identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E48797FFB2FFE540817F24FDCBFEBA.text	03E48797FFB2FFE540817F24FDCBFEBA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Atelopus spumarius Cope 1871	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Atelopus spumarius Cope, 1871</p>
            <p>Figs. 3–4</p>
            <p> Atelopus spumarius Cope, 1871: 222 from “Ambyiacu R.“ (= PERU: Loreto: Río Ampiyacu), syntypes lost (2 specimens at ANSP, leg. John Hauxwell); Lötters et al. 2002a: 168; Lötters et al. 2002b: 97; Lötters et al. 2003: 173; Lötters et al. 2005: 345; Rueda Almonacid et al. 2005: 126; De la Riva et al. 2011: 162; Frost 2024; Lötters 1996: 47; Lötters et al. 2023: Supplementary Table 1; Plewnia et al. 2024b: 241. </p>
            <p> Atelopus spumarius spumarius . — Rivero 1968: 23; Lescure “1973” 1974: 144; Lescure 1981: 894 (designation of a neotype, MNHNP 1979.8382) from “Colonia, bassin de la rivière Ampiyacu (Depto. Loreto), Pérou” (= PERU: Loreto: Río Ampiyacu basin, Colonia); Lescure &amp; Gasc 1986: 717. </p>
            <p>
                  Neotype. PERU: Loreto: Colonia (= Colonia Ancón on the right shore of  
                <a title="Search Plazi for locations around (long -71.98333/lat -3.2)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-71.98333&amp;materialsCitation.latitude=-3.2">Río Zumún</a>
                 close to the juncture with the  
                <a title="Search Plazi for locations around (long -71.98333/lat -3.2)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-71.98333&amp;materialsCitation.latitude=-3.2">Río Yaguasyacu</a>
                 , a tributary to the  
                <a title="Search Plazi for locations around (long -71.98333/lat -3.2)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-71.98333&amp;materialsCitation.latitude=-3.2">Río Ampiyacu</a>
                 ; 3°12’S, 71°59’W, 100 m asl), MNHNP 1979 /8382 (Fig. 3), adult female, leg. May 1978 by Jean Lescure. 
            </p>
            <p> Referred material.   PERU: Loreto: 3 km NE Pebas, AMNH A 103030–103035, 6 adult females (leg. April 1977 by Charles W. Myers and John W. Daly), MUBI 18222–18224, 1 female, 2 males (leg. 26 August 2022 by Philipp Böning, Angel Chujutalli, Jaime Culebras, Christopher H. Heine, Alisha Jung, Karl-Heinz Jungfer, Stefan Lötters and Amadeus Plewnia);  Pebas , NHMW 3886 /1–2, 2 adult females (leg. unknown)  . </p>
            <p> Definition.  Atelopus spumarius  sensu stricto is placed in the genus  Atelopus based on its mt and nc markers nested within  Atelopus (and with the 16S rRNA fragment showing uncorrected p-distance &lt;4% to the genus type species  A. flavescens Duméril and Bibron ), and on having presacrals I and II fused. The species is dinstinguished from all other species by the combination of the following characters: (1) A small-sized species with SVL of adult males 22.2 and 23.4, N = 2, and females 25.8 ± 1.80, N = 10; (2) slender body (SW / SVL = 0.261 and 0.270 in 2 males, 0.287 ± 0.021 in 10 females) with (3) long legs (TIBL / SVL = 0.457 and 0.495 in 2 males, 0.461 ± 0.029 in 10 females) and (4) acuminate to subacuminate snout in males and subacuminate to truncate in females, protruding slightly beyond apex of lower jaw; (5) tympanic membrane absent, tympanic annulus not visible, columella present, ostia pharyngea present; (6) phalangeal formula of hand 1-2-3-3; webbing absent on hand; (7) first finger short (THBL / HAND = 0.423 and 0.455 in 2 males, 0.412 ± 0.020 in 10 females), in adult males with keratinized light brown nuptial pad; (8) phalangeal formula of foot 2-2-3-4-3, webbing formula of toes I 0–0 to 1 / 2 II 0 to 1 / 2 –1 to 11 / 2 III 1 to 11 / 2 –3 IV 3–0 to 11 / 2 V; (9) skin smooth to areolate, dorsally and laterally covered with dense, almost indistinct minute spiculae; they are sometimes aggregated in small groups; ventrally smooth to areolate, cloaca and adjacent thighs most areolate; (10) vertebral column conspicuous, neural processes absent; (11) in life: dorsum black to brown with few yellow, greenish or whitish asymmetric circles, dots or lines; dorsolaterally yellow, greenish or whitish foam-like reticulation forming a band from above eye to groin with numerous black or brown rounded irregular spots (smaller than EYDM); laterally a black or brown band from tip of snout almost to groin, rarely interrupted by yellow, greenish or whitish lines; limbs and dorsal surfaces of hand and foot with yellow, greenish or whitish foam-like reticulation with black or brown spots (smaller or larger than EYDM), thumb and sometimes tips of fingers and toes dorsally yellow to yellowish orange; in males, throat and venter cream, posteriorly becoming orange, sometimes with few tiny brown dots (smaller than or equal to diameter of finger III) on throat and small brown dots (smaller than EYDM) on lateral and posterior vent; in females, throat and chest orange or yellow, venter anteriorly cream yellow, posteriorly orange or yellow, anteriorly with numerous brown or black dots (smaller than or equal to EYDM), sometimes as a continuous chest band; in both sexes palms, plants and ventral surface of cloaca and underside of thighs and upper arm in both sexes and forearm in females bright orange; iris black with a narrow golden ring surrounding horizontally oriented pupil; (12) in preservative: above brown with cream to brownish white pattern; below whitish to cream with brown dots; bright orange life markings faded to pale orange, pale yellow or cream; (13) pulsed call 922 ± 24 ms at dominant frequency 3,404 ± 22 Hz  . </p>
            <p> Diagnosis.  Atelopus spumarius sensu stricto (Figs. 3–4) can be readily distinguished from all other  Atelopus (as far known) by molecular genetics, with an uncorrected p-distance of the 16S rRNA fragment of 3.66% to its closest known described relatives,  A. franciscus Lescure and the  A. hoogmoedi Lescure complex (Appendices 8, 9). It can be morphologically distinguished from all congeners, except from  A. colomai Plewnia, Terán-Valdez, Culebras, Boistel, Paluh, Quezada Riera, Heine, Reyes-Puig, Salazar-Valenzuela, Guayasamin and Lötters ,  A. histrionicus sp. nov. , and  A. manauensis Jorge, Ferrão and Lima, by the combination of small size, dorsal and lateral skin covered with dense, almost indistinct minute spiculae, ventral skin smooth to areolate, presence of a columella and dorsal pattern. From these species, it differs in call duration (922 ± 24 ms vs. 2,049 ± 220 ms in  A. colomai ; 807 ± 80 ms in  A. histrionicus sp. nov. ; 744 ± 84 ms in  A. manauensis ; Jorge et al. 2020b) and in palmar and plantar coloration in life (orange vs. red). It differs from  A. colomai and  A. histrionicus sp. nov. in female ventral coloration and pattern in life (throat and chest orange or yellow, venter anteriorly cream yellow with numerous brown or black dots smaller than or equal to EYDM, sometimes forming a continuous chest band, posteriorly orange or yellow vs. below entirely yellow with large black rounded dots all over the ventral side, except throat, in  A. colomai ; below entirely whitish to cream with large rounded to irregular dark brown markings in the gular region, on chest and venter in  A. histrionicus sp. nov. ). Further, it differs from  A. colomai in having a subacuminate to truncate snout in lateral view in females (vs. acuminate and more protruding). From  A. manauensis ,  A. spumarius sensu stricto differs in having a longer head (males, HLSQ/HDWD = 1.100 –1.151, N = 2, vs. 1.054, N = 11; Jorge et al. 2020b). </p>
            <p> Atelopus spumarius sensu stricto (Figs. 3–4) is similar to its congeners of the flavescens-spumarius clade, and to  A. andinus Rivero ,  A. loettersi De la Riva, Castroviejo-Fisher, Chaparro, Boistel and Padial , and  A. tricolor Boulenger complex (including the junior synonyms  A. rugulosus Noble ,  A. willimani Donoso-Barros ) from the eastern Andean versant of Peru and Bolivia. It differs from the nominal  A. barbotini Lescure ,  A. flavescens Duméril and Bibron (including its junior synonym  A. vermiculatus McDiarmid ),  A. franciscus and the  A. hoogmoedi Lescure complex (including the available name  A. hoogmoedi nassaui Ouboter and Jairam ) in pattern and coloration (dorsally black to brown with few yellow, greenish or whitish asymmetric circles, dots or line; dorsolaterally yellow, greenish or whitish foam-like reticulation forming a band from above eye to groin with numerous black or brown rounded irregular spots; ventrally cream in males, cream and orange or yellow in females, with small dark dots vs. dorsally black with red to pink markings, ventrally reddish pink in  A. barbotini ; dorsally unicolored reddish, brownish or yellow or brown and with olive, brownish reddish and copper-like vermiculation, ventrally reddish pink in males, pink in females in  A. flavescens ; dorsally olive to blackish brown, ventrally reddish in  A. franciscus ; dorsally dark brown or black with yellow, pink, bluish, white, greenish or orange irregular spots and reticulated dorsolateral bands or solid bars, ventrally yellow, pink, bluish, whitish, greenish or orange, in some specimens with dark spots or blotches in the  A. hoogmoedi complex). All these forms, except some populations of the  A. hoogmoedi complex, lack orange palmar and plantar surfaces in life (present in  A. spumarius sensu stricto ). All Guianan harlequin frogs have longer call duration than  A. spumarius sensu stricto (922 ± 24 ms vs. 1,490 ± 140 ms in  A. barbotini ; 1,530 ± 220 ms in  A. flavescens ; 1,490 ±150 ms in  A. franciscus ; 1,190 ± 10 ms in the  A. hoogmoedi complex from French Guiana; 1,060 -1,240 ms in the  A. hoogmoedi complex from Pará, Brazil; Lescure 1981; Cocroft et al. 1990) with the exception of members of the  A. hoogmoedi complex from Amapá, Brazil, (1,160 ± 390 ms; Costa-Campos &amp; Carvalho 2018). In addition,  A. spumarius sensu stricto can be distinguished from  A. barbotini and the  A. hoogmoedi complex by smaller female SVL (25.8 ± 1.80, N = 10, vs. 32.9 ± 2.6, N = 9, in  A. barbotini from the type locality; 34.5 ± 1.9, N = 11, in  A. hoogmoedi from the type locality). It can be distinguished from  A. pulcher Boulenger by smaller male SVL (22.2–23.4, N = 2, vs. 27.27 ± 1.07, N = 13, in  A. pulcher ; Lötters et al. 2002a), ventral coloration in life (females throat and chest orange or yellow, venter anteriorly cream yellow, posteriorly orange or yellow, anteriorly with numerous brown or black dots, sometimes as a continuous chest band vs. vermillion red with irregular dark spots in  A. pulcher ). In addition, it differs from  A. pulcher in having shorter calls (922 ± 24 ms vs. 1,200 ± 100 ms in  A. pulcher ; Lötters et al. 2002a).  Atelopus spumarius sensu stricto differs from  A. harlequin sp. nov. in larger male SVL (22.2 and 23.4, N = 2, vs. 20.6 ± 0.51, N = 4, in  A. harlequin sp. nov. ) and in dorsal coloration in life (black to brown with few yellow, greenish or whitish asymmetric circles, dots or lines, dorsolaterally yellow, greenish or whitish reticulation forming a continuous broad band from above eye to groin bearing numerous black or brown rounded spots vs. yellowish-gold with numerous dark brown, rounded marks and irregular shaped blotches in  A. harlequin sp. nov. ). In addition, it differs from  A. harlequin sp. nov. in having longer calls (922 ± 24 ms vs. 680 ± 30 ms in  A. harlequin sp. nov. ; Asquith &amp; Altig 1987).  Atelopus spumarius sensu stricto differs from  A. seminiferus Cope in smaller female SVL (25.8 ± 1.80, N = 10, vs. 40.0, N = 1), absence of lateral warts (present in  A. seminiferus ) and dorsal coloration in preservative (brown with brownish pale asymmetric circles, dots or line and dorsolateral foam-like reticulation forming a band from above eye to groin with numerous black or brown rounded irregular spots vs. dark brown without pattern in  A. seminiferus ). From  A. andinus ,  A. loettersi and the  A. tricolor complex,  A. spumarius sensu stricto can be distinguished by the presence of a columella (vs. absence; Lötters et al. 2011; authors’ unpubl. data). From  A. loettersi , it can further be distinguished by ventral coloration and pattern in life (females throat and chest orange or yellow, venter anteriorly cream yellow, posteriorly orange or yellow, anteriorly with numerous brown or black dots, sometimes as a continuous chest band vs. mostly red, in some specimens with irregular brown blotches in  A. loettersi ; De la Riva et al. 2011).  Atelopus spumarius sensu stricto can be distinguished from  A. andinus and the  A. tricolor complex by the absence of warts (presence) and by presence of dorsolateral reticulation (absence). </p>
            <p>Description of neotype. Adult female. Body slender (SW/SVL = 0.269), head roughly as long as wide (HLSQ/ HDWD = 1.037), snout protruding below apex of lower jaw, truncate in dorsal view; nostrils directed laterally, barely protuberant, not visible in dorsal view, almost at tip of snout in lateral view, situated above apex of lower jaw; canthus rostralis well defined, concave in dorsal view between eye and nostrils; loreal region concave; lips not flared; top of snout depressed; head slightly concave in lateral view; eyelid flared; tympanum absent, tympanic annulus not visible; supratympanic crest poorly developed; choanae small, rounded; ostia pharyngea present; tongue about twice as long as wide, broadest at its tip (situated posteriorly); limbs long and slender (TIBL/SVL = 0.441); webbing absent on hand; fingers and toes lack lateral fringes; palmar tubercle rounded, poorly developed; supernumerary palmar and plantar tubercles absent; thenar and subarticular tubercles indistinct; tips of digits rounded; phalangeal formula of hand 1-2-3-3; first finger short (THBL/HAND = 0.444); inner metatarsal tubercle indistinct, outer metatarsal tubercle rounded, poorly developed; relative length of toes I&lt;II&lt;III&lt;V&lt;IV; phalangeal formula of foot 2-2-3-4-3, webbing formula of toes I 0–0 II 1 / 2 – 11 / 2 III 1–3 IV 3–1 V; tarsal fold absent; foot roughly three quarters of tibia; skin smooth on entire body, dorsally and laterally covered with dense, almost indistinct minute spiculae; skin areolate on loreal region and ventral surfaces of cloaca and adjacent areas of thighs, venter and throat; cloacal opening in an inconspicuous tube, directed posteriorly; vertebral column visible through the skin, neural processes absent.</p>
            <p>For live coloration, see Remarks.</p>
            <p>In preservative, above brown with brownish pale dorsal spots, circles and dorsolateral irregular reticulation pattern; dorsolateral band posteriorly fused with a narrower band presenting similar reticulation running ventrolaterally from jaws to groin; ventral surfaces of limbs and body pale with brown dots (smaller than or equal to EYDM), which are most dense on lateral and anterior parts of vent and in gular region; smaller dots (smaller than EYDM) on throat and no dots on ventral surface of cloaca and adjacent thighs (Fig. 3).</p>
            <p>Measurements: SVL 29.0, TIBL 12.8, FOOT 11.0, HLSQ 8.4, IOD 2.9, HDWD 8.1, EYDM 2.9, ITNA 2.4, FAL 9.2, HAND 7.2, THBL 3.2, SW 7.9.</p>
            <p>Variation. For meristic variation of characters see Table 3. Specimens from the Pebas area (Fig. 4) correspond to the neotype description. However, they are slightly smaller than the neotype (and other females from Colonia, examined by Lescure 1981). Skin texture varies from smooth to slightly areolate, with almost indistinct minute spiculae dorsally and laterally which can be aggregated in small groups in some specimens. In dorsal view, the snout varies from acuminate to subacuminate in males and from subacuminate to truncate in females.</p>
            <p>In life, dorsal and dorsolateral marks and reticulation pattern varies from yellow to greenish or whitish (Fig. 4).</p>
            <p> Sexual dimorphism is apparent in  A. spumarius sensu stricto with females being larger than males (SVL 25.8 ± 1.80, N = 10, vs. 22.2–23.4, N = 2). In addition, males possess a light brown keratinized nuptial pad on the dorsal surface of first finger and differ in ventral coloration and pattern as indicated above (Fig. 4). </p>
            <p> Skull osteology. General osteological features of two females (MNHNP 1979/8382, neotype, MorphoSource Media ID 000618620; AMNH A103034, MorphoSource Media ID 000436646) are depicted in Figures 5–6 and Appendix 4. The skull of  A. spumarius sensu stricto is triangular in dorsal view. The skull length is 7.8, the skull width is 7.0, and the skull height is 4.8 in AMNH A1033034. The skull roof is slightly rugose. In anterior view, the septomaxilla is U-shaped with medial and lateral rami that extend posteriorly towards the vomer. The lateral ramus is much broader than the medial ramus and bears a nasal process. The ossified sphenethmoid is smooth and underlies the posterior and medial margins of the nasals, as well as the anterior margins of the frontoparietals. The posterior limit of the sphenethmoid is about one-half the length of the margin of the orbit. The prootics are slightly rugose and are fused to the posterolateral edge of the frontoparietals. The exoccipitals encircle the foramen magnum and are fused to the frontoparietals dorsally, the prootics laterally, and the parasphenoid anteroventrally. The otic capsule is well-ossified. Columellae are present, cylindrical in shape. The pars media plectra of the columella is slender and bowed, and the pars interna plectra is broad. The operculum in the fenestra ovalis is cartilaginous. The nasals are triangular and bear an acuminate maxillary process that extends ventrolaterally toward the maxilla and nearly contacts the pars facialis of the maxilla. The nasals are narrowly separated medially. The anterior roof of the braincase has a triangular appearence given by the coarsely rectangular frontoparientals, which bear a short triangular suprarobital flange (more evident in the neotype) with nearly even, non-straight orbital edges. Medial articulation is incomplete in both specimens with the anterior one fourth of the bones lacking articulation in the neotype and the anterior three-fourths of the bones lacking articulation in AMNH A1033034. Suture lines are visible where articulation is present. Occipital groves, which are canals for the carotid artery, are present and partially roofed with bone. In the neotype, the left groove is mostly open, while the right grooves irregularly covered along the midpoint. In AMNH A1033034, the left and right groove are covered only at the anterior and posterior tips. Posteriorly, the frontoparietals are fused posterolaterally to the prootics and posteromedially to the medial anterodorsal margins of exoccipitals. The vomers are small, edentate, medially separated, crescent-shaped, and triradiate with an anterior ramus, prechoanal ramus, and postchoanal ramus. The anterior ramus is directed anterolaterally, pointed at the anterior tip, and is roughly the same size as the postchoanal ramus. The prechoanal ramus is short. The pre- and postchoanal rami form the anteromedial margin of the choana. The prechoanal ramus is directed towards the maxilla, and the postchoanal ramus is directed posterolaterally. The postchoanal ramus is fused to the overlying sphenethmoid. The neopalatine is elongate and triangular. It underlies, and is partially fused to, the sphenethmoid and extends towards the maxilla, making contact with the preorbital process. The neopalatine is narrow and pointed at its medial tip, widens as it approaches the maxilla, and is cylindrical and rounded at its anterior border. The parasphenoid has an inverted T-shape. The cultriform process does not contact the sphenethmoid. The cultriform process extends beyond the midpoint of the orbit. The terminal end of the cultriform process is spear-shaped in the neotype and roughly rounded in AMNH A1033034. The parasphenoid alae are directed posterolateral to the cultriform process and are fused to the overlying otic capsule. The posterior margin of the parasphenoid and a medial posterior process is present (more evident in the neotype). The maxillary arcade is complete and edentate. The paired premaxillae each possess a dorsal alary process that is directed anterolaterally. The lower half of the alary process is wider than the upper half. The pars palatina is biradiate with medial and lateral processes. The lateral process is more than twice as long as the medial process. The medial process is well developed and triangular. A concave border is present where the lateral and medial processes of the pars palatina come together. The posterior tip of the pars dentalis is pointed and articulates with the maxilla. Small, irregular sesamoids are present at the contact zone joint between the pars palatina of the premaxilla and the pars palatina of the maxilla in AMNH A1033034. The external surface of the maxillae is slightly rugose. The pars palatina, which extends along the lingual margin of the maxilla, is narrow, and the posterior half of this shelf articulates with the anterior ramus of the pterygoid. The pars fascialis of the maxilla is directed medially and is short and jagged; its dorsomedial margin nearly contacts the neopalatine, sphenethmoid, and the maxillary process of the nasal. The anterior end of the maxilla is truncate and abuts the premaxilla. The posterior end of the maxilla is pointed and contacts the quadratojugal. The quadratojugal is a small, L-shaped bone that underlies the ventral arm of the squamosal and contacts the posterior process of the maxilla at is anterior margin. The quadratojugal of the neotype is much shorter than that of AMNH A1033034. The paired squamosals possess an otic ramus posterodorsally, a zygomatic ramus anterodorsally, and a ventral ramus. The otic ramus is expanded dorsally and has a broad articulation with the crista parotica of the otic capsule, leaving only the posterior end of the prootic free. The zygomatic ramus is small, triangular, and directly medially. The angle between the dorsal surface of the squamosal and the anterior margin of the ventral ramus is nearly perpendicular. The ventral ramus is flat and blade-like.A squamosal keel is present, extending along the outer surface from the zygomatic ramus to the upper half of the ventral arm. The pterygoid is triradiate, bearing anterior, medial, and posterior rami. The anterior ramus articulates with the pars palatina of the maxilla. The medial and posterior rami are of equal length. The medial ramus nearly contacts the prootic. The posterior ramus is flat and blade-like. The palatoquadrate is cartilaginous and therefore not visible in the microCT dataset. The lower jaw is composed of three ossified elements and Meckel’s cartilage, which is not visible in the microCT dataset. The mentomeckelian is the most anterior element that forms a cartilaginous symphysis at the midline of the jaws. Posterodorsally, the mentomeckelian is fused to the dentary. The dentary is slender, thin, and edentate; this element overlays the anterior half of the angulosplenial. The angulosplenial is the largest mandible element and forms the lingual surface of the lower jaws. The lower jaw length is 6.3 in AMNH A1033034, slightly shorter than the length as the skull from the occipital condyle to the premaxilla. </p>
            <p> The neotype of  Atelopus spumarius sensu stricto differs from the species described herein by the posterior end of the maxilla being free and laterally protruding beyond the lateral limit of the squamosal (vs., in  A. colomai , protruding beyond the lateral limit of the squamosal, but not free; in  A. harlequin sp. nov. and in  A. histrionicus sp. nov. , the squamosal extends laterally beyond the posterolateral limit of the maxilla; Fig. 6), the quadratojugal having a thickened pars glenoidalis and a short pars jugularis (vs., in  A. colomai , pars glenoidalis subrectangular, pars jugularis elongated;  A. harlequin sp. nov. , pars glenoidalis longer but more narrow; in  A. histrionicus sp. nov. , pars glenoidalis triangular; Fig. 6), and the maxilla having a straighter orbital margin (Fig. 6). However, variation in these characters exists between the two populations of  A. spumarius sensu stricto , which might be attributed to imaging conditions or natural variation. </p>
            <p> Further,  A. spumarius sensu stricto can be distinguished from  A. colomai by a slightly rugose skull roof (vs. smooth) and the frontoparietals being fused over two posterior thirds of their total length (vs. completely fused).  Atelopus spumarius sensu stricto differs from  A. harlequin sp. nov. by the posteromedial side of the otic ramus of the squamosal being free (vs. attached along its entire length to the prootic).  Atelopus spumarius sensu stricto can be distinguished from  A. histrionicus sp. nov. by a less rugose skull roof, the lateral margins corresponding to the maxilla being less straight, the posteroventral incisure of the sphenethmoid being less deep, the dorsal end of the dorsal process of the premaxilla (= pars facialis) and the nasals at the same level (vs. protrudes anteriorly beyond the anterior limit of the nasals). For details see Figures 5–6 and Appendix 4. </p>
            <p> A. spumarius A. histrionicus sp. nov. A. harlequin sp. nov. A. colomai</p>
            <p>Squamosal in lateral view</p>
            <p>However, the discriminatory power of osteological features needs to be seen with caution as intraspecific variation, sexual dimorphism and taxonomic value remain poorly understood for the species discussed.</p>
            <p>Vocalization. Calls were recorded by PB from an uncollected male on 27 August 2022, ca. 01:00 pm, at 3 km NE Pebas with a Comica CVM-VM20 directional microphone attached to a Roland R-05 digital receiver. Distance to recorder was 0.4 m and ambient temperature was ca. 25°C. A 51,125 ms recording was available for analysis (CJ ec.cj.aud.22; Fonozoo Sound Code 14654; Fig. 7). It contains 20 pulsed calls with duration 922 ± 24 ms (881–962 ms). Intervals between calls last 1,647 ± 162 ms (1,372 –1,991 ms). Dominant frequency is 3,404 ± 22 Hz (3,373 –3,466 Hz). Amplitude is ascending over the course of the call and slightly decreases over the last 3–6 pulses. Frequency shows harmonics and is modulated with a slight increase towards the end of the call. Each pulsed call consists of 42 ± 3 pulses (37–48), that show a length of 8 ± 4 ms (3–27 ms) each. Pulses slightly increase in length over the course of each call with the last pulse being considerably longer than any other pulse. The first or the first two pulses of each call are detached by a long interval of 122 ± 16 ms (83–147 ms) from all subsequent pulses.</p>
            <p>Two additional recordings display other call types. The first (total recording 2,222 ms; same recording data as above; CJ ec.cj.aud.23; Fonozoo Sound Code 14655; Fig. 8) consists of two pulsed short calls with a duration of 132 and 225 ms, interval 1,110 ms and dominant frequency 2,997 and 3,110 Hz, respectively. Amplitude peaks in the middle of the call with a slight decrease towards the end. Frequency does not show harmonics and is poorly modulated with a minimal increase towards the mid of each call followed by a minimal decrease towards the end and no modulation within pulses. Pulsed short calls consist of 7–16 pulses that show a length of 8 ± 8 ms (3–40 ms) each. The second additional recording (total recording 10,386 ms; same recording data as above; CJ ec.cj. aud.24; Fonozoo Sound Code 14656; Fig. 9) consists of a single pure tone short call with a call duration of 47 ms, dominant frequency of 3,199 Hz and a pulsed short call with a call duration of 68 ms, dominant frequency 3,192 Hz, containing only two pulses with a length of 25 and 38 ms. The interval between the pure tone short call and the pulsed short call is 6,708 ms. In both calls, frequency shows harmonics and is modulated with a decrease towards the end.</p>
            <p> Tadpole. The tadpole of  A. spumarius sensu stricto is unknown. </p>
            <p> Distribution.  Atelopus spumarius sensu stricto is known from three localities in the Río Ampiyacu drainage at ca. 80–100 m asl (Peru, Loreto) (Fig. 2). In addition to the two sites mentioned above, the species has been reported from the nearby Nuevo Peru on the Río Ampiyacu (William Lamar, in litt. 30 July 2022). It is likely present in more localities as the general area is homogenous lowland rain forest and poorly studied, as it is difficult to access. Additional populations from other river drainage systems of unclear taxonomic status may be allocable to  A. spumarius sensu stricto suggesting that the species’ range may be larger (see Remarks). </p>
            <p>Natural history. The species inhabits terra firme lowland rain forest and can be found in close proximity to small streams. In late August 2022, we recorded life history notes at 3 km NE Pebas. All individuals observed were found no more than 2 m away from the edge of a stream ca. 0.5–1 m wide and up to 20 cm deep, characterized by slow-moving clear water, sandy substrate and abundant fallen logs (Fig. 10). Adults were active during the day perching on the steep loamy slopes of the stream or on vegetation (&lt;30 cm above ground). Sex ratio was skewed towards males (12 of 15 adults found). Male-male interaction was observed suggesting home range defence (as described in related taxa, e.g. Ringler et al. 2023): During mid-day, we observed one male approaching a resident (larger) male perching on a trunk in the stream bed (see Plewnia et al. 2024a: Supplementary Video). The resident male started to call, thus showing its bright cream throat towards the other male. In addition, hand waving behavior was performed. The resident male pursued the other male which moved away by climbing up the steep slopes. In the late afternoon, individuals climbed on leaves along the steep slopes, which likely served as sleeping sites at night. Females found in April and August contained well-developed eggs. Amplexus was never observed; juveniles, tadpoles or eggs are unknown.</p>
            <p> Conservation status. We suggest listing  A. spumarius sensu stricto under the range of categories for species considered threatened (Critically Endangered to Vulnerable) based on the following criteria. The known EOO of  A. spumarius sensu stricto is ca. 50 km ² and AOO is 12 km ². While its distributional limits are poorly known it is unlikely to have an EOO greater than 20,000 km ² or to occur at more than 10 threat defined localities. </p>
            <p>Most parts of the Río Ampiyacu drainage are covered in undisturbed primary forest with few human (indigenous) settlements and traditional small-scale agriculture (chacras, slash and burn fields). However, we witnessed deforestation and pollution in the immediate Pebas area, i.e. 3 km NE Pebas. Here, the species was found in high densities in a small forest fragment in August 2022 (15 individuals in approx. 10 person*hours). Our visits to the type locality, Colonia Ancón, and the surroundings (April 2012 and January 2014, SL and collaborators) did not reveal any individuals.</p>
            <p>Presence and potential impact of Bd remain unknown, four individuals sampled August 2022 (3 km NE Pebas) tested negative (prevalence 0 %; CI 0–51.6 %; Appendix 5).</p>
            <p> Atelopus spumarius sensu stricto is further likely to be threatened by climate change in the near future, which might both impact the species habitat as well as interfer and exacerbate existing threats such as disease (Lötters et al. 2023). The species from the western Amazon are among the harlequin frogs likely to be most affected by climate change (Lötters et al. 2023). </p>
            <p>Etymology. Cope (1871) did not provide a derivatio nominis. The epithet is Latin (nominative case, singular) meaning ‘foam maker’, obviously referring to the species’ dorsolateral pattern.</p>
            <p>Remarks. Lescure (1981) collected 34 specimens in 1978 in Colonia Ancón, including 6 females.All specimens were deposited at MNHNP. However, except the neotype, these specimens have not been traced for more than 20 years (Annemarie Ohler, MNHNP, pers. comm. to SL).</p>
            <p> Specimens from Igará-Paraná, Departamento Amazonas, Colombia (BMNH 1905.1.31.10-11) and from Yubineto, Loreto, Peru (MNHNP 1979-8416, AMNH 95815-96005) were additionally assigned to this taxon by Lescure (1981). While these specimens resemble  A. spumarius sensu stricto by external morphology and bioacoustics (Yubineto; cf. Lescure 1981), we are currently unable to draw final conclusions on the taxonomic status of this material. For comments on additional populations potentially conspecific with  A. spumarius sensu stricto see chapter 3.5. </p>
            <p>Lescure (1981) described the neotype in life as above black with yellow dorsolateral pattern; below yellow with black spots. His description of a dorsolateral longitudinal band from orbit to the armpit needs rectification, as this pattern is continued beyond the armpit to the groin. In addition, he refers to the area below cloaca to the inner thighs,</p>
            <p>the palms and plants as vermilion red (Lescure 1981). We are unable to refute this, but this deviation from bright orange coloration as known from specimens in the Pebas area might best be explained by an overinterpretation of this feature. This needs further corroboration once new material becomes available from Colonia Ancón.</p>
            <p>Re-examination of the neotype revealed that “small warts behind the eye” (sensu Lötters et al. 2002a; b) were an interpreation in error; the skin is areolate here (Fig. 11). However, still dorsal and lateral sides are covered with dense, almost indistinct minute spiculae.</p>
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	https://treatment.plazi.org/id/03E48797FFB2FFE540817F24FDCBFEBA	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.		MagnoliaPress via Plazi	Lötters, Stefan;Böning, Philipp;Bailon, Salvador;Castañeda, Jose Daniel Barros;Boistel, Renaud;Catenazzi, Alessandro;Chaparro, Juan C.;Chávez, Germán;Chujutalli, Angel;Coen, Laurent;Coloma, Luis A.;Crawford, Andrew J.;Culebras, Jaime;Martínez, Juan Carlos Cusi;Daza, Juan Manuel;Riva, Ignacio De La;Ellwein, Denise J.;Ernst, Raffael;Flechas, Sandra V.;Fouquet, Antoine;Guayasamin, Juan Manuel;Heine, Christopher;Jorge, Rafael F.;Jung, Alisha;Jungfer, Karl-Heinz;Kaffenberger, Nathalie;Krehenwinkel, Henrik;Marca, Enrique La;Lampo, Margarita;Medina Rangel, Guido F.;Orsen, Ludvig;Paluh, Daniel J.;Gonzalez, José Luis Pérez;Perrin, Jonathan;Quezada Riera, Amanda B.;Reyes-Puig, Juan Pablo;Ross, Bernardo Roca-Rey;C, Daniela;Solano, Luis Alberto Rueda;Salazar-Valenzuela, David;Vazquez, Josefa Celsa Señaris;Sowinski, Morgane;Terán-Valdez, Andrea;Tovar-Ortiz, Angie;Veith, Michael;Venegas, Pablo;May, Rudolf Von;Weitkamp, Timm;Plewnia, Amadeus	Lötters, Stefan, Böning, Philipp, Bailon, Salvador, Castañeda, Jose Daniel Barros, Boistel, Renaud, Catenazzi, Alessandro, Chaparro, Juan C., Chávez, Germán, Chujutalli, Angel, Coen, Laurent, Coloma, Luis A., Crawford, Andrew J., Culebras, Jaime, Martínez, Juan Carlos Cusi, Daza, Juan Manuel, Riva, Ignacio De La, Ellwein, Denise J., Ernst, Raffael, Flechas, Sandra V., Fouquet, Antoine, Guayasamin, Juan Manuel, Heine, Christopher, Jorge, Rafael F., Jung, Alisha, Jungfer, Karl-Heinz, Kaffenberger, Nathalie, Krehenwinkel, Henrik, Marca, Enrique La, Lampo, Margarita, Medina Rangel, Guido F., Orsen, Ludvig, Paluh, Daniel J., Gonzalez, José Luis Pérez, Perrin, Jonathan, Quezada Riera, Amanda B., Reyes-Puig, Juan Pablo, Ross, Bernardo Roca-Rey, C, Daniela, Solano, Luis Alberto Rueda, Salazar-Valenzuela, David, Vazquez, Josefa Celsa Señaris, Sowinski, Morgane, Terán-Valdez, Andrea, Tovar-Ortiz, Angie, Veith, Michael, Venegas, Pablo, May, Rudolf Von, Weitkamp, Timm, Plewnia, Amadeus (2025): A roadmap for harlequin frog systematics, with a partial revision of Amazonian species related to Atelopus spumarius. Zootaxa 5571 (1): 1-76, DOI: 10.11646/zootaxa.5571.1.1, URL: https://doi.org/10.11646/zootaxa.5571.1.1
03E48797FF8BFFF040817AEDFE21FE56.text	03E48797FF8BFFF040817AEDFE21FE56.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Atelopus harlequin Coloma, Plewnia, Boning, Boistel, Ellwein, Lotters, Paluh, Roca-Rey Ross and Venegas 2025	<html xmlns:mods="http://www.loc.gov/mods/v3">
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            <p> Atelopus harlequin Coloma, Plewnia, Böning, Boistel, Ellwein, Lötters, Paluh, Roca-Rey Ross and Venegas sp. nov.</p>
            <p>Species account authors: Luis A. Coloma, Amadeus Plewnia, Philipp Böning, Renaud Boistel, Denise J. Ellwein, Stefan Lötters, Daniel J. Paluh, Bernardo Roca-Rey Ross, Pablo Venegas</p>
            <p>LSID: urn:lsid:zoobank.org:act: 429FD89A-7986-42DF-AA71-5689EAFA8CDB</p>
            <p>Figs. 18–20</p>
            <p> Atelopus “harlequin ”.— Coloma 1997: 39 (conditional name). </p>
            <p> Atelopus spumarius non-Cope.— Asquith &amp; Altig 1987: 32; Lötters et al. 2002a: 169; Lötters et al. 2003: 170; Lötters et al. 2005: 345; Rueda-Almonacid et al. 2005: 126 (by implication). </p>
            <p> Atelopus spumarius spumarius .— Cocroft et al. 1990: 638, Lötters 1996: 47 (by implication). </p>
            <p> A. sp.  cf. spumarius “harlequin ”.— Lötters et al. 2023: Supplementary Table 1. </p>
            <p>
                  Holotype. PERU: Loreto: km 74 on road from Iquitos to Nauta along a tributary of the  
                <a title="Search Plazi for locations around (long -73.52917/lat -4.2855554)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-73.52917&amp;materialsCitation.latitude=-4.2855554">Río Itaya</a>
                 (4°17’08”S, 73°31’45”W, ca. 100 m asl), MUBI 2448 (Fig. 18), adult male, leg. 28 August 2008 by Stefan Lötters and Philine Werner. 
            </p>
            <p>
                  Paratypes. PERU: Loreto:  
                <a title="Search Plazi for locations around (long -75.42833/lat -3.4491668)" href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-75.42833&amp;materialsCitation.latitude=-3.4491668">Pucacuro</a>
                 (3°26’57”S, 75°25’42”W, 150 m asl), FHGO 0177, 7584, 7760, adult males, FHGO 7587, adult female, leg. Jorge Valencia. 
            </p>
            <p> Referred material.   ECUADOR: Pastaza:  Lorocachi , QCAZ 8898, 8899 (cleared and stained)  , 8900, 9447– 9450, 9522, 9544, males, leg. 21–25 February 1996 by Omar Torres Carvajal, María Cecilia Terán and Xavier Cisneros;   PERU: Loreto:  Andoas , CORBIDI 4883  ;   Loreto: Platanoyacu,  Trompeteros , MUBI 7822  . </p>
            <p> Definition.  Atelopus harlequin sp. nov.  is placed in the genus  Atelopus based on its mt and nc markers nested within  Atelopus (and with the 16S rRNA fragment showing uncorrected p-distance &lt;5% to the genus type species  A. flavescens Duméril and Bibron ), and on having presacrals I and II fused. The species is dinstinguished from all other species by the combination of the following characters: (1)A small-sized species with mean SVL of adult males 20.6 ± 0.51, N = 4 and adult female 25.7; (2) slender body (SW / SVL = 0.250 ± 0.003 in 4 males, 0.272 in female) with (3) long legs (TIBL / SVL = 0.479 ± 0.024 in 4 males, 0.490 in female) and (4) acuminate snout, protruding beyond apex of lower jaw; (5) tympanic membrane absent, tympanic annulus not visible, columella present, ostia pharyngea present; (6) phalangeal formula of hand 1-2-3-3; webbing absent on hand; (7) first finger short (THBL / HAND = 0.432 ± 0.021 in 4 males, 0.400 in female), in adult males with keratinized brownish nuptial pad; (8) phalangeal formula of foot 2-2-3-4-3, webbing formula of toes I 0–0 to 1 II 1 / 2 to 1–1 to 2 III 1 to 2–3 IV 3–1 V; (9) skin smooth, dorsally and laterally covered with dense, poorly distinct minute spiculae; ventrally smooth to slightly areolate, cloaca and adjacent thighs areolate; (10) vertebral column conspicuous, neural processes absent; (11) in life: dorsum dark brown with numerous dense yellowish gold irregular circles or lines, in some specimens forming a continuous foam-like reticulation pattern on dorsum; on head, dark brown blotch running from one to another eyelid, centrally connected to a line running to tip of snout forming a irregular T-shaped mark in some specimens; dorsal pattern fused with dorsolateral yellowish golden reticulation forming a band with numerous dark brown irregular spots (smaller than EYDM); laterally a dark brown band from tip of snout almost to groin, sometimes posteriorly interrupted by few yellowish golden lines; limbs and dorsal surfaces of hand and foot with yellowish golden foam-like reticulation with dark brown spots (smaller or larger than EYDM), thumb and sometimes tips of fingers and toes dorsally orange to bright red; in both sexes, throat, chest and venter white to cream, in some specimens with few scattered small brown dots (smaller than EYDM); soles, palms and a triangular blotch below cloaca and adjacent ventral surfaces of thighs bright red; iris black with a broad golden ring surrounding horizontally oriented pupil; (12) in preservative: above brown with cream pattern, below whitish; bright red life markings faded to yellowish white; (13) pulsed call 680 ± 30 ms at frequency range 3,450 –4,050 Hz  . </p>
            <p> Diagnosis.  Atelopus harlequin sp. nov. (Figs. 18–20) can be readily distinguished from all other  Atelopus (as far known) by molecular genetics, with an uncorrected p-distance of the 16S rRNA fragment of 2.37% to its closest known described relative,  A. colomai (Appendices 8, 9). It can be morphologically distinguished from all congeners by the combination of minute size, dorsal and lateral skin covered with dense, poorly distinct minute spiculae, ventral skin smooth to slightly areolate, presence of a columella and dorsal pattern. </p>
            <p> Atelopus harlequin sp. nov. (Figs. 18–20) is similar to its congeners of the flavescens-spumarius clade, to  A. andinus Rivero ,  A. loettersi De la Riva, Castroviejo-Fisher, Chaparro, Boistel and Padial ,  A. reticulatus Lötters, Haas, Schick and Böhme and the  A. tricolor Boulenger complex (including the junior synonyms  A. rugulosus Noble and  A. willimani Donoso-Barros ) from the eastern Andean versant of Peru and Bolivia, and to  A. minutulus Ruíz-Carranza, Hernández-Camacho and Ardila-Robayo from the eastern Andean versant of Colombia. It differs from  A. manauensis Jorge, Ferrão and Lima in dorsal pattern in life (dark brown, with numerous dense yellowish gold irregular circles or lines, fused with dorsolateral yellowish golden reticulation vs. few irregular circles, lines and blotches, not broadly fused with the foam-like reticulated dorsolateral band). It differs from the nominal  A. barbotini Lescure ,  A. flavescens Duméril and Bibron (including its junior synonym  A. vermiculatus McDiarmid ),  A. franciscus Lescure and the  A. hoogmoedi Lescure complex (including the available name  A. hoogmoedi nassaui Ouboter and Jairam ) in pattern and coloration in life (dorsally dark brown with numerous dense yellowish gold irregular circles or lines, sometimes forming a continuous foam-like reticulation pattern on dorsum; dorsolateral yellowish golden reticulation forming a band with numerous dark brown irregular spots; below white to cream, sometimes with few scattered small brown dots with a triangular red blotch below cloaca and adjacent ventral surfaces of thighs vs. dorsally black with red to pink markings, ventrally reddish pink in  A. barbotini ; dorsally unicolored reddish, brownish or yellow or brown and with olive, brownish reddish and copper-like vermiculation, ventrally reddish pink in males, pink in females in  A. flavescens ; dorsally olive to blackish brown, ventrally reddish in  A. franciscus ; dorsally dark brown or black with yellow, pink, bluish, white, greenish or orange irregular spots and reticulated dorsolateral bands or solid bars, ventrally yellow, pink, bluish, white, greenish or orange, in some specimens with dark spots or blotches in the  A. hoogmoedi complex). All these forms, except some populations of the  A. hoogmoedi complex, lack red palmar and plantar surfaces in life (present in  A. harlequin sp. nov. ). Further, with 680 ± 30 ms  A. harlequin sp. nov. has a shorter call than  A. barbotini (1,490 ± 140 ms),  A. flavescens (1,530 ± 220 ms),  A. franciscus (1,490 ±150 ms), the  A. hoogmoedi complex from French Guiana (1,190 ± 10 ms), in the  A. hoogmoedi complex from Amapá, Brazil (1,160 ± 390 ms) and the  A. hoogmoedi complex from Pará, Brazil (1,060 –1,240 ms) (Lescure 1981; Cocroft et al. 1990; Costa-Campos &amp; Carvalho 2018). In addition, with its small SVL (20.6 ± 0.51 in adult males, N = 4), the new species is readily distinguished from the larger  A. barbotini (26.5 ± 0.9, N = 10 from the type locality) and the  A. hoogmoedi complex (25.4 ± 1.4, N = 8, from the type locality).  Atelopus harlequin sp. nov. can be distinguished from  A. spumarius Cope sensu stricto ,  A. colomai Plewnia, Terán-Valdez, Culebras, Boistel, Paluh, Quezada Riera, Heine, Reyes-Puig, Salazar-Valenzuela, Guayasamin and Lötters and  A. histrionicus sp. nov. by call duration (680 ± 30 ms vs. 922 ± 24 ms in  A. spumarius sensu stricto ; 2,049 ± 220 ms in  A. colomai ; 807 ± 80 ms in  A. histrionicus sp. nov. ; Asquith &amp; Altig 1987). Further, the new taxon differs from these species in dorsal pattern (reticulation pattern not densely developed on the dorsum and not broadly fused with dorsolateral bands in  A. spumarius sensu stricto ,  A. colomai and  A. histrionicus sp. nov. ), female ventral pattern (absence in the new species vs. presence of dots) and male SVL (20.6 ± 0.51, N = 4, vs. 22.2 and 23.4, N = 2, in  A. spumarius sensu stricto ; 23.2 ± 1.46, N = 6, in  A. colomai ; 25.0 ± 0.67, N = 7, in  A. histrionicus sp. nov. ). The new species can be distinguished from  A. pulcher Boulenger by shorter calls (680 ± 30 ms vs. 1,200 ± 100 ms; Lötters et al. 2002a).  Atelopus harlequin sp. nov. further differs from  A. pulcher by smaller male SVL (20.6 ± 0.51, N = 4, vs. 27.27 ± 1.07, N = 13; Lötters et al. 2002a) and male coloration in life (dorsally dark brown with numerous dense yellowish gold irregular circles or lines, sometimes forming a continuous foam-like reticulation pattern on dorsum; dorsolateral yellowish golden reticulation forming a band with numerous dark brown irregular spots vs. dark brown dorsum with green spots, circles and blotches and a green broad dorsolateral band, rarely interrupted by small dark brown dots; in females, bright red venter with dark brown spots and a dark brown line on chest in some specimens). The new species differs from  A. seminiferus Cope in smaller SVL (single female 25.7 vs 40.0), the absence of lateral warts (present in  A. seminiferus ) and dorsal coloration (dorsally dark brown with numerous dense yellowish gold irregular circles or lines, sometimes forming a continuous foam-like reticulation pattern on dorsum; dorsolateral yellowish golden reticulation forming a band with numerous dark brown irregular spots vs. dark brown without pattern in  A. seminiferus ). From  A. andinus ,  A. loettersi ,  A. reticulatus and the  A. tricolor complex,  A. harlequin sp. nov. can be distinguished by the presence of a columella (vs. absence; Lötters et al. 2011; Authors’ unpubl. data). From  A. loettersi , it can be distinguished by female ventral coloration and pattern in life (below white to cream, sometimes with few scattered small brown dots with a triangular red blotch below cloaca and adjacent ventral surfaces of thighs vs. mostly red, in some specimens with irregular brown blotches; De la Riva et al. 2011). Further, it can be distinguished from  A. andinus ,  A. reticulatus and the  A. tricolor complex by the absence of warts (presence), by presence of dorsal reticulation (absence), except  A. reticulatus (which shows dark reticulation on a yellow dorsum) and by smaller male SVL (20.6 ± 0.51, N = 4 vs. 26.4 and 27.5, N = 2, in  A. andinus ; 24.7, N = 1, in  A. reticulatus ; 20.8 ± 0.86, N = 17, in the  A. tricolor complex). Based on its small size,  A. harlequin sp. nov. resembles  A. minutulus from which it can be distinguished by the absence of spiculae on the flanks (vs. present in  A. minutulus ) and by coloration and pattern in life (dorsally dark brown with numerous dense yellowish gold irregular circles or lines, sometimes forming a continuous foam-like reticulation pattern on dorsum; dorsolateral yellowish golden reticulation forming a band with numerous dark brown irregular spots vs. numerous green blotches on a brown dorsum and flanks). </p>
            <p>Description of holotype. Adult male. Body slender (SW/SVL = 0.245), head about as long as wide (HLSQ/ HDWD = 1.081), snout protruding beyond apex of lower jaw, acuminate in dorsal view; nostrils directed laterally, slightly protuberant, situated above apex of lower jaw at one fourth distance from tip of snout to eye; canthus rostralis distinct, slightly concave in dorsal view between eye and nostril; loreal region concave; lips not flared; top of snout depressed; head slightly concave in lateral view; interorbital region and occiput flat; eyelid flared; tympanic membrane absent, tympanic annulus not visible; supratympanic crest poorly developed; choanae medium-sized, rounded, widely separated; ostia pharyngea present; tongue three times as long as wide, broadest anteriorly, free for half its length posteriorly; vocal slits present; limbs long and slender (TIBL/SVL = 0.481); webbing absent on hand; fingers and toes lack lateral fringes; palmar tubercle rounded, almost indistinct; supernumerary palmar and plantar tubercles indistinct; thenar and subarticular tubercles indistinct; tips of digits rounded, except on third finger possessing acuminate tip; phalangeal formula of hand 1-2-3-3; first finger short (THBL/HAND = 0.432), dorsally bearing keratinized nuptial pad; inner metatarsal tubercle almost indistinct; outer metatarsal tubercle rounded, poorly developed; relative length of toes I&lt;II&lt;III&lt;V&lt;IV; phalangeal formula of foot 2-2-3-4-3, webbing formula of toes I 0–0 II 1 / 2 –1 III 11 / 2 –3 IV 3–1 V; tarsal fold absent; foot roughly two thirds of tibia; skin smooth on entire body, dorsally and laterally covered with dense, poorly distinct minute spiculae; skin weakly areolate on ventral surfaces of cloaca and adjacent thighs, venter and throat; cloacal opening in an inconspicuous tube, directed posteriorly; vertebral column visible through the skin, neural processes absent.</p>
            <p>In life, dorsum dark brown with dense irregular golden yellow circles and foam like reticulation; dark brown marks (larger than EYDM) and bordered by smaller ones (smaller than EYDM); on head, dark brown blotch running from one to another eyelid, centrally connected to a bar running from tip of snout to posterior corner of head; golden yellow becoming pale toward flanks; dorsolaterally pale-yellow foam-like reticulation, forming a band from above eye to groin with diffuse borders; upper lips and ventrolateral region pale yellow with small brown spots (smaller than or equal to diameter of finger III); laterally a dark brown band from tip of snout to groin; throat and venter whitish cream; limbs dorsally dark brown, covered in golden yellow reticulation, ventrally lower arm and lower leg white with brown blotches, upper arm white, thigh white, towards cloaca bright red; thumb and tips of fingers and toes dorsally orange to red; palms, plants and a triangular blotch on ventral surface of cloaca and adjacent areas of thighs bright red; iris black with a broad yellowish-golden ring around pupil.</p>
            <p>In preservative, above brown with cream pattern, below whitish; bright red markings (in life) yellowish white.</p>
            <p>Measurements: SVL 20.8, TIBL 10.0, FOOT 6.5, HLSQ 6.7, IOD 2.4, HDWD 6.2, EYDM 2.3, EYNO 1.7, ITNA 2.0, FAL 6.3, HAND 4.4, THBL 1.9, SW 5.1.</p>
            <p>Variation. For meristic variation of characters see Table 5. The paratypes (Fig. 19) correspond to the holotype description. Variation exists in the extent of a brown mark on the head which varies from several discontinuous large blotches to a band from eye to eye to a T-shaped mark from tip of snout to a line between the eyes. Sometimes, the lateral band is interrupted or replaced by separated brown spots, particularly posteriorly.</p>
            <p> Sexual dimorphism is apparent in  A. harlequin sp. nov. with the female being larger and lacking vocal slits and nuptial pads. Further, eggs are visible through the ventral skin in the female. </p>
            <p> Skull osteology. General osteological features of the holotype (MUBI 2448) are depicted in Figures 5–6 and Appendix 4. The skull of  A. harlequin sp. nov. (MUBI 2448, MorphoSource media 000618606) is triangular in dorsal view. The skull length is 6.6, the skull width is 5.6, and the skull height is 4.2. The skull roof is moderately rugose. In anterior view, the septomaxilla is U-shaped with medial and lateral rami that extend posteriorly towards the vomer. The lateral ramus is much broader than the medial ramus and bears a nasal process. The ossified sphenethmoid is smooth and underlies the posterior and medial margins of nasals, as well as the anterior and anterolateral margins of the frontoparietals. The posterior limit of the sphenethmoid is about one-third the length of the margin of the orbit. The prootics are slightly rugose and are fused to posterolateral edge of the frontoparietals. The exoccipitals encircle the foramen magnum and are fused to the frontoparietals dorsally, the prootics laterally, and the parasphenoid anteroventrally. The otic capsule is well-ossified. Columellae are present and medially bifurcated. The pars media of the columella is slender and slightly bowed, and the pars interna is broad. The operculum in the fenestra ovalis is not mineralized and therefore not visible in the microCT dataset. The nasals are triangular and bear an acuminate maxillary process that extends ventrolaterally toward the maxilla and nearly contacts the pars facialis of the maxilla. The nasals are widely separated medially. The frontoparietals are coarsely rectangular and the orbital edges are straight. Medial articulation is incomplete with the anterior half of the bones lacking articulation and the posterior half being completely fused with no suture line visible. A supraorbital flange is absent. Occipital groves, which are canals for the carotid artery, are present and nearly entirely open (not roofed with bone), except for the anterior tip. Posteriorly, the frontoparietals are fused to the prootics and exoccipitals. The vomers are small, edentate, medially separated, crescent-shaped, and triradiate with an anterior ramus, prechoanal ramus, and postchoanal ramus. The anterior ramus is directed anterolaterally, rounded at the anterior tip, and is larger than the other rami (nearly reaching the junction of the maxilla and premaxilla). The pre- and postchoanal rami are short, of nearly equal length, pointed, and form the anteromedial margin of the choana. The prechoanal ramus is directed towards the maxilla, and the postchoanal ramus is directed parallel to the midline. The postchoanal ramus of the right vomer is fused to the overlying sphenethmoid, while the two elements nearly contact on the left. The neopalatine is elongate and triangular. It underlies, and is partially fused to, the sphenethmoid and extends towards the maxilla, making contact with the preorbital process. The neopalatine is narrow and pointed at its medial tip, widens as it approaches the maxilla, and is cylindrical and rounded at its anterior border. The parasphenoid has an inverted T-shape and the cultriform process slightly underlies the sphenethmoid. The cultriform process extends beyond the midpoint of the orbit. The terminal end of the cultriform process is irregular and biradiated in shape. The parasphenoid alae are directed posterolateral to the cultriform process and are fused to the overlying otic capsule. The posterior margin of the parasphenoid and a medial posterior process are difficult to discern. The maxillary arcade is complete and edentate. The paired premaxillae each possess a dorsal alary process that is directed anterolaterally. The lower half of the alary process is slightly wider than the upper half. The pars palatina is biradiate with medial and lateral processes. The lateral process is more than twice as long as the medial process. The medial process is well developed and triangular. A concave border is present where the lateral and medial processes of the pars palatina come together. The posterior tip of the pars dentalis is pointed and articulates with the maxilla. The external surface of the maxillae is rugose. The pars palatina, which extend along the lingual margin of the maxilla, is narrow, and the posterior half of this shelf articulates with the anterior ramus of the pterygoid. The pars fascialis of the maxilla is directed medially and is a jagged, irregular shape; its anterior margin is biradiate; its dorsomedial margin contacts the neopalatine and sphenethmoid and nearly contacts the maxillary process of the nasal. The anterior end of the maxilla is truncate and slightly overlaps the premaxilla. The posterior end of the maxilla is pointed and contacts the quadratojugal. The quadratojugal is a small, slender, L-shaped bone that underlies the ventral arm of the squamosal and contacts the posterior process of the maxilla at is anterior margin. The paired squamosals possess an otic ramus posterodorsally, a zygomatic ramus anterodorsally, and a ventral ramus. The otic ramus is expanded dorsally and has a broad articulation with the crista parotica of the otic capsule, leaving only the posterior end of the prootic free. The zygomatic ramus is small and triangular. The angle between the dorsal surface of the squamosal and the anterior margin of the ventral ramus is nearly perpendicular. The ventral ramus is flat and blade-like. A squamosal keel is present, extending along the outer surface from the zygomatic ramus to the upper half of the ventral arm. The pterygoid is triradiate, bearing anterior, medial, and posterior rami. The anterior ramus articulates with the pars palatina of the maxilla. The medial and posterior rami are of equal length. The medial ramus nearly contacts the prootic. The posterior ramus is flat and blade-like. The palatoquadrate is cartilaginous and therefore not visible in the microCT dataset. The lower jaw is composed of three ossified elements and Meckel’s cartilage, which is not visible in the microCT dataset. The mentomeckelian is the most anterior element that forms a cartilaginous symphysis at the midline of the jaws. Posterodorsally, the mentomeckelian is fused to the dentary. The dentary is slender, thin, and edentate; this element overlays the anterior half of the angulosplenial. The angulosplenial is the largest mandible element and forms the lingual surface of the lower jaws. The lower jaw length is 5.5, nearly the same length as the skull from the occipital condyle to the premaxilla. </p>
            <p> Atelopus harlequin sp. nov. differs from the other species (re)described in this paper by having the medial side of the posterodorsal process of the squamosal being attached along its entire length to the prootic (vs. the distal end of the process remaining free; Fig. 6), the frontoparietals being dorsally more convex, and the separation of the occipital condyles being greater; Fig. 6). </p>
            <p> Further, the new species can be distinguished from the neotype of  A. spumarius sensu stricto by the squamosal laterally extending beyond the posterolateral limit of the maxilla and the jugularis process of the quadratojugal thus contacting the posterior process of the maxilla anteriorly (vs. the posterior end of the maxilla being free, laterally protruding beyond the lateral limit of the squamosal), the pars glenoidalis of the quadratojugal being triangular with an enlarged but narrow pars jugularis (vs. pars glenoidalis thickened, pars jugularis short) and a less straight orbital margin of the maxilla. However, variation in these characters exists between the two populations of  A. spumarius sensu stricto , which might be attributed to imaging conditions or natural variation.  Atelopus harlequin sp. nov. differs from  A. colomai by its slightly rugose skull roof (vs. smooth), in having the frontoparietals fused only over two posterior thirds of their total length (vs. completely fused), the squamosal laterally extending beyond the posterolateral limit of the maxilla, and the jugularis process of the quadratojugal contacting the posterior process of the maxilla anteriorly (vs. maxilla protruding beyond the lateral limit of the squamosal, posteriorly, contacting the jugularis process of the quadratojugal) and the pars glenoidalis of the quadratojugal being triangular with an enlarged but narrow pars jugularis (vs. pars glenoidalis subrectangular, pars jugularis elongated).  Atelopus harlequin sp. nov. is distinguished from  A. histrionicus sp. nov. by a less rugose skull roof. On the lateral margins corresponding to the maxilla, a slight curvature is visible on the posterior third in  A. harlequin sp. nov. (vs. lateral margins straight), its posteroventral incisure of the sphenethmoid is less deep, the parasphenoid is more developed, and the dorsal end of the dorsal process of the premaxilla (= pars facialis) remains at the same level (vs. protrudes anteriorly beyond the anterior limit of the nasals). For details see Figures 5–6 and Appendix 4. </p>
            <p>However, the discriminatory power of osteological features needs to be seen with caution as intraspecific variation, sexual dimorphism and taxonomic value remain poorly understood for the species discussed.</p>
            <p> Vocalization. Asquith &amp; Altig (1987) described pulsed calls under the name  A. spumarius from a population close to Nauta (Peru, Loreto) which we assign to  A. harlequin sp. nov. due to its proximity to its type locality and on a color photo of the recorded specimen provided to us by the authors. Their description is based on five calls (recorded in a collection bag at 26° C) with an average call duration of 680 ± 30 ms (670–710 ms). Intervals between calls are described as 2,200 ± 300 ms (5,400 –8,000 ms), where, however, either mean ± SD or range is given in error. Frequency at the beginning of the call is 3,578 ± 98 Hz (3,450 –3,690 Hz) and at the end of the call 3,900 ± 100 Hz (3,770 –4,050 Hz). Each call consists of 23.4 ± 2.4 pulses (20–27), at a pulse rate of 23.4 ± 2.2 (27.5–30.6) at the beginning of the call and 61.3 ± 7.9 (52.2–71.4) at its end. For details see Asquith &amp; Altig (1987). </p>
            <p> In addition, vocalizations were recorded by BR-RR from an uncollected male during handling on 1 February 2020, ca. 01:00 pm, close to the type locality on the Iquitos-Nauta road, with an iPhone 6s. Distance to recorder was 0.05 m, the ambient temperature is unknown. A 16,462 ms recording was available for analysis (CJ ec.cj.aud.18; Fonozoo Sound Code 14650). It consists of 12 pure tone short calls and 9 pulsed short calls (Fig. 21). Pure tone short calls have duration of 16 ± 4 ms (12–25 ms) and a dominant frequency of 3,270 ± 244 Hz (2,724 –3,553 Hz). Frequency shows harmonics and is modulated with a decrease towards the end. Pulsed short calls have call duration of 44 ± 8 ms (26–55 ms) and a dominant frequency of 3,495 ± 77 Hz (3,388 –3,612 Hz). Amplitude peaks with the first pulse and descends over the course of the call. Frequency shows harmonics and is modulated in the last pulse with a decrease towards the end. Pulsed short calls (Fig. 22) consist of 3 ± 1 (2–5) pulses that show a length of 8 ± 7 ms (2–24 ms) each. The last pulse is the longest in all pulsed short calls . </p>
            <p> Tadpole. The tadpole of  A. harlequin sp. nov. is unknown. </p>
            <p> Distribution. In Peru,  A. harlequin sp. nov. is known from various localities between the Río Napo (to its right) and the Río Marañon (to its left) and west of the Amazon River at ca. 100–150 m asl (Fig. 2). As far known, the geographic range is delimited by the latter two rivers, while the distributional limits to the North remain difficult to assess. In Ecuador, this species is known only from Lorocachi in Pastaza Province, situated in the upper Amazon basin at ca. 220 m asl, in the Río Pastaza drainage. It is unknown if the area between Lorocachi and the N and W banks of the Marañon and Amazon Rivers is continuously colonized as the area remains poorly studied and here  A. harlequin sp. nov. is known only from two localities in between (Pucacuro and Andoas). </p>
            <p>Natural history. The species inhabits terra firme lowland rain forest and can be found in close proximity to small streams (Fig. 23). The holotype was collected during the day in disturbed forest in the leaf litter. We observed this diurnal species in proximity to slow-moving streams with sandy substrate that were approximately 1–3 m wide and 30 cm deep. During the day, adult males are mostly found actively walking on the forest floor, close to the stream.</p>
            <p> Conservation status. We suggest listing  A. harlequin sp. nov. as Data Deficient as its population status and its distributional range remain little understood. The species’ EOO is 38,420 km ² and AOO is 40 km ². While most parts of the species’ geographic range are covered in undisturbed primary forest with few human settlements, the populations on the easternmost edge of the species range along the road from Iquitos to Nauta (including its type locality) are highly threatened by habitat loss through large-scale deforestation, agriculture and the construction of roads. Three populations along the Iquitos-Nauta road of the species persisted into the 2010s, but could not be confirmed in 2022 with almost no suitable habitat remaining. In a population a few kilometers further westwards, the species remained abundant despite ongoing deforestation in close proximity to the stream (Fig. 24). Most other populations have not been visited in years hampering a thorough assessment of the species’ conservation status. </p>
            <p>Presence and potential impact of Bd remain unknown.</p>
            <p> Atelopus harlequin sp. nov. is further likely to be threatened by climate change in the near future, which might both impact the species habitat as well as interfer and exacerbate existing threats such as disease (Lötters et al. 2023). The species from the western Amazon are among the harlequin frogs likely to be most affected by climate change (Lötters et al. 2023). </p>
            <p> Etymology. The specific name is a noun in apposition (nominative case, singular) and refers to the widely used common name  “harlequin ” for frogs of the genus  Atelopus that possess striking color patterns and occasionally make awkwarded moves. </p>
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	https://treatment.plazi.org/id/03E48797FF8BFFF040817AEDFE21FE56	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.		MagnoliaPress via Plazi	Lötters, Stefan;Böning, Philipp;Bailon, Salvador;Castañeda, Jose Daniel Barros;Boistel, Renaud;Catenazzi, Alessandro;Chaparro, Juan C.;Chávez, Germán;Chujutalli, Angel;Coen, Laurent;Coloma, Luis A.;Crawford, Andrew J.;Culebras, Jaime;Martínez, Juan Carlos Cusi;Daza, Juan Manuel;Riva, Ignacio De La;Ellwein, Denise J.;Ernst, Raffael;Flechas, Sandra V.;Fouquet, Antoine;Guayasamin, Juan Manuel;Heine, Christopher;Jorge, Rafael F.;Jung, Alisha;Jungfer, Karl-Heinz;Kaffenberger, Nathalie;Krehenwinkel, Henrik;Marca, Enrique La;Lampo, Margarita;Medina Rangel, Guido F.;Orsen, Ludvig;Paluh, Daniel J.;Gonzalez, José Luis Pérez;Perrin, Jonathan;Quezada Riera, Amanda B.;Reyes-Puig, Juan Pablo;Ross, Bernardo Roca-Rey;C, Daniela;Solano, Luis Alberto Rueda;Salazar-Valenzuela, David;Vazquez, Josefa Celsa Señaris;Sowinski, Morgane;Terán-Valdez, Andrea;Tovar-Ortiz, Angie;Veith, Michael;Venegas, Pablo;May, Rudolf Von;Weitkamp, Timm;Plewnia, Amadeus	Lötters, Stefan, Böning, Philipp, Bailon, Salvador, Castañeda, Jose Daniel Barros, Boistel, Renaud, Catenazzi, Alessandro, Chaparro, Juan C., Chávez, Germán, Chujutalli, Angel, Coen, Laurent, Coloma, Luis A., Crawford, Andrew J., Culebras, Jaime, Martínez, Juan Carlos Cusi, Daza, Juan Manuel, Riva, Ignacio De La, Ellwein, Denise J., Ernst, Raffael, Flechas, Sandra V., Fouquet, Antoine, Guayasamin, Juan Manuel, Heine, Christopher, Jorge, Rafael F., Jung, Alisha, Jungfer, Karl-Heinz, Kaffenberger, Nathalie, Krehenwinkel, Henrik, Marca, Enrique La, Lampo, Margarita, Medina Rangel, Guido F., Orsen, Ludvig, Paluh, Daniel J., Gonzalez, José Luis Pérez, Perrin, Jonathan, Quezada Riera, Amanda B., Reyes-Puig, Juan Pablo, Ross, Bernardo Roca-Rey, C, Daniela, Solano, Luis Alberto Rueda, Salazar-Valenzuela, David, Vazquez, Josefa Celsa Señaris, Sowinski, Morgane, Terán-Valdez, Andrea, Tovar-Ortiz, Angie, Veith, Michael, Venegas, Pablo, May, Rudolf Von, Weitkamp, Timm, Plewnia, Amadeus (2025): A roadmap for harlequin frog systematics, with a partial revision of Amazonian species related to Atelopus spumarius. Zootaxa 5571 (1): 1-76, DOI: 10.11646/zootaxa.5571.1.1, URL: https://doi.org/10.11646/zootaxa.5571.1.1
