taxonID	type	description	language	source
74A9ACBE01145958AA398C67EFC6E210.taxon	description	Figs 1, 2, 3	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
74A9ACBE01145958AA398C67EFC6E210.taxon	diagnosis	Diagnosis. Brachypter. The salient features of this new species include the following: dorsally overall dull dark brown color, with pale mottles on apex vertex, front disc, and a pale yellow stripe on frontoclypeal suture extending towards the base of gena. Body broadly depressed and distinctively wide at abdomen. Vertex broad, subquadrate, apical margin broadly rounded, with submedian carina forking dorsally near anterior margin of eyes, carinal branches diverging widely to meet anteriorly just below fastigium which is angled when viewed laterally. Eyes reduced, reddish, slightly emarginate below, barely visible in ventral view. Frons subcircular, short, about as long as wide, with lateral carinae bowed outward, converging both ventrally and dorsally; metatibial spur short and narrow, bearing eight or nine black-tipped sharp teeth on trailing margin. Male terminalia: pygofer short, with small sized outer lobes, inner lobes subtriangular in outline, with broad concavity between them; aedeagus short, bearing dorso-apical horseshoe-like process; anal segments short and wide, unarmed.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
74A9ACBE01145958AA398C67EFC6E210.taxon	description	Description. Brachypterous male (Figs 1 A, B, D, E, 2 A – E). Color (Fig. 1) dull dark brown, with some pale marks. Reddish eyes. Vertex pale along posterior margin, median and Y-shaped carinae, with small yellowish spots on apex between submedian and lateral carina and also between lateral carina and eyes. Frons with continuous row of ~ 10 symmetrical pale dots paralleling median and lateral carinae, and transversal whitish stripe on frontoclypeal suture extending towards the base of gena. Clypeus castaneous on disc, rostrum yellowish. Antennal segments castaneous, slightly pale on anterior surface. Pronotum with pale transversal row of small spots on anterior margin between lateral carinae and several smaller spots on central disc near posterior margin; mesonotum disc with pale longitudinal spots between lateral carinae and some small spots on posterior margin. Tegmen uniformly pale brown. Legs yellowish, darker on base and apex of pro and mesocoxae, annular dark brown stripes near the base and apex of pro- and mesotibiae, apex of pro- and mesofemur, dorsal metafemur, and two annular stripes on metatibiae, one near base and the other on base of apical spines, on dorsal surface of spurs, and base of first tarsomere and apex of third. Abdomen dark brown in dorsal view, with longitudinal bilateral narrow pale stripes on tergites III – VII and ventrally on posterior margins of sternites IV – VII, and laterally around wax pores; anal segment paler, darker on apical margin as well as on anal style. Structure. Body strongly dorsoventrally flattened, suboval in outline. Head narrower than pronotum. Vertex in dorsal view almost as long as wide, rather quadrate, broadly rounded on anterior margin; basal compartment occupying approximately more than basal half of vertex. Median carina present, forked near anterior margin of eyes, arms of fork diverging strongly (angle 170 °) to meet submedian frontal carinae. Submedian frontal carinae arising from the lightly foliate lateral carinae at level of middle of eyes, meeting anteriorly just at the fastigium (Fig. 1 A). In lateral view, head projected downwards in front of the eye, fastigium angled (Fig. 1 E). Frons subcircular, about as long as wide, and as long as clypeus; carinae of frons distinct, evanescent toward apex, lateral carinae bowed outward, converging both ventrally and dorsally; frons widest at antenna level. Frontoclypeal suture ventrally curved. Clypeus sub-triangular with carinae evident, the laterals continuing with genal carina. Rostrum reaching metacoxae, slightly shorter than frons plus clypeus, subapical segment longer than the apical one (1.3: 1). Compound eyes, very reduced, lower margin only slightly incised, barely visible in ventral view. Antennae short, first segment as long as wide, second segment 2 × the first, 2 × longer than wide (Fig. 1 A, B). Pronotum with conspicuous carinae, the laterals divergent, reaching hind margin and ending slightly convex. Mesonotal disc almost as long as pronotum (1.2: 1), carinae conspicuous, lateral ones slightly divergent apically reaching hind margin (Fig. 1 A). Tegmen coriaceous, subquadrate, posterior margin subtruncated to slightly rounded, reaching 4 th segment; veins distinct (Fig. 1 A). Metatibial spur leaf-like, short, narrow apically and concave ventrally, bearing eight or nine black-tipped sharp teeth on trailing margin, almost as long as first segment of metatarsi at notch; first hind tarsomere longer than second plus third (1.7: 1) (Fig. 1 D). Abdomen broadest across segment V, decreasing in width towards apex (Fig. 1 A). Terminalia. Pygofer trapezoidal, with laterodorsal margin slightly truncate, not projected caudad; ventrally ~ 2 × longer than dorsally; dorsally with shallow concave anal margination (Fig. 2 A, B); outer lobes small-sized and rounded, slightly enfolding lateral area of pygofer, in ventral view occupying half the length of pygofer, inner lobes subtriangular in outline, broadly concave between them and with narrow notch between inner and outer lobes, partially closing ventral foramen (Fig. 2 A, B); diaphragm fairly long, regularly narrowed toward middle line and caudally produced in short conical process. Aedeagus short, length-wide ratio: 2.2: 1, regularly tubular and caudally downwardly directed, bearing an apical process projected on dorsal surface to left of phallotreme; this process is forked near base in two long and divergent semi-circularly curved spines (horse-shoe outline), in lateral view extending beyond genital styles (in repose); phallotreme large, near apex on dorsal surface to the right (Fig. 2 D). Suspensorium sclerotized, strap-like, very short, half of aedeagus length (Fig. 2 D). Genital styles (parameres) straight and flattened, widest and divergent in apical 1 / 3, apices truncate and enlarged, with sharp conical process on inner angle, widely expanded and rounded on outer angle, apex reaching dorsal margin of diaphragm at rest (Fig. 2 E). Anal segment broad, collar-like, closely embraced by pygofer, without processes (Fig. 2 C); anal style short (Fig. 2 C). Measurements (n = 3). L., 2.6; b. w., 0.75; M. b. w. at abdominal segment V, 1; t. l., 0.4; v. l., 0.4; v. w., 0.5; f. l., 0.4; M. f. w., 0.4; m. f. w., 0.25; a. l. I, 0.15; a. l. II, 0.2; p. l., 0.3; m. l., 0.4; mti. l., 0.8; mta. l., 0.7; mta. Il., 0.4; s. l., 0.4; t. n., 9. Macropter unknown. Brachypterous female (Figs 1 C, F, 2 F – H). Color. Body coloration pattern and structure similar to male; ovipositor brown with valvifer pale on inner margin; gonapophysis rather pale apically. Terminalia. Ovipositor short, reaching anal segment at base (Fig. 2 F). Valvifer VIII regularly broad, slightly excavated on inner margin near base, with inconspicuous basal projection; separate in repose in ventral view. Gonapophysis VIII wide at base, ventrally projected between valvifers. Gonapophysis IX slightly curved, bearing numerous strong rounded teeth (~ 35) on dorsal margin on> 1 / 2 of its length, a few teeth smaller distally; with three or four ventral teeth (Fig. 2 G, H). Measurements (n = 3). L., 3; b. w., 0.83; M. b. w. at abdominal segment V, 1.265; t. l., 0.4; v. l., 0.35; v. w., 0.5; f. l., 0.4; M. f. w., 0.4; m. f. w., 0.25; a. l. I, 0.15; a. l. II, 0.2; p. l., 0.3; m. l., 0.4; mti. l., 0.85; mta. l, 0.75; mta. Il., 0.5; s. l., 0.4; t. n., 9 – 10. Macropter unknown.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
74A9ACBE01145958AA398C67EFC6E210.taxon	etymology	Etymology. The specific name comes from the Greek letter delta (Δ), which was used to refer to the triangle of fertile land that the Nile forms at its mouth (Nile Delta) and by extension, to other river deltas. In this case, the name refers to the geographical distribution of the species, which is restricted to the region of the Paraná River Delta.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
74A9ACBE01145958AA398C67EFC6E210.taxon	distribution	Distribution. Argentina: Buenos Aires Province (Fig. 3 A, Suppl. material 2).	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
59E2FEDF9DAD51B79D8B3A8797E51088.taxon	distribution	Distribution. Argentina: Formosa, Chaco, and Corrientes provinces (Sosa et al. 2007 b). BRAZIL: Pará and Mato Grosso do Sul States (Muir, 1926). New record: Bolivia.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
AC267F4AFD4453828525F4647ADC8549.taxon	description	Figs 4, 5, 6	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
AC267F4AFD4453828525F4647ADC8549.taxon	diagnosis	Diagnosis. Macropter and brachypter. The salient features of the new species include the following: body mostly dark brown with distinctive yellowish to white marks bordering most of the sclerites of the body with legs paler and lightly marked with dark pigment. Macropters with forewings amber with pale brown veins, with strong dark marks on clavus, along Cu vein, over cross veins, and on last apical cells; brachypter with tegmen amber, brownish transverse marks in middle and claval apex, male pygofer dark brown, with dorsal surface, anal angles, and anal segment pale brown. Vertex narrow, with submedian carina forking dorsally quite far from fastigium; carinal branches closely forming a slender triangle; fastigium rounded in lateral view. Frons long, median carina prominent at or just below fastigium then fine; lateral carinae at base foliated, all carinae fine before the well-defined frontoclypeal suture. Spur large and wide, with 17 or 18 sharp, black-tipped teeth on trailing margin. Male terminalia: pygofer with relatively large lobes, the inner sharpened at apex with rounded external margin and internal sinuous; aedeagus short and tubular, with a short, slender dorso-caudally process curved at apex. Anal segment with two long, sinuous, slender caudally directed processes projecting laterally from the base.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
AC267F4AFD4453828525F4647ADC8549.taxon	description	Description. Macropterous male (Figs 4 A – C, G, 5 A – F). Color (Fig. 4 A – C) dark brown, with distinctive marginal castaneous and yellowish white marks on head, thorax, and abdomen. Head yellowish on posterior compartments of vertex and fovea, only infuscated in concavities, and on both sides of lateral and median carinae of face below fastigium; distinctive whitish marks below the eyes and across frontoclypeal area. Thorax, whitish colored on the pronotal disc between lateral carinae, one spot behind eyes on paranotal disc, a subtriangular shaped spot on each side of postero-lateral margin of mesonotum, metanotum, and scutellum; yellowish, on lateral edges of pronotum, tegula, and a suboval longitudinal median spot on mesonotum disc. Antennal segments pale castaneous with basal segment and proximal 1 / 2 of second segment darkish on dorsal surface. Legs yellow with tarsi dorsally darker, with longitudinal darker spots near base and apex of pro- and mesocoxae, on dorsal surface of femora, apical region of metafemur, and base and apex of metatibiae. Forewing amber, veins pale brown, fuscous along apical veins M 1 + 2 and M 3 + 4 with infuscate areas on central nodal line, last apical cell, along Cu vein, on postclaval angle, and apex of clavus. Abdomen in dorsal view with contrasting white yellowish coloration on drumming segments and two lateral spots on tergites V and VIII, ventrally with most of the segment margined with yellow. Pygofer dark brown on outer lobes and ventral surface; pale castaneous on dorsal surface and inner lobes. Anal segment yellowish and anal style dark brown (Fig. 4 A, B). Structure. Head narrower than pronotum. Vertex rectangular, longer than wide at base (2.1: 1) projecting beyond eyes> 1 / 3 of its length, with rounded frontal inflection; basal compartment slightly concave, occupying approximately more than basal third, stem of Y-shaped carina fine, delimiting shallow depressed areas on both sides; other carinae of head pronounced; submedian carinae forking at dorsal surface of vertex, quite far from fastigium, carinal branches forming a slender triangle, fovea, or areola little> 2 × the length; median carina strongly ridged and prominent at base at or just below fastigium then smooth; lateral carinae foliated at base, removed from eye along the length. (Fig. 4 A). Frons nearly 3 × longer than wide (3: 1.1), strongly narrowed between anterior margins of eyes, maximum width near basal 1 / 3, lateral margins slightly convex at apex; lateral and submedian carinae fine just before frontoclypeal suture, which is arched dorsally. Clypeus subtriangular, longer than wide, median carina weaker at base. Eyes globose, deeply emarginate below to receive the antennae. Rostrum reaching metacoxae, longer than frons, subapical and apical segment subequal. Antennae with the first segment longer than wide, the second segment <2 × the first, length> 2 × its width (Fig. 4 A, B). Pronotum with conspicuous carinae, laterals divergent, slightly convex toward hind margin, reaching it. Mesonotum almost as long as vertex plus pronotum, fine median carina becoming obsolete at apex, lateral carinae inconspicuous, slightly divergent posteriorly, not reaching hind margin (Fig. 4 A). Forewings rather long and slender, rounded at apex, length 3 × their width at subapical region, surpassing distal end of abdomen> ~ 1 / 3 of their length. Metatibial spur, leaf-like, long, and broad, with median rib becoming obsolete at apex which is truncated without teeth, slightly longer than metatarsomere I, with 17 or 18 regular, large, black-tipped teeth on trailing margin; first hind tarsomere longer than second plus third (1.5: 1) (Fig. 4 G). Abdomen regularly wide, compressed dorsoventrally (Fig. 4 A). Terminalia (Fig. 5 A – F). Pygofer in dorsal view, with deeply concave anal emargination, anal angles distinctly projected caudad. In ventral view, expanded, with large, round, kidney-shaped outer lobes enfolding almost the entire lateral surface (Fig. 5 A, B); inner lobes, large, wide in basal 2 / 3, external margin rounded, inner margin sinuous ending apically pointed with deep and broad concavity between their bases; strong emargination between inner and outer lobes (Fig. 5 A, B); diaphragm broad with dorsal margin deeply concave and medially caudad projected into lip-shaped process bearing bunch of rather short, stiff hairs. Aedeagus short, tubular, strongly narrow at base, widening up to the basal 1 / 3 then uniformly tubular and obliquely truncated at apex, with a single short, slender, apically curved process extending shortly beyond oval and apical phallotreme (Fig. 5 C, D). Genital styles (parameres) long, narrow, convergent apically, expanded and gradually tapering basally, broadly rounded along apical 1 / 2 on outer margin, apex hook-like, very curved inward; in ventral view visible between the internal lobes, in almost its entire length (Fig. 5 E). Suspensorium strap-like, connected to the aedeagal base, as long as half length of aedeagus (Fig. 5 C). Connective slightly compressed, almost straight (Fig. 5 C). Anal segment tubular, longer than twice the width, with caudal margin deeply emarginate ventrally; with two long, sinuous, slender, caudally directed processes arising ventrolaterally just below anterior angle which is membranous; anal style slender, twice longer than broad (Fig. 5 F). Measurements (n = 6). L., 3.8; B. L., 2.3; b. w., 1; t. l., 3.2; v. l., 0.4; v. w., 0.2; f. l., 0.6; M. f. w., 0.3; m. f. w., 0.2; a. l. I, 0.15; a. l. II, 0.3; p. l., 0.2; m. l., 0.5; mti. l., 1.13; mta. l., 1; mta. Il., 0.7; s. l., 0.8; t. n., 17 – 18. Brachypterous male. Color similar to macropterous form, with mesonotum paler and uniformly colored; tegmen amber with veins concolorous, with fuscous transverse marks continuous or fragmented, from base of clavus at near axillary region, and one spot at claval apex. Abdomen with similar patterns except tergites V and VI which are more uniformly dark brown contrasting with yellowish segments VII and VIII. Structure. Mesonotum shorter,> 1 / 2 of vertex plus pronotum length. Tegmen slightly longer than wide, rounded on external lateral margins; posterior margin truncate, covering tergite II. Measurements (n = 6). L., 2.3; b. w., 0.8; t. l., 1; v. l., 0.4; v. w., 0.18; f. l., 0.6; M. f. w., 0.25; m. f. w., 0.15; a. l. I, 0.15; a. l. II, 0.3; p. l., 0.2; m. l., 0.25; mti. l., 0.9; mta. l., 0.9; mta. Il., 0.6; s. l., 0.65; t. n., 17 – 18. Macropterous female (Figs 4 E, F, 5 G – I). Color: Head and thorax resemble male. Abdomen, in dorsal view, with contrasting yellowish marks on sides and posterior margins of sclerites, except V and VIII, which are uniformly brown. Abdominal sternites with yellowish margins; pygofer and anal segment yellowish, anal style castaneous; ovipositor dark brown (Fig. 4 E, F). Structure. Resembling male but abdomen is sharply tapered towards genital segments. Forewings surpassing distal end of abdomen ~ 1 / 6 of their length. Anal segment subrectangular; anal style slender. Terminalia (Fig. 5 G – I). Pygofer long, tubular-shaped, tapering toward the apical 1 / 2; in dorsal view exposed shortly beyond tergite VIII. Ovipositor long, strong, slightly sinuous in apical 1 / 2, as long as length of pygofer plus anal segment. Valvifer VIII regularly wide, inner margin rounded at base, separated in repose (Fig. 5 G). Gonapophysis IX, long and slender, overall shape slightly sinuous, apical fifth concave ventrally, with numerous blunt small teeth on most of dorsal surface (Fig. 5 H, I). Measurements (n = 10). L., 4.3; B. L., 3.2; b. w: 0.9; t. l., 3.5; v. l., 0.45; v. w., 0.2; f. l., 0.7; M. f. w., 0.3; m. f. w., 0.2; a. l. I, 0.2; a. l. II, 0.3; p. l., 0.2; m. l., 0.6; mti. l., 1.1; mta. l., 1; mta. Il., 0.7; s. l., 0.6; t. n., 21 – 23. Brachypterous female (Fig. 4 D). Color pattern similar to that of the macropterous female, but tegmina resemble those of brachypterous male. Measurements (n = 5). B. L., 2.9; b. w: 0.8; t. l., 1; v. l., 0.45; v. w., 0.2; f. l., 0.7; M. f. w., 0.3; m. f. w., 0.2; a. l. I, 0.2; a. l. II, 0.3; p. l., 0.2; m. l., 0.3; mti. l., 1; mta. l, 1; mta. Il., 0.6; s. l., 0.7; t. n., 21 – 23.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
AC267F4AFD4453828525F4647ADC8549.taxon	etymology	Etymology. The specific name comes from the Latin, serpentinum (serpentine), referring to the undulated shape of the long and slender male anal processes.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
AC267F4AFD4453828525F4647ADC8549.taxon	distribution	Distribution. Argentina: Misiones, Chaco, Corrientes, Santa Fe, and Buenos Aires provinces. Paraguay: Cordillera department (Fig. 6 A, Suppl. material 2).	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
15A12A3753CA5BD89876235915B05678.taxon	description	Fig. 8	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
15A12A3753CA5BD89876235915B05678.taxon	description	Description. Brachypterous male. Color (Fig. 8 A, B) pattern dark brown lighter on dorsal pygofer and ventrally, similar to macropterous male; tegmina, amber-colored infuscation along claval margin and longitudinal veins, with a white spot on external apical corner. Structure. Tegmen slightly longer than wide, rounded on external lateral and posterior margins, covering tergite II (Fig. 8 A). Measurements (n = 3). L., 2.12; b. w., 0.72; M. b. w. at abdominal segment V, 0.90; t. l., 0.74; v. l., 0.32; v. w., 0.18; f. l., 0.44; M. f. w., 0.25; m. f. w., 0.15; a. l. I, 0.13; a. l. II, 0.17; p. l., 0.19; m. l., 0.28; mti. l., 0.73; mta. l., 0.75; mta. Il., 0.46; s. l., 0.48; t. n., 20.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
15A12A3753CA5BD89876235915B05678.taxon	distribution	Distribution. Argentina: Corrientes province (Sosa et al. 2007 b). Guyana: Demerara River (Muir 1919). New record: Paraguay, Arroyos y Esteros Department.	en	Salinas, Nicolas A., Mariani, Roxana, Marino de Remes Lenicov, Ana M., Rodriguero, Marcela S., Sosa, Alejandro J. (2025): Phylogeny of the planthopper genus Megamelus (Hemiptera, Delphacidae), with the description of two new species from South America. ZooKeys 1224: 29-54, DOI: 10.3897/zookeys.1224.135596
