV 0968) (Fig. 1D ) is grey on dorsum with a few dark brown markings, absence of the X-marking on suprascapular region, the two blotches between the arm and leg insertions are fused, colourations on arm, hand, legs and feet less dark and markings less distinct (crossbands only slightly distinct), tympanum distinct, ca. 50% of eye diameter; ZSM 168/2019 (ACZCV 0966) (Fig. 1C ) dorsally dark brownish, the two blotches between the arm and leg insertions are fused, abdomen, pectoral region and chin darker (with more brown flecks), tympanum distinct, approximately the same size of the eye; ZSM 166/2019 (ACZCV 0939) (Fig. 1B ) dorsally brown-greyish, with a greyish vertebral line, the two blotches between the arm and leg insertions are fused, abdomen, pectoral region and chin darker (with more brown flecks).

Etymology.

The species name is a patronym in the genitive case, honouring Christoph Froschauer (ca. 1490 - April 1564). His family name means "the man from the floodplain full of frogs". Froschauer was the first, and European wide renowned, printer in Zürich and he used to sign his books with a woodcut showing frogs under a tree in a landscape. He was notably known for printing Conrad Gessner’s encyclopaedic " Historia animalium " and the “Zürich Bible", a complete translation into German of the Bible several years before Luther’s Bible appeared. Froschauer published works by Zwingli, Bullinger, Gessner, Erasmus von Rotterdam and Luther during his lifetime. His activity represents the nucleus of the Orell Füssli publishing house, which celebrated its 500th birthday on 9th November 2019, which is the date he was given citizenship in Zürich as a gift for his art.

Distribution, conservation and proposed IUCN Red List status.

This species is known only from north-western Madagascar and apparently restricted to three forest blocks embedded in a matrix of highly degraded habitat: 1) Anketsakely (within Anabohazo Forest block), 2) Ankarafa Forest, and 3) Angorony Forest. The latter locality is assigned to this species based on the DNA sequences deposited in GenBank (accession numbers KC351357 and KC351351) that correspond to specimens DRV6457 and DRV6451 (not examined by us). This forest fragment lays in close proximity to Sahamalaza Peninsula and it is ca. 30 km away from Anketsakely. The range encompasses elevations from 100-340 m above sea level. The suggested conservation status was assessed using the guidelines of the IUCN Red List (IUCN Standards and Petitions Subcommittee 2019). If suitable habitat is considered to be all areas of Ankarafa Forest, Anabohazo Forest (where Anketsakely lies; likely an over-estimate) and Angorony Forest, then the EOO (extent of occurrence) totals 246 km2. If plots with a scale of 2 km2 are used to estimate AOO (area of occupancy), then this species occurs within 6 km2 of habitat. Similar to the recently described Boophis ankarafensis ( Penny et al. 2014) and the other microendemic species of the Sahamalaza peninsula (e.g., Cophyla berara ), this species is likely to be restricted to the typical transitional forest present in this area. Although two of these forest patches are now part of a UNESCO Biosphere Reserve (the Sahamalaza-Îles Radama Biosphere Reserve), forest border patrolling is lacking and forest is still under strong pressure from slash-and-burn activities and timber harvesting ( Penny et al. 2016). Furthermore, the species’ apparent preference for intact forest is likely to limit gene flow between the three known populations. Thus, it is important to establish whether the distribution of this species occurs outside these areas. Habitat loss and fragmentation of these forest fragments is likely the greatest threat to this species’ survival, as indicated by the destruction of the nearby Analavory Forest in 2004 after a man-made fire. Given this on-going destruction of suitable habitat, population declines can be expected to continue unless some remedial action is taken. Thus the species should qualify as Critically Endangered under criterion B (B2ab (i, ii, iii, iv, v) of the IUCN Red List (IUCN Standards and Petitions Subcommittee 2019).

Natural history.

In Anketsakely and Ankarafa this species has been found only in areas with relatively undisturbed forest. Active individuals were found during the day within the leaf-litter on the forest floor, where discreet calling males were also detected.

Call.

Unavailable for analysis. The call of this species is quite inconspicuous and very difficult to locate (G.M. Rosa pers. obs.).

Figures

Figure 1. Life colouration of Stumpffia froschaueri sp. nov. A dorsolateral view of holotype ZSM 169 / 2019 (ACZCV 0940) from Anketsakely (Anabohazo Forest) B dorsolateral view of paratype ZSM 166 / 2019 (ACZCV 0939) from Ankarafa Forest C dorsolateral view of paratype ZSM 168 / 2019 (ACZCV 0966) from Ankarafa Forest D dorsolateral view of paratype ZSM 167 / 2019 (ACZCV 0968) from Ankarafa Forest. Pictures by Goncalo M. Rosa

Figure 3. Voucher colouration of Stumpffia froschaueri sp. nov. A dorsal and B ventral views of the preserved holotype of Stumpffia froschaueri (ZSM 169 / 2019; ACZCV 0940), and ventral surfaces of C hand and D foot. Scale bars: 10 mm; hand and foot not to scale.