taxonID	type	description	language	source
0947107FB2315F538D7E9F781706A0C3.taxon	description	Fig. 13 A – C	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
0947107FB2315F538D7E9F781706A0C3.taxon	diagnosis	Diagnosis. Devario altus sp. nov. is distinguished from its Sri Lankan congeners by the combination of the following characters and character states: anal-fin origin posterior to dorsal-fin origin, beneath 3 rd branched dorsal-fin ray; 1 st infraorbital usually bearing a laterally-directed process; P-stripe continuous, not bifurcated anteriorly; interstripe I + 1 long, extending uninterrupted anterior to vertical through pelvic-fin origin; caudal peduncle long, 19.2 – 21.4 % SL, shallow, 8.8 – 10.5 % SL; body shallow, its depth 21.8 – 26.0 % SL; dorsal-fin height 19.0 – 23.2 % SL; dorsal-fin base length 15.1 – 18.2 % SL; anal-fin base length 16.4 – 20.5 % SL; pelvic fin height 12.9 – 16.6 % SL; pectoral-fin height 20.3 – 23.4 % SL; head depth 17.3 – 19.6 % SL; scales between lateral line and dorsal-fin origin 6 – 7 ½; circumpeduncular scales, 13 – 14; branched dorsal-fin rays 8 ½ - 10 ½. Specifically, D. altus sp. nov. differs from D. malabaricus by possessing (in 73 percent of specimens) a process on the 1 st infraorbital vs lacking (in 95 percent of D. malabaricus); having the P-stripe continuous, not bifurcated anteriorly (vs P-stripe usually bifurcated anteriorly); interstripe I + 1 long, extending uninterrupted anterior to vertical through pelvic-fin origin (vs short, not extending uninterrupted anterior to a vertical through the pelvic-fin origin in D. malabaricus); a caudal peduncle length of 19.2 – 21.4 % SL (vs 12.3 – 19.4 % SL); a body depth of 21.8 – 26.0 % SL (26.1 – 35.4 % SL); and scale rows between lateral line and dorsal-fin origin 6 – 7 ½ (vs 7 ½ - 9). The new species is distinguished from D. memorialis by having the anal-fin origin beneath the 3 rd branched dorsal-fin ray, distinctly posterior to dorsal-fin origin (vs on vertical through dorsal-fin in D. memorialis); the anal-fin base length 16.4 – 20.5 % SL (vs 20.0 – 23.4 % SL); a head depth of 17.3 – 19.6 % SL (vs 14.1 – 15.8 % SL); and 13 or 14 (vs 15 – 18) circumpeduncular scales. Devario altus sp. nov. differs from D. micronema by having the I + 1 interstripe long, extending uninterrupted anterior to a vertical through the pelvic-fin origin (vs short, not extending uninterrupted anterior to a vertical through the pelvic-fin origin in D. micronema); having the P-stripe continuous, uninterrupted anteriorly (vs P-stripe usually interrupted anterior to pelvic-fin origin with bars or blotches); having a longer caudal peduncle, 19.2 – 21.4 % SL (13.2 – 20.0 % SL); having a body depth of 21.8 – 26.0 % SL (vs 23.5 – 36.6 % SL); and having 6 – 7 ½ (7 ½ - 9 ½) scale rows between lateral line and dorsal-fin origin. It is distinguished from Devario pathirana by a shallower body (21.8 – 26.0 % SL, vs 32.4 – 36.9 % SL in D. pathirana); a shallower caudal peduncle depth of 8.8 – 10.5 % SL (vs 11.3 – 13.1 % SL); a shorter dorsal-fin base (15.1 – 18.2 % SL, vs 19.0 – 23.8 % SL); a shorter anal-fin base (16.4 – 20.5 % SL, vs 20.7 – 22.8 % SL); a shorter pelvic fin (12.9 – 16.6 % SL, vs 16.1 – 18.8 % SL); a shorter pectoral fin (pectoral-fin height 20.3 – 23.4 % SL, vs 23.8 – 27.9 % SL); 6 – 7 ½ (vs 8 – 8 ½) scale rows between lateral line and dorsal-fin origin; and 8 ½ - 10 ½ (vs 11 ½) branched dorsal-fin rays. Devario altus sp. nov. is genetically distinct from D. malabaricus, its closest congener, by 1.2 – 1.7 % in the cox 1 DNA barcoding marker and genetically distinct from the remaining species of Sri Lankan Devario by more than 3.6 %. Some populations of Devario in the central hills (‘ D. cf. malabaricus (highland phenotype) ’) possess a color pattern resembling that of D. altus sp. nov.: however, some of the individuals sequenced from these populations contain the mitochondrial haplotype for D. altus sp. nov., while most others contain the D. malabaricus mitochondrial haplotype. Sexual dimorphism is apparent in D. altus sp. nov., with males possessing (and females lacking) tubercles on the branched rays of the pectoral fin.	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
0947107FB2315F538D7E9F781706A0C3.taxon	description	Description. For general appearance, see Fig. 13 A – C; morphometric data are provided in Suppl. material 2. Head and body laterally compressed, elongate. Body depth greatest at pelvic-fin origin. Snout length subequal to eye diameter. Medial margins of dentaries straight, parallel, with an indentation (‘ danionine notch’) anteriorly. Well-developed dermal grooves present along supraorbital shelves. Small, rounded, symphysial knob present, fitting into shallow groove on inner margin of upper jaw. Lower jaw slightly longer than upper. First infraorbital with a process (n = 11) or smooth (n = 4). Tubercles on lower jaw rounded, arranged in a mediodorsal band of 2 – 3 rows, tapering to a single row posteriorly and towards symphysis, present in both sexes. Single row of rounded tubercles on upper margin of upper jaw. Pectoral-fin tubercles present in males. Both maxillary and rostral barbels present. Rostral barbel longer than maxillary barbel, not reaching anterior margin of orbit; maxillary barbel short, not reaching anterior margin of orbit. Lateral line complete, declining steeply for first 7 scales, then curving parallel to ventral body outline, running low on caudal peduncle, terminating on caudal-fin base, with 36 (2), 37 (6), 38 (3), 39 (1), 40 (2), or 41 (1) pored scales on body, plus 1 – 2 on caudal-fin base. Median predorsal scales 15 (3), 16 (10), 17 (1), or 18 (1). Lateral scale rows between origins of dorsal and pelvic fins 6 + 1 + 1 (2), ½ 6 + 1 + 1 (6), 7 + 1 + 1 (2), or ½ 7 + 1 + 1 (5). Circumpeduncular scales 13 (7) or 14 (8). Dorsal fin with 2 – 3 unbranched rays, 8 ½ (1), 9 ½ (10), or 10 ½ (4) branched rays, origin anterior to vertical through anal-fin origin, distal margin straight. Anal fin with 3 unbranched rays, 12 ½ (5), 13 ½ (7), 14 ½ (2), or 15 ½ (1) branched rays, distal margin slightly concave. Pectoral-fin with 1 unbranched and 9 (5), 10 (8) or 11 (2) branched rays, adpressed fin just reaching or surpassing pelvic-fin origin; axial lobe well developed. Pelvic fin with 1 unbranched and 6 (6), or 7 (9) branched rays, origin midway between anal-fin origin and pectoral-fin origin; tip of adpressed pelvic fin not reaching anal-fin origin. Pelvic ‘ axillary’ scale present. Caudal fin with 9 + 8 (15) branched rays, forked, lobes subequal, rounded distally.	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
0947107FB2315F538D7E9F781706A0C3.taxon	etymology	Etymology. The species-name altus is Latin for ‘ high’, an allusion to this species being restricted to the Sri Lankan highlands. Applied as a noun in apposition. Suggested vernacular name in English: Sri Lanka Montane Danio.	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
C2CCC44B8C3B5F479EAD1D5FFDE5DEE8.taxon	diagnosis	Diagnosis. Devario malabaricus is distinguished from Sri Lankan congeners by the combination of the following characters and character states: anal-fin origin posterior to dorsal-fin origin, beneath 3 rd branched dorsal-fin ray (vs anal-fin origin on vertical through dorsal-fin origin in D. memorialis); 1 st infraorbital smooth, with no infraorbital process (vs present in D. micronema, D. pathirana, and most specimens of D. altus); P-stripe usually bifurcated anteriorly (vs P-stripe not bifurcated in D. altus and D. memorialis); interstripe I + 1 short, usually not extending uninterrupted anterior to vertical through pelvic-fin origin (vs long, extending uninterrupted anterior to vertical through pelvic-fin origin in D. altus); caudal peduncle length 12.3 – 19.4 % SL (vs 19.2 – 21.4 % SL in D. altus); caudal peduncle depth 9.8 – 12.9 % SL (vs 8.1 – 9.8 % SL in D. memorialis); body depth 26.1 – 35.4 % SL (vs 21.8 – 26.0 % SL in D. altus; 15.9 – 22.6 % SL in D. memorialis); dorsal-fin base length 16.7 – 23.5 % SL (vs 13.5 – 16.3 % SL in D. memorialis); pelvic-fin height 13.3 – 18.1 % SL (vs 11.6 – 13.3 % SL in D. memorialis); 33 – 39 pored lateral-line scales (vs 38 – 46 in D. memorialis); predorsal scales 14 – 18 (vs 17 – 19 in D. memorialis); 12 – 15 circumpeduncular scales (vs 15 – 18 in D. memorialis); 7 ½ - 9 scales between lateral line and dorsal-fin origin (vs 7 ½ in D. memorialis; 6 – 7 ½ in D. altus); 9 ½ - 12 ½ branched dorsal-fin rays (vs 8 ½ in D. memorialis). Devario malabaricus is genetically distinct from D. altus, its closest congener, by 1.2 – 1.7 % in the cox 1 DNA barcoding marker, and from the remaining Sri Lankan species of Devario by more than 2.8 %. Sexual dimorphism is apparent in D. malabaricus, with the branched rays of the pectoral fin exhibiting tubercles in males but not in females (Sudasinghe 2024).	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
FADBB3695EC4533BB7D322969544F857.taxon	diagnosis	Diagnosis. Devario memorialis is distinguished from its Sri Lankan congeners by the combination of the following characters and character states: anal-fin origin on vertical through dorsal-fin origin (vs anal-fin origin posterior to dorsal-fin origin, beneath 3 rd branched dorsal-fin ray in D. malabaricus, D. micronema, D. altus, and D. pathirana); 1 st infraorbital smooth, with no infraorbital process (vs infraorbital process present in D. micronema, D. pathirana, and most specimens of D. altus); anterior bars on body absent or indistinct (vs present in D. malabaricus, D. micronema, and D. pathirana); preanal length 58.5 – 63.8 % SL (vs 64.9 – 69.8 % SL in D. pathirana); caudal peduncle depth 8.1 – 9.8 % SL (vs 9.8 – 12.9 % SL in D. malabaricus, 11.3 – 13.1 % SL in D. pathirana); body depth 15.9 – 22.6 % SL (vs 26.1 – 35.4 % SL in D. malabaricus, 23.5 – 36.6 % SL in D. micronema, 21.8 – 26.0 % SL in D. altus, 32.4 – 36.9 % SL in D. pathirana); dorsal-fin height 15.7 – 20.0 % SL (vs 19.0 – 23.2 % SL in D. altus, 20.6 – 24.3 % SL in D. pathirana); dorsal-fin base length 13.5 – 16.3 % SL (vs 16.7 – 23.5 % SL in D. malabaricus, 16.6 – 23.3 % SL in D. micronema, 19.0 – 23.8 % SL in D. pathirana); anal-fin base length 20.0 – 23.4 % SL (vs 16.4 – 20.5 % SL in D. altus); pelvic-fin height 11.6 – 13.3 % SL (vs 13.3 – 18.1 % SL in D. malabaricus, 13.8 – 18.5 % SL in D. micronema, 12.9 – 16.6 % SL in D. altus, 16.1 – 18.8 % SL in D. pathirana); pectoral-fin height 19.1 – 23.6 % SL (vs 23.8 – 27.9 % SL in D. pathirana); head depth 14.1 – 15.8 % SL (vs 16.0 – 23.0 % SL in D. micronema, 17.3 – 19.6 % SL in D. altus, 17.6 – 21.7 % SL in D. pathirana); interorbital width 37.2 – 40.7 % HL (vs 40.7 – 45.3 % HL in D. pathirana); rostral barbel 4.0 – 8.9 % HL (vs 8.3 – 14.2 % HL in D. altus, 8.6 – 17.7 % HL in D. pathirana); 38 – 46 lateral-line scales (vs 33 – 39 in D. malabaricus); predorsal scales 17 – 19 (vs 14 – 18 in D. malabaricus, 13 – 17 in D. micronema, 14 – 15 in D. pathirana); 15 – 18 circumpeduncular scales (vs 12 – 15 in D. malabaricus, D. micronema, 13 – 14 in D. altus, D. pathirana); 7 ½ scales between lateral line and dorsal-fin origin (vs 7 ½ – 9 in D. malabaricus, 7 ½ – 9 ½ in D. micronema, 6 – 7 ½ in D. altus, 8 – 8 ½ in D. pathirana); 8 ½ branched dorsal-fin rays (vs 9 ½ – 12 ½ in D. malabaricus, D. micronema, 8 ½ – 10 ½ in D. altus, 11 ½ in D. pathirana); 14 ½ – 15 ½ branched anal-fin rays (vs 13 ½ – 14 ½ in D. pathirana). Devario memorialis is genetically distinct from the remaining Sri Lankan species of Devario by an uncorrected p-distance of more than 3.9 % in the cox 1 DNA barcoding marker.	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
D2B5A728E3825FE6B3E9747659499FD6.taxon	diagnosis	Diagnosis. Devario micronema is distinguished from Sri Lankan congeners by the combination of the following characters and character states: anal-fin origin posterior to origin of dorsal fin, beneath 3 rd branched dorsal-fin ray (vs anal-fin origin on vertical through dorsal-fin origin in D. memorialis); 1 st infraorbital with an infraorbital process (vs infraorbital process absent in D. malabaricus and D. memorialis); interstripe I + 1 short, usually not extending uninterrupted anterior to vertical through pelvic-fin origin (vs long, extending uninterrupted anterior to pelvic-fin origin in D. altus); caudal peduncle length 13.2 – 20.0 % SL (vs 19.2 – 21.4 % SL in D. altus); body depth 23.5 – 36.6 % SL (vs 15.9 – 22.6 % SL in D. memorialis); dorsal-fin base length 16.6 – 23.3 % SL (vs 13.5 – 16.3 % SL in D. memorialis); pelvic-fin height 13.8 – 18.5 % SL (vs 11.6 – 13.3 % SL in D. memorialis); head depth 16.0 – 23.0 % SL (vs 14.1 – 15.8 % SL in D. memorialis); predorsal scales 13 – 17 (vs 17 – 19 in D. memorialis); circumpeduncular scales 12 – 15 (vs 15 – 18 in D. memorialis); scales between lateral line and dorsal-fin origin 7 ½ - 9 ½ (vs 7 ½ in D. memorialis); branched dorsal-fin rays 9 ½ - 12 ½ (vs 8 ½ in D. memorialis). Devario micronema is genetically distinct from D. pathirana, its closest congener by 1.2 – 1.6 % in the cox 1 DNA barcoding marker, and from the remaining Sri Lankan species of Devario by over 2.8 %. Among these, D. micronema shows, across its range, the greatest phenotypic diversity in terms of color-pattern and habitus (Fig. 14) as well as greatest genetic diversity among the Sri Lankan species of Devario. Several of our specimens are suggestive of mitonuclear discordance and / or phenotypes that are intermediate between D. malabaricus and D. micronema. These specimens are considered in this study as potential hybrids. Sexual dimorphism in D. micronema is apparent, with males exhibiting tubercles on the branched rays of the pectoral fin, a character absent in females (Sudasinghe 2024).	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
4ED80C87D23A5943B0FA3154A640AA30.taxon	diagnosis	Diagnosis. Devario pathirana is distinguished from Sri Lankan congeners by the combination of the following characters and character states: it is the only completely barred Devario in the island, with 7 – 11 bars on the side of the body (vs fewer than 4 bars in D. malabaricus, D. micronema, D. memorialis, D. altus); P-stripe short, ending above the caudal peduncle (P-stripe long, extending uninterrupted above the caudal peduncle in D. malabaricus, D. micronema, D. memorialis, D. altus); anal-fin origin posterior to origin of dorsal fin, beneath 3 rd branched dorsal-fin ray (vs anal-fin origin on vertical through dorsal-fin origin in D. memorialis); 1 st infraorbital with an infraorbital process (vs infraorbital process absent in D. malabaricus, D. memorialis); preanal length 64.9 – 69.8 % SL (vs 58.5 – 63.8 % SL in D. memorialis); caudal peduncle length 15.7 – 19.6 % SL (vs 19.2 – 21.4 % SL in D. altus); caudal peduncle depth 11.3 – 13.1 % SL (vs 8.1 – 9.8 % SL in D. memorialis, 8.8 – 10.5 % SL in D. altus); body depth 32.4 – 36.9 % SL (15.9 – 22.6 % SL in D. memorialis; 21.8 – 26.0 % SL in D. altus); dorsal-fin height 20.6 – 24.3 % SL (vs 15.7 – 20.0 % SL in D. memorialis); dorsal-fin base length 19.0 – 23.8 % SL (vs 13.5 – 16.3 % SL in D. memorialis, 15.1 – 18.2 % SL in D. altus); anal-fin base length 20.7 – 22.8 % SL (vs 16.4 – 20.5 % SL in D. altus); pelvic-fin height 16.1 – 18.8 % SL (vs 11.6 – 13.3 % SL in D. memorialis; 12.9 – 16.6 % SL in D. altus); pectoral-fin height 23.8 – 27.9 % SL (vs 19.1 – 23.6 % SL in D. memorialis, 20.3 – 23.4 % SL in D. altus); head depth 17.6 – 21.7 % SL (vs 14.1 – 15.8 % SL in D. memorialis); interorbital width 40.7 – 45.3 % HL (vs 37.2 – 40.7 % HL in D. memorialis); rostral barbel 8.6 – 17.7 % HL (vs 4.0 – 8.9 % HL in D. memorialis); 14 – 15 predorsal scales (vs 17 – 19 in D. memorialis); 13 – 14 circumpeduncular scales (vs 15 – 18 in D. memorialis); 8 – 8 ½ scales between lateral line and dorsal-fin origin (vs 7 ½ in D. memorialis; 6 – 7 ½ in D. altus); 11 ½ branched dorsal-fin rays (vs 8 ½ in D. memorialis); 13 ½ – 14 ½ branched anal-fin rays (vs 14 ½ – 15 ½ in D. memorialis). Devario pathirana is genetically distinct from D. micronema, its closest congener by 1.2 – 1.6 % for the cox 1 DNA barcoding marker and genetically distinct from rest of the Sri Lankan species of Devario by over 3.6 %.	en	Sudasinghe, Hiranya, Pethiyagoda, Rohan, Meegaskumbura, Madhava (2025): Species without borders: the diversification of giant danios in Sri Lanka (Teleostei, Danionidae, Devario). Evolutionary Systematics 9 (2): 229-258, DOI: 10.3897/evolsyst.9.162251
