identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
2791B92FE8DC53B995AFC3D593B5D87F.text	2791B92FE8DC53B995AFC3D593B5D87F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne anatiala Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne anatiala sp. nov.</p><p>Figures 2, 4 –7, 30, 31, 46, 47</p><p>Remark.</p><p>MRSN A 5477, a specimen housed in the same jar as MRSN A 5524, the specimen of R. maraorao sp. nov. discussed above, likewise allegedly comes from Tsingy de Bemaraha in western Madagascar, but agrees morphologically with R. anatiala . We are confident that this specimen, too, was collected in northeastern Madagascar. However, as for R. maraorao, we do not include the mislocated specimen in the diagnosis of the species (also because we lack DNA sequence data on it), in order to avoid introducing errors, but recognise that it is probably referrable to R. anatiala sp. nov. nonetheless.</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 1 ’ by Vieites et al. (2009) and ‘ Rhombophryne sp. Ca 01 ’ by Scherz et al. (2016 b) and Belluardo et al. (2022), and ‘ Rhombophryne sp. 1 ’ and ‘ UCS “ Sp. 1 ” ’ by Carné and Vieites (2024).</p><p>Justification of the new species.</p><p>The new species was found to be the sister species of R. sonaliae sp. nov. (described above) with high support based on a multigene phylogeny (Belluardo et al. 2022). The genetic divergences between these two lineages is&gt; 3.8 % uncorrected p distance in the 16 S 3 ’ fragment, and thus at a level typically indicative of species-level differentiation in Malagasy frogs (Vieites et al. 2009). We found absence of haplotype sharing between R. anatiala sp. nov. and R. sonaliae sp. nov. and between R. anatiala sp. nov. and any other Rhombophryne species in the analysed fragment of the nuclear gene RAG 1 (Fig. 4). Morphologically, the two lineages differ in relative toe length (see Diagnosis below). Taken together, and based on other example of pairs of species of Malagasy frogs allopatrically distributed in the North East vs. Northern Central East of Madagascar, we consider the hypothesis of these two lineages representing two distinct species as most likely, although we acknowledge the need for in-depth study of their contact zone. In the highly unlikely case future studies will find the two nomina to be subjective synonyms, we recommend as first reviser action to give priority to sonaliae over anatiala.</p><p>Holotype.</p><p>MRSN A 2696 (RJS 0597) an adult female, collected on 15 February 2002 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.9403&amp;materialsCitation.latitude=-15.4177" title="Search Plazi for locations around (long 49.9403/lat -15.4177)">‘Camp 7’ Antsiranana Faritany</a> (15.4177°S, 49.9403°E, ca. 800 m a. s. l.), Amparihy, Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by J. E. Randrianirina (Figs 30, 31, 47) .</p><p>Paratypes.</p><p>Two specimens: MRSN A 4544 (FAZC 10314; Fig. 46), and MRSN A 4600 (FAZC 10304), two ovigerous adult females, collected on 11 and 12 December 1999 (respectively) at ‘ Camp 5 ’ Menamalona (15.3812°S, 49.9878°E, ca 800 m a. s. l.), Ilampy Corridor, Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by F. Andreone and J. E. Randrianirina.</p><p>Assignment of specimens.</p><p>The paratype MRSN A 4544 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ FJ 559295; Vieites et al. 2009). COI is so far not available for this species. The 16 S 3 ’ sequence of the holotype is identical to that of this specimen (OL 780571) and has also been sequenced for 16 S 5 ’ (OL 780600). Both paratypes conform well with the holotype in morphology (MRSN A 4544 could only be examined from photographs; Fig. 46). The assignment of MRSN A 4600 is further confirmed by molecular taxonomic identification (Fig. 2). MRSN A 2696 and MRSN A 4544 are present in the RAG 1 haplotype network (Fig. 4).</p><p>A further specimen, MRSN A 2584 (RJS 0348), is recovered by molecular taxonomic identification as conspecific, but could not be examined morphologically. Database records indicate that this should be a Spinomantis specimen, indicating some kind of specimen or metadata mix-up. To avoid confusion, we have not designated this specimen as a paratype.</p><p>Definition.</p><p>Within the R. laevipes group, this species is sister to R. sonaliae sp. nov. (described above) based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium to large body size, adult SVL 34.3–37.9 mm; (2) head wider than long (HW / HL 1.56–1.76); (3) moderately small eyes (ED / HL 0.31–0.35); (4) relatively long hindlimbs (HIL / SVL 1.51–1.58); (5) small inner metatarsal tubercle (IMTL / FOL 0.11–0.13); (6) outer metatarsal tubercle absent; (7) second finger subequal to fourth; (8) third toe distinctly longer than fifth, reaching the second phalange of the fourth toe; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of large white inguinal spots on a black background, and a light-brown dorsolateral stripe broadening from the eye towards the inguinal region; (13) presence of large white or yellowish spots on a black background on the posterior thigh and ventral surfaces of the shanks; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne anatiala sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ G’ at site 259, ‘ C’ at site 654, and ‘ G’ at site 739. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. anatiala sp. nov. can usually be distinguished from all species except R. sonaliae sp. nov. described above by the presence of a light brown dorsolateral stripe expanding from the eyes towards the inguinal region. However, as this trait is not always present (Fig. 46 A), additional traits are necessary for a reliable diagnosis.</p><p>Rhombophryne anatiala sp. nov. can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by smooth skin (vs rough), absence of chin barbels (vs general presence), longer hindlimbs (HIL / SVL 1.51–1.58 vs 1.10–1.41); from R. mangabensis and R. savaka by larger body size (SVL 34.3–37.9 mm vs 18.7–23.2 mm), and presence of multiple large light-coloured inguinal spots (vs black inguinal ocelli in R. savaka); from R. mavokely sp. nov., R. ornata, and members of the R. ellae species group by the absence of reddish orange colouration on the posterior thigh and in the inguinal region (vs presence); from members of the R. minuta species group by shorter hindlimb length (HIL / SVL 1.51–1.58 vs 1.73–1.84), and less gracile habitus; from R. proportionalis by much larger adult body size (SVL 34.3–37.9 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence).</p><p>Within the R. laevipes species group, R. anatiala sp. nov. can be distinguished from R. laevipes by shorter hindlimbs (HIL / SVL 1.51–1.58 vs 1.62–1.86), and larger inner metatarsal tubercle (IMTL / FOL 0.11–0.13 vs 0.08–0.11); from R. botabota by larger body size (SVL 34.3–37.9 mm vs 24.2–32.2 mm), and presence of large light-coloured spots in the inguinal region and on the posterior thigh and ventral shank (vs absence); and from R. nilevina by smaller body size (SVL 34.3–37.9 mm vs 41.2–57.2 mm). As far as is known, Rhombophryne anatiala sp. nov. can only be distinguished morphologically from R. sonaliae sp. nov. by the longer third toe, reaching the midpoint of the second phalange of the fourth toe (vs reaching or barely surpassing the joint of the first and second phalange of the fourth toe). For distinction from the last newly described member of this species group, see its description below.</p><p>Description of holotype.</p><p>Specimen in a moderately good state of preservation, though slightly soft (Fig. 30). An incision made along the left flank to assess the sex, and part of the tongue taken as a tissue sample. Small ova, ovaries, and fallopian tubes clearly visible.</p><p>Body robust and rotund. Head wider than long (HW / HL 1.76). Eyes quite small (ED / HL 0.31), slightly depressed, pupils small, oval. Snout rounded in dorsal and slightly truncate in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril equidistant between tip of snout and eye, slightly protuberant. Tympanum indistinct, round, TDH / ED 0.59. Supratympanic fold distinct, slightly raised, dark brown, running straight from posterior corner of eye over tympanum, curving gently ventrally posterior to the tympanum to above the forearm’s insertion. Lateral head similar in colour to dorsal head, except for the supratympanic fold. Superciliary spines absent, one subtle light dot above each eye. Dorsal skin smooth in preservative. Tongue partly taken as tissue sample, originally broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth curved, without pronounced height variation, separated at the midline by a small gap. Choanae round and surrounded by pigment, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.380), strongly built. Fingers without webbing, moderately long, relative lengths 1 &lt;2 &lt;4 &lt;3 on left hand, 1 &lt;4 &lt;2 &lt;3 on right, in both cases distinctly so (Fig. 31). Finger tips not expanded, rounded. Finger subarticular tubercles distinct, whitish, flat; inner metacarpal tubercle broad and rounded, outer metacarpal / palmar tubercle indistinct, greyish. Hind limb strong, tibiotarsal articulation reaching between tympanum and eye when adpressed anteriorly along the body. TIBL / SVL 0.428. Inner metatarsal tubercle present, oblong, whitish, distinctly raised. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third, third toe extending to the middle of the second phalange of the fourth toe; subarticular tubercles slightly distinct; toe tips slightly enlarged, rounded (Fig. 31).</p><p>After 19 years in preservative, the base colour is a coffee brown across the dorsum, and a darker brown on the limbs. The dorsum is mottled brown, with a clear V-shaped marking over the suprascapular region. Behind the eyes, running towards the inguinal region, are lighter brown wavy patterns riddled with tiny white points, very similar to R. sonaliae sp. nov. No distinct colour difference between the lateral and dorsal head except on the supratympanic fold, which is dark brown. The inguinal region bears several large white spots which also extend onto the anterior thigh, on a dark brown background. There are also several white spots on the posterior thigh, ventral shank, and dorsal foot. The ventral thigh is mostly cream, mottled with light brown. The ventral feet are a light brown. A light annulus is present before the terminal phalange of each finger and toe. The legs have distinct crossbands on the thigh, shank, tarsus, and foot. The arms have a single crossband on the lower arm. The palmar surface is grey-brown. The chin is light brown, the ventral abdomen mostly cream, with a few brown flecks. The cloaca region is unmarked, and there is no blackish bar above it.</p><p>Variation.</p><p>SVL 34.3–37.9 mm (n = 2). For variation in other measurements see Table S 6. MRSN A 4600 has a distinctly different skin texture in preservative, and is also much less mottled than the holotype. It lacks the V-shaped marking of the dorsum and the crossbands on the legs of the holotype. MRNS A 4544 likewise lacks the V-shaped marking on the dorsum, but does have the crossbands on its legs.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>The Malagasy adjective anatiala meaning ‘ in the forest’, in reference to the habitat of this species in the rainforests of Masoala. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is only known from (1) Amparihy (type locality) and (2) Menamalona on the Masoala Peninsula. Elevation: 778–800 m a. s. l.</p><p>Natural history.</p><p>Little is known of the ecology of this species, but we presume that it is similar to its sister species, R. sonaliae sp. nov., in its ecology and behaviour; see the account of that species, above.</p></div>	https://treatment.plazi.org/id/2791B92FE8DC53B995AFC3D593B5D87F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
401DF3D9C1BD556CB560AB4F04F56228.text	401DF3D9C1BD556CB560AB4F04F56228.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne Boettger 1880	<div><p>Rhombophryne Boettger, 1880</p><p>Type species.</p><p>Rhombophryne testudo Boettger, 1880</p><p>Synonymy.</p><p>Mantiphrys Mocquard, 1895 . Type species: Mantiphrys laevipes Mocquard, 1895 .</p><p>Definition.</p><p>A cophyline microhylid frog genus characterised by the following suite of characters: (1) a procoelous vertebral column, (2) knobbed or unornamented terminal phalanges (i. e., not Y- or T-shaped), (3) unexpanded or weakly expanded terminal discs of fingers and toes, (4) presence (very occasional absence) of a curved clavicle, and (5) divided vomer usually bearing teeth on the postchoanal portion.</p><p>Diagnosis.</p><p>Within Madagascar, only Scaphiophryne, Stumpffia, and Plethodontohyla can be confused with Rhombophryne . Scaphiophryne are stouter and rounder in general, often with expanded terminal discs (vs knobbed or unornamented in Rhombophryne), and have an undivided vomer (vs divided into pre- and post-choanal portions). Stumpffia have straight or absent clavicles (vs curved or absent), typically lack teeth on the maxillae (vs generally present), lack teeth on the postchoanal vomers (vs generally present). Several members of the genus Plethodontohyla are arboreal and can easily be distinguished by presence of Y- or T-shaped terminal phalanges on fingers and toes; terrestrial Plethodontohyla can be distinguished by the combination of absence of clavicles and absence of frontoparietal-nasal contact (vs frontoparietal-nasal contact present in all Rhombophryne lacking clavicles) (see also the key provided in the appendices of Scherz et al. 2016 b).</p><p>Distribution.</p><p>Limited primarily to the northern half of Madagascar, with the southern-most occurrence in Ivohibe in South Central Eastern Madagascar (R. sp. ‘ Ivohibe’). Apparently absent from western and southern Madagascar.</p></div>	https://treatment.plazi.org/id/401DF3D9C1BD556CB560AB4F04F56228	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
99EB634C94B15AF6AE376825A2250205.text	99EB634C94B15AF6AE376825A2250205.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne botabota Scherz 2016	<div><p>Rhombophryne botabota Scherz et al., 2016</p><p>Figures 2, 4 –7, 30, 31, 36 –38</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 1 ’ by Wollenberg et al. (2008), ‘ Rhombophryne sp. 2 ’ by Vieites et al. (2009), and ‘ Rhombophryne sp. Ca 2 ’ by Perl et al. (2014). Carné and Vieites (2024) treated specimens from Makira as ‘ DCL “ Makira ” ’.</p><p>Holotype.</p><p>ZSM 358/2005 (FGZC 2896), an adult male, collected on 18 February 2005 at ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.7433&amp;materialsCitation.latitude=-14.4364" title="Search Plazi for locations around (long 49.7433/lat -14.4364)">Camp Simpona’</a> (14.4364°S, 49.7433°E, 1326 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by F. Glaw, M. Vences, and R. D. Randrianiaina (Figs 30, 31, 37) .</p><p>Paratypes.</p><p>Six specimens: ZSM 538/2009 (ZCMV 11473), ZSM 539/2009 (ZCMV 11474), two presumably immature males, collected on 21 June 2009 at ‘ Camp 1 ’ Angozongahy, (15.4370°S, 49.1186°E, 1009 m a. s. l.), western side of the Makira Plateau, Analanjirofo Region, northeastern Madagascar, by M. Vences, D. R. Vieites, F. Ratsoavina, R. Randrianiaina, E. Rajeriarison, T. Rajofiarison, and J. Patton; MRSN A 2956 (FN 7164), MRSN A 2640 (FN 7238), MRSN A 2954 (FN 7281), MRSN A 2955 (FN 7300), two males and two females, collected on 9–15 December 1997 at ‘ Camp 2 ’ Andranomadio (14.5304°S, 49.4383°E, 860 m a. s. l.), Ambolokopatrika-Betaolana Forest, Sava Region, northeastern Madagascar, by F. Andreone, G. Aprea, and J. E. Randrianirina.</p><p>Referred specimens.</p><p>UADBA -A- ZCMV 15158 (Fig. 36 L – N), ZSM 515/2016 (ZCMV 15241; Fig. 36 J, K), and ZSM 513/2016 (ZCMV 15159; Fig. 36 O – Q), three juveniles, and ZSM 514/2016 (ZCMV 15160; Fig. 36 D – F), ZSM 523/2016 (ZCMV 15216; Fig. 36 G – I), and ZSM 524/2016 (ZCMV 15279; Fig. 36 A – C), three unsexed adults, all collected on 18–20 November 2016 in the vicinity of ‘ Camp Simpona’ (ca 14.4366°S, 49.7434°E, 1325 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, A. Rakotoarison, M. Bletz, M. Vences, J. Razafindraibe, and A. Razafimanantsoa; MNCN - uncatalogued (DRV 5836), an unsexed individual, collected on 25 June 2009 at ‘ Camp 1 ’ Angozongahy, (15.4370°S, 49.1186°E, 1009 m a. s. l.), western side of the Makira Plateau, Analanjirofo Region, northeastern Madagascar, by M. Vences, D. R. Vieites, F. Ratsoavina, R. Randrianiaina, E. Rajeriarison, T. Rajofiarison, and J. Patton.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ EU 341102 / FJ 559297; 16 S 5 ’ EU 341102; Vieites et al. 2009) and COI barcoding (COI KF 611585; Perl et al. 2014). All paratypes and referred specimens conform well with the holotype in morphology. MRSN A 2640, MRSN A 2956, ZSM 513/2016, ZSM 514/2016, ZSM 515/2016, ZSM 524/2016, ZSM 538/2009, ZSM 539/2009, MNCN - DRV 5836, and UADBA -A- ZCMV 15158 are further confirmed by molecular taxonomic identification (Fig. 2). MRSN A 2956 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. laevipes group, this species appears to be sister to a clade consisting of R. sonaliae sp. nov. and R. anatiala sp. nov. (described below) based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 24.2–32.2 mm; (2) head wider than long (HW / HL 1.44–1.69); (3) moderately small eyes (ED / HL 0.33–0.38); (4) moderately long hindlimbs (HIL / SVL 1.41–1.75); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.13); (6) outer metatarsal tubercle absent; (7) second finger shorter than or equal in length to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of distinct inguinal spots (sometimes dark brown lines converging anterodorsally); (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 4 analysed calls by one individual) tonal, with (17) call duration 161–201 ms, (18) inter-call interval 2529–2716 ms, and (19) dominant frequency 1256–1291 Hz.</p><p>Rhombophryne botabota is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ G’ at site 304, ‘ T’ at site 568, and ‘ A’ at site 682. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. botabota can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by smooth skin (vs rough), absence of chin barbels (vs general presence), longer hindlimbs (HIL / SVL 1.41–1.75 vs 1.10–1.41); from R. mangabensis by larger body size (SVL 24.2–32.2 mm vs 19.6–23.2 mm), and smooth skin (vs finely granular with scattered warts); from R. savaka by larger body size (SVL 24.2–32.2 mm vs 18.8–20.9 mm), absence of a diastema in the vomerine teeth (vs presence), and absence of inguinal spots (vs occasional presence); from R. mavokely sp. nov., R. ornata, and members of the R. ellae species group by the absence of reddish orange colouration on the posterior thigh and in the inguinal region (vs presence); from members of the R. minuta species group by generally shorter hindlimb length (HIL / SVL 1.41–1.75 vs 1.73–1.84), and less gracile habitus; from R. proportionalis by much larger adult body size (SVL 24.2–32.2 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence).</p><p>Within the R. laevipes species group, see the account of R. laevipes above for distinction from that species. Rhombophryne botabota can be distinguished from R. nilevina by smaller body size (SVL 24.2–32.2 mm vs 41.2–57.2 mm), absence of white spots in the inguinal region and on the posterior thigh and ventral shank (vs sometimes present), and tonal (vs pulsed) advertisement call that is shorter (call duration 161–201 ms vs 538–598 ms) and higher in dominant frequency (1256–1291 Hz vs 526–534 Hz). For distinction from newly described members of this species group, see their respective descriptions below.</p><p>Variation.</p><p>SVL ranges from 24.2–32.2 mm; for variation in other measurements, see Table S 6. Skin texture of the newly collected material was surprisingly variable in life, ranging from smooth to tubercular (Fig. 36). The supratympanic fold was consistently highly pronounced and always dark in colour. Colouration was noted to vary substantially in the type series by Scherz et al. (2016 a), most importantly including the occasional presence of spots on the posterior thigh, and presence of a dark inguinal ‘ stripe’ in almost all individuals. Many also possessed a pair of dark spots on top of the head behind the eyes (Fig. 36 J, M, P).</p><p>Call.</p><p>The allocation of the analysed recording to R. botabota is slightly tentative, as the recorded individual was not seen calling. However, R. botabota was the only Rhombophryne species found at the place of call emission (see Scherz et al. 2016 a). The advertisement call recorded on 24 June 2009 (12: 00 h) at Angozongahy site, Makira (estimated air temperature 22–24 ° C), consists of a single tonal note of moderate duration repeated at regular intervals, but apparently not arranged in regular call series. Calls lack any frequency modulation, but exhibit moderate amplitude modulation, with call energy increasing from the beginning of the call and reaching its maximum at approximately half of the call’s duration (Fig. 38). Numerical parameters of 4 analysed calls are as follows: call duration (= note duration) 161–201 ms (180 ± 19 ms); inter-call intervals 2529–2716 ms (2638 ± 97 ms); dominant frequency 1256–1291 Hz (1274 ± 16 Hz); prevalent bandwidth 1100–3200 Hz; harmonic frequency bands recognizable at about 630, 1910, 2580 and 3160 Hz. Calls from the same recording have been described by Scherz et al. (2016 a) using a slightly different terminological scheme. Slight differences in temporal measurements when compared to Scherz et al. (2016 a) result from different recognition of the ‘ tapering tail’ in calls, which we here consider to likely represent an echo effect.</p><p>Etymology.</p><p>The Malagasy adjective botabota meaning ‘ chubby’, in reference to the body shape of this species. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>Known from (1) Marojejy National Park (type locality), (2) Ambolokopatrika-Betaolana, and (3) Makira in northeastern Madagascar (Scherz et al. 2016 a). Elevation: 860–1326 m a. s. l. Records from 750 ± 200 m a. s. l. by Rakotoarimalala and Raselimanana (2023) cannot be verified as we have not examined material and have no DNA sequences, so they are not included here.</p><p>Natural history.</p><p>Rhombophryne botabota is a semi-fossorial or detriticolous species found in mid-elevation humid forest. In the vicinity of ‘ Camp Simpona’ in November 2016, we found R. botabota to be moderately abundant during surveys. Specimens were routinely found by raking over quadrats of leaf litter, or collecting the quadrats in rice bags and sorting through them by hand. Previously, most specimens had been collected using pitfall traps (Scherz et al. 2016 a). Some calls were heard, but no individuals could be observed calling. Paratype ZSM 538/2009 was recorded by Scherz et al. (2016 a) to have a subulinid snail in its gut, making it the only record of gastropod consumption by a Rhombophryne species.</p></div>	https://treatment.plazi.org/id/99EB634C94B15AF6AE376825A2250205	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
4167246972065DB8ABD633B37DF6B936.text	4167246972065DB8ABD633B37DF6B936.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne coronata (Vences & Glaw 2003)	<div><p>Rhombophryne coronata (Vences &amp; Glaw, 2003)</p><p>Figures 2, 4 –7, 67 –69, 71, 72, 74</p><p>Plethodontohyla coronata Vences &amp; Glaw, 2003</p><p>Rhombophryne coronata — Glaw and Vences (2007)</p><p>Remark</p><p>. Belluardo et al. (2022) reported the candidate species Rhombophryne sp. Ca 13, based on specimen KU 340732 (CRH 457; Fig. 71 A), an adult female (containing 12 eggs), collected on 5 January in 2015 in Vohidrazana Private Reserve (18.9761°S, 48.4993°E, ca. 1150 m a. s. l.), Alaotra-Mangoro Region, central eastern Madagascar, by C. R. Hutter and S. Lambert. We here refrain from treating the taxonomy of that candidate species, because (1) it is very closely related to R. coronata, (2) we have not been able to identify any morphological features that distinguish these lineages (they are extremely similar both externally and osteologically; compare Figs 72 and 73), and (3) we lack bioacoustic data from R. sp. Ca 13. Based on molecular phylogenetic data Rhombophryne coronata and R. sp. Ca 13 form a highly supported clade (Fig. 2). Our ASAP species delimitation retained R. sp. Ca 13 as a distinct species. However, data from the nuclear gene RAG 1 reveal haplotype sharing between Rhombophryne coronata and R. sp. Ca 13 (Fig. 4). Altogether, this is weak evidence for species-level distinction, and we need additional material from more sites in this area in order to clarify the taxonomy of these frogs.</p><p>Holotype.</p><p>ZFMK 57459, an adult male, collected on 19 February 1994 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.033&amp;materialsCitation.latitude=-19.167" title="Search Plazi for locations around (long 48.033/lat -19.167)">Ankeniheny</a> (19.167°S, 48.033°E, 900 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by F. Glaw, N. Rabibisoa, and O. Ramilison .</p><p>Paratypes.</p><p>Two specimens: ZSM 694/2001 (FGMV 2001.199), and UADBA -A- FGMV 2001.200, two adult males, collected on 16 February 2001 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.933&amp;materialsCitation.latitude=-18.917" title="Search Plazi for locations around (long 47.933/lat -18.917)">Mandraka</a> (18.917°S, 47.933°E, 1220 m a. s. l.), Analamanga Region, North Central East Madagascar, by M. Vences and D. R. Vieites .</p><p>Referred specimens.</p><p>ZSM 473/2005 (ZCMV 2222) and ZSM 474/2005 (ZCMV 2223), two unsexed adults, collected in February 2005 in a pitfall trap in Andasibe (18.9362°S, 48.4122°E, 939 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by R. Dolch and collaborators.</p><p>Assignment of specimens.</p><p>The paratype ZSM 694/2001 (FGMV 2001.199) is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341103; Vieites et al. 2009). A tissue sample with specimen number MV 2001. A 1 (corresponding to either ZSM 694/2001 or UADBA -A- FGMV 2001.200) is used as the reference for this species in COI barcoding (COI KF 611586; Perl et al. 2014). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of specimens ZSM 473/2005, and ZSM 474/2005 is further confirmed by molecular taxonomic identification (Fig. 2). MV 2001. A 1, ZSM 473/2005, and ZSM 474/2005 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is sister to the unconfirmed candidate species R. sp. Ca 13 in a clade sister to all other nominal species based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 19.4–23.2 mm; (2) head wider than long (HW / HL 1.51–1.55); (3) moderately large eyes (ED / HL 0.41–0.46); (4) relatively short hindlimbs (HIL / SVL 1.28–1.47); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.13); (6) outer metatarsal tubercle absent; (7) second finger subequal to fourth; (8) third toe distinctly longer than fifth, first toe reduced to a nub in some specimens; (9) dorsal skin finely granular; (10) absence of chin barbels; (11) presence of three small, equal-sized superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne coronata is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 275, ‘ A’ at site 288, and ‘ A’ at site 381; in 16 S 3 ’, ‘ A’ at site 967, ‘ T’ at site 1182, and ‘ A’ at site 1322.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. coronata can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. Within the R. serratopalpebrosa species group, it is unique in possession of a reduced first toe. For further distinction from R. serratopalpebrosa and R. guentherpetersi, see the diagnosis of those species, above. It can further be distinguished from R. vaventy by much smaller adult body size (SVL 19.4–23.2 mm vs&gt; 50 mm), and finely granular dorsal skin (vs rugose); from R. ornata by much smaller adult body size (SVL 19.4–23.2 mm vs up to 33.0 mm), three superciliary spines (vs two), and absence of reddish colouration on the thighs and inguinal region; from R. tany by slightly smaller adult body size (SVL 19.4–23.2 mm vs 24.6 mm), indistinct supratympanic fold (vs distinct), and shorter hindlimb (HIL / SVL 1.28–1.43 vs 1.56); from R. regalis by absence of S-shaped dermal postnasal fold, superciliary spines being of roughly equal size (vs three spines differing in size: 1 st large, 2 nd medium, 3 rd diminutive), and shorter relative tibia length (TIBL / SVL 0.35–0.39 vs 0.47–0.56); from R. diadema by superciliary spines being of roughly equal size (vs posterior-most spine being diminutive compared to the other two), smaller relative tympanum size (TDH / ED 0.37–0.59 vs 0.59–0.64), and shorter relative tibia length (TIBL / SVL 0.35–0.39 vs 0.44–0.46). For distinction from new species described herein, see accounts below.</p><p>A detailed description of the species is given in Vences and Glaw (2003) but we here add more data on the variation and distribution of the species, as well as a redescription of the call.</p><p>Variation.</p><p>SVL 19.4–23.2 mm (n = 4). For variation in other measurements, see Table S 10. Specimens are very consistent in anatomy, except one specimen (ZSM 474/2005, ZCMV 2223), which differs as follows: the fourth finger is longer than the second, but only slightly; in this specimen, the first finger is also visibly reduced, to a greater extent than in other specimens examined, whereas the first toe is not reduced, unlike other specimens examined.</p><p>Our measurements of the holotype of R. coronata differ in some points quite substantially from the original measurements of Vences and Glaw (2003), especially HL (5.7 mm vs 7.3 mm), NSD (1.3 vs 1.6 mm), TDH (1.4 vs 1.7 mm), HAL (4.7 vs 5.0 mm), FORL (13.2 vs 12.6 mm), FOTL (14.7 vs 14.4 mm), HIL (33.0 vs 30.6 mm), and IMCL (0.9 vs 1.1 mm). These differences are undoubtedly due to differences in measurement style; hence, we emphasise that all measurements reported in this paper were taken by the same measurer (MDS) based on the measurement scheme shown in Fig. 1.</p><p>Rhombophryne sp. Ca 13 is extremely similar to R. coronata morphologically (Fig. 71). The one ovigerous adult female specimen we examined (KU 340732, CRH 457) differed from specimens of R. coronata by a longer forelimb (FORL / SVL 0.71 vs 0.32–0.59; FARL / SVL 0.48 vs 0.37–0.43), and hindlimb (TIBL / SVL 0.45 vs 0.35–0.39; HIL / SVL 1.63 vs 1.28–1.47); tibiotarsal articulation reaching the eye (vs the tympanum) when adpressed forward along the body. The skin texture and colour pattern, as well as the size distribution of superciliary spines, is, however, practically identical.</p><p>Call.</p><p>The advertisement call recorded on 19 February 1994 (18: 30–19: 00 h) at Ankeniheny (air temperature 23.5 ° C) consists of a short tonal note emitted in regular call series of variable duration (Fig. 74). Most calls (= notes) are composed of two recognizable pulses. These two-pulse calls exhibit a remarkable and complex pattern of frequency modulation, with a distinct upward sweep corresponding to the initial pulse, followed by a downward sweep corresponding to the second pulse. In addition, an upward shift in dominant frequency is recognizable in the second pulse (Fig. 74). Numerical parameters of 15 analysed calls are as follows: call duration (= note duration) 68–95 ms (83 ± 7 ms); inter-call intervals within regular call series 284–331 ms (308 ± 13 ms); number of calls per call series 5–12 (9 ± 3); duration of regular call series 1641–4468 ms (3162 ± 1119 ms); dominant frequency 3488–3660 Hz (3588 ± 81 Hz), with a second prominent peak present at 7200–7400 Hz; prevalent bandwidth 1800–9500 Hz; harmonic frequency bands recognizable with approximately 3500 Hz spacing up to 19,000 Hz. Calls from the same recording (Vences et al. 2006 CD 3, track 29) have been described by Vences and Glaw (2003) using a slightly different terminological scheme.</p><p>Etymology.</p><p>Latin feminine nominative singular adjective meaning ‘ crowned’, referring to the superciliary spines.</p><p>Distribution.</p><p>This species is known from (1) Ankeniheny (type locality), (2) Mandraka, (3) Andasibe, (4) Berano (18.8371°S, 48.3185°E, 1126 m a. s. l.) (Rakotoarisoa 2012) and (5) various sites locations in the Ambatovy-Analamay area (Raselimanana 2010). Records of the species from Zahamena (IUCN SSC Amphibian Specialist Group 2016 c; Goodman et al. 2018) and other locality records given in Goodman et al. (2018) are in need of confirmation, since we have not seen any voucher specimens and there are no DNA sequences available. If R. sp. Ca 13 is considered conspecific, Vohidrazana will be added to the localities known for this species. Elevation: 900–1220 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne coronata is a semi-fossorial or detriticolous species found in mid-elevation humid forest. Adult males were found calling from the leaf litter (not inside burrows) at night in rainforest.</p></div>	https://treatment.plazi.org/id/4167246972065DB8ABD633B37DF6B936	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
23EF76B41D1357FD8C57FDD5E4326341.text	23EF76B41D1357FD8C57FDD5E4326341.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne coudreaui (Angel 1938)	<div><p>Rhombophryne coudreaui (Angel, 1938)</p><p>Figures 2, 4 –9, 13, 14</p><p>Plethodontohyla coudreaui Angel, 1938</p><p>Rhombophryne coudreaui — Frost et al. (2006)</p><p>Holotype.</p><p>MNHN 1937.19, an adult female, collected in January 1937 from ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.2&amp;materialsCitation.latitude=-17.9" title="Search Plazi for locations around (long 49.2/lat -17.9)">Réserve de Betampona</a> (forêt orientale, près de Moramanga, à mi-chemin, entre Tamatave et Tananarive) ’ (Angel 1938: 261), today the Réserve Naturelle Intégrale de Betampona, Antsinanana Region, eastern Madagascar (approx. coordinates 17.9°S, 49.2°E), by M. Coudreau (Figs 8, 9, 14) .</p><p>Paratypes.</p><p>None.</p><p>Referred specimens.</p><p>MRSN A 6280 (FAZC 13600), an unsexed adult, collected on 12 February 2007 at Sahambendrana (17.8984°S, 49.2154°E, 458 m a. s. l.), Betampona Strict Nature Reserve, Antsinanana Region, eastern Madagascar, by G. de Sousa Miranda Rosa and F. Andreone (Fig. 13 A – C); MRSN A 6347 (FAZC 13887), an adult male, collected on 17 November 2007 at Maintimbato (17.8940°S, 49.2283°E, 255 m a. s. l.), Betampona Strict Nature Reserve, Antsinanana Region, eastern Madagascar, by G. de Sousa Miranda Rosa, F. Andreone, and J. Noel; MRSN A 6267 (FAZC 13624) and MRSN A 6271 (FAZC 13653; Fig. 13 D – G), two unsexed individuals, collected on 19 February 2007 at Maintimbato (17.8940°S, 49.2283°E, 255 m a. s. l.), Betampona Strict Nature Reserve, Antsinanana Region, eastern Madagascar, by G. de Sousa Miranda Rosa.</p><p>Assignment of specimens.</p><p>The specimen MRSN A 6347 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ FJ 559299 / HM 364772; Vieites et al. 2009). All referred material agrees morphologically with the holotype, and is topotypical. Assignment of MRSN A 6271, MRSN A 6267 and MRSN A 6280 is further confirmed by molecular taxonomic identification (Fig. 2). An additional tissue sample from a topotypical individual (not collected), DLR 550 (tissue number ACZC 7731) is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. testudo group, this species is sister to R. maraorao sp. nov. (described below) based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small to medium body size, adult SVL 21.5–26.4 mm; (2) head much wider than long (HW / HL 1.65–1.85); (3) small eyes (ED / HL 0.29–0.31); (4) short hindlimbs (HIL / SVL 1.17–1.30); (5) large inner metatarsal tubercle (IMTL / FOL 0.15–0.18); (6) generally indistinct outer metatarsal tubercle; (7) second finger equal in length to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin rough to rugose with converging longitudinal rows of tubercles; (10) presence of few, small chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) broad but not fortified skull, without expansion of squamosal, maxilla, or posterior process of angulosplenial; and (15) clavicles absent.</p><p>Rhombophryne coudreaui is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 259, and ‘ T’ at site 329 and ‘ T’ at site 348. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. coudreaui can easily be distinguished from all other species except R. testudo, R. matavy, and R. maraorao sp. nov. described below by its flattened, rotund body and short, broad head. Among these species, see the account of R. testudo above for distinction from that species; it can be distinguished from R. matavy by smaller body size (SVL 21.5–26.4 mm vs 40–50 mm), rougher dorsal skin, and less fortified skull; for distinction from R. maraorao sp. nov., see the description of that species, below. Rhombophryne coudreaui may also differ from all other Rhombophryne species by the absence of carpal 2 in the hand (vs presence); see the ‘ Osteology of the Rhombophryne testudo species group’ below for discussion.</p><p>Redescription of the holotype.</p><p>A specimen in a mediocre state of preservation, with its arms and legs stretched straight and several broken bones (Figs 8, 9, 14; see osteology, below). An incision made along the left flank makes ova visible and has torn the stomach, which evidently contains the remains of some arthropods, but which we have not emptied.</p><p>Body robust and rotund, stout. Head very wide and short, much wider than long (HW / HL 1.72). Eyes small, pupils round. Snout rounded in dorsal view, rounded in lateral view. A few small chin barbels. Canthus rostralis distinct, concave. Loreal region concave, slightly oblique. Nostril openings small, angled laterally, closer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, round. TDH / ED 0.824. Supratympanic fold distinct, raised, from posterior corner of eye straight over tympanum, then slightly curving downward. There are a few tiny bumps above each eye, which do not form distinct superciliary spines. Maxillary and vomerine teeth present, vomerine teeth distinct, each straight row bearing a diastema in its middle, separated from the contralateral at the midline by a small gap. Choanae oblong. Tongue very broad, round, attached anteriorly, posteriorly free.</p><p>Forelimbs relatively short and thick, the upper arm hardly distinguishable from the body cavity (FARL / SVL 0.321). Fingers without webbing, all short, relative lengths 1 &lt;2 = 4 &lt;3. Finger subarticular tubercles distinct, large and flat; inner metacarpal tubercle large (IMCL / HAL 0.289); outer metacarpal / palmar tubercle paired. Hind limb thick, tibiotarsal articulation extent when adpressed anteriorly along the body cannot be assessed due to the many broken bones in the legs. TIBL / SVL 0.302. Inner metatarsal tubercle present, large (IMTL / FOL 0.178). Outer metatarsal tubercle absent. Toes not webbed; first toe strongly reduced, second toe moderately reduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles indistinct, single and round; toe tips not enlarged.</p><p>The dorsal skin is rather rough, with notable tubercles outlined in dark brown above the suprascapular and sacral regions. The ventral skin is as rough as the dorsal. Along the lower lip there are a series of small barbels, especially two on either side of the medial symphysis a little behind the line of the jaw.</p><p>After at least 83 years in preservative, the colouration of the specimen is pale creamy brown, with darker areas on the forearms and the areas of the body that are covered in the fixed specimen, such as the axillae. The ventral surface is a more homogeneous light brown. Distinctive dark brown markings surround the tubercles on the suprascapular and sacral regions, but otherwise the dorsum is rather homogeneous in colour. The hindlimbs have lost most colouration, but a distinctive cream tubercle surrounded in brown is present on each shank. The forelimbs are ventrally dark brown on the lower arm, with a brown crossband anteriorly; pale and indistinctly patterned on the upper arm. The colour of the specimen in life is unknown.</p><p>Variation.</p><p>SVL 21.5–26.4 mm (n = 5); full measurements are presented in Table S 5. The base colouration of all recently collected specimens is a dark chocolatey brown (see Fig. 13 for an idea of colour variation in life). Much of the body patterning is consistent, especially the dark suprascapular spots which define the short edge of a trapezium where the eyes mark the long edge. MRSN A 6271 has a chin marbled with cream and also has cream spots on its anterior thigh and interior shank; no other examined specimen shares these patterns.</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>Eponym for Mr. Coudreau, ‘ Conservateur des Réserves naturelles de Madagascar’ (Angel 1938).</p><p>Distribution.</p><p>This species is confined to low-elevation forests in eastern Madagascar, and is with certainty known only from (1) Réserve Naturelle Intégrale de Betampona (type locality), where it has been found at Sahambendrana, Maintimbato and Sahaindrana (Rosa et al. 2012) as well as at Sahabefoza (17.9142°S, 49.2078°E, 353 m a. s. l.). However, S. Andrianarivo provided MDS photographs clearly assignable to this species from (2) Ampasina, a highly degraded forest just 8 km east of Betampona (17.918°S, 49.323°E, ca 80 m a. s. l.), taken on 2 October 2022. This indicates that the species is present and may be able to persist in strongly anthropogenically altered forest. Elevation: ca 80–200 m a. s. l.</p><p>The record from Ambolokopatrika-Betaolana (Andreone 2003) unambiguously refers to R. maraorao sp. nov. (described below), as it is based on the type material of that species. The specimen MNHN 1973.576 from Marojejy in the northeast of Madagascar was assigned to R. coudreaui by Andreone and Randrianirina (2008), but the material used for comparison to R. coudreaui was also from the northeast and is described as a new species, R. maraorao sp. nov., below. Based on the locality of this specimen and its apparent affinity with R. maraorao sp. nov., we consider it likely that MNHN 1973.576 is referrable to that species, and therefore remove Marojejy from the locality record of R. coudreaui . The same applies also to other records from Marojejy (Raselimanana et al. 2000) and Maroantsetra (IUCN SSC Amphibian Specialist Group 2016 d).</p><p>Natural history.</p><p>Rhombophryne coudreaui is a fossorial species restricted to low-elevation humid forest that is seldom encountered and poorly known. This species exhibits a death-feigning behaviour beyond normal tonic immobility wherein it straightens and stiffens all four limbs (Fig. 13 F). One specimen was found at Sahaindrana campsite (Betampona Strict Nature Reserve) walking on the leaf litter in the early hours of the morning (around 6 am). Two individuals were found underneath wet leaf litter and rotten logs in almost-closed-canopy humid forest on the ridge at 460 m at 10: 40 am in Sahabefoza.</p></div>	https://treatment.plazi.org/id/23EF76B41D1357FD8C57FDD5E4326341	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
D8B31A62B11457C2BF04E51BC1588E3D.text	D8B31A62B11457C2BF04E51BC1588E3D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne diadema Scherz 2017	<div><p>Rhombophryne diadema Scherz et al., 2017</p><p>Figures 2, 4 –7, 67 –69</p><p>Remark.</p><p>This species is only known from the type series.</p><p>Holotype.</p><p>ZSM 1629/2012 (FGZC 3604), an adult female, collected between 26 and 30 November 2012 on the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.4411&amp;materialsCitation.latitude=-13.6817" title="Search Plazi for locations around (long 49.4411/lat -13.6817)">Sorata Massif</a> (13.6817°S, 49.4411°E, 1339 m a. s. l.), Sava Region, northern Madagascar, by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina, and A. Razafimanantsoa .</p><p>Paratypes.</p><p>Two specimens: ZSM 1628/2012 (FGZC 3731), an adult male, and UADBA -A 60289 (FGZC 3611), an ovigerous adult female, collected between 26 and 30 November 2012 at a creek above the campsite on the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.4404&amp;materialsCitation.latitude=-13.678" title="Search Plazi for locations around (long 49.4404/lat -13.678)">Sorata Massif</a> (13.6780°S, 49.4404°E), Sava Region, northern Madagascar, by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina, and A. Razafimanantsoa .</p><p>Referred specimens.</p><p>ZSM 341/2016 (AEA 041), an unsexed adult, collected on 25 May 2016 in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.5307&amp;materialsCitation.latitude=-13.7483" title="Search Plazi for locations around (long 49.5307/lat -13.7483)">Andravory</a> (13.7483°S, 49.5307°E, 1168 m a. s. l.), Sava Region, northeastern Madagascar, by S. Megson and co-workers . This specimen is referred on morphological grounds and has not been genetically verified.</p><p>Assignment of specimens.</p><p>The paratype ZSM 1628/2012 (FGZC 3731) is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KU 724171; Scherz et al. 2016 a; 16 S 5 ’ KY 748097; Scherz et al. 2017 a). Paratype UADBA -A 60289 (FGZC 3611) is used as the reference specimen for this species in COI barcoding (COI OL 790125; Belluardo et al. 2022). The holotype is identical to the 16 S 3 ’ reference sequence, except one insertion and an undetermined stretch (16 S 3 ’ PZ 418419) and differs by one mutation in the 16 S 5 ’ fragment (16 S 5 ’ PZ 405231). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of the paratypes is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 1629/2012 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is placed in a clade containing R. guentherpetersi, R. ornata, and R. tany based on molecular phylogenetic data, but the exact relationships within that clade are poorly resolved (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 21.5–23.4 mm; (2) head wider than long (HW / HL 1.40–1.59); (3) moderately large eyes (ED / HL 0.38–0.44); (4) relatively long hindlimbs (HIL / SVL 1.66–1.89); (5) small inner metatarsal tubercle (IMTL / FOL 0.07–0.11); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin granular to rugose; (10) absence of chin barbels; (11) presence of three superciliary spines, the posterior-most being diminutive; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne diadema is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 266, ‘ A’ at site 640, and ‘ T’ at site 740; in 16 S 3 ‘ A’ at site 965, ‘ T’ at site 1083, ‘ T’ at site 1086, and ‘ T’ at site 1329. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. diadema can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. For distinction from R. serratopalpebrosa, R. guentherpetersi, R. coronata, R. vaventy, R. ornata, R. tany, and R. regalis, see diagnoses of those species above. For distinction from the new species described herein, see accounts below.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>Latin feminine noun in apposition meaning ‘ diadem’, in reference to the superciliary spines.</p><p>Distribution.</p><p>This species is only known from (1) Sorata Massif (type locality). Elevation: 1339–1407 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne diadema is a semi-fossorial or detriticolous species found in mid-elevation humid forest. Individuals were found during the day in the leaf litter. The holotype contained at least 13 well-developed, yellow eggs (diameter 2.45 ± 0.25 mm) (Scherz et al. 2017 a).</p></div>	https://treatment.plazi.org/id/D8B31A62B11457C2BF04E51BC1588E3D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
5E6BE6897A795CEA85BDC3D3E31FC2D3.text	5E6BE6897A795CEA85BDC3D3E31FC2D3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne ellae Scherz 2020	<div><p>Rhombophryne ellae species group</p><p>Rhombophryne ellae, R. kilonjy sp. nov., R. quentini sp. nov. (sp. Ca 07)</p><p>Summary of morphology.</p><p>A group of medium-sized (SVL 22.1–30.1 mm), terrestrial or detriticolous Rhombophryne species. All have a rotund habitus, very similar to the R. laevipes species group. They have bright-orange colouration on the posterior side of the limbs in life, and R. ellae and R. quentini sp. nov. have black inguinal ocelli outlined in white (absent in R. kilonjy sp. nov.). Teeth are present on the maxilla, premaxilla, and postchoanal vomer. Clavicles are present. Skin texture is granular.</p><p>Calls.</p><p>Unknown.</p><p>Distribution.</p><p>This group is restricted to the north of Madagascar (Fig. 7; Table S 4), in Montagne d’Ambre ( R. ellae), Tsaratànana ( R. quentini sp. nov.), and the Sahamalaza Peninsula ( R. kilonjy sp. nov.).</p><p>Ecology.</p><p>Species found in humid and subhumid tropical forest.</p></div>	https://treatment.plazi.org/id/5E6BE6897A795CEA85BDC3D3E31FC2D3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
A5568CCC911C5136BC397755C1AC9A69.text	A5568CCC911C5136BC397755C1AC9A69.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne ellae Scherz 2020	<div><p>Rhombophryne ellae Scherz, 2020</p><p>Figures 2, 4 –7, 50 –53</p><p>Holotype.</p><p>ZSM 76/2018 (MSZC 0534), an adult or subadult female, collected on 28 December 2017 in ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.1746&amp;materialsCitation.latitude=-12.5066" title="Search Plazi for locations around (long 49.1746/lat -12.5066)">Montagne d’Ambre National Park</a> (12.5066°S, 49.1746°E, 892 m a. s. l.) ’, Diana Region, northern Madagascar by M. D. Scherz, J. H. Razafindraibe, A. Razafimanantsoa, O. Randriamalala, S. M. Rasolonjavato, R. Tiavina, E. Z. Lattenkamp, and A. Rakotoarison (Figs 50, 51, 52 A – D, 53) .</p><p>Paratypes.</p><p>None.</p><p>Referred specimens.</p><p>ZSM 903/2003 (FGMV 2002.0918), an unsexed adult, collected on 18 February 2003 in Montagne d’Ambre National Park (no precise coordinates available), Diana Region, northern Madagascar by F. Glaw, R. D. Randrianiaina, and A. Razafimanantsoa (Fig. 52 E, F); ZSM 222/2004 (FGZC 432), a juvenile specimen, collected on 20 February 2004 in Montagne d’Ambre National Park (no precise coordinates available), Diana Region, northern Madagascar, by F. Glaw, M. Puente, R. Randrianiaina, and A. Razafimanantsoa.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ MT 371794; Scherz 2020) and COI barcoding (COI MT 372330; Scherz 2020), and is present in the RAG 1 haplotype network (Fig. 4). No other specimens of the species have yet been sequenced. Both of the newly referred specimens conform well with the holotype in morphology.</p><p>Definition.</p><p>Within the R. ellae group, this species is sister to a clade consisting of R. quentini sp. nov. and R. kilonjy sp. nov. (described below) based on molecular phylogenetic data. It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 24.9 mm; (2) head wider than long (HW / HL 1.47); (3) moderately small eyes (ED / HL 0.38); (4) relatively long hindlimbs (HIL / SVL 1.71); (5) small inner metatarsal tubercle (IMTL / FOL 0.9); (6) outer metatarsal tubercle weakly pronounced; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of large black inguinal ocelli bordered with white; (13) presence of bright orange colouration on the hidden surfaces of the legs in life; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne ellae is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 3 ’, ‘ G’ at site 1025, and ‘ A’ at site 1074 (16 S 5 ’ not available for this species).</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. ellae can be distinguished from all species except R. quentini sp. nov. described below by the combination of distinctive white-outlined black inguinal ocelli with reddish orange colouration on the hidden surfaces of the legs. For distinction from R. quentini sp. nov. and R. kilonjy sp. nov., see the description of those species, below.</p><p>Variation.</p><p>In the original description, Scherz (2020) reported on the single specimen known to him at that time. ZSM 903/2003 (Fig. 52 E, F) and ZSM 222/2004, agree very well with the original description both in morphology and in colouration. The dorsal skin of ZSM 903/2003 is slightly more granular than the type specimen, and the skin on the underside of the legs is darker grey. ZSM 222/2004 appears faded in colour. Morphometric measurements are provided in Table S 7.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>Eponym for Ella Z. Lattenkamp, partner of MDS.</p><p>Distribution.</p><p>This species is only known from (1) Montagne d’Ambre National Park (type locality).</p><p>Natural history.</p><p>Rhombophryne ellae is a semi-fossorial or detriticolous species found in mid-elevation humid forest; its ecology is poorly known. Gut contents of the holotype included ‘ three whole ants and one ant head, seemingly belonging to two different species (one of the whole ants is diminutive), the head of a jumping spider ( Salticidae), and the elytra and other body parts of a beetle. ’ (Scherz 2020: 319).</p></div>	https://treatment.plazi.org/id/A5568CCC911C5136BC397755C1AC9A69	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
0D2454BCF93050B0A9FF9DA503FC166A.text	0D2454BCF93050B0A9FF9DA503FC166A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne guentherpetersi (Guibe 1974)	<div><p>Rhombophryne guentherpetersi (Guibé, 1974)</p><p>Figures 2, 4 –7, 67 –69</p><p>Mantipus guentherpetersi Guibé, 1974</p><p>Plethodontohyla guentherpetersi — Blommers-Schlösser and Blanc (1991)</p><p>Rhombophryne guentherpetersi — Glaw and Vences (2007)</p><p>Plethodontohyla guntherpetersi — Raxworthy et al. (2008), misspelling</p><p>Remark.</p><p>A detailed redescription of the species was given by Scherz et al. (2017 a).</p><p>Holotype.</p><p>MNHN 1953.165, an adult female, collected on an unknown date on the ‘ Tsaratànana Massif’ (no precise coordinates, at ‘ 2600 m’ a. s. l.), northern Madagascar (Guibé 1973: 1181). Scherz et al. (2017 a) stated that the collectors of this specimen were unknown, but the MNHN catalogue records J. Millot and J. Guibé as the collectors.</p><p>Paratypes.</p><p>Four specimens: MNHN 1953.165 A, a subadult female, MNHN 1953.165 B, an adult female, with the same collection data as the holotype. MNHN 1973.592, a subadult female, and MNHN 1973.593, a juvenile, collected on 1 November 1966 on Mission ORSTOM by C. P. Blanc from the same locality as the holotype (Guibé 1973: 1181). Note: the MNHN catalogue records R. Paulian as the collector.</p><p>Referred specimens.</p><p>ZSM 606/2014 (DRV 6220), an adult female, ZSM 607/2014 (DRV 6223; Fig. 69 B), an adult male, UADBA -A 60775 (DRV 6210) and UADBA -A 60776 (ZCMV 12401), two unsexed adults, and UADBA -A 60782 (ZCMV 12435), a subadult, all collected on 16 June 2010 at ‘ Camp 4 ’ Andranomadio (14.0801°S, 48.9854°E, 2503 m a. s. l.), Tsaratànana Massif, Diana Region, northern Madagascar by D. Vieites, M. Vences, R. D. Randrianiaina, F. M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, F. Randrianasolo, F. Randrianasolo, and T. Rajoafiarison; ZSM 608/2014 (DRV 6231), an adult female, collected on 18 June 2010 at 14.0665°S, 48.9832°E, 2732 m a. s. l., Tsaratànana Massif, Diana Region, northern Madagascar, by D. Vieites, M. Vences, R. D. Randrianiaina, F. M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, F. Randrianasolo, F. Randrianasolo, and T. Rajoafiarison.</p><p>Assignment of specimens.</p><p>The specimen ZSM 606/2014 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KU 724178; Scherz et al. 2016 b; 16 S 5 ’ KY 748098; Scherz et al. 2017 a) and COI barcoding (COI OL 790126; Belluardo et al. 2022). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of specimen UADBA -A 60776 is further confirmed by molecular taxonomic identification (Fig. 2). UADBA -A 60776 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is placed in a clade containing R. diadema, R. ornata, and R. tany based on molecular phylogenetic data, but the exact relationships within that clade are poorly resolved (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium to large body size, adult SVL 27.3–35.7 mm; (2) head wider than long (HW / HL 1.35–1.42); (3) moderately large eyes (ED / HL 0.35–0.41); (4) relatively short hindlimbs (HIL / SVL 1.33–1.45); (5) small inner metatarsal tubercle (IMTL / FOL 0.10–0.13); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin slightly granular, with pronounced glandular swellings in the dorsolateral area of the trunk and on the tibia; (10) absence of chin barbels; (11) presence of two small superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne guentherpetersi is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 369, ‘ G’ at site 714, and ‘ C’ at site 735; in 16 S 3 ’. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. guentherpetersi can easily be distinguished from all other species (including all species newly described herein) by the presence of large, swollen tibial glands.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>Eponym for Günther Peters, German herpetologist.</p><p>Distribution.</p><p>This species is only known from (1) high elevation on the Tsaratànana Massif (type locality). Elevation: 2503–2732 m a. s. l. Raxworthy et al. (2008) reported the species from 1450–2700 m a. s. l., but their identifications have not been verified by us, and the lower elevation records may refer to other species.</p><p>Natural history.</p><p>Rhombophryne guentherpetersi is a semi-fossorial or detriticolous species found in montane forest and ericoid thicket. No further information on its natural history is available.</p></div>	https://treatment.plazi.org/id/0D2454BCF93050B0A9FF9DA503FC166A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
8CEE2CFB49A452838467D502DA4CAB66.text	8CEE2CFB49A452838467D502DA4CAB66.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne kilonjy Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne kilonjy sp. nov.</p><p>Figures 2, 5 –7, 50, 51, 56, 57</p><p>Informal names.</p><p>This species was referred to as ‘ Plethodontohyla sp. ’ by Andreone et al. (2001), ‘ Rhombophryne sp. aff. alluaudi ’ by Penny et al. (2016), and ‘ Rhombophryne sp. (UCS) ’ by Penny et al. (2017). It has not been sequenced previously.</p><p>Justification of the new species.</p><p>The new species is sister to R. quentini sp. nov. (described above), and these two species are together sister to R. ellae, as part of the R. ellae group that is phylogenetically distinct from other Rhombophryne (Belluardo et al. 2022) . Museomic sequencing yielded 0.4 × coverage for 117 bp of the 16 S 3 ’ marker, and 0.9 × coverage for 272 bp of the 16 S 5 ’ marker, but even these fragmentary data are sufficient to assign this species to this group, and identify it as the sister of R. quentini sp. nov. The genetic distance towards other members of the species group ranges from 5.13 % in 16 S 3 ’ to 12.4 % in 16 S 5 ’ (which might be exaggerated based on the fragmented state of the DNA; Table S 3) and all members share a diagnostic ‘ G’ at site 338 in the concatenated 16 S alignment. It is morphologically distinct from both R. ellae and R. quentini sp. nov. by lacking inguinal ocelli (vs presence), and several osteological features relating to bones of the skull, and the humerus (see the diagnosis, below), and also from R. ellae by a distinct outer metatarsal tubercle (vs weakly pronounced).</p><p>Holotype.</p><p>MRSN A 4627 (FAZC 10505), an adult female, collected on 12 February 2000 in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.92&amp;materialsCitation.latitude=-14.3" title="Search Plazi for locations around (long 47.92/lat -14.3)">Berara</a> (ca 14.3°S, 47.92°E, ca 170 m a. s. l.), Sahamalaza Peninsula, Sofia Region, northwestern Madagascar, by F. Andreone, J. E. Randrianirina, and M. Vences (Figs 51, 52, 57, 58) .</p><p>Paratypes.</p><p>Three specimens: MRSN A 4601 (FAZC 10557), MRSN A 4610, and MRSN A 4611 (FAZC 10558), three juveniles, all with the same collection data as the holotype but collected on 14–16 February 2000 .</p><p>Assignment of specimens.</p><p>The three paratypes were collected at the type locality, and can be assigned to the species morphologically. There are no known Rhombophryne from the same locality with which these specimens could be confused, and they also could not be confused with any other cophyline microhylid species known from this area. The species does not currently have a reference specimen for DNA barcoding, and has not been sequenced for the RAG 1 marker.</p><p>Definition.</p><p>Within the R. ellae group, this species is sister to R. quentini sp. nov. based on molecular phylogenetic data. It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 27.3 mm; (2) head wider than long (HW / HL 1.65); (3) moderately small eyes (ED / HL 0.42); (4) relatively long hindlimbs (HIL / SVL 1.40); (5) small inner metatarsal tubercle (IMTL / FOL 0.11); (6) outer metatarsal tubercle present but indistinct; (7) second finger shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli; (13) presence of reddish orange colouration on the hidden surfaces of the legs in life; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne kilonjy sp. nov. is furthermore distinguished from all other Rhombophryne species by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042), which were identified visually, as inclusion in the MolD analysis would have distorted the results due to insufficiently complete sequences: in 16 S 5 ’ ‘ T’ at site 446, ‘ C’ at site 468, ‘ C’ at site 580, ‘ G’ at site 600, ‘ G’ at site 601, and ‘ T’ at site 670; in 16 S 3 ’ ‘ T’ at site 964, and ‘ T’ at site 1004.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. kilonjy sp. nov. can be distinguished from all other Rhombophryne except R. ellae, R. quentini sp. nov., R. kotrobaratra sp. nov., R. mavokely sp. nov., and R. ornata by the presence of reddish orange colouration on the hidden surfaces of the legs. From R. ellae and almost all other Rhombophryne species, it can be distinguished by the pronounced outer metatarsal tubercle in adults (vs weakly pronounced), except R. quentini sp. nov. It can be distinguished from R. quentini sp. nov. and R. ellae by the absence of inguinal ocelli. Superficially it is rather similar to R. mavokely sp. nov., but it differs from that species by absence of a black spot on the supratympanic fold (vs presence), larger body size (SVL 27.3 mm vs 21.2–21.3 mm), and presence of a distinct outer metatarsal tubercle (vs indistinct). Based on the specimens examined, the skeleton of R. kilonjy sp. nov. also differs from those of R. ellae and R. quentini sp. nov. in the following traits: broader prootic, columella with a dorsal extension to the pars interna plectri (vs without a dorsal extension), parasphenoid cultriform process narrowing anteriorly (vs not narrowing anteriorly), higher coronoid process of the angulosplenial (vs lower), humeral crista lateralis distinct (vs weakly developed).</p><p>Description of holotype.</p><p>Specimen in an excellent state of preservation (Fig. 50). An incision made along the left flank to assess the sex, and part of the left thigh muscle taken as a tissue sample. Developing ova are clearly visible in the body cavity.</p><p>Body rotund. Head wider than long (HW / HL 1.65). Eyes moderately small (ED / HL 0.43), pupils small, oval. Snout somewhat boxy in dorsal and rounded in lateral view, with a small protrusion at the snout tip. Canthus rostralis indistinct, concave. Loreal region weakly concave, slightly oblique. Nostril closer to eye than to tip of snout, slightly protuberant. Tympanum indistinct, oval, TDH / ED 0.47. Supratympanic fold indistinct in preservative, visible only as melanised curve, but in life highly distinct, arcing strongly from posterior corner of eye over tympanum (Fig. 56). Lateral head similar in colour to dorsal head. Superciliary spines absent. Dorsal skin smooth in preservative, but finely granular in life (Fig. 56). Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth with a lateral diastema, the medial elements distinct, separated from one another at the midline by a small gap. Choanae oblong, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.366), moderately slender. Fingers without webbing, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger distinctly shorter than fourth (Fig. 51). Finger tips not expanded, rounded. Finger subarticular tubercles distinct, cream, flat; inner metacarpal tubercle slender, outer metacarpal / palmar tubercle indistinct, cream, single. Hind limb strong, tibiotarsal articulation reaching approximately to the tympanum when adpressed anteriorly along the body. TIBL / SVL 0.367. Inner metatarsal tubercle present, oblong, whitish, distinctly raised. Outer metatarsal tubercle distinct, rounded and raised. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles weak, single; toe tips slightly enlarged, especially of fourth toe, rounded to subacuminate (Fig. 51).</p><p>After 24 years in preservative, the base colour was a grey-brown with darker brown speckles throughout, and a darker brown area over the mid-dorsum in a broad strip. The dorsum is marked with two poorly defined darker brown chevrons, specifically one over the suprascapular region and a second over the iliac region. In each chevron are a small number of symmetrical white spots surrounded by dark brown. The inguinal region lacks any trace of ocelli. No distinct colour difference between the lateral and dorsal head. The supratympanic fold is dark brown from the eye to the tympanum, fading posteriorly. The dorsal hindlimbs bear indistinct crossbands that line up with the leg folded, in the same colours as the dorsum and chevrons. The ventral foot surface is dark brown. The anterior and posterior thigh is pale cream in preservative (this was reddish orange in life; Fig. 56), as is the ventral thigh and shank. A distinct, dark brown trapezoid is present around the cloaca. The dorsal foot is mottled cream and light brown. A light annulus is present before the terminal phalange of each finger and toe. The arms have a single crossband on the antebrachium. The dorsal hand is lighter medially than laterally; the ventral hand is cream. The chin is mottled light brown with cream flecks, fading to completely cream over the sternal region into the immaculate abdomen.</p><p>Variation.</p><p>The holotype is the only known adult, and measures 27.3 mm in SVL; measured juveniles (n = 3) ranged from 12.3–15.7 mm. For variation in other measurements, see Table S 7. Juveniles are morphologically consistent with the holotype. The outer metatarsal tubercle is consistently distinct. The development of vomerine teeth is variable among the examined juveniles, with one specimen (MRSN A 4611) lacking vomerine teeth (but the postchoanal portion of the vomer is clearly visible in the palate), whereas the other specimens have distinct and rather large vomerine teeth. All specimens lack inguinal spots. All specimens have at least a faint dark chevron on the dorsum.</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>The Malagasy noun kilonjy meaning ‘ pebble’, in reference to the small, rotund shape of this species, reminiscent of a smooth pebble. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is only known from (1) Berara on the Sahamalaza Peninsula (type locality). Elevation: 170–210 m a. s. l. (Andreone et al. 2001).</p><p>Natural history.</p><p>Rhombophryne kilonjy sp. nov. is a semi-fossorial or detriticolous species known only from low-elevation western dry forest, drier than the localities of all other known Rhombophryne . One specimen was collected in a pitfall trap. No further specific details are known of its ecology.</p></div>	https://treatment.plazi.org/id/8CEE2CFB49A452838467D502DA4CAB66	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
A3AD2B8B74CD5649B0423A095D57DEAD.text	A3AD2B8B74CD5649B0423A095D57DEAD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne kotrobaratra Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne kotrobaratra sp. nov.</p><p>Figures 2, 4 –7, 30, 31, 48, 49</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 10 ’ by Vieites et al. (2009) and ‘ Rhombophryne sp. Ca 10 ’ by Scherz et al. (2015 b), Lambert et al. (2017), Scherz (2020), and Belluardo et al. (2022), and ‘ Rhombophryne sp. 10 ’ and ‘ UCS “ Sp. 10 ” ’ and ‘ UCS “ Sp. 20 ” ’ by Carné and Vieites (2024). A specimen of this species (MRSN A 2610) was mistakenly figured as the specimen MRSN A 2620 of ‘ R. sp. Ca 19 ’ by Belluardo et al. (2022).</p><p>Justification of the new species.</p><p>The new species was found to be the sister species of R. sp. Ca 15 from Betampona of which no further data are available (see account on this candidate species below). Together, the clade of these two lineages is sister to all other species of the R. laevipes group based on a multigene phylogeny (Belluardo et al. 2022) which by itself already excludes conspecificity with any of the other nominal species. Genetic divergence of R. kotrobaratra sp. nov. to all other nominal species of Rhombophryne is&gt; 7.5 % uncorrected p distance in the 3 ’ - terminal fragment of 16 S and thus much higher than the 3 % level typically indicative of species-level differentiation in Malagasy frogs (Vieites et al. 2009). There is no haplotype sharing in the analysed fragment of the nuclear gene RAG 1 between R. kotrobaratra sp. nov. and any other species of the Rhombophryne genus in the analysed dataset (Fig. 4). Furthermore, it also differs from other nominal species in the R. laevipes group by morphological characters (see Diagnosis below), and its status as distinct species is therefore beyond doubt. More research is necessary to understand the status of R. sp. Ca 15 (see below).</p><p>Holotype.</p><p>MRSN A 2990 (RJS 0581), an adult female, collected on 17 February 2002 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.9403&amp;materialsCitation.latitude=-15.4177" title="Search Plazi for locations around (long 49.9403/lat -15.4177)">‘Camp 7’ Antsiranana Faritany</a> (15.4177°S, 49.9403°E, ca. 778 m a. s. l.), Amparihy, Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by J. E. Randrianirina (Figs 30, 31) .</p><p>Paratypes.</p><p>Nine specimens: MRSN A 2610 (FAZC 10312; Fig. 48 A – C), an adult male, and MRSN A 2599 (FAZC 10307), an adult female, collected on 11 and 13 December 1999 (respectively) at ‘ Camp 5 ’ Menamalona (15.3812°S, 49.9878°E, ca 800 m a. s. l.), Ilampy Corridor, Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by F. Andreone and J. E. Randrianirina; MRSN A 1845 (FN 6310), an adult female, collected on 3 December 1996 in Antsarahan’ny Tsararano (ca 14.9067°S, 49.6867°E, ca 700 m a. s. l.), Sava Region, northeastern Madagascar, by F. Andreone (Fig. 48 E, F); MRSN A 2609 (FN 7519; Fig. 49), an adult male, and MRSN A 2601 (FN 7560), an adult female, collected on 23 and 24 November 1998 (respectively) at ‘ Camp 2 ’ Beanjada (15.2638°S, 49.9583°E, ca 1000 m a. s. l.), Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by F. Andreone and J. E. Randrianirina; UADBA -A- APR 12594, UADBA -A- APR 12541, and UADBA -A- APR 12563, a putative male and two small juveniles (respectively), collected on 16–19 May 2016 at mid-elevation in Marojejy National Park (14.4559°S, 49.7749°E, 765–780 m a. s. l.), Sava Region, northeastern Madagascar, by A. P. Raselimanana; UADBA -A- APR 12765, an adult female, collected on 1 June 2016 at mid-elevation in southwestern Marojejy National Park (14.5074°S, 49.6153°E, 835 m a. s. l.), Sava Region, northeastern Madagascar, by A. P. Raselimanana.</p><p>Assignment of specimens.</p><p>The paratype MRSN A 2610 (FAZC 10312) is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 5 ’ AY 594073; Andreone et al. 2005; 16 S 3 ’ AY 594111; Vieites et al. 2009). The holotype is used as the reference specimen for COI barcoding (COI OL 790134; Belluardo et al. 2022). The 16 S sequence of the holotype differs from the 16 S reference sequence by eight mutations and three gaps in the 16 S 3 ’ fragment (98 % identical; OL 780580). All paratypes and referred specimens conform well with the holotype in morphology; no further specimens are sequenced. MRSN A 2990 and MRSN A 2610 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. laevipes group, this species along with R. sp. Ca 15 represents the sister clade of all other group members based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 24.1–33.0 mm; (2) head wider than long (HW / HL 1.46–1.64); (3) moderately small eyes (ED / HL 0.34–0.44); (4) long hindlimbs (HIL / SVL 1.81–1.92); (5) small inner metatarsal tubercle (IMTL / FOL 0.09–0.11); (6) outer metatarsal tubercle sometimes present; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin granular with scattered warts in life; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of distinct inguinal spots; (13) presence of bright orange colouration on the hidden surfaces of the legs in life; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne kotrobaratra sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 288, ‘ T’ at site 355, and ‘ T’ at site 402. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. kotrobaratra sp. nov. is highly distinctive, and can be distinguished from all other species by the combination of orange colouration on the hidden surfaces of the legs (otherwise present only in the R. ellae species group and R. mavokely sp. nov.), absence of inguinal ocelli (present in R. ellae and R. quentini sp. nov.), and long legs (HIL / SVL 1.81–1.92 vs 1.50–1.61 in R. mavokely sp. nov., 1.40–1.80 in the R. ellae species group). It also typically has a distinctive dark chevron dorsally, which can be useful to distinguish it from most other species. These chevrons and in general the presence of a rather tubercular dorsal skin appear to distinguish the species from all other nominal species in the R. laevipes group.</p><p>Description of holotype.</p><p>An ovigerous adult female in an excellent state of preservation (Fig. 30). Right arm removed as a tissue sample, and a slit made along the right flank to check the sex. The abdomen is full of highly developed ova.</p><p>Body robust. Head wider than long (HW / HL 1.55), distinctly narrower than abdomen. Eyes moderately small (ED / HL 0.41), slightly depressed, pupils lenticular. Snout rounded in dorsal and lateral view. Canthus rostralis indistinct, concave. Loreal region weakly concave, oblique. Nostril closer to tip of snout than to eye, very slightly protuberant. Tympanum indistinct, round, TDH / ED 0.55. Supratympanic fold indistinct, weakly raised, curving slightly from posterior corner of eye to above the forearm’s insertion. Lateral head similar in colour to dorsal head. Superciliary spines absent. Dorsal skin smooth in preservative. Tongue partly taken as tissue sample, originally broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth curved, without height variation, separated at the midline by a small gap. Choanae oval, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.407) and narrow. Fingers without webbing, moderately long, relative lengths 1 &lt;4 &lt;2 &lt;3, second finger distinctly longer than fourth (Fig. 31). Finger tips not expanded, rounded. Finger subarticular tubercles distinct, whitish, flat; inner metacarpal tubercle distinct and rounded, outer metacarpal / palmar tubercle distinct and oblong, cream. Hind limb strong with particularly thick and robust thighs, tibiotarsal articulation reaching the snout when adpressed anteriorly along the body. TIBL / SVL 0.520. Inner metatarsal tubercle present, oblong, whitish, distinctly raised. Outer metatarsal tubercle present as a faint lightening of the general area. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles distinct; toe tips slightly enlarged, rounded (Fig. 31).</p><p>After 21 years in preservative, the base colour is a mottled light brown across the dorsum and limbs. The dorsum bears one very distinct dark brown chevron over the suprascapular region, and a suggestion of a second one over the hip. There are darker brown blotches within these patterns, giving an overall mottled appearance. An interocular brown band is present with a tan anterior edge. Behind it, two small tan spots are present, surrounded by dark brown. No distinct colour difference between the lateral and dorsal head. The supratympanic fold is not marked by colour. The inguinal region lacks dark or light spots. The hindlimbs have distinct dark brown crossbands on thigh, shank, and foot. The posterior and anterior thigh is light cream mottled with brown; the ventral thigh is immaculate medially and becomes more mottled distally. The cloaca region is slightly darker brown than the rest of the posterior thigh. Mottling continues on the ventral shank and medial foot. The ventral feet are light brown. A cream annulus is present before the terminal phalange of each finger and toe. The arms have a single dark brown crossband on the lower arm. The palmar surface is light brown except for the cream tubercles. The chin is cream in base colour but mottled with light brown, fading posteriorly to immaculate cream over the scapular region go the abdomen.</p><p>Variation.</p><p>Sexual size dimorphism quite pronounced: SVL of males 24.1–26.4 mm (n = 2), females 27.0–33.0 mm (n = 5); for variation in other measurements see Table S 6. No obvious dichromatism; males have only slightly darker chins than females. The dorsal pattern consistently includes the dark chevron marking over the suprascapular region. The posterior chevron varies from highly distinct (MRSN A 2609), to its least distinct in the holotype (Fig. 30). The legs of APR 12765 and APR 12594 are distinctly narrower than the rest of the type series. These specimens are also greyer in colour, possibly as a result of differences in preservation method. The ventral surface of APR 12765 is covered in tiny round dark markings that are clearly distinct from melanosomes. These structures are absent from all other specimens.</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>The Malagasy noun kotrobaratra meaning ‘ thunder and lightning’, in reference to the typical conditions under which this frog emerges, during stormy weather with heavy rain. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is known from (1) the western Masoala Peninsula: Amparihy (type locality), Menamalona, and Beanjada, and (2) Antsarahan’ny Tsararano, and (3) Marojejy (Fig. 48). It occurs at intermediate elevations (ca 600–1000 m a. s. l.).</p><p>Natural history.</p><p>Rhombophryne kotrobaratra sp. nov. is a semi-fossorial or detriticolous species found in low- to mid-elevation humid forest. Specimens collected by APR were active at night on wet thick leaf litter covering the forest floor, after a heavy rain late in the afternoon. All observed individuals were found in almost-closed-canopy forest.</p></div>	https://treatment.plazi.org/id/A3AD2B8B74CD5649B0423A095D57DEAD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
90686EC754335D9090B4A8DC8D59D299.text	90686EC754335D9090B4A8DC8D59D299.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne laevipes (Mocquard 1895)	<div><p>Rhombophryne laevipes species group</p><p>Contents.</p><p>Rhombophryne anatiala sp. nov. (sp. Ca 01), R. botabota, R. kotrobaratra sp. nov. (sp. Ca 10), R. laevipes, R. nilevina (including sp. Ca 03), R. sonaliae sp. nov. (sp. Ca 19), R. sp. Ca 15</p><p>Summary of morphology.</p><p>A group of large-bodied, detriticolous to terrestrial Rhombophryne species. All have a rotund habitus but less flattened than members of the R. testudo species group. This group has the largest body size in the genus (SVL 24.1–57.2 mm). Almost all species have whitish spots on the legs and inguinal region (absent in R. botabota, R. kotrobaratra sp. nov., and some specimens of R. nilevina). Teeth are present on the maxilla, premaxilla, and postchoanal vomer. Clavicles are present. Skin texture is smooth to granular.</p><p>Calls.</p><p>Calls are known from R. nilevina, R. laevipes, R. botabota and R. sonaliae sp. nov. They consist of single low frequency, pulsed or tonal notes emitted at long intervals.</p><p>Distribution.</p><p>This group is widely distributed (Fig. 7; Table S 4), with members in the north ( R. laevipes, R. nilevina), northeast ( R. botabota, R. nilevina, R. anatiala sp. nov., and R. kotrobaratra sp. nov.), and east ( R. nilevina, R. sonaliae sp. nov., R. sp. Ca 15) of Madagascar.</p><p>Ecology.</p><p>Species found in humid and subhumid tropical forest.</p></div>	https://treatment.plazi.org/id/90686EC754335D9090B4A8DC8D59D299	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
2EE70DCAA0BD58B188B233A576C2C2BF.text	2EE70DCAA0BD58B188B233A576C2C2BF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne laevipes (Mocquard 1895)	<div><p>Rhombophryne laevipes (Mocquard, 1895)</p><p>Figures 2, 4 –7, 30 –35</p><p>Mantiphrys laevipes Mocquard, 1895</p><p>Mantophrys laevipes — Mocquard (1909), subsequent incorrect spelling</p><p>Mantipus laevipes — Guibé (1946)</p><p>Plethodontohyla laevipes — Blommers-Schlösser and Blanc (1991)</p><p>Rhombophryne laevipes — Frost et al. (2006)</p><p>Remarks.</p><p>Type species of Mantiphrys Mocquard, 1895 . Parker (1934) considered R. laevipes to be a synonym of Mantipus hildebrandti Peters, 1883, and it was, in essence, described as such by Mocquard; the description given is of a specimen assigned to M. hildebrandti, but in a footnote Mocquard notes the numerous points of difference from Peters’ description, and states that, if Peters’ description is found to be accurate, then he would propose to name the new form Mantiphrys laevipes (Mocquard 1895) . The name was resurrected by Guibé (1946). Today, Mantipus hildebrandti is considered a junior synonym of Plethodontohyla inguinalis Boulenger, 1882 . We have verified this based on re-examination of the holotype of M. hildebrandti, ZMB 10440, which was reported lost by Glaw and Vences (1992) but was rediscovered by Bauer et al. (1996).</p><p>Some specimens formerly thought to be juveniles of this species have been reidentified in this study as R. ellae .</p><p>Holotype.</p><p>MNHN 1893.285, an adult male, collected on an unknown date on ‘ Montagne d’Ambre’, Diana Region, northern Madagascar by Mr Alluaud and Mr Belly (Figs 30, 31, 34) .</p><p>Paratypes.</p><p>None.</p><p>Referred specimens.</p><p>ZSM 902/2003 (FGMV 2002.0916), a juvenile specimen, collected on 19 February 2003, and ZSM 925/2003 (FGMV 2002.0998), an adult female, collected on 19–20 February 2003 on Montagne d’Ambre (no precise coordinates), Diana Region, northern Madagascar, by F. Glaw, R. D. Randrianiaina, and A. Razafimanantsoa; ZSM 200/2004 (FGZC 387) and ZSM 218/2004 (FGZC 423), a juvenile and an unsexed individual, collected on 19 February 2004 on the Voie des Milles Arbres (12.5200°S, 49.1756°E, 1052 m a. s. l.), Montagne d’Ambre National Park, Diana Region, northern Madagascar, by F. Glaw, M. Puente, R. Randrianiaina, A. Razafimanantsoa; ZSM 2188/2007 (FGZC 1283), a juvenile, collected on 13 March 2007 near the Cascade Antakarana (ca 12.52°S, 49.17°E, ca 1030 m a. s. l.), Montagne d’Ambre National Park, Diana Region, northern Madagascar, by P. Bora; ZSM 77/2018 (MSZC 0648), adult male (call voucher), collected at 14: 56 h on 19 November 2017 near the Gîte d’Étape (12.5204°S, 49.1718°E, 1066 m a. s. l.), Montagne d’Ambre National Park, Diana Region, northern Madagascar, by M. D. Scherz, J. H. Razafindraibe, A. Razafimanantsoa, O. Randriamalala, S. M. Rasolonjavato, R. Tiavina, and A. Rakotoarison (Fig. 32 A – C, 33 A – C); UADBA -A 62340 (MSZC 0526), a subadult, collected at night on 25 December 2017 at 12.5205°S, 49.1890°E, 1028 m a. s. l. in Montagne d’Ambre National Park, Diana Region, northern Madagascar, by M. D. Scherz, J. H. Razafindraibe, A. Razafimanantsoa, O. Randriamalala, S. M. Rasolonjavato, R. Tiavina, and A. Rakotoarison (Fig. 32 D – F); UADBA -A 62493 (MSZC 0654), a juvenile, collected on 19 November 2017 at 12.5206°S, 49.1862°E, 1008 m a. s. l. in Montagne d’Ambre National Park, Diana Region, northern Madagascar, by M. D. Scherz, J. H. Razafindraibe, A. Razafimanantsoa, O. Randriamalala, S. M. Rasolonjavato, R. Tiavina, and A. Rakotoarison (Fig. 32 G – I); UADBA -A- FGZC 1052, individual of unknown sex or maturity, collected on 25 February 2007 on Montagne d’Ambre in a pitfall trap (precise coordinates not available), Diana Region, northern Madagascar, by F. Glaw, P. Bora, H. Enting, J. Köhler, and I. Knoll.</p><p>The specimen ZSM 853/2003 (FGMV 2002.823), an adult female, collected on 3 February 2003 at ‘ Camp 1 ’ (13.9770°S, 48.4220°E, 751 m a. s. l.), Manongarivo Massif, Diana Region, northern Madagascar, by F. Glaw, R. D. Randrianiaina, and M. Vences (Fig. 33 H – I), is phylogenetically sister to all other specimens collected so far (Fig. 2), and differs genetically from all other specimens of R. laevipes by 1.24–2.46 % in the 16 S 3 ’ marker, and 4.90–5.45 % in the 16 S 5 ’ marker, but is consistently recovered as conspecific in species delimitation, and is therefore assigned to this species. A second specimen from the other side of this mountain, BMNH 1987.2300, an unsexed adult, collected on 25 February 1988 at ‘ Camp B’ (14.067°S, 48.283°E, 350 m a. s. l.), Manongarivo Massif, Sofia Region, northern Madagascar, by C. J. Raxworthy, agrees morphologically, and is consequently also assigned to this species.</p><p>Assignment of specimens.</p><p>ZSM 218/2004 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341104; Vieites et al. 2009) and COI barcoding (COI KF 611587; Perl et al. 2014). All of the referred material except ZSM 853/2003 is collected at the general type locality. The adult specimens conform with the holotype in overall morphology, and especially in the presence of white spots on the hidden surfaces of the legs. Assignment of UADBA -A- FGZC 1052 (not examined morphologically), UADBA -A 62340 (MSZC 0526), UADBA -A 62493 (MSZC 0654), ZSM 77/2018, and ZSM 853/2003 is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 77/2018 has been sequenced with the FrogCap hybrid capture probe set and is included in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. laevipes group, this species has a rather isolated position, and is probably sister to a subclade containing various other species based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) large body size, adult SVL 42.8–50.8 mm; (2) head wider than long (HW / HL 1.57–1.84); (3) moderately small eyes (ED / HL 0.30–0.34); (4) relatively long hindlimbs (HIL / SVL 1.62–1.86); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.11); (6) outer metatarsal tubercle absent; (7) second finger shorter than or equal in length to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin smooth with very few raised bumps in some individuals; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of white inguinal spots on a dark brown background; (13) presence of white spots on the posterior thigh and ventral surface of the shanks; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 12 analysed calls by one individual) highly pulsed, with (17) call duration 208–254 ms, (18) inter-call interval 1563–2186 ms, and (19) dominant frequency 612–631 Hz.</p><p>Rhombophryne laevipes is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 354, ‘ G’ at site 395, and ‘ C’ at site 611; in 16 S 3 ’, ‘ G’ at site 1075, ‘ T’ at site 1086, and ‘ G’ at site 1165.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. laevipes can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by smooth skin (vs rough), absence of chin barbels (vs presence), much longer hindlimbs (HIL / SVL 1.62–1.86 vs 1.10–1.41), and presence of white spots in the inguinal region and on the ventral shank (vs absence); from R. mangabensis, R. mavokely sp. nov., R. savaka, R. proportionalis, and members of the R. ellae and R. minuta species groups by much larger adult body size (SVL 42.8–50.8 mm vs 11.0– 30.1 mm), and presence of white spots in the inguinal region and on the ventral shank (vs absence); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence), and presence of white spots in the inguinal region and on the ventral shank (vs absence).</p><p>Within the R. laevipes species group, R. laevipes may not be distinguishable from R. nilevina morphologically (traits formerly thought to be diagnostic of the latter species are now known to be variable and overlap with R. laevipes; see the account, below), but can be distinguished by its shorter advertisement call (call duration 208–254 ms vs 526–534 ms), much shorter inter-call interval (1.5– 2.2s vs 42–126 s), and higher dominant frequency (612–631 Hz vs 526–534 Hz). It can be distinguished from R. botabota by the presence of white spots on the inguinal region, posterior thigh, and ventral shank (vs black inguinal ocelli or absence of inguinal spots, and no remarkable colour on the hindlimbs), and pulsed (vs tonal) advertisement call that is longer (call duration 208–254 ms vs 161–201 ms) and lower in dominant frequency (612–631 Hz vs 1256–1291 Hz). For distinction from newly described members of this species group, see their respective descriptions below.</p><p>Redescription of the holotype.</p><p>An adult male specimen in a good state of preservation, though strongly faded (Figs 30, 31). An L-shaped incision made in the dorsal skin, a further such incision made over the pectoral girdle to expose the pectoral girdle, and a straight incision along the right flank. The rictus is ripped on both sides, and a small pinhole through the middle of the chin.</p><p>Body robust. Head wider than long (HW / HL 1.62). Eyes quite large (ED / HL 0.31), pupils large, round. Snout rounded in dorsal and lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum distinct, oval, TDH / ED 0.57. Supratympanic fold distinct, strongly raised, running straight from posterior corner of eye over tympanum, curving ventrally posterior to the tympanum towards the insertion of the arm. Superciliary spines absent. Dorsal skin smooth. Ventral skin very finely granular. Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, varying in height, separated at the midline by a small gap. Choanae ovoid, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively long (FARL / SVL 0.403), strongly built. Fingers without webbing, moderately long, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger slightly shorter than fourth. Finger tips not expanded, rounded. Finger subarticular tubercles distinct, whitish, flat; inner metacarpal tubercle broad and flat, outer metacarpal / palmar tubercle indistinct. Hind limb strong, tibiotarsal articulation reaching tympanum when adpressed anteriorly along the body. TIBL / SVL 0.524. Inner metatarsal tubercle present, oblong, somewhat bulging. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles distinct, slightly raised; toe tips not enlarged, rounded.</p><p>After at least 128 years in preservative, the colouration of the specimen is largely lost and the skin rather translucent, especially ventrally. The only traces of colouration that remain are on the legs: The posterior thighs are dark brown with several large white spots on the ‘ hidden’ areas (not symmetrical). The left shank and foot are dark brown dorsally, and ventrally the shanks are also dark brown with round whitish spots. Colouration in life unknown.</p><p>Variation.</p><p>SVL 42.8–50.8 mm (n = 5). For variation in other measurements, see Table S 6. A dark brown chevron is often present over the scapular region (Figs 32, 33), and the side of the head is usually dark brown below the supratympanic fold, but lighter brown ventrally along the mouth and rictus (Figs 32 A, 32 D, 32 G, 33 A, 33 D, 33 F, 33 H). The inguinal region is dark brown with white spots, and the posterior thigh (e. g., Fig. 33 A, B, F, H) and ventral shank also have distinctive white or orange spots. These are present also in juveniles (e. g., Fig. 32 I). The dorsal colouration can be highly reticulated dark patterns (e. g., Fig. 33 G, H), or just scattered dark spots (e. g., Fig. 33 F). The ventral abdomen is highly variable, usually with numerous dark spots, and large unpigmented areas (Figs 32 C, 32 F, 32 I, 33 E, 33 I). The dorsal thigh and shank showed dark brown crossbands in all adult specimens we examined, but these were only very weakly visible in the juvenile UADBA -A 62493 (MSZC 0654) (Fig. 32 G – I). Juveniles and subadults tend to be quite orange in colour (Fig. 32 D – I, see also image captioned ‘ Rhombophryne sp., Montagne d’Ambre’ on page 448 of Glaw and Vences 2007), often with an X-shaped marking (which can be indistinct, cf. Fig. 32 I) on the dorsum. The areas that lack dark colouration appear to correspond to those that appear white in adults. Round spots on the ventral shank are less distinct in juveniles, and apparently become more distinct with age (Fig. 32). The dark colouration of the lateral head is always present.</p><p>Call.</p><p>The advertisement call recorded on 19 November 2017 (14: 30 h) near the Gîte d’Étape (air temperature 21.3 ° C; voucher specimen ZSM 77/2018, MSZC 0648) consists of a low-pitched, pulsed note of moderate duration, usually emitted in short call series containing 2 or 3 calls, although one single call was also recorded (Fig. 35). Calls (= notes) exhibit distinct amplitude modulation, with call energy starting at low level, rapidly increasing after about a fifth of the call’s duration, reaching its maximum at about half of the call’s duration, followed by a decrease towards the call’s end. Pulses are very narrowly spaced and very short in duration, with each pulse apparently shorter than 3 ms, but difficult to measure exactly. It is also not possible to count the number of pulses per note (= call), but pulse repetition rate can roughly be calculated in major parts of the note. Numerical parameters of 12 analysed calls are as follows: call duration (= note duration) 208–254 ms (228 ± 13 ms); inter-call intervals within regular call series 1563–2186 ms (1758 ± 231 ms); number of calls per call series 2 or 3 (2.1 ± 0.3); duration of regular call series 2008–4167 ms (2632 ± 899 ms); pulse repetition rate within calls about 310–320 pulses / second; dominant frequency 612–631 Hz (619 ± 6 Hz); prevalent bandwidth 320–3000 Hz; numerous parallel frequency bands recognizable with approximately 320 Hz spacing up to 9200 Hz, which represent a reflection of pulse rate (see Köhler et al. 2017).</p><p>Etymology.</p><p>Latin feminine nominative singular adjective derived from the words ‘ laevis’, meaning smooth or slippery, and ‘ pes’ meaning ‘ foot’.</p><p>Distribution.</p><p>This species is known from mid-elevation rainforests of (1) Montagne d’Ambre National Park (type locality). In an intensive survey on Montagne d’Ambre in 2017 and 2018, we did not find it above 1100 m a. s. l. on this mountain, nor did we find it below 900 m a. s. l. This matches observations by Raxworthy and Nussbaum (1994), who recorded this species to occur 900–1200 m a. s. l. Additionally, we here confirm its occurrence (2) on the Manongarivo Massif (at 350–751 m a. s. l.). Elevation: 350–1200 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne laevipes is a semi-fossorial or detriticolous species found in low- to mid-elevation humid forest. Males call during and shortly after rain, especially at night, from concealed positions within the leaf litter. Between November 2017 and January 2018, calling activity was greatest at the start of November, and diminished as rain frequency increased. This suggests that mating may take place at the start of the rainy season. When handled, captured males that had been calling struggled to deflate their vocal sacs, and emitted distress calls. Juvenile and subadult individuals were more commonly encountered than adults, usually active at night. After capture, we spent a short time observing the behaviour of ZSM 77/2018. Its initial escape attempts were long leaps. After calming down, when placed in leaf litter, it did not attempt to dig backwards, as is thought to be the dominant mode of digging in Rhombophryne . Instead, it attempted to hide by pushing itself under objects head-first, using its strong hindlimbs to push its body forward (Fig. 33 C). Eudeline et al. (2015) reported on the predation of this species by Compsophis albiventris .</p></div>	https://treatment.plazi.org/id/2EE70DCAA0BD58B188B233A576C2C2BF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
E3D7EAF3A773541F9F086BD7DBFBC844.text	E3D7EAF3A773541F9F086BD7DBFBC844.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne longicrus Scherz 2015	<div><p>Rhombophryne longicrus Scherz et al., 2015</p><p>Figures 2, 4 –7, 50, 51, 65, 66</p><p>Holotype.</p><p>ZSM 1630/2012 (FGZC 3653), an adult female with immature oocytes, collected on 28 November 2012 in the montane forest of the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.439&amp;materialsCitation.latitude=-13.675" title="Search Plazi for locations around (long 49.439/lat -13.675)">Sorata Massif</a> (ca 13.675°S, 49.439°E, ca 1580 m), Sava Region, northern Madagascar, by A. Rakotoarison, A. Razafimanantsoa, T. Rajoafiarison, F. M. Ratsoavina, O. Hawlitschek, and F. Glaw (Figs 65 A, 65 B, 67) .</p><p>Paratypes.</p><p>One specimen: UADBA -A 60271 (FGZC 3651; Fig. 65 C, D), an adult male, with the same collection data as the holotype .</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KR 025898; Scherz et al. 2015 a; 16 S 5 ’ KY 748101; Scherz et al. 2017 a). Paratype UADBA -A 60271 is used as the reference specimen for this species in COI barcoding (COI OL 790127; Belluardo et al. 2022). The paratype conforms well with the holotype in morphology, and is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 1630/2012 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>This species is sister to R. minuta, the only other species of the R. minuta group, based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small to medium body size, adult SVL 23.8–28.0 mm; (2) head wider than long (HW / HL 1.22–1.39); (3) moderately small eyes (ED / HL 0.40–0.41); (4) relatively long hindlimbs (HIL / SVL 1.81–1.83); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.09); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne longicrus is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 451, and ‘ T’ at site 547; in 16 S 3 ’, ‘ C’ at site 1096, ‘ G’ at site 1126, ‘ T’ at site 1163 and ‘ A’ at site 1315.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. longicrus can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by its smooth skin (vs rough), absence of chin barbels (vs general presence), and much longer hindlimbs (HIL / SVL 1.81–1.83 vs 1.10–1.41); from R. mangabensis and R. savaka by its much longer hindlimbs (HIL / SVL 1.81–1.83 vs 1.37–1.60); from R. mavokely sp. nov., R. kotrobaratra sp. nov., and members of the R. ellae species group by the absence of orange colouration on the posterior thigh and in the inguinal region (vs presence); from R. kotrobaratra sp. nov. further by absence of a dark chevron dorsally (vs presence); from R. laevipes, R. nilevina, R. sonaliae sp. nov., and R. anatiala sp. nov. by smaller adult body size SVL 23.8–28.0 mm vs 33.0– 57.2 mm), and absence of white spots in the inguinal region and on the posterior thigh (vs presence in most specimens); from R. botabota by more gracile body (vs rotund) and longer hindlimbs (HIL / SVL 1.81–1.83 vs 1.41–1.75); from R. proportionalis by larger adult body size (SVL 23.8–28.0 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence). For distinction from its sister species, R. minuta, see the account of that species, above. A detailed description of the species is given in Scherz et al. (2015 a).</p><p>Variation.</p><p>Both known specimens are highly similar to one another, but differ in size (SVL of male paratype 23.8 mm, female holotype 28.0 mm) and colouration, with the male being much yellower and having melanised flecks across its venter. For variation in morphometric measurements, see Table S 9.</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>Invariable Latin noun derived from the words longus, meaning ‘ long’, and crus meaning ‘ leg’, referring to the unusually long legs of this species.</p><p>Distribution.</p><p>This species is only known from (1) high elevation on the Sorata Massif (type locality). Elevation: ca 1580 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne longicrus is a scansorial species found in high-elevation humid forest. Scherz et al. (2015 a) documented the presence of small insects (mostly beetles) and a spider (probably Salticidae), along with moss, in the guts of the type specimens. The holotype contained&gt; 20 immature oocytes, the largest ranging from 1.3 to 1.6 mm in size. The authors speculated that this species may have a more saltatorial locomotion and lifestyle than its congeners. One of the type specimens was collected by A. Rakotoarison in a tree, sitting at the corner of a branch; this detail was forgotten from the original description, but could also suggest climbing behaviour, which has not previously been documented in this genus.</p></div>	https://treatment.plazi.org/id/E3D7EAF3A773541F9F086BD7DBFBC844	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
196EC850EEE25924A03B00697CA2951F.text	196EC850EEE25924A03B00697CA2951F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne mangabensis Glaw, Kohler & Vences 2010	<div><p>Rhombophryne mangabensis Glaw, Köhler &amp; Vences, 2010</p><p>Figures 2, 4 –9, 18 –20</p><p>Rhombophryne minuta [partim] – Glaw and Vences (1992)</p><p>Remark.</p><p>This species was referred to as R. minuta by Glaw and Vences (1992), where its call and morphology were first described.</p><p>One sequence belonging to this species, KU 724182 in GenBank, is erroneously ascribed to collection number ZCMV 891 (corresponding to ZSM 463/2005, a Platypelis grandis); the actual R. mangabensis specimen to which this sequence belongs is ZCMV 890.</p><p>Informal names.</p><p>The call was published as ‘ Plethodontohyla sp. (Nosy Mangabe) ’ by Vences et al. (2006). This species was listed as ‘ Rhombophryne sp. aff. minuta ’ by Glaw and Vences (2007), as ‘ Rhombophryne sp. 2 Nosy Mangabe’ by Wollenberg et al. (2008), and as ‘ Rhombophryne sp. 5 ’ by Vieites et al. (2009).</p><p>Holotype.</p><p>ZSM 470/2005 (ZCMV 886), an adult male, collected on 22 February 2005 on ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.77&amp;materialsCitation.latitude=-15.5" title="Search Plazi for locations around (long 49.77/lat -15.5)">Nosy Mangabe island’</a> (ca 15.50°S, 49.77°E, ca 50–100 m a. s. l.), Analanjirofo Region, northeastern Madagascar, by F. Glaw, R. D. Randrianiaina, and M. Vences (Figs 8, 9, 19) .</p><p>Paratypes.</p><p>Eight specimens: ZSM 471/2005 (ZCMV 2118), ZSM 472/2005 (ZCMV 2154), UADBA -A- ZCMV 2119, and UADBA -A- ZCMV 890, three adult specimens (probably males) and one subadult, with the same collection data as the holotype (Fig. 18 A – C); ZFMK 52769 –52771, three adult males, and ZFMK 52772, one subadult specimen, all collected on 13 March 1991 at the type locality, by F. Glaw and M. Vences (ZFMK 52769 is shown in Fig. 18 D).</p><p>Referred specimens.</p><p>ZSM 1573/2009, a subadult, collected in May / June 2003 on Nosy Mangabe (15.4959°S, 49.7693°E), Analanjirofo Region, northeastern Madagascar, by N. Lutzmann; ZSM 262/2016 (FGZC 5417), an unsexed adult, collected on 10 August 2016 near the ‘ Eco-Lodge chez Arol’ (ca 15.712°S, 49.964°E, 21 m a. s. l.), Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by F. Glaw, D. Prötzel, J. Forster, K. Glaw, and T. Glaw; MRSN A 2557, an unsexed adult, collected on 8 February 2002 at ‘ Camp 2 ’ Mahalevona (ca 15.4178°S, 49.9410°E, ca 669 m a. s. l.), Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by J. E. Randrianirina; UADBA - A-APR 12051 (Fig. 18 E), an unsexed specimen, collected on 25 November 2015 at Ambohitsitondroina (15.5694°S 50.0034°E, 609 m a. s. l.), Masoala National Park, Analanjirofo Region, northeastern Madagascar, by A. Raselimanana.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ EU 341109 / KU 724181; 16 S 5 ’ EU 341109; Vieites et al. 2009) and COI barcoding (COI KF 611588 / KU 937775; Perl et al. 2014). All paratypes and referred specimens conform well with the holotype in morphology. The paratypes are all topotypical. Assignment of specimens ZSM 471/2005, UADBA -A- ZCMV 2119, MRSN A 2557, and ZSM 262/2016 is further confirmed by molecular taxonomic identification (Fig. 2). MRSN A 2557 and ZSM 262/2016 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. testudo group, this species is sister to R. savaka based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 19.6–23.2 mm; (2) head wider than long (HW / HL 1.60–1.86); (3) moderately small eyes (ED / HL 0.34–0.38); (4) short hindlimbs (HIL / SVL 1.54–1.60); (5) small inner metatarsal tubercle (IMTL / FOL 0.09–0.12); (6) indistinct outer metatarsal tubercle; (7) second finger roughly equal in length to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular with a few scattered warts, especially a suprascapular pair outlined in black; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles reduced, thin, curving; and (16) advertisement call (based on 20 analysed calls by one individual) tonal, with (17) call duration 180–237 ms, (18) inter-call interval 419–822 ms, and (19) dominant frequency 3300–3550 Hz.</p><p>Rhombophryne mangabensis is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 383, ‘ G’ at site 567 and ‘ C’ at site 641; in 16 S 3 ’, A’ at site 917, ‘ A’ at site 1088, ‘ G’ at site 1090, and ‘ A’ at site 1127.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. mangabensis can easily be distinguished from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence); from members of the R. minuta species group by less gracile body and much shorter limbs; from R. proportionalis by its much larger body size (SVL 19.6–23.2 mm vs 11.0– 12.3 mm); from members of the R. ellae species group by the absence of inguinal spots and reddish orange colouration on the hindlimbs (vs presence); from members of the R. laevipes species group by the absence of inguinal spots or ocelli (vs presence in most species), and smaller adult body size (SVL 19.6–23.2 mm vs 24.1–57.2 mm). Within the R. testudo species group, see accounts of R. testudo, R. coudreaui, and R. matavy above for distinction from those species. From its sister species, R. savaka, it can be distinguished by the less pronounced diastema in vomerine teeth, slightly longer relative tibia length (TIBL / SVL 0.44–0.46 vs 0.37–0.42), absence of inguinal spots (vs presence), and by advertisement call with slightly longer call duration (180–237 ms vs 121–142 ms), generally longer inter-call interval (419–822 ms vs 302–451 ms), and lower dominant frequency (3300–3550 Hz vs 4144–4210 Hz). For distinction from newly described species, see accounts of those species, below.</p><p>Variation.</p><p>SVL 19.6–23.2 mm (n = 4 adults; two presumed subadults measure 16.5 and 18.9 mm, respectively); full measurements are presented in Table S 5. Measurements are largely consistent with those given in the original description by Glaw et al. (2010 b). All specimens are consistent in morphology and colouration, including the newly referred specimens from mainland Madagascar (ZSM 262/2016 and MRSN A 2557). Two raised black markings with small light brown points in the suprascapular region are consistently present as part of the variably complete anterior chevron marking; a posterior chevron running from the inguinal region anteriorly over the dorsum is sometimes incomplete at the midline but is usually indicated (Fig. 18 A, C – E). Shanks often exhibit a broad middle stripe and two narrow stripes proximal and distal to this stripe (Fig. 18 A, C – E).</p><p>Call.</p><p>The advertisement call recorded on 13 March 1991 at Nosy Mangabe (air temperature unknown) consists of a tonal note of moderate duration emitted in regular call series containing a variable number of calls (2–5) repeated at rather regular succession (Fig. 20). Each call exhibits some irregular amplitude modulation, with 2 or 3 peaks recognizable within each call, with the terminal peak having the highest energy. Slight upward frequency modulation is evident in each call until the last third of the call’s duration, followed by a terminal drop in frequency. Numerical parameters of 20 analysed calls are as follows: call duration (= note duration) 180–237 ms (198 ± 15 ms); inter-call intervals within regular call series 419–822 ms (484 ± 113 ms); duration of regular call series 820–2895 ms (1779 ± 925 ms); dominant frequency 3300–3550 Hz (3434 ± 85 Hz); prevalent bandwidth 2800–7500 Hz; weak frequency bands recognizable at approximately 5250 and 6760 Hz. Calls from the same recording (Vences et al. 2006, CD 3, track 31) have been described by Glaw et al. (2010 b) using a slightly different terminological scheme.</p><p>Etymology.</p><p>Toponym referring to Nosy Mangabe, the island where the species was originally discovered.</p><p>Distribution.</p><p>This species was originally known only from (1) Nosy Mangabe (type locality), but its presence on the Masoala Peninsula was reported by Raherilalao et al. (2016) and confirmed by Belluardo et al. (2022). The extent of its range on the peninsula constitutes three further distinct sites: (2) Tampolo (under which we include the nearby ‘ Eco-Lodge chez Arol’), (3) Mahalevona, and (4) Ambohitsitondroina (15.5694°S 50.0034°E, 609 m a. s. l.; Fig. 18 E). Elevation: ca 15–669 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne mangabensis is a semi-fossorial or detriticolous species found in low-elevation humid forest. Specimens were collected at night during heavy rain, during which many calls were heard in an intense chorus. Males called from concealed positions in the leaf litter of ‘ apparently primary low-altitude rainforests’ (Glaw et al. 2010 b: 240). Calling has not been directly observed, but the call described above certainly belongs to this species, as specimens caught while searching for the source of calling were inflated with air, typical for calling Rhombophryne . Active specimens were encountered on the forest floor during the day, in the evening, and at night. One individual was captured by pitfall traps set up in a valley with closed-canopy humid forest associated with thick leaf litter and many rotten logs.</p><p>Karyotype.</p><p>Aprea et al. (2007) reported on the karyotype of ‘ Rhombophryne sp. aff. minuta ’ from Masoala, but the specimen number they reported, MRSN A 3656, is from Manarikoba in Tsaratànana, and is therefore not referrable to this species, but in fact belongs to Rhombophryne quentini sp. nov., described below. The karyotype of R. mangabensis is therefore unknown.</p></div>	https://treatment.plazi.org/id/196EC850EEE25924A03B00697CA2951F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
E6C1FEBE943E58CAAC2A3F3CCF8B3DC8.text	E6C1FEBE943E58CAAC2A3F3CCF8B3DC8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne maraorao Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne maraorao sp. nov.</p><p>Figures 2, 4 –9, 24, 25</p><p>Remark.</p><p>Specimens assignable to this species were attributed to Rhombophryne testudo by Blommers-Schlösser and Blanc (1991) and Guibé (1978), as R. coudreaui by Andreone (2003), as both R. coudreaui and R. testudo by Glaw and Vences (2007), and as R. coudreaui by Andreone and Randrianirina (2008).</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 9 ’ by Vieites et al. (2009), ‘ Rhombophryne sp. Ca 09 ’ by Belluardo et al. (2022), and ‘ Rhombophryne sp. 9 ’ and ‘ CCS “ Sp. 9 ” ’ by Carné and Vieites (2024).</p><p>Justification of the new species.</p><p>According to current knowledge, this is the allopatric sister lineage of R. coudreaui, which it also closely resembles morphologically. However, it is genetically highly divergent (10 % uncorrected p distance in the 16 S 3 ’ fragment), there is no haplotype sharing with R. coudreaui or any other Rhombophryne species in the nuclear gene RAG 1 (Fig. 4), and it has subtle but recognizable morphological differences (see Diagnosis below). We are therefore convinced that it represents an evolutionarily independent lineage.</p><p>Holotype.</p><p>MRSN A 2115 (FN 7705), an adult male, collected on 2 December 1998 in the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=50.02&amp;materialsCitation.latitude=-15.3" title="Search Plazi for locations around (long 50.02/lat -15.3)">Ambatoledama Corridor</a> (Andasin’i Governera) (ca 15.30°S, 50.02°E, 650 m a. s. l.), Masoala Peninsula, Sava Region, northeastern Madagascar, by F. Andreone and J. E. Randrianirina (Figs 8, 9, 24 D, 25) .</p><p>Paratypes.</p><p>Three specimens: MRSN A 2496 (no field number) and MRSN A 2497 (RJS 0496), an adult female and a juvenile, respectively, collected on 8 February 2002 at ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.9403&amp;materialsCitation.latitude=-15.4177" title="Search Plazi for locations around (long 49.9403/lat -15.4177)">Camp 7</a> ’ Antsiranana Faritany (15.4177°S, 49.9403°E, ca. 800 m a. s. l.), Amparihy, Masoala Peninsula, Analanjirofo Region, northeastern Madagascar, by J. E. Randrianirina ; MRSN A 6929 (FN 7292), an adult male, collected on 13 December 1997 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.43&amp;materialsCitation.latitude=-14.5433" title="Search Plazi for locations around (long 49.43/lat -14.5433)">‘Camp 3’ Antsinjorano</a> (14.5433°S, 49.4300°E, ca 950 m a. s. l.), Ambolokopatrika-Betaolana Forest, Sava Region, northeastern Madagascar by F. Andreone, G. Aprea &amp; J. E. Randrianirina (Fig. 24 A – C) . Note: MRSN A 6929 formerly bore the collection number MRSN A 2066 (Andreone 2003), but this number was apparently duplicated, so a new number was issued.</p><p>Referred specimens.</p><p>MNHN 1973.576, an unsexed specimen, collected on 19 December 1972 at ‘ Sambava, km 17 route d’Andapa’ near the Marojejy Massif, Sava Region, northeastern Madagascar, by C. Blanc. This specimen was assigned morphologically to ‘ R. coudreaui ’ by Andreone and Randrianirina (2008), but the material used for comparison to R. coudreaui in that work constitutes the type series of R. maraorao sp. nov., and does not include any material of R. coudreaui sensu stricto . Although we were not able to re-examine this specimen, its assignment to R. maraorao sp. nov. is partly supported by the textual description from Andreone and Randrianirina (2008: 47), ‘ this specimen (likely a female 34.9 mm long) does not show an evident tympanum, lacks protruding jaw whiskers and shows the typical rhomboidal drawing on the head, and has a rather dark belly’. This assignment remains tentative, however, and requires confirmation by either morphological inspection or museomic sequencing.</p><p>MRSN A 5524, originally reported by Andreone and Randrianirina (2008) as being from Tsingy de Bemaraha in western Madagascar, was found by Belluardo et al. (2022) to be genetically identical with R. maraorao sp. nov. Belluardo et al. (2022) showed that its locality was probably confused, and it is most likely from somewhere in northeastern Madagascar. It is referred to this species on genetic and morphological grounds, but its incorrect former locality is not considered as part of the distribution of the species. We also do not include it in the diagnosis of the species, but comment on how its morphology expands our knowledge of the species in the Variation section, below.</p><p>BMNH 1987.331, an unsexed possible subadult, collected on 31 August 1986 in Anandrivola Forest (15.775°S, 49.600°E, ca 580 m a. s. l.), Analanjirofo Region, northeastern Madagascar, by C. J. Raxworthy, is tentatively assigned to this species based on its small size, rotund body and short head, rough skin, and the second finger being slightly shorter than the fourth (the last character being the key feature distinguishing it from R. coudreaui; see Diagnosis, below).</p><p>Specimens previously attributed to R. coudreaui from Marojejy (Raselimanana et al. 2000) and Maroantsetra (IUCN SSC Amphibian Specialist Group 2016 d) may be referable to R. maraorao sp. nov., as these two species are only now being recognised as distinct, with R. maraorao sp. nov. apparently occurring north of R. coudreaui . However, these records may also refer to R. mavokely sp. nov. (described below), which may look similar in its flat body profile and broad head, so carefuly scrutiny is warranted, and we do not include these locations in the distribution of this species.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 5 ’ AY 594072; Andreone et al. 2005; 16 S 3 ’ AY 594110; Vieites et al. 2009). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of MRSN A 2497 is further confirmed by molecular taxonomic identification. MRSN A 2115 and MRSN A 2497 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. testudo group, this species is sister to R. coudreaui based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small to medium body size, adult SVL 22.5–28.9 mm; (2) head much wider than long (HW / HL 1.69–1.88); (3) small eyes (ED / HL 0.25–0.27); (4) short hindlimbs (HIL / SVL 1.10–1.28); (5) well-developed inner metatarsal tubercle (IMTL / FOL 0.13–0.19); (6) indistinct outer metatarsal tubercle; (7) second finger slightly to distinctly longer than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin rough to rugose with longitudinal rows of tubercles; (10) presence of pronounced, pointed chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) broad but not fortified skull, without expansion of squamosal, maxilla, or posterior process of angulosplenial; and (15) clavicles absent.</p><p>Rhombophryne maraorao sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 406, ‘ T’ at site 459, and ‘ T’ at site 500; in 16 S 3 ’, ‘ A’ at site 1057, ‘ A’ at site 1075, and ‘ A’ at site 1092.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. maraorao sp. nov. can easily be distinguished from all other species except R. testudo, R. coudreaui, and R. matavy by its rotund body and short, broad head. Among these species, it can be distinguished from R. testudo by smaller body size (SVL 22.5–28.9 mm vs 33.0– 43.1 mm), longer head (HW / HL 1.69–1.88 vs 2.00–2.60), rougher skin, smaller chin barbels, and much less fortified skull; and from R. matavy by smaller body size (SVL 22.5–28.9 mm vs 40–50 mm), longer head (HW / HL 1.69–1.88 vs 1.88–2.05), presence of chin barbels (vs absence), and less fortified skull. Rhombophryne maraorao sp. nov. is most closely related to, and most similar to, R. coudreaui . These species are difficult to distinguish morphologically. The most reliable character for distinguishing these species is the presence of pronounced pointed barbels along the lower lip in R. maraorao sp. nov., versus fewer, less pronounced barbels in R. coudreaui . They may also differ in eye size, with R. coudreaui having larger relative eye size (ED / HL 0.29–0.31) than R. maraorao sp. nov. (ED / HL 0.25–0.27); and in relative finger length, with the second finger at least slightly longer than the fourth in R. maraorao sp. nov., vs equal in length in R. coudreaui .</p><p>Description of holotype.</p><p>An adult male specimen in a moderately good state of preservation, though fairly soft (Figs 8, 9). An incision made along the right flank to check the sex. Testes distinct.</p><p>Body robust and rotund. Head wider than long (HW / HL 1.76). Eyes quite small (ED / HL 0.25), pupils large, round. Snout rounded in dorsal and lateral views. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, round, TDH / ED 0.594. Supratympanic fold moderately distinct, raised but tubercular and so uneven in height, forming a sigmoid curve from the line of the top of the eye straight over the tympanum, curving down behind the tympanum, and then posteriorly to the top of the arm. A row of five (right) or three (left) small superciliary spines above each eye, as well as small granules distributed over the top of the eye and head. A row or pronounced tubercles (barbels) along the lower lip. Tongue partly removed as a tissue sample. It was attached anteriorly, broad, and unlobed. Maxillary and vomerine teeth present, vomerine teeth present in two curving rows separated at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short and thick, the upper arm hardly distinguishable from the body cavity (FARL / SVL 0.323). Fingers without webbing, all short, relative lengths 1 &lt;2 = 4 &lt;3 (Fig. 9). Finger subarticular tubercles distinct, round, whitish; inner metacarpal tubercle broad and flat, outer metacarpal / palmar tubercle distinct and round. Hind limb thick, tibiotarsal articulation reaching insertion of arm when adpressed anteriorly along the body. TIBL / SVL 0.304. Inner metatarsal tubercle present, large. Outer metatarsal tubercle faint. Toes not webbed; first toe distinctly reduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles distinct, broad and round; toe tips not enlarged (Fig. 9).</p><p>After 25 years in preservative, the colouration of the specimen is largely faded. It is beige in base colour, with light brownish symmetrical markings. These border the back of the head region, the scapular region, and from the inguinal region to the posterior third of the back, forming a beige cross-like shape over the central dorsum. The legs too are beige above with faint light brown crossbands on the thigh, more distinct crossbands on the shank, and a dark posterior tarsus. The hidden surfaces of the thighs are whitish, but there is a trapezoidal brown marking around the vent. The dorsal and lateral foot are beige with very fine brown specks. The ventral body has a beige base-colour, intensely speckled with brown (more distinct than above), especially over the chin and pectoral girdle. This pattern extends onto the ventral arms and legs. Colouration in life (Fig. 24 D) was much browner than the specimen is in preservative. The outline of the markings on the dorsum in beige-cream was particularly distinctive.</p><p>Variation.</p><p>SVL 22.5–28.9 mm (n = 3; a putative subadult, MRSN A 2497, measures 20.9 mm, and the referred specimen with uncertain collection locality, MRSN A 5524, measures 29.8 mm). For variation in measurements, see Table S 5. All specimens have pronounced tubercles along the lower lip. Most individuals are a brownish cream colour dorsally in preservative, much lighter in colour than most R. coudreaui, but MRSN A 5524 is much browner and more mottled, and the colour of this species in life is also dark brown (Fig. 24); differences in colour of the specimens may be due to preservation fluids or other factors, and are not taken to be diagnostic. The referred specimen BMNH 1987.331 is rather dark, but has also been stained to have blue-green bones, which may have altered its skin colour as well. Much of the body patterning is consistent, especially the dark suprascapular spots which define the short edge of a trapezoid where the eyes form the ends of the long edge. The sacral region also has dark markings, though less consistently than in R. coudreaui . In four of the five specimens (not MRSN 2497) the chin is marbled brown and cream. The first toe of MRSN A 2496 is less distinctly reduced than other specimens, and is connected to the second toe with distinct webbing on the left side but not on the right. MRSN A 5524 has a substantially larger relative inner metatarsal tubercle than the other individuals (IMTL / FOL 0.23 vs 0.13–0.19).</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>The Malagasy adjective maraorao meaning ‘ rough to the touch’, in reference to the rugose skin of this species in life and in preservative. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>Currently known from (1) Ambatoledama Corridor (Andasin’i Governera); (2) Amparihy, (3) Ambolokopatrika-Betaolana, and possibly (4) Anandrivola. Referral of material from Marojejy to this species remains too uncertain to include in its distribution at present. Elevation: ca 650–950 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne maraorao sp. nov. is a fossorial species found in low- to mid-elevation humid forest; its ecology is poorly known, but presumably much like R. coudreaui . All of the individuals from Masoala and Ambolokopatrika-Betaolana were unearthed and collected whilst digging in the soil to install pitfall traps. The gut of MRSN A 6929 contained a beetle, an amphipod, a small ant, pieces of an arthropod, and detritus (several leaf and bark fragments).</p></div>	https://treatment.plazi.org/id/E6C1FEBE943E58CAAC2A3F3CCF8B3DC8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
B217C77146DA50B7A7C2C031B7BF9F63.text	B217C77146DA50B7A7C2C031B7BF9F63.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne matavy D'Cruze, Kohler, Vences & Glaw 2010	<div><p>Rhombophryne matavy D’Cruze, Köhler, Vences &amp; Glaw, 2010</p><p>Figures 2, 4 –9, 15 –17</p><p>Plethodontohyla matavy — Peloso et al. (2016), ex-error due to sample confusion</p><p>Rhombophryne matavy — Scherz et al. (2016 b)</p><p>Remark.</p><p>This species is only known from the type series.</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. nov. ’ by D’Cruze et al. (2008), and ‘ Rhombophryne sp. 8 ’ by Vieites et al. (2009).</p><p>Holotype.</p><p>ZSM 1628/2008 (FGZC 1888), an adult male, collected on 27 February 2008, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.2233&amp;materialsCitation.latitude=-12.4628" title="Search Plazi for locations around (long 49.2233/lat -12.4628)">Forêt d’Ambre Special Reserve</a> (now part of Montagne d’Ambre National Park) (12.4628°S, 49.2233°E, 482 m a. s. l.), Diana Region (part of former Antsiranana Province), northern Madagascar, by N. D’Cruze, F. Glaw, J. Köhler, and local guides (Figs 8, 9, 15, 16) .</p><p>Paratypes.</p><p>Three specimens: ZSM 1629/2008 (FGZC 1889), UADBA -A- FGZC 1890, and UADBA -A- FGZC 1891, three adult males with the same collection data as the holotype except that UADBA -A- FGZC 1891 was collected on 28 February 2008.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ FJ 559298; Vieites et al. 2009) and COI barcoding (COI KF 611589; Perl et al. 2014). All paratypes agree morphologically with the holotype, and are topotypical. Assignment of UADBA -A- FGZC 1890 is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 1628/2008 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. testudo group, this species is sister to R. testudo based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) large body size, adult SVL 40–50 mm (see ‘ Variation’ below for discussion of these values); (2) head much wider than long (HW / HL 1.88–2.05); (3) small eyes (ED / HL 0.24–0.28); (4) short hindlimbs (HIL / SVL 1.17–1.25); (5) large inner metatarsal tubercle (IMTL / FOL 0.21–0.26); (6) pronounced outer metatarsal tubercle; (7) second finger rough equal in length to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin with scattered warts and ridges; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) broad and slightly fortified skull, but without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles absent; and (16) advertisement call (based on 15 analysed calls by one individual) tonal, with (17) call duration 161–209 ms, (18) inter-call interval 267–353 ms, and (19) dominant frequency 1588–1688 Hz.</p><p>Rhombophryne matavy is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 474, ‘ T’ at site 475, and ‘ C’ at site 538; in 16 S 3 ’, ‘ A’ at site 1018, ‘ C’ at site 1072, and ‘ T’ at site 1078.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. matavy can easily be distinguished from all other species except R. testudo, R. coudreaui, and R. maraorao sp. nov. described below by its rotund body and short, broad head. Among these species, see the accounts of R. testudo (its sister species) and R. coudreaui above for distinction from those species; for distinction from R. maraorao sp. nov., see the description of that species, below.</p><p>Variation.</p><p>Adult male SVL 44.6–45.1 mm (n = 2); full measurements are presented in Table S 5. D’Cruze et al. (2010) measured the same specimens and reported their SVLs to be substantially larger (46.7–47.6 mm). The reason for the discrepancy in measured size is not clear. D’Cruze et al. (2010) also measured two paratypes, UADBA -A- FGZC 1890 and 1891, as 39.4 and 49.4 mm, respectively. For this reason, we have given the SVL in the diagnosis above as between 40 and 50 mm.</p><p>Call.</p><p>The advertisement call of the holotype (ZSM 1628/2008), recorded on 27 February 2008 at Forêt d’Ambre Special Reserve (air temperature unknown), consists of a tonal note of moderate duration emitted in regular call series containing 5 calls repeated at regular succession (n = 3; Fig. 17). Each call exhibits considerable amplitude modulation, with highest energy present at approximately the last third of the call’s duration. Slight upward frequency modulation is evident in each call, followed by a fast terminal drop in frequency. Numerical parameters of 15 analysed calls are as follows: call duration (= note duration) 161–209 ms (183 ± 14 ms); inter-call intervals within regular call series 267–353 ms (299 ± 22 ms); number of calls per call series 5 (5.0 ± 0.0); duration of regular call series 2091–2202 ms (2130 ± 48 ms); dominant frequency 1588–1688 Hz (1652 ± 16 Hz); prevalent bandwidth 750–6800 Hz; frequency bands recognizable at approximately 880, 2500, 3360, 4160, 5060, and 5840 Hz. Calls from the same recording have been described by D’Cruze et al. (2010) using a slightly different terminological scheme.</p><p>Etymology.</p><p>The Malagasy adjective matavy meaning ‘ fat’, in reference to the body shape of the species, especially when inflated (D’Cruze et al. 2010). Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is only known from (1) the low-elevation parcel of forest in Montagne d’Ambre National Park, formerly known as Forêt d’Ambre Special Reserve (type locality). Elevation: ca 482 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne matavy is a fossorial species found in low-elevation humid forest; its ecology is poorly known, but presumably much like its sister species, R. testudo . Highest calling activity (in choruses) was noticed during and just after rain, and the specimens were collected at night, mostly up to 5 cm deep in the leaf litter, though one was found active on top of it (D’Cruze et al. 2010). D’Cruze et al. (2010) noted a defensive behaviour involving the stretching of the limbs and arching the back, like R. testudo . They reported the faeces of one individual (ZSM 1628/2008) to include the remains of several insects, including beetles.</p></div>	https://treatment.plazi.org/id/B217C77146DA50B7A7C2C031B7BF9F63	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
50752E50712652C78456F5659A1CA078.text	50752E50712652C78456F5659A1CA078.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne mavokely Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne mavokely sp. nov.</p><p>Figures 2, 5 –9, 26 –29</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. Ca 17 ’ by Belluardo et al. (2022), and ‘ Rhombophryne sp. 17 ’ and ‘ CCS “ Sp. 17 ” ’ by Carné and Vieites (2024).</p><p>Justification of the new species.</p><p>The new species is placed in a clade containing R. mangabensis, R. savaka, and an undescribed candidate species (R. sp. Ca 16) from the Andapa area for which only genetic data are known. It differs, however, from all nominal species of Rhombophryne by very high genetic divergences: 10.3 % and 10.9 % uncorrected p distance in the 16 S 3 ’ fragment from R. mangabensis and R. savaka, respectively; and&gt; 11 % from all other species (Belluardo et al. 2022). It furthermore differs from R. mangabensis and R. savaka by the presence of reddish-orange colouration on the posterior thigh, and from the syntopic R. savaka by bioacoustic and further morphological characters (see Diagnosis below). In addition, it is phylogenetically sister to the R. mangabensis / R. savaka subclade (Belluardo et al. 2022) and thus cannot be considered conspecific with either of these species. Its status as distinct species is therefore beyond doubt.</p><p>Holotype.</p><p>ZSM 521/2016 (MSZC 0269), an adult male (call voucher), collected on 22 November 2016 in low elevation primary rainforest within <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.7782&amp;materialsCitation.latitude=-14.44" title="Search Plazi for locations around (long 49.7782/lat -14.44)">Marojejy National Park</a> (14.4400°S, 49.7782°E, 402 m a. s. l.), Sava Region, northeastern Madagascar, by M. D. Scherz, J. H. Razafindraibe, and A. Razafimanantsoa (Figs 8, 9, 26 A – C, 28) .</p><p>Paratypes.</p><p>Three specimens: ZSM 517/2016 (ZCMV 15088), an unsexed adult, collected on the morning of 16 November 2016 at ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.785&amp;materialsCitation.latitude=-14.446" title="Search Plazi for locations around (long 49.785/lat -14.446)">Camp 0</a> ’ (ca 14.446°S, ca 49.785°E, ca 310 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, M. Vences, A. Rakotoarison, M. C. Bletz, J. H. Razafindraibe, and A. Razafimanantsoa (Fig. 26 D – G) ; ZSM 518/2016 (MSZC 0310), a juvenile, and UADBA -A- MSZC 0307 (Fig. 26 H – K), an unsexed adult, both collected in the evening of the 26 November 2016 from the same locality as ZSM 517/2016, by M. D. Scherz, J. H. Razafindraibe, A. Razafimanantsoa, and C. R. Hutter .</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 5 ’ OL 780607; Belluardo et al. 2022). All paratypes conform well with the holotype in morphology, and are from near the type locality. Assignment of specimens ZSM 518/2016, ZSM 517/2016, and UADBA -A- MSZC 0307 is further confirmed by molecular taxonomic identification (Fig. 2). RAG 1 sequence of this species is so far not available.</p><p>Definition.</p><p>Within the R. testudo group, this species is most closely related to R. mangabensis, R. savaka, and its possible sister lineage R. sp. Ca 16 based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 21.2–21.3 mm; (2) head wider than long (HW / HL 1.60–1.68); (3) moderately small eyes (ED / HL 0.36–0.38); (4) short hindlimbs (HIL / SVL 1.50–1.61); (5) small inner metatarsal tubercle (IMTL / FOL 0.12–0.13); (6) outer metatarsal tubercle absent; (7) second finger distinctly longer than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular with a few scattered warts, especially a suprascapular pair outlined in black; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) presence of reddish orange in the inguinal region and on the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 16 analysed calls by one individual) tonal, with (17) call duration 231–262 ms, (18) inter-call interval 623–808 ms, and (19) dominant frequency 3178–3265 Hz.</p><p>Rhombophryne mavokely sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 315, ‘ C’ at site 326, and ‘ T’ at site 331; in 16 S 3 ’, ‘ G’ at site 937, ‘ G’ at site 938, and ‘ A’ at site 956.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. mavokely sp. nov. can easily be distinguished from all species except R. ellae, R. ornata, and two other species described below ( R. quentini sp. nov. and R. kotrobaratra sp. nov.), by the presence of reddish orange colouration on the posterior thigh. Among these species, it can be distinguished from R. ellae by a more dorsoventrally compressed habitus, smaller body size (SVL 21.2–21.3 mm vs 24.9 mm), wider head (HW / HL 1.60–1.68 vs 1.47), and shorter hindlimbs (HIL / SVL 1.50–1.61 vs 1.71); and from R. ornata by the absence of superciliary spines (vs presence). For distinction from R. quentini sp. nov. and R. kotrobaratra sp. nov., see the descriptions of those species, below. When the orange of the limbs is not visible there is also some resemblance to R. coudreaui and R. maraorao, which however differ by generally larger adult body size (SVL 21.5–28.9 vs 21.2–21.3 mm), much shorter hindlimbs (HIL / SVL 1.10–1.30 vs 1.50–1.61), and presence of chin barbels (vs absence).</p><p>Rhombophryne mavokely sp. nov. occurs syntopically or in near sympatry with the closely related R. savaka . Aside from colouration, R. mavokely sp. nov. can be distinguished from R. savaka by its rougher dorsal skin with more ridges (vs relatively smooth), and by the longer advertisement call duration (231–262 ms vs 121–142 ms), longer inter-call interval (623–808 ms vs 221–395 ms), and slightly lower dominant frequency (3178–3265 Hz vs 3714–3861 Hz).</p><p>Description of holotype.</p><p>A specimen in a good state of preservation (Fig. 8). A tissue sample was taken from its left thigh for genetic analysis.</p><p>Body rotund. Head wider than long (HW / HL 1.68). Eyes quite small (ED / HL 0.36), pupils small, round. Snout rounded in dorsal view, truncate in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, round, TDH / ED 0.62. Supratympanic fold indistinct, demarcated at its anterior end by a dark comma-shaped marking. In life it was more distinct, curving from the posterior corner of the eye over the tympanum and towards the insertion of the arm (Fig. 26 A). Superciliary spines absent, but two white dots above each eye. Small granules and ridges distributed over the body in life are largely flat in preservative but mostly still marked by a lighter colouration. Most prominent of these are a pair of markings in the suprascapular region, and a series of three (left) and two (right) bumps along the ilia and in the sacral region. Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth present in two curving rows separated at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.417), fairly slender. Fingers without webbing, all short, relative lengths 1 &lt;4 &lt;2 &lt;3, second finger slightly longer than fourth (Fig. 9). Finger tips not expanded, slightly pointed. Finger subarticular tubercles indistinct, whitish; inner metacarpal tubercle broad and flat, outer metacarpal / palmar tubercle indistinct, greyish. Hind limb thick, tibiotarsal articulation reaching the eye when adpressed anteriorly along the body. TIBL / SVL 0.461. Inner metatarsal tubercle present, oblong, whitish. Outer metatarsal tubercle visible only as a slightly lightened area of the outer metatarsus. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles distinct, greyish, rather flat; toe tips slightly enlarged, slightly pointed (Fig. 9).</p><p>After seven years in preservative, the colouration of the specimen is quite vibrant, though lighter than in life (Fig. 8). The base colour is a soft pink, becoming more reddish posteriorly and mauve anteriorly, especially over the head. The dorsal surfaces of the eyes are dark mauve. There are several dark brown markings on the body, particularly the comma-shaped markings in the supratympanic region, the black spots with mauve centres on the suprascapular region and iliac / sacral region, and crossbands on the forelimbs. Additionally, the ventral surfaces of the feet, and especially the ventral portion of the posterior thigh are almost black. The darkened area of the thigh is not trapezoidal as in some other Rhombophryne species, but really extends over the entire length of the posterior thigh. It is separated from the dorsal colouration by areas that are cream in preservative. The dorsal surfaces of the legs have faint brown crossbands, two on the thigh, two on the shank, and two or three on the feet. The dorsal surface of the foot is laterally pinkish, but over most of its surface is mottled brown, with a light annulus before each toe tip. The dorsal surfaces of the hands are similar. The flank colouration merges seamlessly with the venter (i. e., there is no dorsolateral colour border, nor indeed a lateral head colour border). The ventral colouration is largely cream mottled with a greyish brown. The chin is darkest, with the posterior extent of the vocal sac clearly demarcated. The ventral arms are a light cream, less yellow than over the hind limbs. Colouration in life was like in preservative but generally more orange.</p><p>Variation.</p><p>SVL 21.2–21.3 mm (n = 2). For variation in other measurements, see Table S 5. Morphology is highly consistent. All specimens collected were characterised by highly similar colouration and patterning, except that in ZSM 517/2016 the posterior thigh is bright pink where it is cream in ZSM 521/2016, the thigh is more clearly striated, and the dark colouration around the cloaca is more restricted in distribution. All known specimens possess a dark suprascapular spot with a light middle, and a dark, almost comma-shaped marking in the supratympanic region (Figs 26, 27), similar to R. mangabensis (Fig. 18), and distinct from R. savaka (Fig. 21).</p><p>Call.</p><p>The holotype (ZSM 521/2016) was collected after recording a call in primary low-altitude rainforest, under leaf litter, during dry weather. It was not seen calling, but was collected from the location of the call emission, and the call is similar to but distinctly different from that of R. savaka, and we are therefore confident that it is assignable to this species and this individual. The advertisement call, recorded on 23 November 2016 (20: 20 h) at an unknown air temperature, consists of a tonal note of moderate duration emitted in regular call series (Fig. 29). Each call exhibits some amplitude modulation with a very short intermittent drop in energy occurring after about one third of the call’s duration. This drop in energy is also visible in the audiospectrogram, with frequencies slightly being shifted in the last two thirds of the call’s duration when compared to the first third and showing a somewhat different structure of harmonics. Overall, slight upward frequency modulation is evident within each call, with a short terminal drop. Numerical parameters of one complete call series and 16 analysed calls are as follows: call duration (= note duration) 231–262 ms (245 ± 8 ms); inter-call intervals within regular call series 623–808 ms (701 ± 59 ms); number of calls per call series 14; duration of regular call series 12,583 ms; dominant frequency 3178–3265 Hz (3219 ± 28 Hz); prevalent bandwidth 1500–8400 Hz; frequency bands in the first third of the call recognizable at approximately 2050, 3100, 4200, 5200 and 6140 Hz, in the last two thirds of the call at approximately 1650, 3250, 4800, 6400, and 8100 Hz.</p><p>Etymology.</p><p>The Malagasy adjective mavokely meaning ‘ pink’, in reference to the pinkish colouration of this species in life and in preservative, especially on the legs. ‘ Mavokely’ is also a linguistic quirk of the Malagasy language, as it is derived from the juxtaposition of the components mavo meaning ‘ yellow, and kely meaning ‘ small’. This name thus forms a double entendre, by referring also to the small stature and yellowish colouration of some individuals in life. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is currently definitively known only from (1) Marojejy National Park (type locality). However, photographs of specimens tentatively referrable to this species from (2) Anjanaharibe-Sud (14.72°S, 49.46°E, ca 1430 m a. s. l.) and (3) Simpona Lodge in Makira Natural Park (near 15.1992°S, 49.6208°E, ca 410 m a. s. l.) have been sent to us by Éric Mathieu and Lucas Pasteur, respectively (Fig. 27), suggesting that it may have a somewhat larger distribution in the rainforests of northeastern Madagascar. Elevation: ca 310–402 m a. s. l., but the record from Anjanaharibe-Sud would add ca 1000 m elevation to ca 1430 m a. s. l., if verified.</p><p>Natural history.</p><p>Rhombophryne mavokely is a semi-fossorial or detriticolous species found in low- to mid-elevation humid forest. All specimens were found in the leaf litter, and this species probably spends most of its life in the interstitial layer between the surface of the leaflitter and the soil. ZSM 517/2016 was collected less than a metre from a curled-up snake, Alluaudina bellyi, which is likely to prey upon this and other leaf-litter dwelling frog species (Hutter et al. 2018). We found Rhombophryne mavokely sp. nov. at consistently lower elevation than R. savaka, but it is not clear if there is a contact zone between the two species at around 400–450 m a. s. l., and records from higher elevations mentioned above remain to be verified.</p></div>	https://treatment.plazi.org/id/50752E50712652C78456F5659A1CA078	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
8D54AD27A18F58208E4AA2FC6D7DFF8C.text	8D54AD27A18F58208E4AA2FC6D7DFF8C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne minuta (Guibe 1975)	<div><p>Rhombophryne minuta species group</p><p>Contents.</p><p>Rhombophryne longicrus, R. minuta</p><p>Summary of morphology.</p><p>A group of small to medium-sized (SVL 16.9–28.0 mm) terrestrial, and likely scansorial, Rhombophryne species. Both species have a gracile habitus, long, slender limbs, and lack inguinal markings. Teeth are present on the maxilla, premaxilla, and postchoanal vomer. Clavicles are present. Skin texture is rather smooth.</p><p>Calls.</p><p>Calls are only known from R. minuta . It emits a high-pitched two-pulsed call at regular, long intervals.</p><p>Distribution.</p><p>This group is found only in the northeast (Marojejy, R. minuta) and north (Sorata, R. longicrus) of Madagascar (Fig. 7; Table S 4).</p><p>Ecology.</p><p>Species found in humid high-elevation tropical forest.</p></div>	https://treatment.plazi.org/id/8D54AD27A18F58208E4AA2FC6D7DFF8C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
E18B16D329F65435BDA525BDD79ECC82.text	E18B16D329F65435BDA525BDD79ECC82.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne minuta (Guibe 1975)	<div><p>Rhombophryne minuta (Guibé, 1975)</p><p>Figures 2, 4 –7, 50, 51, 61 –64</p><p>Mantipus minutus Guibé, 1975</p><p>Plethodontohyla minuta — Blommers-Schlösser and Blanc (1991)</p><p>Rhombophryne minuta — Glaw and Vences (2007)</p><p>Remark.</p><p>Several references to R. minuta actually refer to other species of Rhombophryne, including those from Manongarivo (Rakotomalala 2002) and Ambolokopatrika (Andreone et al. 2000).</p><p>The identity of this species was clarified by Glaw and Vences (2007) and Glaw et al. (2010 b), and it was discussed further by Scherz et al. (2015 a).</p><p>Holotype.</p><p>MNHN 1975.15, an adult female, collected in November 1972 on the ‘ Marojejy Massif, at altitude’, Sava Region, northeastern Madagascar, by C. P. Blanc (Figs 50, 51, 63) .</p><p>Paratypes.</p><p>Eight specimens: MNHN 1975.16 –23, all with the same collection data as the holotype (Guibé 1975).</p><p>Referred specimens.</p><p>ZSM 476/2005 (ZCMV 2003; Fig. 61 A, B), ZSM 477/2005 (ZCMV 2004; Fig. 61 C), two unsexed adults, and ZSM 359/2005 (FGZC 2897), ZSM 360/2005 (FGZC 2899), and UADBA -A- FGZC 2898, three unsexed presumed subadults, all collected on 17 February 2005 above ‘ Camp Simpona’ (14.4408°S, 49.7399°E, ca 1700–1800 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by F. Glaw, M. Vences, and R. D. Randrianiaina.</p><p>Assignment of specimens.</p><p>The specimen ZSM 360/2005 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341106; Vieites et al. 2009) and COI barcoding (COI KF 611591; Perl et al. 2014). Assignment of specimens ZSM 359/2005 and UADBA -A- FGZC 2898 is further confirmed by molecular taxonomic identification (Fig. 2), based on museomic sequences for 16 S 3 ’ produced from the holotype (see Museomics section, above). The assignment of these specimens to this name, as well as ZSM 476/2005, ZSM 477/2005, is also supported by morphological comparison with the holotype. All of the referred specimens have been collected at the type locality. UADBA -A- FGZC 2898 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>This species is sister to R. longicrus, the only other species of the R. minuta group, based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 16.9–21.8 mm; (2) head wider than long (HW / HL 1.36–1.43); (3) moderately small eyes (ED / HL 0.42–0.45); (4) relatively long hindlimbs (HIL / SVL 1.73–1.82); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.09); (6) outer metatarsal tubercle absent or indistinct; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence or absence of faint dark oblong markings in the inguinal region; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 10 analysed calls by one individual) containing two pulses, with (17) call duration 114–162 ms, (18) inter-call interval 4660–8050 ms, and (19) dominant frequency 4030–4230 Hz.</p><p>Rhombophryne minuta is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ G’ at site 355, ‘ T’ at site 566, and ‘ C’ at site 592. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. minuta can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by its generally smaller size (SVL 16.9–21.8 mm vs 21.5–50), smooth skin (vs rough), absence of chin barbels (vs general presence), and much longer hindlimbs (HIL / SVL 1.73–1.82 vs 1.10–1.41); from R. mangabensis and R. savaka by its much longer hindlimbs (HIL / SVL 1.73–1.82 vs 1.37–1.60), and pulsed (vs tonal) advertisement call; from R. mavokely sp. nov. and members of the R. ellae species group by the absence of orange colouration on the posterior thigh and in the inguinal region (vs presence); from all members of the R. laevipes species group by smaller adult body size SVL 16.9–21.8 mm vs 24.1–57.2 mm), and absence of white spots in the inguinal region and on the posterior thigh (vs presence in most species); from R. proportionalis by larger adult body size (SVL 16.9–21.8 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence).</p><p>Rhombophryne minuta is most similar to its sister species, R. longicrus . It can be distinguished from that species by its smaller size (SVL 16.9–21.8 mm vs 23.8–28.0 mm), slightly smaller hand (HAL / SVL 0.28–0.28 vs 0.29–0.30), generally shorter tibia (TIBL / SVL 0.46–0.49 vs 0.49–0.52), and lack of dark colouration on the supratympanic region (vs presence).</p><p>Redescription of holotype.</p><p>A specimen in a good state of preservation (Fig. 50). Throat and pectoral girdle skin reflected to reveal pectoral girdle. A small incision along the right flank made to assess sex. Ovigerous female with ova at various stages of development. Part of the liver was taken through this incision for genetic analysis in 2021.</p><p>Body gracile. Head wider than long (HW / HL 1.36). Eyes fairly large (ED / HL 0.43), pupils small, round. Snout rounded in dorsal view, obtuse in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, small, round, TDH / ED 0.36. Supratympanic fold distinct, raised, demarcated by a dark line, running from posterior corner of eye over tympanum and curving posteriorly to end above the insertion of the arm. Superciliary spines absent. A low sagittal fold runs from the tip of the snout to the middle of the body. Tongue relatively narrow, attached anteriorly, posteriorly free, rounded distally. Maxillary and vomerine teeth present, vomerine teeth forming two straight rows separated at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively long (FARL / SVL 0.50) and slender. Fingers without webbing, all long, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger distinctly shorter than fourth (Fig. 51). Finger tips not expanded, rounded. Finger subarticular tubercles indistinct, whitish; inner metacarpal tubercle small, slightly rounded, outer metacarpal / palmar tubercle indistinct. Hind limb gracile, tibiotarsal articulation reaching the eye when adpressed anteriorly along the body. TIBL / SVL 0.46. Inner metatarsal tubercle present, distinctly raised, oblong, light brown. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles indistinct, greyish, flat, rather oblong; toe tips simple and round (Fig. 51).</p><p>After approximately 50 years in preservative, the colouration of the specimen is brown above and light tan below. Marbled brown and cream patterns of the venter that were illustrated by Guibé (1975) are still faintly visible. The dorsum does not have any distinctive patterning to it, but there are hints of dark patches in the inguinal region, and a dark spot on the left flank. One dark brown crossband is present on the lower arm. No crossbands on the hindlimbs.</p><p>Variation.</p><p>SVL of adults 16.9–21.8 mm (n = 3), subadults 15.4–15.6 (n = 2). Insufficient sexed specimens exist to assess sexual size dimorphism. For variation in other measurements, see Table S 9. Available specimens are largely phenotypically homogeneous. Colouration in life is somewhat variable (Fig. 61). Unlike the holotype, there are no inguinal markings in the other material we have examined.</p><p>Call.</p><p>The advertisement call recorded on 17 February 2005 (21: 00 h) near Camp Simpona, Marojejy National Park (air temperature &lt;20 ° C), consists of a tonal note of moderate duration emitted at long and somewhat irregular intervals, not arranged in regular call series (Fig. 64). Each call (= note) exhibits distinct amplitude modulation, with two clearly separated pulses being evident, the second being slightly longer in duration and exhibiting higher call energy. Very slight overall upward frequency modulation is recognizable within each call, with a slight medium frequency drop between the two pulses. Numerical parameters of 10 analysed calls are as follows: call duration (= note duration) 114–162 ms (138 ± 17 ms); inter-call intervals 4660–8050 ms (6520 ± 1416 ms); dominant frequency 4030–4230 Hz (4143 ± 59 Hz); prevalent bandwidth 3500–6400 Hz; weak frequency band recognizable at approximately 6200 Hz. Calls from the same recording (Vences et al. 2006 CD 3, track 30) have been described by Glaw et al. (2010 b) using a slightly different terminological scheme.</p><p>Etymology.</p><p>Latin feminine nominative singular adjective meaning ‘ very small’, presumably in reference to the comparatively small size of this species relative to other relatives.</p><p>Distribution.</p><p>This species is only known from (1) high elevation on the Marojejy Massif (type locality). Elevation: 1700–1800 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne minuta is a scansorial species found in high-elevation humid forest. A scansorial species living in and on ericoid and lichenous vegetation near the tree line. Males call in the rainy season at night on low vegetation, up to 1 m above the ground (Glaw and Vences 2007).</p></div>	https://treatment.plazi.org/id/E18B16D329F65435BDA525BDD79ECC82	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
4D7F02B3447D5B948FA05E71CEF8D155.text	4D7F02B3447D5B948FA05E71CEF8D155.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne nilevina Lambert, Hutter & Scherz 2017	<div><p>Rhombophryne nilevina Lambert, Hutter &amp; Scherz, 2017</p><p>Figures 2, 4 –7, 30, 31, 39 –42</p><p>Remark.</p><p>Lambert et al. (2017) mistakenly gave the holotype’s collection number as KU 340893 in their tables 1 and 2 and figure 3. The correct number is KU 340897.</p><p>Informal names.</p><p>The candidate species ‘ Rhombophryne sp. 3 ’ of Vieites et al. (2009) and ‘ Rhombophryne sp. Ca 3 ’ of Perl et al. (2014) and Belluardo et al. (2022) from Tsaratànana is here included in R. nilevina due to its small uncorrected p distances (Tables S 2, S 3) and the results of our species delimitation analysis.</p><p>Carné and Vieites (2024) treated specimens from Marojejy and Anjanaharibe-Sud as ‘ R. sp. cf. nilevina ’ referring to them as ‘ DCL “ North ” ’, and treated R. sp. Ca 03 as an unconfirmed candidate species ‘ UCS “ Sp. 3 ” ’.</p><p>Holotype.</p><p>KU 340897 (CRH 798), an adult male, collected at mid-day on 8 February 2015 from ‘ montane rainforest near the former village of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=47.5054&amp;materialsCitation.latitude=-21.1287" title="Search Plazi for locations around (long 47.5054/lat -21.1287)">Andemaka in Ranomafana National Park’</a> (ca 21.1287°S, 47.5054°E, 1240 m a. s. l.), Vatovavy-Fitovinany Region, South Central East Madagascar, by S. M. Lambert, E. Rajeriarison, and R. J. Fulgence (Figs 30, 31, 40) .</p><p>Paratypes.</p><p>One specimen: UADBA -A- CRH 799, an adult male, collected on the morning of 7 February 2015 from the same locality as the holotype, by S. M. Lambert and R. J. Fulgence (Fig. 39 A – C).</p><p>Referred specimens.</p><p>ZSM 516/2016 (ZCMV 15215), an unsexed adult, collected on 18 November 2016 near ‘ Camp Simpona’ (ca 14.4366°S, 49.7434°E, 1325 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, A. Rakotoarison, M. Bletz, M. Vences, J. Razafindraibe, and A. Razafimanantsoa; MRSN A 4598 (FN 7252), an adult male, collected on 12 December 1997 near ‘ Camp 2 ’ Andranomadio (ca 14.7180°S, 49.5124°E, 820 m a. s. l.), Ambolokopatrika-Betaolana corridor, Sava Region, northeastern Madagascar, by F. Andreone.</p><p>According to our species delimitation, specimens formerly assigned to R. sp. Ca 03 are also included in R. nilevina: MRSN A 2631 (FAZC 11002), an adult female, ZSM 664/2001 (FGMV 2001.124), a subadult, ZSM 667/2001 (FGMV 2001.131), a &lt;10 mm juvenile (the original genetic reference specimen for the candidate species), MRSN A 3016 (FAZC 11135), an adult male (sexed by incision), and MRSN A 3249 (FAZC 11086), a juvenile, all collected between 4 and 9 February 2001 at Antsahamanara (ca 14.05°S, 48.79°E, ca 1000 m a. s. l.), Tsaratànana Massif, Diana Region, Madagascar, by F. Andreone, F. Mattioli, J. E. Randrianirina, and M. Vences.</p><p>Assignment of specimens.</p><p>The holotype is to be used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KY 288475; Lambert et al. 2017), but by inclusion of R. sp. Ca 03, so too is ZSM 667/2001 (16 S 3 ’ FJ 559296; Vieites et al. 2009). ZSM 667/2001 (MV 2001. F 5; KF 611592) and MVTIS 2001. E 50 (KF 611593) (both originally R. sp. Ca 03) are used as references for COI barcoding (Perl et al. 2014). The paratype and referred material conform well with the holotype in morphology, but see Variation section for comments on colour variation within the species. The paratype is topotypical. Assignment of specimens ZSM 516/2016, ZSM 667/2001, MRSN A 2631, MRSN A 4598, and MRSN A 3016 (as well as tissue sample MV 2001 - E 50) is confirmed by molecular taxonomic identification (Fig. 2). KU 340897, MRSN A 4598, MRSN A 2631, and MVTIS 2001. E 50 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. laevipes group, this species forms a subclade with R. botabota, R. sonaliae sp. nov. and R. anatiala sp. nov. (described below) based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) large body size, adult SVL 41.2–57.2 mm; (2) head wider than long (HW / HL 1.42–1.81); (3) moderately small eyes (ED / HL 0.31–0.34); (4) relatively long hindlimbs (HIL / SVL 1.52–1.83); (5) small inner metatarsal tubercle (IMTL / FOL 0.10–0.14); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence or absence of white inguinal spots on a blackish background; (13) presence or absence of white spots on the posterior thigh and ventral shank; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 7 analysed calls by one individual) pulsed, with (17) call duration 538–598 ms, (18) inter-call interval 42–126 s, and (19) dominant frequency 526–534 Hz.</p><p>Rhombophryne nilevina is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ G’ at site 389, ‘ T’ at site 401, and ‘ T’ at site 555; in 16 S 3 ’, ‘ T’ at site 956, ‘ T’ at site 1075, ‘ A’ at site 1131, and ‘ A’ at site 1157.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. nilevina can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by smooth skin (vs rough), absence of chin barbels (vs general presence), longer hindlimbs (HIL / SVL 1.52–1.83 vs 1.10–1.41); from R. mangabensis and R. savaka by much larger body size (SVL 41.2–57.2 mm vs 18.7–23.2 mm), longer relative forearm length (FARL / SVL 0.39–0.42 vs 0.33–0.38), well-ossified clavicles (vs poorly ossified), absence of black inguinal spots (vs presence in R. savaka); from R. mavokely sp. nov., R. ornata, and members of the R. ellae species group by the absence of reddish orange colouration on the posterior thigh and in the inguinal region (vs presence); from members of the R. minuta species group by usually shorter relative hindlimb length (HIL / SVL 1.52–1.81 vs 1.73–1.84), and less gracile habitus; from R. proportionalis by much larger adult body size (SVL 41.2–57.2 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence).</p><p>Within the R. laevipes species group, see the accounts of R. laevipes and R. botabota above for distinction from those species. Note that several traits thought to distinguish R. nilevina from R. laevipes and R. botabota (e. g., absence of stark colour boundary between lateral head and dorsum, and absence of white spots in the inguinal region; Lambert et al. 2017) are now known to vary within the species (see Variation, below) and cannot be used as diagnostic traits anymore. For distinction from newly described members of this species group, see their respective descriptions below.</p><p>Description of MRSN A 2631.</p><p>(a specimen of R. sp. Ca 03 now included in R. nilevina): An adult female specimen in a good state of preservation. The right half of the body cavity is opened with the organs exposed; a piece of liver tissue was taken for genetic analysis. A small number of well-developed eggs lies in the dorsal part of the body cavity, which is taken up mostly with the thick-walled stomach.</p><p>Body robust. Head wider than long (HW / HL 1.71). Eyes quite small (ED / HL 0.32), pupils large, round. Snout rounded in dorsal and lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum distinct, round, TDH / ED 0.75. Supratympanic fold distinct, slightly raised, running straight from posterior corner of eye over tympanum, curving gently ventrally posterior to the tympanum, demarcating the boundary of the dark brown lateral head colouration. Superciliary spines absent, but one white dot above each eye. Dorsal skin smooth in preservative, was slightly granular in life with a few larger granules (Fig. 41). Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth present, curved, with small lateral diastemata, separated at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.443), strongly built. Fingers without webbing, moderately long, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger distinctly shorter than fourth. Finger tips not expanded, rounded. Finger subarticular tubercles distinct, whitish, flat; inner metacarpal tubercle broad and flat, outer metacarpal / palmar tubercle indistinct, greyish. Hind limb strong, tibiotarsal articulation reaching between eye and nostril when adpressed anteriorly along the body. TIBL / SVL 0.537. Inner metatarsal tubercle present, oblong, whitish, somewhat bulging. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles distinct, cream, slightly raised; toe tips not enlarged, rounded.</p><p>After 22 years in preservative, the colouration of the specimen remains quite vibrant, though less strongly contrasting than in life (Fig. 41). The base colour is a coffee brown across the dorsum, and venter except that it takes on a purplish tone over the abdomen, and a richer brown on the chin. On the dorsum, a broad pattern of darker brown is present, across which is strewn almost black markings, especially in an interocular line, a small trianguloid marking on the head, and a V-shaped area on the back. The lateral surface of the head is almost black, strongly separated from the dorsal body colouration. The inguinal region has a large blackish splotch and a series of small white spots. The dorsal surfaces of the hindlimbs have a series of dark brown crossbands; the posterior surfaces of the thighs are dark chocolate brown with a few light brown or cream spots. The cloaca marks the top of a blackish trapezoid, and above it there is a small blackish bar. The ventral thighs are coffee brown with small cream flecks, the ventral shanks are dark chocolate brown with cream spots. The dorsal foot is spotted with cream, its ventral surface dark brown. The ventral abdomen is cream and a purplish brown. Each finger and toe has a cream annulus before the distalmost phalange. The arm has blackish disjointed crossbands dorsally, but is coffee brown below, including on the palmar surface. Colouration in life was like in preservative but with a grey, rather than a brown, base colour (Fig. 41).</p><p>Variation.</p><p>SVL ranges from 41.2–57.2 mm; for variation in other measurements, see Table S 6. The newly examined material from northeastern Madagascar generally agrees well with the type series in morphology, despite extending the range of the species by well over 800 km. Unlike the types, northern specimens have distinct white spots in the inguinal region and on the posterior thigh and ventral shank. Many specimens have at least a spot, generally a bar, above the cloaca (absent from MRSN 4598), and all have a distinct colour border between their lateral head and dorsal head. MRSN A 4598 has distinct vocal slits that confirm its sex as male.</p><p>Call.</p><p>The advertisement call recorded on 8 February 2015 (12: 00 h) near Andemaka, Ranomafana National Park (air temperature estimated to be around 20 ° C; presumably emitted by the holotype KU 340897), consists of a pulsed note of moderate to long duration, emitted at irregular and long intervals. Calls (= notes) exhibit distinct amplitude modulation, with call energy starting at low level, moderately increasing to about one fourth of the call’s duration, rapidly increasing after half of the call’s duration, reaching its maximum in its last quarter, followed by a decrease towards the call’s end. The second half of the call is characterized by a short intermittent drop in call energy. Pulses are very narrowly spaced in the second half of the call and very short in duration, with each pulse being apparently shorter than 2 ms, but almost impossible to measure exactly. In the first half of the call, pulses are longer in duration (ca. 9 ms) and repeated less rapidly. It is not possible to count the number of pulses per note (= call), but pulse repetition rate can roughly be calculated for each section of the note. Numerical parameters of 7 analysed calls are as follows: call duration (= note duration) 538–598 ms (561 ± 21 ms); inter-call intervals rather long and irregular, varying from 42–126 s (81 ± 41 s); pulse repetition rate within first half of the call 100 pulses / second, within second half of the call approximately 260–270 pulses / second; dominant frequency 526–534 Hz (530 ± 3 Hz); prevalent bandwidth 250–4500 Hz; numerous parallel frequency bands recognizable within the second half of the call, representing a reflection of pulse rate (see Köhler et al. 2017). Calls from the same recording have been described by Lambert et al. (2017) using a different terminological scheme.</p><p>Etymology.</p><p>The Malagasy adjective nilevina meaning ‘ buried’, in reference to the habit of this species of calling from burrows that must be excavated to catch the frogs. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>Records from northern Madagascar (Belluardo et al. 2022) dramatically increased the range of R. nilevina . Currently known from (1) Andemaka in Ranomafana National Park (type locality), (2) Ambolokopatrika, (3) Marojejy, and (4) Antsahamanara on the Tsaratànana Massif. Elevation: 1000–1325 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne nilevina is a semi-fossorial or detriticolous species found in low- to mid-elevation humid forest. Lambert et al. (2017) reported that R. nilevina called during the day from burrows within the soil, and its calling activity was heightened by inclement weather. The newly referred specimens from the north were collected active above the ground in largely undisturbed primary rainforest.</p><p>Karyotype.</p><p>The karyotype of a specimen assignable to this candidate species (FAZC 11063 from Tsaratànana) was described by Aprea et al. (2007) under the name ‘ Plethodontohyla laevipes ’. Its karyotype was 2 n = 26, with a subtelocentric fourth chromosome pair. Nucleolar organiser regions were located in a peritelomeric position on the long arm of the second chromosome pair. Centromeric bands were CMA 3 and DAPI negative.</p></div>	https://treatment.plazi.org/id/4D7F02B3447D5B948FA05E71CEF8D155	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
05E4174CB1B358E2B96E59A1C7049FEB.text	05E4174CB1B358E2B96E59A1C7049FEB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne ornata Scherz 2015	<div><p>Rhombophryne ornata Scherz et al., 2015</p><p>Figures 2, 4 –7, 67 –69</p><p>Holotype.</p><p>ZSM 1816/2010 (ZCMV 12384), an adult male, collected on 13 June 2010 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.9573&amp;materialsCitation.latitude=-14.1526" title="Search Plazi for locations around (long 48.9573/lat -14.1526)">‘Camp 2’ Matsabory Maiky</a> (14.1526°S, 48.9573°E, 2021 m a. s. l.), Tsaratànana Massif, Diana Regions, northern Madagascar, by M. Vences, D. R. Vieites, R. D. Randrianiaina, F. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison .</p><p>Paratypes.</p><p>Five specimens: ZSM 1815/2010 (ZCMV 12382), UADBA -A 60834 (ZCMV 12381), UADBA -A 60835 (ZCMV 12383), UADBA -A 60734 (ZCMV 12455), and ZSM 2859/2010 (DRV 06156), five juvenile or subadult specimens, with same collecting data as the holotype except UADBA -A 60734, which was collected 15–20 June 2010 .</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KP 895584; Scherz et al. 2015 b; 16 S 5 ’ KY 748102; Scherz et al. 2017 a) and COI barcoding (COI KF 611583; Perl et al. 2014). All paratypes are topotypical and conform well with the holotype in morphology. Assignment of ZSM 1815/2010 and ZSM 2859/2010 is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 1816/2010 is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is placed in a clade containing R. diadema, R. guentherpetersi, and R. tany based on molecular phylogenetic data, but the exact relationships within that clade are poorly resolved (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters (based on a set of four subadults and one adult): (1) medium body size, adult SVL up to 33 mm; (2) head wider than long (HW / HL 1.39–1.51); (3) moderately large eyes (ED / HL 0.40–0.45); (4) moderately long hindlimbs (HIL / SVL 1.46–1.64); (5) small inner metatarsal tubercle (IMTL / FOL 0.09–0.10); (6) outer metatarsal tubercle absent; (7) second finger subequal to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin somewhat rugose; (10) absence of chin barbels; (11) presence of two distinct superciliary spines; (12) presence of reddish colouration in inguinal region; (13) presence of reddish colouration on anterior and posterior surface of thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne ornata is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 315, ‘ G’ at site 345, ‘ T’ at site 682, and ‘ C’ at site 721. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. ornata can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. Within the R. serratopalpebrosa species group, it can be distinguished from all species by the presence of reddish orange colouration in the inguinal region and on the thigh (vs absent).</p><p>A detailed description of the species is given in Scherz et al. (2015 b).</p><p>Call.</p><p>The call of this species is not known.</p><p>Etymology.</p><p>Latin feminine nominative singular adjective meaning ‘ ornate’, in reference to the colourful appearance of this species, especially in contrast to other Rhombophryne .</p><p>Distribution.</p><p>This species is only known from (1) Matsabory Maiky on the Tsaratànana Massif (type locality). Elevation: 2021 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne ornata is a semi-fossorial or detriticolous species found in high-elevation humid forest. Specimens were collected hiding under logs and stones in forest; one specimen contained a frog in its stomach, as well as insect and plant remains (Scherz et al. 2015 b).</p></div>	https://treatment.plazi.org/id/05E4174CB1B358E2B96E59A1C7049FEB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
934722B69CF05981B8E5FAAF3621AC8F.text	934722B69CF05981B8E5FAAF3621AC8F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne proportionalis Scherz 2019	<div><p>Rhombophryne proportionalis Scherz et al., 2019</p><p>Figures 2, 4 –7, 50, 51, 58 –60</p><p>Remark.</p><p>This species is only known from the type series.</p><p>Informal names.</p><p>This species was listed as ‘ Stumpffia sp. Ca 34 ’ by Perl et al. (2014), Scherz et al. (2016 b, 2017 b), and Peloso et al. (2017).</p><p>Holotype.</p><p>ZSM 1826/2010 (ZCMV 12404), an adult male (seen calling, not recorded), collected on 16 June 2010 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.9782&amp;materialsCitation.latitude=-14.1182" title="Search Plazi for locations around (long 48.9782/lat -14.1182)">‘Camp 3’ Bepia</a> (14.1182°S, 48.9782°E, 2294 m a. s. l.), Tsaratànana Massif, Diana Region, northern Madagascar by M. Vences, D. R. Vieites, R. D. Randrianiaina, S. Rasamison, and E. Rajeriarison (Figs 50, 51, 58, 59) .</p><p>Paratypes.</p><p>Two specimens: ZSM 1840/2010 (ZCMV 12405), an adult male specimen with the same collection data as the holotype; and ZSM 636/2014 (DRV 6224), an adult presumed male, collected on 16 June 2010 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.9854&amp;materialsCitation.latitude=-14.0801" title="Search Plazi for locations around (long 48.9854/lat -14.0801)">‘Camp 4’ Andranomadio</a>, (14.0801°S, 48.9854°E, 2503 m a. s. l.), Tsaratànana Massif, Diana Region, northern Madagascar by M. Vences, D. R. Vieites, R. D. Randrianiaina, S. Rasamison, and E. Rajeriarison .</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 5 ’ KC 351480; Klages et al. 2013; 16 S 3 ’ KU 937808; Scherz et al. 2016 b) and COI barcoding (COI KF 611640; Perl et al. 2014). Both paratypes agree morphologically with the holotype and are topotypical. Assignment of specimen ZSM 1840/2010 is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 1826/2010 and ZSM 1840/2010 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>This species is the sole member of the R. proportionalis group and phylogenetically distinct from all other Rhombophryne based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) diminutive body size, adult SVL 11.0– 12.3 mm; (2) head wider than long (HW / HL 1.56–1.66); (3) comparatively large eyes (ED / HL 0.40–0.48); (4) relatively short hindlimbs (HIL / SVL 1.21–1.33); (5) small inner metatarsal tubercle (IMTL / FOL 0.07–0.14); (6) outer metatarsal tubercle variably present or absent; (7) second finger equal in length to fourth; (8) third toe distinctly longer than fifth, first toe highly reduced; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of black inguinal spots or lines; (13) absence of distinctive markings along the posterior thigh; (14) paedomorphic skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles absent; and (16) advertisement call (based on 63 analysed calls by one individual) tonal, emitted in rapid, regular series, with (17) call duration 27–60 ms, (18) inter-call interval within series 45–88 ms (9–17 calls per series), and (19) dominant frequency 5162–5738 Hz.</p><p>Rhombophryne proportionalis is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 496, ‘ C’ at site 511, and ‘ G’ at site 547; in 16 S 3 ’, ‘ T’ at site 966, ‘ C’ at site 967, ‘ A’ at site 1013 and ‘ T’ at site 1165.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. proportionalis can easily be distinguished from all other species by its much smaller body size (SVL 11.0– 12.3 mm vs&gt; 16 mm). It also exhibits a host of miniaturised skeletal features, described in detail by Scherz et al. (2019 b), that also set it apart from all congeners. Additionally, the advertisement call of R. proportionalis is unique for the genus in consisting of regular series of calls emitted in rapid succession (all other species with known calls emit single calls at much longer intervals).</p><p>Variation.</p><p>As reported by Scherz et al. (2019 b), adult SVL ranges from 11.0– 12.3 mm; for other morphometric measurements, see Table S 8. The type series is rather homogeneous, with a little variation in the shape of the head and supratympanic fold. Colouration is also rather consistent, though dorsolateral lines can be present or absent.</p><p>Call.</p><p>The advertisement call recorded on 15 June 2010 (11: 40 h) at Camp Bepia, Tsaratànana (air temperature unknown) from several individuals, consists of a high-pitched tonal note of short duration emitted in regular call series at regular and fast succession (Fig. 60). Each call exhibits slight amplitude modulation, with highest energy present at approximately the middle of the call’s duration, with few calls at the end of call series exhibiting two separated energy peaks. Slight upward frequency modulation is evident in each call. Numerical parameters of 63 analysed calls are as follows: call duration (= note duration) 27–60 ms (45 ± 8 ms); inter-call intervals within regular call series 45–88 ms (63 ± 9 ms); number of calls per call series 9–17 (13 ± 3); duration of regular call series 893–1765 ms (1323 ± 291 ms); dominant frequency 5162–5738 Hz (5461 ± 123 Hz); prevalent bandwidth 5000–6300 Hz. Calls from the same recording have been described by Scherz et al. (2019 b) using the note-centred terminological scheme.</p><p>Etymology.</p><p>Latin feminine nominative singular adjective meaning ‘ proportional’, in reference to the proportional dwarfism of the species.</p><p>Distribution.</p><p>This species is only known from (1) Bepia on the Tsaratànana Massif (type locality), and (2) the nearby locality Andranomadio. Elevation: 2294–2503 m a. s. l. A record from Marojejy at 1300 m a. s. l. (Rakotoarimalala and Raselimanana 2023) requires verification.</p><p>Natural history.</p><p>Rhombophryne proportionalis is a terrestrial or detriticolous species found in high-elevation humid forest. Males emit advertisement calls during the day. Nothing else is known of its natural history.</p><p>Osteology.</p><p>The osteology of Rhombophryne proportionalis (Fig. 59) was described in detail by Scherz et al. (2019 b). It is highly divergent from all other Rhombophryne species, as a result of its strongly miniaturised morphology.</p></div>	https://treatment.plazi.org/id/934722B69CF05981B8E5FAAF3621AC8F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
DC3E17C9065F5A6AA817B731D4899531.text	DC3E17C9065F5A6AA817B731D4899531.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne proportionalis Scherz, Hutter, Rakotoarison, Riemann, Rödel, Ndriantsoa, Glos, Roberts & Crottini 2019	<div><p>Rhombophryne proportionalis species group</p><p>Contents.</p><p>Rhombophryne proportionalis</p><p>Summary of morphology.</p><p>A highly divergent, proportionally miniaturised species (SVL 11.0– 12.3 mm), characterised by its small body size, paedomorphic skull with anteriorly displaced mandibular articulation and verticalized lateral elements, and the absence of ossified clavicles. Teeth are present on the maxilla, premaxilla, and postchoanal vomer; those on the vomer are diminutive. Skin is fairly smooth.</p><p>Calls.</p><p>A trill-like call, consisting of high-pitched tonal notes of short duration emitted in regular call series at regular and fast succession, unlike that of any other known Rhombophryne species.</p><p>Distribution.</p><p>This group is currently only known from Tsaratànana (Fig. 7; Table S 4).</p><p>Ecology.</p><p>Known only from humid tropical forest in northern Madagascar.</p></div>	https://treatment.plazi.org/id/DC3E17C9065F5A6AA817B731D4899531	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
C59A551A640658A8AAB8EE4C3F429A08.text	C59A551A640658A8AAB8EE4C3F429A08.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne quentini Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne quentini sp. nov.</p><p>Figures 2, 4 –7, 50, 51, 54, 55</p><p>Remark.</p><p>This species was figured as possibly representing Rhombophryne guentherpetersi by Glaw and Vences (2007).</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 3 Tsaratanana’ by Wollenberg et al. (2008), ‘ Rhombophryne sp. 7 ’ by Vieites et al. (2009), ‘ “ Rhombophryne ” sp. Ca 07 Tsaratanana’ by Scherz et al. (2016 b), ‘ Rhombophryne sp. Ca 07 ’ by Belluardo et al. (2022), ‘ Plethodontohyla sp. 2 ’ by Andreone et al. (2009), and ‘ Rhombophryne sp. 7 ’ and ‘ UCS “ Sp. 7 ” ’ by Carné and Vieites (2024).</p><p>Justification of the new species.</p><p>The new species is sister to R. kilonjy sp. nov., and these two species are together sister to R. ellae, as part of the R. ellae group that is phylogenetically distinct from other Rhombophryne (Belluardo et al. 2022) . From R. ellae, the new species differs by a high genetic divergence of 6.3 % uncorrected p distance in the 16 S 3 ’ fragment and thus distinctly higher than the 3 % level typically indicative of species-level differentiation in Malagasy frogs (Vieites et al. 2009). There is no haplotype sharing between R. quentini sp. nov. and R. ellae in the analysed fragment of the nuclear gene RAG 1 (Fig. 4). Furthermore, it also consistently differs from R. ellae by various morphological and osteological characters, especially its pronounced metatarsal tubercle (see Diagnosis below), and its status as distinct species is therefore beyond doubt. See also the Justification section for R. kilonjy sp. nov. below.</p><p>Holotype.</p><p>MRSN A 4616 (FAZC 11134), an adult female, collected on 9 February 2001 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.784&amp;materialsCitation.latitude=-14.045" title="Search Plazi for locations around (long 48.784/lat -14.045)">‘Camp 1’ Antsahamanara</a> (ca 14.045°S, 48.784°E, ca 1000 m a. s. l.), Manarikoba, Réserve Naturelle Intégrale de Tsaratànana, Diana Region, northern Madagascar, by F. Andreone, F. Mattioli, J. E. Randrianirina, and M. Vences (Figs 50, 51, 54 A, B) .</p><p>Paratypes.</p><p>12 specimens: MRSN A 4615 (FAZC 11129; Fig. 55), MRSN A 4617 (FAZC 11133), and MRSN A 3248 (FAZC 11161 = FGMV 2001. G 46 tissue sample), MRSN A 4638 (FAZC 11169), four unsexed adults, MRSN A 4628 (FAZC 11060), a juvenile, MRSN A 4604 (FAZC 11048), an unsexed individual of uncertain maturity, MRSN A 3656 (FAZC 11062; Fig. 54 C, D), an adult female, MRSN A 4629 (FAZC 11059; Fig. 54 E, F), a subadult female, MRSN A 4595 (FAZC 11151), an adult probable female (ambiguous genitalia), and MRSN A 4630 (FAZC 11146) and MRSN A 4631 (FAZC 11084), and UADBA -A 28193 (MV 2001.134), three adult males, all with same collection data as the holotype, but collected on 4–13 February 2001</p><p>Referred specimens.</p><p>BMNH 1987.2297 (field number ‘ 9 ’), an unsexed adult, collected on 12 February 1988 at ‘ Camp B’ in Manongarivo Special Reserve (14.067°S, 48.283°E, 350 m a. s. l.) by C. J. Raxworthy conforms morphologically with this species, but is not genetically verified. Due to its lower elevational distribution and geographical separation from other records, it is here tentatively included in the circumscription of R. quentini sp. nov.</p><p>Assignment of specimens.</p><p>The paratype MRSN A 3248 (FAZC 11161 = FGMV 2001. G 46) is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341108; Vieites et al. 2009). The holotype is used as the reference specimen for this species in COI barcoding (COI OL 790133; Belluardo et al. 2022). The sequence of the holotype is identical to the sequence of the reference specimen for 16 S in 16 S 3 ’ (OL 780577), and differs by one mutation in 16 S 5 ’ (99.8 % identical; OL 780602). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of specimens MRSN A 4615, MRSN A 3248, MRSN A 4604 and MRSN A 4617 is further confirmed by molecular taxonomic identification (Fig. 2). Unsequenced, morphologically identified specimens MRSN A 4638, MRSN A 4628, MRSN A 3656, MRSN A 4629, MRSN A 4595, MRSN A 4630, MRSN A 4631, UADBA -A 28193 were also collected at the type locality, and can be assigned to the species morphologically. MRSN A 4604, MRSN A 4615, and MRSN A 3248 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. ellae group, this species is sister to R. kilonjy sp. nov. (described below) based on molecular phylogenetic data (not shown). It is characterised by the possession of the following unique suite of characters: (1) small to medium body size, adult SVL 21.4–30.1 mm; (2) head wider than long (HW / HL 1.31–1.66); (3) moderately small eyes (ED / HL 0.32–0.43); (4) relatively long hindlimbs (HIL / SVL 1.44–1.80); (5) small inner metatarsal tubercle (IMTL / FOL 0.6–0.12); (6) outer metatarsal tubercle very distinct; (7) second finger subequal, shorter than, or seldom longer than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of large black inguinal ocelli bordered with white; (13) presence of reddish orange colouration on the hidden surfaces of the legs in life; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne quentini sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 264, ‘ C’ at site 301, and ‘ T’ at site 369 (but note that its sister species, R. ellae, has not been analysed for this fragment); in 16 S 3 ’, ‘ C’ at site 1138, ‘ T’ at site 1139, and ‘ T’ at site 1216.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. quentini sp. nov. can be distinguished from all other Rhombophryne except R. ellae by the combination of distinctive white-outlined black inguinal ocelli with reddish orange colouration on the hidden surfaces of the legs. From R. ellae and almost all other Rhombophryne species, it can be distinguished by the very pronounced outer metatarsal tubercle in adults (vs weakly pronounced). It can further be distinguished from R. ellae by smaller relative tympanum size (TDH / ED 0.43–0.61 vs 0.62). The skeleton of R. quentini sp. nov. also seems to differ from that of R. ellae in the following traits: posterior edge of nasal with a bend in it (vs continuous in R. ellae), less acute angle between zygomatic and otic rami of squamosal as seen in dorsal view, anterior ramus of pterygoid lying abreast of the dorsal edge of the maxilla (vs running along the lingual edge), coracoids medially only weakly flared (vs strongly), shorter transverse processes on presacral vertebrae 5–8, and oblique groove and dorsal prominence weak (vs distinct). For osteological distinction from R. kilonjy sp. nov., see the description of that species, below.</p><p>Description of holotype.</p><p>Specimen in a moderately good state of preservation, though slightly soft (Figs 50). An incision made along the right flank to assess the sex, and part of the right foot taken as a tissue sample. Small ova, ovaries, and fallopian tubes clearly visible.</p><p>Body robust and rotund. Head wider than long (HW / HL 1.43). Eyes fairly small (ED / HL 0.33), pupils small, round. Snout slightly pointed in dorsal and slightly truncate in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril approximately equidistant between tip of snout and eye, slightly protuberant. Tympanum distinct, round, TDH / ED 0.52. Supratympanic fold indistinct, slightly raised, partly melanised, running straight from posterior corner of eye over tympanum, curving gently ventrally posterior to the tympanum to anterior to the forearm’s insertion. Lateral head similar in colour to dorsal head. Superciliary spines absent. Dorsal skin smooth in preservative. Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth with a strong diastema, the medial elements distinct, separated from one another at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.382), moderately slender. Fingers without webbing, relative lengths 1 &lt;4 &lt;2 &lt;3, second finger distinctly longer than fourth (Fig. 51). Finger tips not expanded, rounded. Finger subarticular tubercles distinct, greyish, flat; inner metacarpal tubercle slender and rounded, outer metacarpal / palmar tubercle distinct, greyish, paired. Hind limb strong, tibiotarsal articulation reaching eye when adpressed anteriorly along the body. TIBL / SVL 0.443. Inner metatarsal tubercle present, oblong, whitish, distinctly raised. Outer metatarsal tubercle highly distinct, rounded and raised. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles highly distinct, elongated, grey; toe tips distinctly enlarged, especially of fourth toe, rounded to subacuminate (Fig. 51).</p><p>After 22 years in preservative, the base colour is a dark greyed brown across the dorsum and flanks, including the dorsal limbs. The dorsum is marked with darker brown chevrons, specifically one over the suprascapular region and a second over the iliac region. Between the chevrons are additional patches of this darker brown. There are distinct, oblong inguinal ocelli that are a rich dark brown in colour with a very thin whitish edge. No distinct colour difference between the lateral and dorsal head. The supratympanic fold is dark brown from the eye to the tympanum, and then the colour stops. The dorsal hindlimbs bear indistinct crossbands that line up with the leg folded, in the same colours as the dorsum and chevrons. The ventral foot surface is almost black but for the tubercles (metatarsal and subarticular), which are light grey. The anterior and posterior thigh is pale cream in preservative (this was bright orange in life; Fig. 54 A, B), as is the ventral thigh and shank. A distinct, almost black trapezoid is present around the cloaca. The dorsal foot is mottled cream and coffee brown. A light annulus is present before the terminal phalange of each finger and toe. The arms have a single crossband on the antebrachium. The dorsal hand is lighter medially than laterally; the ventral hand is light brown. The chin is mottled light brown with cream flecks, fading to completely cream over the sternal region into the immaculate abdomen.</p><p>Variation.</p><p>SVL of adults 22.1–30.1 mm (n = 10); among sexed adults, males 22.1–25.8 mm (n = 3), females 24.9–26.5 mm (n = 3); thus, very little sexual size dimorphism. For variation in other measurements, see Table S 7. This species is based on an exceptionally large series of specimens, which are however mostly phenotypically homogeneous. The pronounced outer metatarsal tubercle is always present, though it is less obvious in the smallest specimens. All specimens have pale posterior and anterior thighs and ventral shanks in preservative, where they were orange in life; distinct blackish inguinal ocelli with narrow whitish outline in most individuals; and a dark brown trapezoid around the cloaca. Most specimens have a series of dark chevrons on the dorsum, especially one in the suprascapular region. Subadult female MRSN A 4629 has a highly distinct colouration, with a broad vertebral stripe from the head to the cloaca (Fig. 54 E, F) not seen in any other individuals, but otherwise matches the other specimens well. The chins of almost all specimens are lighter and more flecked with cream than the holotype. The adult male UADBA -A 28193 has a rich coffee brown chin. This specimen also has more patterning on its ventral shanks than most others. In most individuals, the second finger is shorter than the fourth, unlike the holotype; this is not a diagnostic character.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>MDS wishes to dedicate this new frog species to his son, Quentin Artemis Lattenkamp, in the hope that it will continue to encourage his love for animals.</p><p>Distribution.</p><p>This species is only known from (1) Manarikoba on the western side of the Tsaratànana Strict Nature Reserve (type locality). A putative record from the southern side of Manongarivo Special Reserve (BMNH 1987.2297) at 350 m a. s. l. should be validated with further material from this area. Elevation: ca 1000 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne quentini sp. nov. is a semi-fossorial or detriticolous species found in mid- and possibly also low-elevation humid forest. The gut of MRSN A 3656 contained pieces of a blattodean that must have been relatively large in life, a diminutive ant, a flatbug (family Aradidae), and several unidentifiable arthropod pieces. According to Andreone et al. (2009), Rhombophryne quentini sp. nov. was one of the most frequently encountered species in pitfall traps in Manarikoba; they captured 22 individuals in pitfall traps.</p><p>Karyotype.</p><p>The karyotype of MRSN A 3656 was described by Aprea et al. (2007) under the name ‘ Plethodontohyla sp. aff. minuta ’, with the incorrect locality ‘ Masoala’. It was described to have 2 n = 26, with a telocentric eighth chromosome pair. Nucleolar organiser regions were located on the long arms of the sixth chromosome pair. Centromeric bands were CMA 3 and DAPI negative.</p></div>	https://treatment.plazi.org/id/C59A551A640658A8AAB8EE4C3F429A08	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
BE33DAEFCBE45CAC8BE6E51675D7B9A6.text	BE33DAEFCBE45CAC8BE6E51675D7B9A6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne regalis Scherz 2017	<div><p>Rhombophryne regalis Scherz et al., 2017</p><p>Figures 2, 5 –7, 67 –69</p><p>Holotype.</p><p>MRSN A 4602 (FN 7292), an adult male, collected on 13 December 1997 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.43&amp;materialsCitation.latitude=-14.5433" title="Search Plazi for locations around (long 49.43/lat -14.5433)">‘Camp 3’ Antsinjorano</a> (14.5433°S, 49.4300°E, ca 980 m a. s. l.), Ambolokopatrika-Betaolana Forest, Sava Region, northeastern Madagascar, by F. Andreone, J. E. Randrianirina, and G. Aprea .</p><p>Paratypes.</p><p>Five specimens: MRSN A 4603 (FN 7146), an adult female, same collection data as the holotype; MRSN A 4619, a subadult, and MRSN A 4620, an adult female, collected on 6 December 1997 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.4383&amp;materialsCitation.latitude=-14.54" title="Search Plazi for locations around (long 49.4383/lat -14.54)">‘Camp 2’ Andranomadio</a> (14.5400°S, 49.4383°E, ca 860 m a. s. l.), Ambolokopatrika-Betaolana Forest, Sava Region, northeastern Madagascar, by F. Andreone, J. E. Randrianirina, and G. Aprea ; MRSN A 4618, an adult female, collected on 26 June 1996 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.5958&amp;materialsCitation.latitude=-14.8283" title="Search Plazi for locations around (long 49.5958/lat -14.8283)">Ambinanin’antsahamaloto</a> (14.8283°S, 49.5958°E, ca 800 m a. s. l.), Besariaka Forest, Sava Region, northeastern Madagascar, by F. Andreone and J. E. Randrianirina ; MRSN A 6058, an adult female, collected in January 1996 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.4634&amp;materialsCitation.latitude=-14.7783" title="Search Plazi for locations around (long 49.4634/lat -14.7783)">‘Camp W 1’ west slope of Anjanaharibe-Sud</a> (Analabe; 14.7783°S, 49.4634°E, ca 1050 m a. s. l.), Sava Region, northeastern Madagascar, by F. Andreone, J. E. Randrianirina, and H. Randriamahazo .</p><p>Referred specimens.</p><p>UADBA -A 14174 (APR 00650), UADBA -A 14205 (APR 00477), UADBA -A 14172 (APR 00476), an adult female, an unsexed adult, and a juvenile, respectively, collected between 23 and 29 October 2001 in Marojejy National Park (14.4183°S, 49.6014°E, 810 m a. s. l.), Sava Region, northeastern Madagascar, by A. P. Raselimanana.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (under the incorrect name R. serratopalpebrosa) (16 S 3 ’ EU 341111; Vieites et al. 2009). It has not been sequenced for COI. All paratypes conform well with the holotype in morphology. RAG 1 sequence of this species is so far not available.</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, the precise phylogenetic relationships of this species are poorly resolved, but it is possibly closely related to R. serratopalpebrosa and R. vaventy based on the existing but fragmentary DNA sequence data (Fig. 2). It is characterised by the possession of the following unique suite of characters: (1) small to medium body size, adult SVL 20.2–26.5 mm; (2) head wider than long (HW / HL 1.21–1.46); (3) moderately large eyes (ED / HL 0.35–0.40); (4) moderately long hindlimbs (HIL / SVL 1.64–1.84); (5) small inner metatarsal tubercle (IMTL / FOL 0.06–0.12); (6) outer metatarsal tubercle absent; (7) second finger slightly to distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin shagreened; (10) absence of chin barbels; (11) presence of three or four pronounced superciliary spines, the longest being anterior to the eye; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne regalis is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ G’ at site 302, ‘ A’ at site 334, ‘ G’ at site 598, ‘ T’ at site 601, ‘ C’ at site 603, ‘ A’ at site 607, and ‘ C’ at site 615 (noteworthy that the sequence was retrieved from museomics and some sites may result from low quality sequence data); in 16 S 3 ’, ‘ A’ at site 964, ‘ T’ at site 982, ‘ G’ at site 1199.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. regalis can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. Within the R. serratopalpebrosa species group, it can be distinguished from all species except R. serratopalpebrosa by the presence of an S-shaped dermal postnasal fold. For distinction from R. serratopalpebrosa, see the diagnosis of that species, above. For distinction from new species described herein, see accounts below.</p><p>Variation.</p><p>The newly examined material from Marojejy (referred specimens, above) are clearly assignable to this species based on the presence of S-shaped postnasal fold, most enlarged first superciliary spine, dorsal skin texture. Measurements are given in Table S 10.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>Latin feminine nominative singular adjective meaning ‘ regal’, in reference to the superciliary spines.</p><p>Distribution.</p><p>Rhombophryne regalis was originally described from (1) Anjanaharibe-Sud, (2) Ambolokopatrika-Betaolana forest (type locality), and (3) Besariaka (Scherz et al. 2017 a). The newly referred material expands the range of the species to (4) Marojejy, supporting the findings of Rakotoarimalala and Raselimanana (2023), who reported it from ca 1300 m a. s. l. Elevation: ca 800–1300 m a. s. l. (the upper elevation being rather uncertain).</p><p>Natural history.</p><p>Rhombophryne regalis is a semi-fossorial or detriticolous species found in mid-elevation humid forest. Specimens were found active on the ground during rainy weather at night (Scherz et al. 2017 a).</p></div>	https://treatment.plazi.org/id/BE33DAEFCBE45CAC8BE6E51675D7B9A6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
4525CE3251BF5EA1B514FAAAB58C1FC6.text	4525CE3251BF5EA1B514FAAAB58C1FC6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne savaka Scherz 2016	<div><p>Rhombophryne savaka Scherz et al., 2016</p><p>Figures 2, 4 –9, 21 –23</p><p>Informal names.</p><p>This species was listed as ‘ Rhombophryne sp. 4 ’ by Vieites et al. (2009) and Glaw et al. (2010 b), and ‘ Rhombophryne sp. Ca 4 ’ by Perl et al. (2014).</p><p>Holotype.</p><p>ZSM 468/2005 (ZCMV 2065), an adult male, collected on 18 February 2005 at ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.7605&amp;materialsCitation.latitude=-14.435" title="Search Plazi for locations around (long 49.7605/lat -14.435)">Camp Marojejia’</a> (14.4350°S, 49.7605°E, 746 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by F. Glaw, M. Vences, and R. D. Randrianiaina (Figs 8, 9, 22) .</p><p>Paratype.</p><p>One specimen: UADBA -A- ZCMV 2079, a specimen of unknown age and sex with the same collection data as the holotype.</p><p>Referred specimens.</p><p>ZSM 519/2016 (MSZC 0319; Fig. 21 D – F) and KU 347465 (CRH 1653; Fig. 21 A – C), two adult males (call vouchers), collected at night on 2 December 2016 near ‘ Camp Mantella’ (ca 14.438°S, 49.776°E, 450 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, J. H. Razafindraibe, C. R. Hutter, and A. Razafimanantsoa; ZSM 520/2016 (MSZC 0318; Fig. 21 G – I), an adult female, collected at night on 1 December 2016 near ‘ Camp Marojejia’ (14.4340°S, 49.7610°E, ca. 750 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, J. H. Razafindraibe, C. R. Hutter, and A. Razafimanantsoa.</p><p>Assignment of specimens.</p><p>The holotype is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KU 724176; Vieites et al. 2009) and COI barcoding (COI KF 611594; Perl et al. 2014). The paratype and referred material conform well with the holotype in morphology, and were collected at or near the type locality. Assignment of specimens ZSM 519/2016, ZSM 520/2016, and UADBA -A- ZCMV 2079 is further confirmed by molecular taxonomic identification (Fig. 1). KU 347465 has been sequenced with the FrogCap hybrid capture probe set; it is included in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. testudo group, this species is sister to R. mangabensis based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) small body size, adult SVL 18.8–20.9 mm; (2) head wider than long (HW / HL 1.66–1.80); (3) moderately small eyes (ED / HL 0.35–0.43); (4) short hindlimbs (HIL / SVL 1.37–1.55); (5) small inner metatarsal tubercle (IMTL / FOL 0.11–0.14); (6) moderately distinct outer metatarsal tubercle; (7) second finger roughly equal in length to fourth, except in one individual where it is much shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular with a few scattered warts, especially a suprascapular pair outlined in black; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) occasional presence of small black inguinal ocelli without light outline; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 28 analysed calls by one individual) tonal, with (17) call duration 121–142 ms, (18) inter-call interval 221–395 ms, and (19) dominant frequency 3714–3861 Hz.</p><p>Rhombophryne savaka is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 355, ‘ G’ at site 664 and ‘ T’ at site 687; in 16 S 3 ’, ‘ A’ at site 937, ‘ T’ at site 1094, and ‘ T’ at site 1155.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. savaka can easily be distinguished from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence); from members of the R. minuta species group by less gracile body and much shorter limbs; from R. proportionalis by its much larger body size (SVL 18.8–20.9 mm vs 11.0– 12.3 mm); from members of the R. ellae species group by the absence of reddish orange colouration on the hindlimbs (vs presence); from members of the R. laevipes species group by the absence of white outline on inguinal spots or whitish inguinal spots (vs presence in most species), and smaller adult body size (SVL 18.8–20.9 mm vs 24.1–57.2 mm). Within the R. testudo species group, see accounts of R. testudo, R. coudreaui, R. matavy, and its sister species R. mangabensis above for distinction from those species. For distinction from newly described species, see accounts of those species, below.</p><p>Variation.</p><p>SVL 18.7–20.9 mm (n = 3). For variation in other measurements, see Table S 5.</p><p>A detailed description of the species is given in Scherz et al. (2016 a), but it was based on just a single individual. Additional specimens collected in 2016 are very similar to the holotype (Fig. 21). The largest specimen is an ovigerous female, ZSM 520/2016 (Fig. 21 G – I), but this specimen is only 0.5 mm larger than the male holotype. The skin was consistently rather smooth in all individuals. ZSM 519/2016 shows a large diastema in its vomerine teeth, consistent with the holotype. All specimens are consistent in overall colouration, but differ in details of patterning. ZSM 520/2010 has an extensive near-black bar on its lower posterior thigh, where the other specimens have only a small trapezoid. This specimen also has blackish spots above its suprascapular region and faint crossbands on its hindlimbs, which are lacking form the other specimens. Inguinal spots were present only in the holotype; ZSM 520/2016 has a diminutive inguinal spot on the right side but not the left. The posterior thigh is solid cream in colour. The abdomen is also immaculate cream in all specimens, and very translucent in life (Fig. 21 C, F, I). At least one specimen (ZSM 520/2016) has dark markings above the suprascapular region, but these are not as distinctive or dark as in R. mangabensis or in R. mavokely sp. nov. described below.</p><p>Call.</p><p>The advertisement call recorded on 2 December 2016 at Camp 1, Marojejy National Park (call voucher ZSM 519/2016 [MSZC 0319]; air temperature 22.6 ° C) during rainfall, consists of a tonal note of short duration emitted in short regular call series (Fig. 23). Inter-call intervals within call series tend to be slightly longer in duration towards the end of a series. Each call exhibits some slight amplitude modulation, with call energy moderately fast increasing towards the middle of the call, then keeping the same level of energy until the last quarter of the call’s duration when it quickly drops towards the call’s end. Slight upward frequency modulation is evident in each call until two thirds of the call’s duration, followed by a slight terminal drop in frequency. Numerical parameters of 28 analysed calls are as follows: call duration (= note duration) 121–142 ms (132 ± 5 ms); inter-call intervals within regular call series 221–395 ms (296 ± 45 ms); number of calls per call series 4–5 (4.3 ± 0.5); duration of regular call series 1342–1883 ms (1483 ± 187 ms); dominant frequency 3714–3861 Hz (3807 ± 47 Hz); prevalent bandwidth 1600–7800 Hz; frequency bands recognizable at approximately 1900, 5700, and 7680 Hz.</p><p>Additional calls recorded on 2 December 2016 at the same locality (voucher specimen KU 347465 [CRH 1653]) completely agree in general characteristics with the call described above, except for a slightly higher dominant frequency explainable by smaller body size of the calling male, and have the following numerical parameters (3 call series and 13 calls analysed): call duration (= note duration) 110–122 ms (115 ± 3 ms); inter-call intervals within regular call series 302–451 ms (340 ± 46 ms); number of calls per call series 4–5 (4.3 ± 0.6); duration of regular call series 1461–1932 ms (1631 ± 261 ms); dominant frequency 4144–4210 Hz (4175 ± 27 Hz); prevalent bandwidth 1800–8500 Hz.</p><p>Etymology.</p><p>The Malagasy noun savaka meaning ‘ diastema’, in reference to the diastema in the vomerine teeth that were thought to be diagnostic of this species. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>Rhombophryne savaka is known only from (1) Marojejy National Park (type locality). Elevation: 450–750 m a. s. l. (based on newly collected material). Records from 1300 ± 200 m a. s. l. by Rakotoarimalala and Raselimanana (2023) cannot be verified as we have not examined material and have no DNA sequences, so they are here not included.</p><p>Natural history.</p><p>Rhombophryne savaka is a semi-fossorial or detriticolous species found in low- to mid-elevation humid forest. Calling activity was triggered by rain in November and December 2016, especially at night. Locally the species seemed rather common, though it was very difficult to localise calling individuals. Males emit advertisement calls from within or on the leaf litter; ZSM 519/2016 was observed calling with a leaf resting on its back. Females likewise live within the leaf-litter; ZSM 520/2016 was collected whilst attempting to find a calling male from within the same quadrat of leaf litter (the male could not be found).</p></div>	https://treatment.plazi.org/id/4525CE3251BF5EA1B514FAAAB58C1FC6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
3580AF705DF75E288069D8056FDC694E.text	3580AF705DF75E288069D8056FDC694E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne serratopalpebrosa (Guibe 1975)	<div><p>Rhombophryne serratopalpebrosa species group</p><p>Contents.</p><p>Rhombophryne coronata, R. diadema, R. guentherpetersi, R. ornata, R. regalis, R. serratopalpebrosa, R. tany, R. sp. Ca 13, R. sp. ‘ Ivohibe’, R. vaventy .</p><p>Summary of morphology.</p><p>A group of small to large-sized (SVL 19.4–58.6 mm), mostly terrestrial, possibly semi-detriticolous or even semi-fossorial frogs. They have a gracile to rotund habitus. The group is characterised by the possession of superciliary spines (otherwise only present on a few members of the R. testudo species group, which however differ strongly in other aspects of morphology, making confusion impossible). Most lack inguinal markings. Teeth are present on the maxilla, premaxilla, and postchoanal vomer. Clavicles are present. Skin texture varies from smooth to rugose.</p><p>Calls.</p><p>Calls are only known from R. coronata . It makes a high-pitched short tonal note emitted in regular call series of variable duration. Most calls are composed of two recognizable pulses. Its frequency modulation — containing upward and downward frequency sweeps within each note — is unique among all known Rhombophryne calls.</p><p>Distribution.</p><p>This group is widely distributed, with members in the north ( R. guentherpetersi, R. ornata, R. tany, R. diadema), northeast ( R. serratopalpebrosa, R. vaventy, R. regalis), and east ( R. coronata, R. sp. ‘ Ivohibe’, R. sp. Ca 13) (Fig. 7; Table S 4) of Madagascar. See Scherz et al. (2017 a) for detailed discussion of the biogeography of this clade.</p><p>Ecology.</p><p>Species found in humid and subhumid tropical forest.</p></div>	https://treatment.plazi.org/id/3580AF705DF75E288069D8056FDC694E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
C89A974B363A5969ABB2A18044667E86.text	C89A974B363A5969ABB2A18044667E86.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne serratopalpebrosa (Guibe 1975)	<div><p>Rhombophryne serratopalpebrosa (Guibé, 1975)</p><p>Figures 2, 4 –7, 67 –70</p><p>Mantipus serratopalpebrosus Guibé, 1975</p><p>Plethodontohyla serratopalpebrosa — Blommers-Schlösser and Blanc (1991)</p><p>Rhombophryne serratopalpebrosa — Glaw and Vences (2007)</p><p>Remarks.</p><p>All references to Rhombophryne serratopalpebrosa between 2005 and 2017 that did not refer specifically to the holotype of the species, were in fact referring to R. regalis (Scherz et al. 2017 a) . Previous references to R. serratopalpebrosa referred to any of a number of different members of the R. serratopalpebrosa species group; all of these references should therefore be carefully checked against the distribution and morphology of the other species mentioned here.</p><p>Until now, this species has only been known from the holotype, which is stained green and is in generally poor condition; it was redescribed by Scherz et al. (2014). A specimen collected by APR at high elevation on the Marojejy Massif is the first individual discovered that morphologically matches the holotype. We obtained small fragments of the 5 ’ fragment of 16 S rRNA through museomics methods, which confirm identity with the holotype (Fig. 2). We therefore provide a detailed description of this specimen here.</p><p>Holotype.</p><p>MNHN 1975.24, an adult female, collected on 29 November 1972 in the ‘ thickets at the summit of Marojejy’, Sava Region, northeastern Madagascar, by C. P. Blanc (Guibé 1975: 1088) .</p><p>Paratypes.</p><p>None.</p><p>Referred specimen.</p><p>UADBA 18258 (APR 04222), an adult female, collected at 11: 00 am on 27 October 1996 at high elevation on the Marojejy Massif (14.4339°S, 49.7347°E, 1550 m a. s. l.), Sava Region, northeastern Madagascar, by A. P. Raselimanana. As stated above, the assignment of this specimen to this species is confirmed by molecular taxonomic identification (Fig. 2), in addition to be topotypical and conforming well morphologically with the holotype. RAG 1 sequence of this species is so far not available.</p><p>Definition.</p><p>The precise phylogenetic relationships of this species are poorly resolved, but it is possibly closely related to R. regalis and R. vaventy based on the existing but fragmentary DNA sequence data (Fig. 2). It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 28.5–30.8 mm; (2) head wider than long (HW / HL 1.41–1.51); (3) moderately large eyes (ED / HL 0.38–0.38); (4) relatively long hindlimbs (HIL / SVL 1.87–1.88); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.09); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin granular; (10) absence of chin barbels; (11) presence of four pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. serratopalpebrosa can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. Within the R. serratopalpebrosa species group, it can be distinguished from all species except R. regalis by the presence of an S-shaped dermal postnasal fold. It can be distinguished from R. regalis by its substantially larger size (SVL 28.5–30.8 mm vs 20.2–26.5 mm), strong supratympanic fold (vs weak), four superciliary spines (vs three), typically larger relative tympanum size (TDH / ED 0.72–0.75 vs 0.47–0.75), longer relative forearm length (FARL / SVL 0.55–0.60 vs 0.42–0.51), parasphenoid cultriform process not broadening anteriorly (vs broadening), absence of contact between exoccipitals ventromedially (vs presence), and narrow (vs broad) contact between squamosal and quadratojugal. Based on the newly studied specimen, we can rule out the ossification of the pubis and stepping of the anterior edge of the ventral ramus of the squamosal as diagnostic traits to distinguish these two species, suggested by Scherz et al. (2017 a). For distinction from new species described herein, see accounts below.</p><p>Description of UADBA 18258.</p><p>An adult female in an excellent state of preservation (Fig. 69 A). Slightly soft on the abdomen. Tissue sample taken from the right thigh, and a slit made in the right flank to assess the sex. Large yellowish ova present.</p><p>Overall extremely closely resembling MNHN 1975.24 as it looked before it was stained green. Body rhomboid. Head very wide, much wider than long (HW / HL 1.41). Eyes large and protuberant (ED / HL 0.37), pupils small, round. Snout rounded in dorsal and lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique, a distinct S-shaped postnasal fold. Nostril nearer to tip of snout than to eye, slightly protuberant. Tympanum distinct, large, oval, TDH / ED 0.75. Supratympanic fold distinct, raised, not melanised, running in a slightly S-shaped curve, straight over the tympanum, curving ventrally behind it, and then posteriorly again toward the dorsal insertion of the forelimb. Lateral head similar in colour to dorsal head. Five superciliary spines of varying hight and distinction present above each eye. Dorsal skin slightly shagreened, distinctly different in texture from flank and ventral skin. Ventral skin smooth. Tongue broad, attached anteriorly, posteriorly free, with a distinct raised, perpendicular ridge-like structure on its anterodorsal surface. Maxillary and vomerine teeth present, vomerine teeth straight, not varying in height, separated from one another at the midline by a small gap. Choanae round, placed laterally in the roof of the mouth.</p><p>Forelimbs long (FARL / SVL 0.552), slender. Fingers without webbing, long, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger much shorter than fourth. Finger tips not expanded, elongated. Finger subarticular tubercles indistinct, cream, flat; inner metacarpal tubercle small and rounded, outer metacarpal / palmar tubercle indistinct, cream, single. Hind limb slender and long, tibiotarsal articulation reaching snout when adpressed anteriorly along the body. TIBL / SVL 0.493. Inner metatarsal tubercle present, oblong, cream, weakly raised. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, long, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles indistinct; toe tips elongated, not expanded.</p><p>After 29 years in preservative, the base colour is coffee brown, mottled across the whole body with a mixture of lighter and darker brown. The limbs are slightly lighter than the trunk in dorsal colouration. Flanks with small cream spots, but otherwise largely lacking noteworthy patterning: there are no inguinal spots or other markings; the hindlimbs are largely homogeneous, being more pigmented above than below, and forelimbs the same. No distinct colour difference between the lateral and dorsal head. The upper lip has a thin white line running from below the eye to the nostril, and an oblique light marking from the posterior corner of the eye to the rictus anterior to the tympanum. There is a light patch lacking pigment on the left inguinal area, but this may be due to how tags were tied before they were attached at the right knee. Ventrally, the chin is light brown, and this colour fades gently to grey-brown over the abdomen. The ventral legs have a few small cream spots. The ventral hands and feet are cream-brown. There is no distinctive colouration around the cloaca.</p><p>Variation.</p><p>SVL 28.5–30.8 mm (n = 2). For variation in other measurements, see Table S 10. The newly examined specimen UADBA 18258 agrees in almost all particulars with the holotype.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>Latin feminine nominative singular compound adjective, derived from serratus meaning ‘ serrated’, and palpebra meaning ‘ eyelid’, and the ending - osa, meaning ‘ full of’, in reference to the superciliary spines of this species.</p><p>Distribution.</p><p>This species is only known from (1) high elevation on the Marojejy Massif (type locality). Elevation: ca 1550 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne serratopalpebrosa is a presumably scansorial species found in high-elevation humid forest, but its ecology is largely unknown. The newly collected specimen UADBA 18258 was collected jumping on the forest floor during a morning rain.</p></div>	https://treatment.plazi.org/id/C89A974B363A5969ABB2A18044667E86	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
FA1B8FD870D259729BBC9D4C3BEB37DD.text	FA1B8FD870D259729BBC9D4C3BEB37DD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne sonaliae Scherz & Glaw & Petzold & Andreone & Köhler & Raselimanana & Hutter & Belluardo & Rakotoarison & Hofreiter & Vences & Crottini 2026	<div><p>Rhombophryne sonaliae sp. nov.</p><p>Figures 2, 4 –7, 30, 31, 43 –45</p><p>Remarks.</p><p>This species has been incorrectly assigned to Plethodontohyla alluaudi since at least 1975 (Blommers-Schlösser 1975, 1976; Blommers-Schlösser and Blanc 1991), but the identity of that nomen was recently clarified (Bellati et al. 2018), leaving this relatively well-known species in need of a new name.</p><p>The vast majority of mentions of ‘ Mantipus alluaudi ’, ‘ Plethodontohyla alluaudi ’, and ‘ Rhombophryne alluaudi ’ in publications between 1976 and 2018 in fact refer to this species, especially in all genetic analyses (Blommers-Schlösser 1976; Vences and Glaw 2003; Vences et al. 2003 b, 2006; Vallan et al. 2004; Veith et al. 2004; Andreone et al. 2005; van der Meijden et al. 2005; Frost et al. 2006; Aprea et al. 2007; van der Meijden et al. 2007; Wollenberg et al. 2008; Vieites et al. 2009; D’Cruze et al. 2010; Glaw et al. 2010 b; Raselimanana 2010; Perl et al. 2014; Scherz et al. 2014, 2015 a, 2015 b, 2016 a, 2016 b, 2017 a, 2017 b). The following exceptions are noted: records from the north of Madagascar, including Ambolokopatrika (Andreone et al. 2000), Manongarivo (Rakotomalala 2002), and Bemanevika (Rabearivony et al. 2010) probably refer to other species of Rhombophryne . Records from southeastern Madagascar likely refer to the true Plethodontohyla alluaudi (Ramanamanjato et al. 2002; Andrianarimisa et al. 2009; Ramanamanjato and Soanary 2011). Some records, e. g., that from Andringitra (Raxworthy and Nussbaum 1996) are uncertain, and are omitted from our treatment here, pending verification with voucher specimens.</p><p>Belluardo et al. (2022) mistakenly figured specimen MRSN A 2610 of R. sp. Ca 10 as specimen MRSN A 2620 of R. sp. Ca 19 in figure 1 of their paper.</p><p>Informal names.</p><p>This species was referred to as ‘ Rhombophryne sp. ’ by Bellati et al. (2018), ‘ Rhombophryne sp. ex-alluaudi’ by Scherz (2020), ‘ Rhombophryne sp. Ca 19 ’ by Belluardo et al. (2022), and ‘ Rhombophryne sp. 19 ’ and ‘ CCS “ Sp. 19 ” ’ by Carné and Vieites (2024).</p><p>Justification of the new species.</p><p>The new species is placed in a clade containing R. botabota, R. nilevina, and a further new species, R. anatiala sp. nov. (described below), which was recovered as its sister species with high support (Belluardo et al. 2022). Genetic divergences among these lineages are less significant than those observed in the R. testudo group, but they all consistently differ among each other by&gt; 3.5 % uncorrected p distance in the 16 S 3 ’ fragment (3.8 % between R. sonaliae sp. nov. and R. anatiala sp. nov.) and thus at a level typically indicative of species-level differentiation in Malagasy frogs (Vieites et al. 2009). Furthermore, the new species differs from R. botabota by larger body size, from R. nilevina by advertisement call differences, and from both these species by colour pattern (many individuals have a light brown dorsolateral stripe and large light spots in the inguinal region and posteroventrally on hindlimbs; see Diagnosis below). There is no detected haplotype sharing with any other Rhombophryne species in the nuclear gene RAG 1 (Fig. 4). These differences are substantial and together with the molecular divergence strongly support the hypothesis of R. sonaliae sp. nov. constituting a new species. See also the Justification section for R. anatiala sp. nov. below.</p><p>Carné and Vieites (2024) recovered two deep conspecific lineages of this species, from Iampirano (misspelled ‘ Lampirano’) and Tsararano. Our species delimitation based on 16 S 3 ’ analyses also recovered these as conspecific.</p><p>Holotype.</p><p>ZSM 3/2002 (FGMV 2001.1271) an adult male, collected on 3 December 2001 in Andasibe (no precise coordinates), Alaotra-Mangoro Region, North Central East Madagascar, by M. Vences (Figs 30, 31, 44) .</p><p>Paratypes.</p><p>Seven specimens: ZSM 467/2005 (ZCMV 2221), a juvenile, and ZSM 466/2005 (ZCMV 2209), an ovigerous adult female, collected in February 2005 in pitfall traps in Andasibe (ca 18.9361°S, 48.4122°E, 939 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by R. Dolch and Mitsinjo colleagues; MRSN A 2620 (FN 6426), an adult male (sexed by incision), collected on 10 December 1996 in Tsararano (ca 14.907°S, 49.687°E, ca 700 m a. s. l.), Sava Region, northeastern Madagascar, by F. Andreone; MRSN A 5034 (RJS 0883), an adult male, collected on 15 January 2005 in Iampirano (16.8218°S, 49.1843°E, ca 500 m a. s. l.), Analanjirofo Region, North Central East Madagascar, by J. E. Randrianirina; ZFMK 52765, an adult presumed male, collected on 16 February 1991 in Andasibe (ca 18.9245°S, 48.4149°E, ca 940 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by F. Glaw and M. Vences (Fig. 43 A – D); UADBA 20308 (ZCMV 968), an adult male, collected on 21 February 2004 in Torotorofotsy (ca 18.874°S, 48.373°E, ca 960 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by M. Vences, E. Edwards, and C. Woodhead; MRSN A 7143 (FAZC 15560), an adult male, collected on 6 March 2017 in Maromizaha (18.9653°S, 48.4652°E, ca 1000 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar, by E. Coppola (Fig. 43 H, I).</p><p>Referred specimens.</p><p>BMNH 1986.28, an unsexed adult, collected on 8 July 1985 by C. J. Raxworthy at ‘ Camp 5 ’ in Zahamena Reserve (ca 17.67°S, ca 48.83°E, elevation unknown), Analanjirofo Region, North central east Madagascar, by C. J. Raxworthy. UADBA -A 62371 (MSZC 1480), collected on 13 March 2026 by Sandratra Rakotomanga, Alice Petzold, and Mark D. Scherz near Torotorofotsy (18.76965°S, 48.43532°E, 953 m a. s. l.), Alaotra-Mangoro Region, North Central East Madagascar; this specimen has been partly dissected in order to sequence and annotate its genome.</p><p>Assignment of specimens.</p><p>The paratype UADBA 20308 (ZCMV 968) is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341105; Vieites et al. 2009) and COI barcoding (COI KF 611584; Perl et al. 2014). The sequence of the holotype is identical to the sequence of this specimen in 16 S 3 ’ (AY 594112), and it differs by two mutations in 16 S 5 ’ (99.7 % identical; AY 594074), and by five mutations in COI (99.1 % identical; EF 396063). All paratypes and referred specimens conform well with the holotype in morphology. Assignment of specimens ZSM 466/2005, MRSN A 2620, MRSN A 5034, and MRSN A 7143 is further confirmed by molecular taxonomic identification (Fig. 2). ZSM 3/2002, MRSN A 2620, and MRSN A 7143 are present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. laevipes group, this species is sister to R. anatiala sp. nov. (described below) based on molecular phylogenetic data (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium to large body size, adult SVL 33.0– 44.7 mm; (2) head wider than long (HW / HL 1.59–1.71); (3) moderately small eyes (ED / HL 0.28–0.36); (4) relatively long hindlimbs (HIL / SVL 1.44–1.53); (5) small inner metatarsal tubercle (IMTL / FOL 0.12–0.15); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth, reaching or barely surpassing the joint of the first and second phalange of the fourth toe; (9) dorsal skin smooth; (10) absence of chin barbels; (11) absence of pronounced superciliary spines; (12) presence of large white or yellowish inguinal spots on a black background, and a light-brown dorsolateral stripe broadening from the eye towards the inguinal region; (13) presence of large white or yellowish spots on a black background on the posterior thigh and ventral surfaces of the shanks; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; (15) clavicles present, thin, curving; and (16) advertisement call (based on 4 analysed calls by one individual) pulsed, with (17) call duration 207–228 ms, (18) inter-call interval long (not measured), and (19) dominant frequency 785–828 Hz.</p><p>Rhombophryne sonaliae sp. nov. is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 392, ‘ T’ at site 445, and ‘ A’ at site 740; in 16 S 3 ’, ‘ A’ at site 1027, ‘ T’ at site 1076, ‘ C’ at site 1104, and ‘ T’ at site 1172.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. sonaliae sp. nov. can usually be distinguished from all species except R. anatiala sp. nov. described below by the presence of a light brown dorsolateral stripe expanding from the eyes towards the inguinal region. However, as this trait is not always present (Fig. 43 E, F), additional traits are necessary for a reliable diagnosis.</p><p>Rhombophryne sonaliae sp. nov. can easily be distinguished from R. testudo, R. coudreaui, R. matavy, and R. maraorao sp. nov. by smooth skin (vs rough), absence of chin barbels (vs general presence), longer hindlimbs (HIL / SVL 1.44–1.53 vs 1.10–1.41); from R. mangabensis and R. savaka by larger body size (SVL 33.0– 44.7 mm vs 18.7–23.2 mm), presence of multiple large light-coloured inguinal spots (vs black inguinal ocelli in R. savaka), and pulsed (vs tonal) advertisement call; from R. mavokely sp. nov., R. ornata, and members of the R. ellae species group by the absence of reddish orange colouration on the posterior thigh and in the inguinal region (vs presence); from members of the R. minuta species group by shorter hindlimb length (HIL / SVL 1.44–1.53 vs 1.73–1.84), and less gracile habitus; from R. proportionalis by much larger adult body size (SVL 33.0– 44.7 mm vs 11.0– 12.3 mm); and from all members of the R. serratopalpebrosa species group by the absence of superciliary spines (vs presence).</p><p>Within the R. laevipes species group, R. sonaliae sp. nov. can be distinguished from R. laevipes by shorter hindlimbs (HIL / SVL 1.44–1.53 vs 1.62–1.86), larger inner metatarsal tubercle (IMTL / FOL 0.12–0.15 vs 0.08–0.11), and advertisement call with a higher dominant frequency (785–828 Hz vs 612–631 Hz); from R. botabota by larger body size (SVL 33.0– 44.7 mm vs 24.2–32.2 mm), presence of large light-coloured spots in the inguinal region and on the posterior thigh and ventral shank (vs absence), and pulsed (vs tonal) advertisement call that is slightly longer (call duration 207–228 ms vs 161–201 ms) and lower in dominant frequency (785–828 Hz vs 1256–1291 Hz); and from R. nilevina by generally shorter hindlimbs (HIL / SVL 1.44–1.83 vs 1.52–1.69), and advertisement call that is shorter (call duration 207–228 ms vs 538–598 ms) and higher in dominant frequency (785–828 Hz vs 526–534 Hz). For distinction from other newly described members of this species group, see their respective descriptions below.</p><p>Description of holotype.</p><p>An adult male specimen in a moderately good state of preservation (Fig. 30). The abdomen is opened with the organs exposed. Tip of right third finger missing.</p><p>Body robust and rotund. Head broader than trunk, wider than long (HW / HL 1.59). Eyes quite small (ED / HL 0.28), pupils small, round. Snout rounded in dorsal and slightly truncate in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique. Nostril closer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, round, TDH / ED 0.64. Supratympanic fold distinct, slightly raised, running straight from posterior corner of eye over tympanum, curving gently ventrally posterior to the tympanum. Slight colour border between lateral head and dorsal head, with the lateral head being lighter in colour. Superciliary spines absent, no white dots above the eyes. Dorsal skin smooth in preservative. Tongue broad, attached anteriorly, posteriorly free. Maxillary and vomerine teeth present, vomerine teeth curved, with a dent each row marginally closer to midline than to lateral end, separated at the midline by a small gap. Choanae round and surrounded by pigment, placed laterally in the roof of the mouth.</p><p>Forelimbs relatively short (FARL / SVL 0.386), strongly built. Fingers without webbing, moderately long, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger distinctly shorter than fourth (Fig. 31). Finger tips not expanded, rounded. Finger subarticular tubercles distinct, whitish, flat; inner metacarpal tubercle broad and rounded, outer metacarpal / palmar tubercle indistinct, greyish. Hind limb strong, tibiotarsal articulation reaching between tympanum and eye when adpressed anteriorly along the body. TIBL / SVL 0.407. Inner metatarsal tubercle present, oblong, whitish, distinctly raised. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third, third toe just exceeding the base of the second phalange of the fourth toe; subarticular tubercles indistinct; toe tips slightly enlarged, rounded (Fig. 31).</p><p>After 20 years in preservative, the base colour is a coffee brown across the dorsum, and a lighter brown on the limbs. The central dorsum is immaculate, bordered either side by greyish wavy patterns riddled with tiny white points, giving an overall ‘ milky way’ impression. There is a slight colour border between the dorsal colour and the lateral head, with the lateral head lighter in colour than the dorsal surface. The inguinal region bears several large white spots which also extend onto the anterior thigh. There are also several white spots on the posterior thigh, ventral shank, and dorsal foot. The ventral thigh is light brown flecked with cream. The ventral feet are a light chocolate brown. A light annulus is present before the terminal phalange of each finger and toe. The legs lack crossbands dorsally. The arms have a single crossband on the lower arm. The palmar surface is grey-brown. The chin is light brown, the ventral abdomen flecked with cream. The cloaca region is unmarked, and there is no blackish bar above it. The colouration in life was not recorded.</p><p>Variation.</p><p>Type series SVL 33.0– 44.7 mm (n = 5 adults). For variation in other measurements, see Table S 6. Morphology is highly consistent. Third toe consistently reaching the second phalangeal articulation of the fourth toe or barely exceeding it. The most distinctive and constant feature is the diffuse light brown to grey stripe running from the eye toward the inguinal region flecked with tiny white specks, and distinct whitish or yellowish spots on the anterior and posterior thigh, inguinal region, and ventral shank. Only in MRSN A 2620 are these spots not clearly defined and rather small. In all other specimens they are extremely strongly defined. Ventral colour likewise consistent, but varying over the posterior abdomen from covered in flecks (ZSM 467/2005) to almost immaculate (MRSN A 5034). Chin dark in all available specimens. The referred specimen, BMNH 1986.28, is somewhat smaller than the rest of the type series, and its dorsal colouration is more mottled.</p><p>Call.</p><p>The advertisement call recorded on 21 February 2004 (15: 20 h) at Torotorofotsy (air temperature 19.1 ° C) consists of a single pulsed note of moderate duration repeated at long irregular intervals (Fig. 45). Calls (= notes) exhibit distinct amplitude modulation, with call energy starting at low level, constantly increasing to about two thirds of the call’s duration where energy reaches its maximum, followed by a decrease towards the call’s end. Pulses are more narrowly spaced in the last two thirds of the call’s duration and very short in duration, with each pulse having around 1 ms duration, but impossible to measure exactly. In the first third of the call, pulses are longer in duration (5–7 ms) and repeated less rapidly. It is not possible to count the number of pulses per note (= call), but pulse repetition rate can roughly be calculated for each section of the note. Numerical parameters of 4 analysed calls from 2 individuals are as follows: call duration (= note duration) 207–228 ms (219 ± 10 ms); inter-call intervals apparently long and irregular and not measured (probably poorly motivated calling; see Köhler et al. 2017); pulse repetition rate within first half of the call approximately 130–190 pulses / second, within second half of the call approximately 370–420 pulses / second; dominant frequency 785–828 Hz (805 ± 23 Hz); prevalent bandwidth 350–4000 Hz; numerous parallel frequency bands recognizable within the last two thirds of the call, representing a reflection of pulse rate (see Köhler et al. 2017). The call recording analysed herein has been published on the audio CD by Vences et al. (2006) as ‘ Plethodontohyla alluaudi ’ (CD 3, track 28). Assignment to this species is tentative, as the calling individual was not vouchered.</p><p>Etymology.</p><p>We dedicate this species to Sonali Garg, in recognition of her substantial contributions to knowledge of taxonomy and biogeography of South and Southeast Asian frogs, especially microhylids (e. g., Garg et al. 2018, 2019, 2022; Garg and Biju 2019).</p><p>Distribution.</p><p>Evidently widespread in eastern to northeastern Madagascar, but seldom encountered. Definitive records come from (1) Andasibe (type locality), (2) Torotorofotsy, (3) Iampirano, (4) Tsararano, (5) Maromizaha (Ratsimbazafy et al. 2008), and (6) Mandraka (Blommers-Schlösser 1975). We have examined a referrable specimen from (7) Zahamena (BMNH 1986.28). It has also been recorded in the (8) Ambatovy-Analamay region (Raselimanana 2010), and (9) An’Ala (Vallan et al. 2004). Elevation: ca 500–1200 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne sonaliae sp. nov. is a semi-fossorial or detriticolous species found in mid-elevation humid forest. Calling activity occurs mostly during or shortly after rain, often during the day but probably also at night. The captured calling male from Torotorofotsy was calling during the day from a burrow in the forest floor under the leaf litter. The gut contents of MRSN A 2620 included four large termites (each ca 9 mm long), the remains of a katydid (shanks 10.8 mm), a whole spider, a staphylinid rove beetle (ca 9 mm), and three small ants, as well as two parasitic worms. There was only little detritus other than these arthropods in the gut. Those of MRSN A 5034 contained a beetle tentatively assigned to Elateridae (click beetles), measuring 10.6 mm, a beetle elytrum, pieces of an ant, and a few large pieces of leaf, as well as two small parasitic worms. Field notes on ZFMK 52765 indicate that it had a caterpillar, among other things, in its stomach. Vallan et al. (2004) recorded a slight increase in detection of this species after logging of forest in An’Ala, but this may have been coincidental. Blommers-Schlösser (1975) found a female under a fallen tree trunk at 1200 m a. s. l. Referred specimen UADBA -A 62371 (MSZC 1480) was collected calling (including in response to playbacks) after rain at 10 h 13 in the morning on 13 March 2026. It was calling from under ca 10 cm of soil at the base of a tree; whilst excavating it, we encountered an adult Plethodontohyla notosticta, an adult Paradoxophyla cf. palmata, and a putative adult Madascincus cf. nanus .</p><p>Karyotype.</p><p>The karyotype was originally described by Blommers-Schlösser (1976) based on a female from Mandraka (as ‘ Mantipus alluaudi ’). It consists of 10 large and 16 small pairs of chromosomes (2 n = 26). Pair 4 subtelocentric, pairs 3, 7, 10, and 13 submetacentric. Aprea et al. (2007) provided a new description based on the holotype (ZSM 3/2002, as ‘ Plethodontohyla alluaudi ’). They reported both the fourth and the eleventh chromosome pairs to be subtelocentric. Nucleolar organiser regions were located in a peritelomeric position on the long arm of the second chromosome pair. Centromeric bands were DAPI positive.</p></div>	https://treatment.plazi.org/id/FA1B8FD870D259729BBC9D4C3BEB37DD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
07B2DE06B0FF55E2B1F544322E3E8D93.text	07B2DE06B0FF55E2B1F544322E3E8D93.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne tany Scherz 2015	<div><p>Rhombophryne tany Scherz et al., 2015</p><p>Figures 2, 4 –7, 67 –69</p><p>Remark.</p><p>This species is only known from the holotype.</p><p>Holotype.</p><p>ZSM 1814/2010 (ZCMV 12359), an adult male, collected on 13 June 2010 at <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.95728&amp;materialsCitation.latitude=-14.15256" title="Search Plazi for locations around (long 48.95728/lat -14.15256)">‘Camp 2’ Matsabory Maiky</a> (14.15256°S, 48.95728°E, 2021 m a. s. l.), Tsaratanana Massif, Diana Region, northern Madagascar, by M. Vences, D. Vieites, R. D. Randrianiaina, F. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, and T. Rajoafiarison .</p><p>Paratypes.</p><p>None.</p><p>Assignment of specimens.</p><p>The holotype and sole known specimen of the species is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ KP 895585; Scherz et al. 2015 b; 16 S 5 ’ KY 748104; Scherz et al. 2017 a) and COI barcoding (COI KF 611582; Perl et al. 2014). It is present in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is placed in a clade containing R. diadema, R. ornata, and R. guentherpetersi based on molecular phylogenetic data, but the exact relationships within that clade are poorly resolved (Fig. 2; Belluardo et al. 2022). It is characterised by the possession of the following unique suite of characters: (1) medium body size, adult SVL 24.6 mm; (2) head wider than long (HW / HL 1.53); (3) moderately large eyes (ED / HL 0.36); (4) relatively short hindlimbs (HIL / SVL 1.56); (5) small inner metatarsal tubercle (IMTL / FOL 0.07); (6) outer metatarsal tubercle absent; (7) second finger equal to fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin finely granular; (10) absence of chin barbels; (11) presence of two distinct superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne tany is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 504, ‘ T’ at site 557, ‘ T’ at site 578, and ‘ G’ at site 721. No diagnostic sites were identified for 16 S 3 ’.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. tany can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. For distinction from R. serratopalpebrosa, R. guentherpetersi, R. coronata, R. vaventy, and R. ornata, see the diagnosis of those species, above. It can be distinguished from R. regalis by the absence of an S-shaped dermal postnasal fold, two (vs three) superciliary spines, and stronger supratympanic fold; and from R. diadema by two (vs three) superciliary spines, stronger supratympanic fold, slightly shorter relative tibia length (TIBL / SVL 0.43 vs 0.44–0.46), prootics not in contact with parasphenoid alae (vs in contact), parasphenoid cultriform process with parallel edges (vs broadening anteriorly), nasals anterolaterally displaced (vs not displaced), quadratojugal-squamosal contact narrow (vs broad), anterior edge of ventral ramus of squamosal weakly (vs distinctly) stepped, dorsal prominence of iliac shafts strong (vs weak), and possibly unossified pubis (vs partially ossified). For distinction from new species described herein, see accounts below.</p><p>A detailed description of the species is given in (Scherz et al. 2015 b).</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>The Malagasy adjective tany meaning ‘ brown’, in reference to the overall brown colour of this species. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>This species is only known from (1) Matsabory Maiky on the Tsaratànana Massif (type locality). Elevation: 2021 m a. s. l.</p><p>Natural history.</p><p>Rhombophryne tany is a semi-fossorial or detriticolous species so far known only from high-elevation bamboo forest (Scherz et al. 2015 b).</p></div>	https://treatment.plazi.org/id/07B2DE06B0FF55E2B1F544322E3E8D93	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
2CB7C119692A506FA56A67305ED88382.text	2CB7C119692A506FA56A67305ED88382.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne testudo Boettger 1880	<div><p>Rhombophryne testudo species group</p><p>Rhombophryne coudreaui, R. mangabensis, R. maraorao sp. nov. (sp. Ca 09), R. matavy, R. mavokely sp. nov. (sp. Ca 17), R. savaka, R. sp. Ca 16, R. testudo</p><p>Summary of morphology.</p><p>A group of small to large (SVL 18.7–45.1 mm), detriticolous to truly fossorial Rhombophryne species. All have relatively short limbs and a rather flat and round habitus. Fossorial species ( R. testudo, R. matavy, R. coudreaui, and R. maraorao sp. nov.) lack clavicles entirely. The remaining species are detriticolous and retain clavicles (though reduced in R. mangabensis). Skin texture ranges from smooth ( R. mangabensis) to rugose and tubercular ( R. coudreaui). Maxillary, premaxillary, and vomerine teeth are present or absent.</p><p>Calls.</p><p>Calls known from all species except R. coudreaui and R. maraorao sp. nov. The call of R. testudo is a rather long, deep, pulsed note. That of R. matavy is a shorter tonal call, emitted in a series. The calls of R. mangabensis, R. savaka, and R. mavokely sp. nov. consist of high-pitched whistles, emitted in series.</p><p>Distribution.</p><p>This group is widely distributed (Fig. 7; Table S 4), with members in the north ( R. matavy and R. testudo), northeast ( R. mangabensis, R. maraorao sp. nov., R. mavokely sp. nov., R. savaka, R. sp. Ca 16), and east ( R. coudreaui) of Madagascar.</p><p>Ecology.</p><p>Species found in humid and subhumid tropical forest. Some present on both mainland and off-shore islands.</p></div>	https://treatment.plazi.org/id/2CB7C119692A506FA56A67305ED88382	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
043ECBCAAEBD55E48D2E862ACACDE045.text	043ECBCAAEBD55E48D2E862ACACDE045.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne testudo Boettger 1880	<div><p>Rhombophryne testudo Boettger, 1880</p><p>Figures 2, 4 –12</p><p>Holotype.</p><p>SMF 4241 (formerly 1177.2 a), an adult female, collected on an unknown date on ‘ insula Nossi Bé’ (<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=48.4&amp;materialsCitation.latitude=-13.4" title="Search Plazi for locations around (long 48.4/lat -13.4)">Nosy Be</a>; approx. coordinates 13.2– 13.4°S, 48.2– 48.4°E), Diana Region, Madagascar, by Carl Ebenau (erroneously later designated as a lectotype by Mertens 1967) (Figs 8, 9, 11).</p><p>Paratypes.</p><p>None. Boettger (1880 a) states explicitly that the species is based on a single specimen.</p><p>Referred specimens.</p><p>Numerous specimens claimed to be syntypes / paralectotypes exist as a result of the erroneous designation of SMF 4241 as lectotype by Mertens (1967). We are not aware of an exhaustive list, but Barbour and Loveridge (1929) stated there were seven in the Museum of Comparative Zoology of Harvard, namely MCZ 2163 (one specimen), 2236 (one), 2731 (three), plus 9317 and 9318 (two). At least one of the specimens in MCZ 2731 was exchanged to the Field Museum of Natural History, where it is archived as FMNH 73363 and incorrectly listed as a ‘ co-type’. A further specimen (EHT-HMS 44690, ex- SMF 4267) is listed as present in the type collection of the University of Illinois Museum of Natural History (Phillips 2015). This is also not a type. Four specimens formerly under catalogue number ZMB 10059 (now divided into ZMB 10059 [one specimen] and 51041–51043) were also originally incorrectly included in the catalogue of the Museum für Naturkunde, Berlin as types of R. testudo, but this was corrected in the digital catalogue.</p><p>We here examined SMF 4247 –4263 (16 specimens), which Mertens (1967) incorrectly considered paralectotypes — none of these specimens in fact has type status. Additionally, we examined ZSM 474/2000 (FGMV 2000.276) and ZSM 475/2000 (FGMV 2000.277), two adult specimens of uncertain sex, collected on 11 March 2000 on the east coast of Lokobe National Park (13.4133°S, 48.3342°E, 110 m a. s. l.), Nosy Be, Diana Region, northern Madagascar, by F. Glaw, K. Schmidt, and M. Vences; UADBA -A- FGZC 5620, an adult male, collected on 29 April 2019 on the Kindro trail, Ampasindava (13.4099°S, 48.3051°E, ca 80 m a. s. l.), Lokobe National Park, Nosy Be, Diana Region, northern Madagascar, by F. Glaw, M. D. Scherz, and A. Razafindraibe (Fig. 10); and ZMB 51041, an adult female, collected in 1887 on Nosy Be (no precise collection data), Diana Region, northern Madagascar, by C. Reuter (originally part of ZMB 10059). We further examined photographs of the false ‘ syntypes’, MCZ 2163, MCZ 9317, and MCZ 9318, to confirm their identity and morphological agreement with the holotype.</p><p>Assignment of specimens.</p><p>All referred specimens are unambiguously assigned to the species based on their overall morphology (the rotund body, short limbs, and chin barbels are unmistakable) and collection around the type locality. ZSM 475/2000 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ EU 341110 / AY 594125; 16 S 5 ’ EU 341110 / AY 594087; Vieites et al. 2009); a sample from either this specimen or ZSM 474/2000, with the tissue number MV 2000 - B 59, was used as the reference for COI barcoding (COI KF 611596 / MH 940140; Perl et al. 2014). These specimens cluster in our mitochondrial phylogeny (Fig. 2). Both specimens are included in the RAG 1 haplotype network (Fig. 4).</p><p>Definition.</p><p>Type species of the genus Rhombophryne . Within the R. testudo group, this species is sister to R. matavy (Fig. 2; Belluardo et al. 2022). It is characterised by having the following unique suite of characters: (1) medium to large body size, adult SVL 33.0– 43.1 mm; (2) head much wider than long (HW / HL 2.00–2.60); (3) small eyes (ED / HL 0.23–0.39); (4) short hindlimbs (HIL / SVL 1.18–1.41); (5) large inner metatarsal tubercle (IMTL / FOL 0.15–0.24); (6) generally pronounced outer metatarsal tubercle; (7) second finger slightly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin rough with converging paravertebral ridges; (10) presence of chin barbels; (11) absence of pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) highly robust skull with very broad squamosal, maxilla, and posterior process of angulosplenial; (15) clavicles absent; and (16) advertisement call (based on 4 analysed calls by one individual) highly pulsed (162–177 pulses per call), with (17) call duration 823–894 ms, (18) inter-call interval 6071–10186 ms, and (19) dominant frequency 508–524 Hz.</p><p>Rhombophryne testudo is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 456, ‘ T’ at site 651 and ‘ C’ at site 734; in 16 S 3 ’, ‘ C’ at site 1099, ‘ C’ at site 1179, and ‘ G’ at site 1216.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. testudo can easily be distinguished from all other species by the presence of several distinct chin barbels and its fortified skull, and from all species except R. coudreaui, R. matavy, and R. maraorao sp. nov. described below by its rotund body. From R. matavy it can further be distinguished by its pulsed (vs tonal) advertisement call that is much longer (call duration 823–894 ms vs 161–209 ms), and with a lower dominant frequency (508–524 Hz vs 1588–1688 Hz).</p><p>Redescription of the holotype.</p><p>A specimen in a good state of preservation, quite stiff and hard, somewhat desiccated (Figs 8, 9). Developing eggs evident in a micro-CT scan of the specimen.</p><p>Body robust, rhomboid. Head very wide and short, much wider than long (HW / HL 2.45). Eyes small, pupils small, round. Snout obtuse, slightly protruding and rounded in dorsal view, angular in lateral view, with five barbels on it, two slightly above and between the nostrils, two directly below these, and a central one below these over the commissure of the jaws. Canthus rostralis distinct, convex. Loreal region concave, slightly oblique. Nostril openings small, angled anterolaterally, equidistant between tip of snout and eye, slightly protuberant. Tympanum indistinct, vertically oval. TDH / ED 0.827. Supratympanic fold distinct, weakly raised but tubercular and so uneven in height, forming a gentle curve from the line of the top of the eye over the tympanum, sloping down to the posterior of the top of the arm. Superciliary spines are absent but there is a small light-coloured spot centrally above each eye, with several small tubercles on the dorsal surface of the eye.</p><p>The jaw is firmly shut, such that it is impossible to open without damaging the specimen irreparably. The original description described the specimen as lacking maxillary teeth, having small and weak palatal bones (interpreted as the neopalatines), a simple unfringed ‘ fold’ on the palate, which is interpreted as being the vomerine ridge, as the species lacks proper vomerine teeth (see Variation below), and having a large tongue that is wider than it is long, attached anteriorly and posteriorly free with a longitudinal furrow but without lateral emargination. Given that the jaw is so tightly shut, it is not clear how this information was obtained.</p><p>Forelimbs relatively short and thick, the upper arm hardly distinguishable from the trunk (FARL / SVL 0.397). Fingers without webbing, all short, relative lengths 1 &lt;2 = 4 &lt;3 (Fig. 9). Finger subarticular tubercles indistinct, single, round; inner metacarpal tubercle large, slightly shorter than first finger (IMCL / HAL 0.278); outer metacarpal / palmar tubercle indistinct, single. Hind limb thick, tibiotarsal articulation extent when adpressed anteriorly along the body cannot be assessed due to strong fixation. TIBL / SVL 0.259. Inner metatarsal tubercle present, large, longer than first toe, roughly as long as second toe (IMTL / FOL 0.193). Outer metatarsal tubercle faint. Toes not webbed; first toe distinctly reduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles indistinct, single and round; toe tips not enlarged. Finger and toe tips without circum-marginal grooves (Fig. 9).</p><p>Dorsal skin smooth, with numerous tubercles on the flanks and limbs and along the jaw, and especially a number of barbels on the bottom lip. The ventral skin is wrinkled but smooth. Symmetrical converging ridges are present from behind the exoccipital region of the head running toward the cloaca, with a pair of less distinct, less extensive ridges between them from the posterior head to roughly the level of the suprascapulae.</p><p>After at least 137 years in preservative, the colouration of the specimen is still brown and does not appear to have faded except perhaps in a blemished area on the right side of the dorsum at roughly the level of the ilia. The brown base colour has darker-brown symmetrical markings, which consist of a chevron running from the medial inguinal region to the centre of the dorsum and two dark irregular patches posterior to the suprascapular region. The tubercles and digits of the hands and feet are cream, and there is a fine cream line defining the fold between the dorsal and ventral posterior thigh. A single dark brown crossband is present on the shank, one on the tarsus, and one on the lateral foot that matches with that of the shank when the leg is fully folded. The ventral colour is as the dorsum, but lighter brown over the chin and with a curious translucent spot on the ventral abdomen. The colour of the specimen in life is unknown.</p><p>Variation.</p><p>SVL 33.0– 43.1 mm (n = 18); full measurements are presented in Table S 5. Most specimens could not be sexed because we did not have permission to dissect them, and the jaws of most specimens were so firmly fixed shut that we could not open the mouth and search for traces of vocal slits. Overall morphology is highly consistent. The tongue is broad and disc-like, posteriorly free. Maxillary teeth are absent. The vomerine teeth ridges form two straight transversal rows posterior to the rounded choanae, not meeting medially. The second and fourth fingers can vary in which is longer. There is a little variation in colouration, with some individuals uniformly brown in dorsal colour and others with dark-brown markings (e. g., Fig. 10).</p><p>Call.</p><p>The advertisement call recorded in January or February 1992 at Nosy Be (air temperature unknown) consists of a long, pulsed note emitted at irregular intervals (Fig. 12); calls are not arranged in call series. Pulses within calls are generally narrowly spaced and partly fused, but pulse repetition is distinctly faster in the second half of each call and results in respective parallel frequency bands reflecting pulse rate. Amplitude within calls is constantly increasing from the beginning towards the end where it drops to zero quickly. Numerical parameters of 4 analysed calls are as follows: call duration (= note duration) 823–894 ms (854 ± 30 ms); inter-call intervals 6071–10,186 ms (8370 ± 2100 ms); pulses per call 162–177 (169 ± 7); pulse repetition rate within first half of call approximately 135 pulses / second, within second half approximately 280 pulses / second; dominant frequency 508–524 Hz (519 ± 7 Hz); prevalent bandwidth 500–3200 Hz. Calls from the same recording (Vences et al. 2006, CD 3, track 33) have been described by Glaw and Vences (1992), D’Cruze et al. (2010), and Lambert et al. (2017) using slightly different terminological schemes.</p><p>Etymology.</p><p>Feminine nominative Latin noun meaning ‘ turtle’ or ‘ tortoise’, presumably in reference to the short limbs and rotund body shape of this species.</p><p>Distribution.</p><p>Confirmed from (1) Nosy Be (type locality), and (2) Nosy Komba (Glaw and Vences 1992; Andreone et al. 2003; Hyde Roberts and Daly 2014). Elevation: 0–450 m a. s. l. (the peak of Mt Lokobe on Nosy Be).</p><p>Records from Réunion (Guibé 1978) resulted from a clerical error when information were being copied from one catalogue to another, and were corrected by Glaw and Vences (2002). A record from ‘ Imerina’ by Parker (1934) is based on BMNH 1895.10. 29.30, collected by Richard Baron. The location Imerina could refer to anywhere across the high plateau of central Madagascar (the area where Baron served as a reverend; Laurence 1987). Baron was one of the most widely travelled explorers of that time, and the lack of precise collection date means that refinement of locality is challenging. However, it should be noted that Baron undertook an enormous journey from central-eastern Madagascar, along the coast north to Antsiranana, and then down the west coast to Nosy Be (a journey of almost 2000 km), collecting plants along the way (JSTOR Global Plants correspondence KADC 2664 from Library and Archives at Royal Botanic Gardens, Kew). It is therefore conceivable that the location is an error; that Baron collected the specimen in Nosy Be at the end of his journey and sent it along with the consignment of plants to London. Examination of this specimen revealed it to be an adult male. It does not have any evident morphological differences from other R. testudo specimens. However, as stated above, a museomic sample of this specimen confirmed it to have moderately strong genetic differentiation from other R. testudo specimens. A BLASTn search of every contig in NCBI, however, consistently identified Rhombophryne testudo ZSM 475/2000 as closest match, and with a sequence coverage of 4.9 × and 2.9 × for 16 S 3 and 16 S 5, respectively, it remains possible that a genetically divergent lineage is present in Imerina (or at some other location along Baron’s travel routes), which would be in need of re-discovery to verify. Records from Marojejy and Andapa-Sambava (Guibé 1978; Blommers-Schlösser and Blanc 1991; probably deriving from the same specimen, see Glaw and Vences 2002) referred to R. coudreaui by Andreone and Randrianirina (2008), but are here referred to R. maraorao sp. nov., described below.</p><p>Natural history.</p><p>Rhombophryne testudo is a fossorial species found in primary and secondary low-elevation humid forest, where it inhabits burrows 5–10 cm deep beneath the leaf litter (Glaw and Vences 1994). Calls are emitted from January to April, at highest intensity before and during heavy rain, irrespective of the time of day (Glaw and Vences 1994). Calling can occur without rain, but a series of two or three calls is then followed by a long pause of several minutes (Glaw and Vences 1994). Köhler et al. (1997) reported finding a 10 cm wide, 15 cm deep nest beneath a rock beside a small creek, containing two adults and 18 juveniles measuring 9–11 mm in SVL, in December 1995. This suggests development occurs in subterranean burrows with parental care, perhaps by both parents (Köhler et al. 1997). No tadpoles or eggs of the species have yet been observed.</p><p>A defensive behaviour is known wherein specimens stretch their hindlimbs to prevent them being turned onto their backs (Glaw and Vences 1994). We have also observed individuals puff themselves up significantly when caught, presumably to make them more difficult to swallow — common behaviours in burrowing microhylid frogs.</p><p>Keefe and Blackburn (2020) compared the humerus of Rhombophryne testudo to other burrowing frogs. They noted its similarity to forward-burrowers, suggesting that it may also be able to burrow forwards, but highlighting the need for behavioural observations of this species.</p><p>The gut of the holotype contains a great deal of soil, as evident from our micro-CT scans (Fig. 11). We examined the contents of the digestive system of an adult female specimen (with well-developed eggs), ZMB 51041. The stomach contained mostly detritus, especially tiny fragments of wood and leaves. Fragments of two insects were present in this detritus: the 4.7 mm abdomen of a small rove beetle ( Staphylinidae), and the thorax and partial abdomen of an ant. The intestines contained mostly a fine mud with tiny fragments of wood and leaf, the head of an ant, and the thorax and head of the same rove beetle. By volume, the vast majority of gut contents was non-animal material. This suggests that most of the ingested matter of these frogs might be substrate, not insects. Although unlikely, more ecological research is needed to understand if these animals may be partly detritivorous or geophagous. However, it is more likely that insect larvae, annelid worms, or other soft-bodied organisms are the primary prey of these frogs, and are digested quickly and thus absent from the digestive system of preserved frogs.</p><p>Karyotype.</p><p>The karyotype of this species was described by Aprea et al. (2007), with 2 n = 26 largely metacentric chromosomes (second and third pairs submetacentric, fourth pair subtelocentric / telocentric). Nucleolar organiser regions were in a pericentromeric position on the short arm of the second chromosome pair. Heterochromatin was indistinct, and Chromomycin A 3 (CMA 3) and 4 ', 6 - diamidino- 2 - phenylindole (DAPI) negative, i. e., lacking particularly GC- or AT-rich regions.</p></div>	https://treatment.plazi.org/id/043ECBCAAEBD55E48D2E862ACACDE045	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
7BE09820F9BD54298918590634365864.text	7BE09820F9BD54298918590634365864.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne undefined-Ca-15	<div><p>Rhombophryne sp. Ca 15</p><p>Remark.</p><p>This candidate species was identified by Belluardo et al. (2022) based on the specimen DLR 620 (extraction number ACP 3194), which has been sequenced for the mitochondrial markers 16 S 3 ’ (OL 780581), 16 S 5 ’ (OL 780604), 12 S (OL 780590), COB (OL 853700), and COI (OL 790135), and the nuclear markers BDNF (OL 853691), POMC (OL 780557), RAG 1 (PZ 377035), RAG 2 (OL 780561), SACS-A (OL 853704), and KIAA 1239 (OL 780539). It has also been referred to as ‘ R. sp. 15 ’ and ‘ UCS “ Sp. 15 ” ’ by Carné and Vieites (2024). Other than its collection locality (Betampona Strict Nature Reserve, central eastern Madagascar), we have no further data on this specimen. Based on its phylogenetic position, it appears to be the closest relative of R. kotrobaratra . No specimens were available to us at this time and it was not possible to assess this species taxonomically, but it is distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ T’ at site 288, ‘ T’ at site 413, and ‘ C’ at site 642; in 16 S 3 ’, ‘ G’ at site 899, ‘ A’ at sit 1026, ‘ G’ at site 1267, and ‘ T’ at site 1276.</p></div>	https://treatment.plazi.org/id/7BE09820F9BD54298918590634365864	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
9D9C27F1A70857B59CB2BCD5D270A342.text	9D9C27F1A70857B59CB2BCD5D270A342.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne undefined-Ca-16	<div><p>Rhombophryne sp. Ca 16</p><p>Remark.</p><p>This candidate species was identified by Belluardo et al. (2022) based exclusively on GenBank sequences of the specimen AMNH A 181903 (RAX 10368), a specimen registered as ‘ Rhombophryne cf. minuta ’ on the AMNH online database portal (https://emu-prod.amnh.org, accessed 25 January 2023). It was referred to as ‘ Rhombophryne sp. 16 ’ by Belluardo et al. (2022) and ‘ UCS “ Sp. CA 16 ” ’ by Carné and Vieites (2024). It has been sequenced for mitochondrial markers 16 S 3 ’ and 16 S 5 ’ (KM 509192), COI (KM 509860), and nuclear genes BDNF (KM 509332), TYR (KM 509984), SIAH 1 (KM 509728), and C-MYC (KM 509466). It was collected in the Andapa area (given as a county in the AMNH online database, without precise coordinates) of the Sava Region, northeastern Madagascar by C. J. Raxworthy. Based on its phylogenetic position (Fig. 2; Belluardo et al. 2022), it appears to be a member of the R. mangabensis species complex, but we have not been able to examine this specimen, and therefore are not able to assess it taxonomically. Genetically, it differs by&gt; 7.95 % uncorrected p distance in the 16 S 3 ’ fragment from all other Rhombophryne species (Table S 2). It is distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ A’ at site 293, ‘ G’ at site 483, and ‘ A’ at site 539. No diagnostic sites were identified for 16 S 3 ’.</p></div>	https://treatment.plazi.org/id/9D9C27F1A70857B59CB2BCD5D270A342	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
422CE28145ED529AB112D8A075DE237B.text	422CE28145ED529AB112D8A075DE237B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne undefined-Ivohibe’	<div><p>Rhombophryne sp. ‘Ivohibe’</p><p>Figures 2, 4 –7, 67 –69, 76</p><p>Remark.</p><p>This population has been reported as ‘ Plethodontohyla serratopalpebrosa ’ by Raxworthy and Nussbaum (1996) and Raselimanana (1999). No genetic or bioacoustic data are available from it, and despite some morphological differences to all known species in the genus, we here refrain from describing it as a new species until new material is collected and its genetic affinities have been clarified.</p><p>Available material.</p><p>UADBA 17419 (APR 01644), UADBA 17420 (APR 1660), two adult females, and UADBA 17421 (APR 01663), UADBA 17422 (APR 01677), UADBA 17423 (APR 01770), UADBA 17424 (APR 01771), UADBA 17430 (APR 01784), five unsexed adults, all collected between 14 October and 2 November 1997 in the Reserve Spécial Pic d’Ivohibe (22.4833°S, 46.9694°E, ca 1200 m a. s. l.), Ihorombe Region, South Central Eastern Madagascar, by A. P. Raselimanana; UADBA 17425 (APR 01853), an unsexed adult, collected at 16: 00 h on 2 November 1997 in the corridor between Andringitra and Pic d’Ivohibe (22.4175°S, 46.8858°E, ca 1200 m a. s. l.), Ihorombe Region, central eastern Madagascar, by A. P. Raselimanana.</p><p>Circumscription.</p><p>Morphologically, this species shows the following suite of characters: (1) small to medium body size, adult SVL 21.6–25.8 mm; (2) head wider than long (HW / HL 1.34–1.59); (3) moderately large eyes (ED / HL 0.33–0.43); (4) relatively long hindlimbs (HIL / SVL 1.66–1.89); (5) small inner metatarsal tubercle (IMTL / FOL 0.08–0.10); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin shagreened; (10) absence of chin barbels; (11) presence of three to four pronounced superciliary spines, the longest being centrally over the eye; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Thus, this species can easily be distinguished from all species except other Rhombophryne except members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. It differs from R. serratopalpebrosa and R. regalis by the absence of an S-shaped dermal postnasal fold; from R. regalis by the longest superciliary spine being over the centre of the eye (vs the anterior-most spine in R. regalis); from R. guentherpetersi by the absence of large, swollen tibial glands (vs presence); from R. coronata by longer hindlimb (HIL / SVL 1.66–1.89 vs 1.28–1.43), first toe unreduced (vs usually reduced), superciliary spines varying in length (vs small and equally sized), and shagreened dorsal skin (vs finely granular), and sphenethmoid not appreciably extending anterior to the postchoanal vomer (vs extending anteriorly); from R. vaventy by much smaller adult body size (SVL 21.6–25.8 mm vs&gt; 50 mm); from R. ornata by smaller adult body size (SVL 21.6–25.8 mm vs up to 33 mm), three to four superciliary spines (vs two), and probable absence of reddish orange colouration in the inguinal region and on the thigh (vs presence); from R. tany by longer hindlimb (HIL / SVL 1.66–1.89 vs 1.56), and three to four superciliary spines (vs two); and from R. diadema possibly by more shagreened dorsal skin and more distinct supratympanic fold, by the longest superciliary spine being over the centre of the eye (vs anterior two spines equal in length and posterior-most diminutive).</p><p>Generalised morphological description.</p><p>SVL 21.6–25.8 mm (n = 8); for variation in other measurements, see Table S 10. Body rhomboid, head rather triangular. Head wider than long, eyes fairly large, pupils small, round. Snout slightly pointed in dorsal and truncate in lateral view. Canthus rostralis distinct, concave. Loreal region concave, oblique, with no trace of an S-shaped curve posterior to the nostril. Nostril nearer to tip of snout than to eye, slightly protuberant. Tympanum indistinct, round. Supratympanic fold distinct, strongly raised, partly melanised, curving gently from posterior corner of eye over tympanum toward anterior insertion of the forearm. Lateral head similar in colour to dorsal head. Three to four distinct superciliary spines above each eye, the longest over the centre of the eye. Dorsal skin shagreened in most specimens. Ventral skin smooth. Tongue quite narrow, broadening posteriorly, attached anteriorly, posteriorly free, not lobed. Maxillary and vomerine teeth present, vomerine teeth rather straight, without any variation in height, separated from one another at the midline by a small gap. Choanae oval, placed laterally in the roof of the mouth, slightly melanised.</p><p>Forelimbs rather long, slender. Fingers without webbing, relative lengths 1 &lt;2 &lt;4 &lt;3, second finger distinctly shorter than fourth. Finger tips not expanded, rounded. Finger subarticular tubercles indistinct; inner metacarpal tubercle small, weakly developed, greyish, outer metacarpal / palmar tubercle indistinct, greyish, single. Hind limb slender. Inner metatarsal tubercle present, oblong, greyish, weakly raised. Outer metatarsal tubercle absent. Toes not webbed; toes unreduced, toe lengths 1 &lt;2 &lt;5 &lt;3 &lt;4, fifth toe distinctly shorter than third; subarticular tubercles indistinct, grey; toe tips not enlarged, slightly pointed, especially fourth toe.</p><p>The base colour is a dark brown across the dorsum and flanks, including the dorsal limbs. The dorsum has several nearly black markings on it in all individuals, but vary in intensity; some individuals have lines that run slightly discontinuously from the suprascapular region toward the inguinal region roughly parallel to one another. No distinct inguinal spots present, no traces of cream-bordered markings present. No distinct colour difference between the lateral and dorsal head. The supratympanic fold is blackish above the tympanum, but otherwise dark brown. The hindlimbs lack distinct crossbands, but a concentration of these blackish markings on the mid-shank gives the impression of a crossband. The posterior thigh is not distinctly coloured. The foot is laterally dark brown, but medially creamy brown. A cream annulus is present before the terminal phalange of each finger and toe. The ventral foot is dark brown, fading to grey along the toes. No distinct colouration around the cloaca. The arms have several blacking markings, including a moderately large marking in the middle of the antebrachium. The dorsal hand is lighter medially than laterally; the ventral hand is light brown. The chin is mottled light brown with tan flecks, fading to completely tan over the sternal region into the immaculate abdomen.</p><p>Distribution.</p><p>Specimens referred to this putative candidate species are from (1) the Pic d’Ivohibe Special Reserve (type locality), and (2) the corridor between Ivohibe and Andringitra. Raxworthy and Nussbaum (1996) recorded specimens probably referrable to this species at 1625 m a. s. l. in Andringitra National Park, but also gave it a range from 1240 to 1300 m a. s. l. in the same table; it is not clear which range is correct, but the latter better matches the specimens discussed here. Elevation: ca 1200–1275 m a. s. l.</p><p>Natural history.</p><p>This is a semi-fossorial or detriticolous species found in mid-elevation humid forest. Specimens of this species were caught in pitfall traps and on surveys by Raselimanana (1999). During the day they hide under thick leaf litter or under rotten logs. An individual (UADBA 17422, APR 01677) was found under a rotten log with tadpoles at 16: 00 h in primary rain forest on a ridge. Regrettably, both the tadpoles and data pertaining to them have been lost.</p></div>	https://treatment.plazi.org/id/422CE28145ED529AB112D8A075DE237B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
4815557DA5885227BAB855AF3F821664.text	4815557DA5885227BAB855AF3F821664.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhombophryne vaventy Scherz, Ruthensteiner, Vences & Glaw 2014	<div><p>Rhombophryne vaventy Scherz, Ruthensteiner, Vences &amp; Glaw, 2014</p><p>Figures 2, 4 –7, 67 –69, 75</p><p>Remark.</p><p>The 3 ’ fragment of the 16 S gene sequences of specimen AMNH A 167315 (referred to as ‘ R. cf. vaventy ’ in Belluardo et al. 2022 and ‘ R. sp. cf. vaventy ’ and ‘ DCL “ Sorata ” ’ by Carné and Vieites 2024) from Sorata is poorly differentiated from the homologous sequence of paratype UADBA -A- FGZC 2842, and our species delimitation results confirm that these are conspecific.</p><p>Holotype.</p><p>ZSM 357/2005 (FGZC 2876), an adult male, collected on 17 February 2005 at ‘ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=49.7434&amp;materialsCitation.latitude=-14.4367" title="Search Plazi for locations around (long 49.7434/lat -14.4367)">Camp Simpona’</a> (14.4367°S, 49.7434°E, 1326 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by F. Glaw, M. Vences, R. D. Randrianiaina .</p><p>Paratypes.</p><p>One specimen: UADBA -A- FGZC 2842, an unsexed probable adult, with same collection data as the holotype but collected on 16 February 2005 .</p><p>Referred specimens.</p><p>ZSM 522/2016 (ZCMV 15278), an adult female, collected on 20 November 2016 near ‘ Camp Simpona’ (14.4366°S, 49.7434°E, 1325 m a. s. l.), Marojejy National Park, Sava Region, northeastern Madagascar, by M. D. Scherz, A. Rakotoarison, M. Bletz, M. Vences, and J. Razafindraibe (Fig. 75).</p><p>Based on our species delimitation analysis, AMNH A 167315, a specimen of unknown sex and age, collected at Camp 1, Vohemar, Sorata Mountain (13.6858°S, 49.4419°E, ca 1300 m a. s. l.), Sava Region, northeastern Madagascar, is also attributable to this species. However, that specimen has not been examined, and this assignment remains tentative at present.</p><p>Assignment of specimens.</p><p>The paratype UADBA -A- FGZC 2842 is used as the reference specimen for this species in 16 S rRNA barcoding (16 S 3 ’ and 16 S 5 ’ EU 341107; Vieites et al. 2009) and COI barcoding (COI KF 611595; Perl et al. 2014). The holotype is identical to this specimen in 16 S 3 ’ (GenBank: PZ 418427) and differs by three gaps in a repetitive string of cytosines in 16 S 5 ’ (GenBank: PZ 405243; Fig. 2). The paratype and ZSM 522/2016 conform well with the holotype in morphology, and are topotypical. ZSM 357/2005 and UABA-A- FGZC 2842 are present in the RAG 1 haplotype network (Fig. 1).</p><p>Definition.</p><p>Within the R. serratopalpebrosa group, this species is possibly closest related to R. regalis and R. serratopalpebrosa based on molecular phylogenetic data (Fig. 2). It is characterised by the possession of the following unique suite of characters: (1) large body size, adult SVL 51.9–58.6 mm; (2) head wider than long (HW / HL 1.56–1.70); (3) moderately large eyes (ED / HL 0.43–0.45); (4) relatively long hindlimbs (HIL / SVL 1.88–1.95); (5) small inner metatarsal tubercle (IMTL / FOL 0.11–0.14); (6) outer metatarsal tubercle absent; (7) second finger distinctly shorter than fourth; (8) third toe distinctly longer than fifth; (9) dorsal skin rugose; (10) absence of chin barbels; (11) presence of four pronounced superciliary spines; (12) absence of inguinal ocelli or spots; (13) absence of distinctive markings along the posterior thigh; (14) unfortified skull without expansion of maxilla or posterior process of angulosplenial; and (15) clavicles present, thin, curving.</p><p>Rhombophryne vaventy is furthermore distinguished from all other Rhombophryne species analysed by the following robust diagnostic nucleotide combination in the mitochondrial 16 S rRNA gene (positions relative to the full 16 S sequence of Anilany helenae MZ 751042): in 16 S 5 ’, ‘ C’ at site 309, ‘ C’ at site 385, ‘ G’ at site 642; in 16 S 3 ’, ‘ C’ at site 920, ‘ A’ at site 1177, and ‘ A’ at site 1217.</p><p>Diagnosis.</p><p>Within the genus Rhombophryne, R. vaventy can easily be distinguished from all species except other members of the R. serratopalpebrosa species group by the presence of distinct superciliary spines. Within the R. serratopalpebrosa species group, it can be distinguished from all species by its much larger body size (SVL 51.9–58.6 mm vs &lt;36 mm), and its rugose dorsal skin texture (vs smooth to granular).</p><p>A detailed description of the species is given in Scherz et al. (2014), but see also notes in Scherz et al. (2017 a).</p><p>Variation.</p><p>Adult SVL 51.9–58.6 mm; for variation in other measurements see Table S 10. The newly collected female specimen is 6.7 mm larger than the male holotype; there may be some degree of sexual size dimorphism. In all other respects, it agrees closely with the types.</p><p>Call.</p><p>The call of this species is unknown.</p><p>Etymology.</p><p>The Malagasy adjective vaventy meaning ‘ large’, in reference to the exceptionally large size of this species. Used as an invariable noun in apposition.</p><p>Distribution.</p><p>Originally known only from (1) the Marojejy Massif (Rakotoarimalala and Raselimanana 2023 reported it from 1300 ± 200 m a. s. l. and 1875 ± 200 m a. s. l.), but here expanded to encompass (2) Sorata as well by the assignment of AMNH A 167315. Elevation: ca 1300–1875 m a. s. l. (the upper elevation being rather uncertain).</p><p>Natural history.</p><p>Rhombophryne vaventy is a semi-fossorial or detriticolous species found in mid- to high-elevation humid forest. The newly collected adult female specimen ZSM 522/2016 was found active on the ground during the day by local guides.</p></div>	https://treatment.plazi.org/id/4815557DA5885227BAB855AF3F821664	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Scherz, Mark D.;Glaw, Frank;Petzold, Alice;Andreone, Franco;Köhler, Jörn;Raselimanana, Achille P.;Hutter, Carl;Belluardo, Francesco;Rakotoarison, Andolalao;Hofreiter, Michael;Vences, Miguel;Crottini, Angelica	Scherz, Mark D., Glaw, Frank, Petzold, Alice, Andreone, Franco, Köhler, Jörn, Raselimanana, Achille P., Hutter, Carl, Belluardo, Francesco, Rakotoarison, Andolalao, Hofreiter, Michael, Vences, Miguel, Crottini, Angelica (2026): A wealth of riches: A comprehensive revision of diamond frogs, genus Rhombophryne Boettger, 1880 (Microhylidae: Cophylinae), with the description of seven new species. Vertebrate Zoology 76: 485-610, DOI: 10.3897/vz.76.e172567
