taxonID	type	description	language	source
03A087A043324A1CFF7AE87A90935A91.taxon	diagnosis	Diagnosis. Imago. Forewing (Figs 1, 22) crossveins generally reduced in number, comprised between 45 – 65, trimmed with violet or brown especially in the costal and subcostal fields. Costal and subcostal fields with few crossveins (<12), at least two rows of crossveins in the radial field and one crossing the radial-median fields. Male foreleg with first tarsal segment reduced in length, ca 4 times shorter than the second one (Figs 3, 4). Styliger plate straight or slightly convex, last two segments of the gonopods subequal in length (Figs 6, 23). Male genitalia with inner median titillators, penis lobes with apical, subapical and basal spines; lateral sclerite broad and basal sclerite weakly developed (Figs 7, 24).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043324A1CFF7AE87A90935A91.taxon	description	Nymph. Head unicolorous, not thickened, with sometimes lateral markings but without whitish spots on the fore margin (Figs 79 – 80). Distal dentisetae on maxillae with inner one entire and outer one fringed (Figs 43 – 44). Scattered setae on the maxillae fimbriate (Figs 45 – 46). Labial glossae rhomboid with outer margin convex (Fig. 12). Supracoxal spurs present but rounded or bluntly developed (Figs 79 – 80). Spines on the upper face of femora rounded or truncate (Figs 14, 62). Inner margin of femora without long and thin setae (Figs 13, 61). Gills generally tinted with greyish-purple (Figs 16 - 18, 79 – 80). All gills with fibrillous and plate like structures, except gill VII only plate-like. Hind tibiae with two rows of fine setae (Figs 13, 15, 61). Tarsal claw with 3 - 4 subapical teeth. Caudal filaments with whorls of spine like setae on each segment, without long and thin setae. Eggs. Chorion covered by polygonal ridges; one pole with KCT’s larger than those in equatorial area; micropyle with double margin (Figs 20 – 21).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043324A1CFF7AE87A90935A91.taxon	discussion	Species included. Compsoneuria spectabilis Eaton, 1881 (imagos, nymphs): Java, Sumatra Compsoneuria lieftincki (Ulmer, 1939) (imagos): Java Discussion. At the imaginal stage, the combination of the forewing crossveins, the male foretarsi proportions and genitalia will distinguish the genus from all its relatives. At the nymphal stage, Compsoneuria is distinguished from its relatives by the shape of the supracoxal spurs, by the presence of two rows of setae on the hind tibiae, by the shape of the outer distal dentiseta, and by the shape and coloration of the gills. Using the key provided by Webb & McCafferty (2008), Compsoneuria will key out at couplet 15 leading to “ Afronurus in part ”.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043324A15FF7AEF2A91415C20.taxon	materials_examined	Material examined. Specimens in ethanol 1 male imago, Buitenzorg, 13.2.32. Lieftinck leg. [ZMH] 1 male (one fore- and one hind legs on slide) and 1 female imagos, Buitenzorg, XI. 1919, Karny leg. [ZMH] 1 female subimago, Buitenzorg, VII. 1929, Lampe, Thienemann leg. [ZMH] 2 female imago and 1 male subimago, Buitenzorg, I. 1929, Lampe, Thienemann leg. [ZMH] 4 nymphs (two mounted on slides), Indonesia, Java, Buitenzorg [Bogor], in Tjiliwung [Ciliwung] River, 25. V. 1929, FB 3, Prof. Thienemann leg [ZMH, MZL] (sub. nom. C. thienemanni) 4 nymphs, Indonesia, West Java, Fish ponds south of the Puntjak pass, 13. IX. 1928, B 13, Prof. Thienemann leg [ZMH] (sub. nom. C. thienemanni) 1 nymph, Indonesia, West Java, Buitenzorg, Pakantjilan River, 16. IX. 1928, B 7, Prof. Thienemann leg [ZMH] (sub. nom. C. thienemanni) 8 nymphs (one mounted on slide), Indonesia, Sumatra Barat, Ombilin River, Talawi, 277 m., 00 ° 34.147 ’ S 100 ° 43.543 ’ E, 8. XI. 2011, UN 4, M. Balke leg [MZL, ZMH, LIPI] 1 nymph, Indonesia, Sumatra Barat, stream near Lubuk Sikaping, 440 m., 0 ° 10.839 ’ S 100 ° 08.533 ’ E, 29. IX. 2009, SUM 12, M. Balke & D. Amran leg. [MZL] Pinned specimen. One male imago, Buitenzorg, Java. K. Kraepelin leg. 24. II – 12. III. 1904 Comments on the material examined of C. spectabilis. The pinned specimen is the one observed and mentioned by Klapálek (1905, p. 107) and illustrated by Ulmer (1924, fig. 53 A, fig. 55 C). In figure 53 A the segment V of Klapálek’s specimen does not possess a sagittal dark banding, i. e. these marks are present only on segments III – IV and VI – VII. This drawing is thus an interpretation by Ulmer (1924, p. 86): “ auf Tergit V manchmal undeutlich oder fehlend ”. Genitalia and legs of the 1919 male imago have been illustrated by Ulmer (1924, figs 56 – 58); the attribution of the specimen to coll. Jacobson in the legends is a probable mistake since the original label bears the name of Karny. Sequence data. One specimen has been used for the study by Vuataz et al. (2013) under the name “ Heptageniidae 2 ” in figures and “ Heptageniidae sp. 2 ” in table S 1, with one mitochondrial (CO 1) and two nuclear genes (H 3, wg) sequenced. Access numbers in GenBank are for CO 1: HF 536606, for wg: HF 536599, for H 3: HF 536592.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043324A15FF7AEF2A91415C20.taxon	description	Description. Male imago description based on the 1932 specimen. General coloration faded, with legs yellowish, thorax and compound eyes brownish-orange, wings translucent, costal and subcostal fields milky, abdomen translucent with ventral nerve cord whitish, gonopods yellowish, cerci whitish, some junctions between segments lightly tinted in orange, suggesting cerci originally banded with darker color. Measurements: Body length: ca 6.5 mm; Forewing length: 6.9 mm; Gonopods length: 0.9 mm; Cerci length: ca. 14 mm Head: Compound eyes large, meeting in middle at distance equal to distance between lateral ocelli; lateral ocelli two times larger than front one. Frons translucent with well-marked carina. Thorax: Transversal suture on mesothorax present but barely visible; medial depression of furcasternum of mesothorax subparallel. Forewing (Fig. 1) with few crossveins, arranged roughly in several rows; costal field with ca. 10 crossveins, subcostal field with only 6; hindwing as in Fig. 2. Foreleg (Fig. 3) with tibia ca 1.3 x length of femur, and tarsi subequal or slightly longer than tibia; tarsal segment 1 very short, 4.5 x shorter than second; segment 3 and 4 subequal, 0.8 x length of segment 2, segment 5 0.5 x length of segment 4; tarsal claw dissimilar, one obtuse, one hooked. Hind leg with tibia subequal to femur, tarsi shorter than tibia, tarsal segments 1 and 2 subequal, segment 4 the shortest, segments 3 and 5 subequal and slightly shorter than segments 1 and 2. Abdomen (genitalia): styliger plate (Fig. 6) almost straight, slightly emarginated adjacent to bases of gonopods.; basal segment rounded, 2 x longer than wide; segment 1 short and broad, segments 3 and 4 subequal in length, together about ⅓ length of segment 2; segment 2 with small rounded inner projection at base. Penis lobes separated by V shape in ventral view (Fig. 7) and U shape in dorsal view; lobes expanded laterally; apical sclerite long, extending to half length of lobe, with distinct subapical spine; base of apical sclerite tinted in orange brown, suggesting strong chitinization. In dorsal view, lateral sclerite broad and rounded, not constricted at apex, basal sclerite weakly developed, with small and acute spine directed inwards on outer margin. Titillators small but present, visible in ventral view, but barely visible in dorsal view. Nymph. (Note: The nymph of C. spectabilis has been described by Ulmer (1939) under the name Compsoneuriella thienemanni (figs 442 – 454). It is redescribed here anyway for comparative purposes.) Size: body length between 5.7 and 7.9 mm. Coloration (Fig. 79): head uniformly dark brown, except white spot around ocelli and transverse white line starting from lateral ocelli and reaching margin of head in front of compound eyes in perpendicular orientation; prothorax medium brown with two whitish spots, one proximal, one distal, on median part of each hemi-thorax, sometimes linked together with light line; dorsal face of femora greyish brown, with three elongated maculae on outer margin, sometimes less visible on inner margin, apices of femora and tibiae dark brown, tibiae uniformly medium brown, sometimes slightly banded; abdominal tergites medium brown, with markings, as follows: segments III, IV, VI and VII with double sagittal blackish line crossing whole segment, on segment V, diffuse yellowish macula in middle, on segments VIII and IX, elongate transverse yellowish macula crossed in middle by sagittal dark brown line, sometimes incomplete; posterior margin of tergites II – VII with blackish line enlarging when reaching sides; abdominal sternites uniformly whitish; cerci medium brown with dark brown banding every two segments. Head: Labrum (Fig. 8) greatly extended laterally, with narrow tips slightly turned posteriorly; anterior margin almost straight, covered by numerous long and simple setae; ventral surface with row of small stout setae; dorsal surface with scattered long and simple setae directed medially. Mandibles strongly arched, both with lateral margin covered with several rows of long and thin setae; right mandible (Fig. 9) with outer incisor saw-like on its inner margin, composed of ca. 9 teeth, and with subapical tooth in ventral position; inner incisor trifid; presence of 4 fimbriate setae below incisors and 4 long and simple setae below mola; left mandible (Fig. 10) with outer incisor saw-like on its inner margin, composed of ca. 6 teeth, and with subapical tooth in ventral position; inner incisor slender and pointed, bearing 4 – 5 setae-like structures arising from ventral face; 4 fimbriate setae present below incisors, followed by tuft of short setae above mola, and 8 long and simple setae below mola. Maxillae with row of ca. 13 comb-shaped setae on anterior margin, medial comb-shaped setae bearing 12 – 13 teeth (Fig. 47); ventral surface of galea covered by scattered fimbriate setae (Figs 45 – 46); proximal dentiseta bifid and fimbriate, first (outer) distal one fimbriate, second (inner) distal one entire (Figs 43 – 44). Hypopharynx (Fig. 11) with welldeveloped superlinguae, densely covered with long and simple setae on lateral margin, ending below rounded apex of lobes; lingua with tuft of small setae. Labium (Fig. 12) characterized by well-separated glossae, somewhat rhomboid; paraglossae very large, and extending laterally, more than two times longer than wide. Thorax: Leg ornamentation rather similar among fore-, mid- and hindlegs. Supracoxal spurs weakly developed on forelegs, apparent on mid- and hindlegs, rounded apically. Hindleg (Figs 13, 60) with femur bearing row of long and simple setae on its outer margin; inner margin with row of stout spines directed toward apex, and an incomplete submarginal row of stout spines; bristles on upper face of femora elongated (Figs 14, 62), with rounded apex; tibiae each with two rows of long and very thin setae, one on outer margin, one on lateral ridge (Figs 13, 61); inner margin with only 5 – 6 short spines regularly arranged. Tarsal claw hooked, with 3 – 4 subapical denticles (Fig. 63). Abdomen: Posterolateral projections weakly developed on segments I – IV, more pronounced on segments V – IX. Gills broadly oval (Figs 16 – 18), fibrillate part well-developed on gills I – VI, absent on gill VII; plate gills with tracheation barely visible, strongly colored with purple gray on most of plate, except in apical region and close to margins. Posterior margin of tergites (Fig. 19) with few but very large spines, with 2 – 3 intercalary small pointed spines. Cerci with whorl of stout and pointed setae on each segment. Eggs (from mature nymph and female): Size: 115 – 130 µm x 80 – 95 µm; chorion smoothly granulated, covered by polygonal ridges (Fig. 21 a), and KCTs arranged in polygonal mesh with ridges joining them. KCTs on one pole regularly arranged and three times larger than those on other part of egg (Fig. 21 b). Micropyles located in equatorial area, with prominent double margins (Fig. 21 c).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043324A15FF7AEF2A91415C20.taxon	discussion	Discussion. The specimens housed in ZMH completely fit the diagnosis proposed by Eaton (1885) and Ulmer (1924), especially concerning wing venation and leg proportions. Fig. 1 of the forewing is almost identical to Fig. 55 A in Ulmer (1924) of a female imago also from Buitenzorg. Figs 3 and 4 of the legs are identical to those proposed by Ulmer (1924) in Fig. 58 for another male imago from Buitenzorg. Genitalia are basically comparable in the general shape; Ulmer’s drawing (Ulmer 1924, Fig. 56) exhibits the characteristic apical sclerite, although he did not draw the spine in ventral view, but mentioned it in the text and illustrated it in lateral view (Ulmer 1924, Fig. 57 B). Major differences between the two drawings are the stem of the lobes which are clearly convex in our specimen but concave in Ulmer’s drawing, and the shape of the styliger plate which is not flat but concave with a pointed process in the middle. As the specimen drawn by Ulmer (1924, Figs 56 – 58) is also deposited in the collections of ZMH, it has been re-studied and a complementary description and interpretation is given. The contour of the genitalia of the 1919 specimen in ventral and dorsal view is in accordance with Ulmer’s drawing, except the stem of the penis which is clearly convex. The shape of the styliger plate is undoubtedly different but results of a partial drying of the specimen which has folded the plate, hence making this “ pointed process ” apparent but not real. The basal segments of the gonopods are not in the same plane as the styliger because of the folding. Penis lobes are somewhat distorted and the lateral sclerites are not well visible but the two small spines at the base of the sclerites are noticeable as are those of the apical sclerites in lateral view (Ulmer 1924, Fig. 57 B). Therefore, there is no clear difference between these two specimens, taking into account that the 1919 specimen is less well preserved than the 1932 one. The major difference is the presence of concealed and hard to see titillators in C. spectabilis, as well as the presence of subapical spines on the ventral side of the penis lobes; these two structures were evidently overlooked by Ulmer (1924) in his redescription. The slide preparation of fore- and hind legs made by Ulmer is presented in Figs 4 and 5. Foreleg (Fig. 4) with coloration patterns very characteristic; the dorsal face of the femur is covered by small and dark maculae in the distal part; apex of the femur with two dark maculae near the inner and the outer margins; proximal part of the tibia dark brown, becoming lighter distally on a short distance; apex of tibia medium brown, as are the distal outer part of each tarsal segments. Hindleg (Fig. 5) with coloration patterns as in the foreleg. There is a third male imago available in the ZMH collections, pinned and mentioned by Klapálek (1905), and also coming from Buitenzorg. The observation of the genitalia is difficult because of the twisted gonopods, but the general shape is in accordance with the two others, the apical sclerite with the median spine is present, the lateral sclerite has the same shape, but the minute spine at the base of the sclerite cannot be seen. Wing venation is in agreement with the one of the described specimen. This specimen is therefore also considered as belonging to C. spectabilis. It is the only one that gives an idea of the color patterns of the thorax and the abdomen. The nymph of C. spectabilis described here is the only one known to belong to the genus Compsoneuria at the moment (but see discussion, and Fig. 80). The generic diagnosis therefore applies to the species C. spectabilis.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433B4A14FF7AEABA95D35ACB.taxon	materials_examined	Material examined. Two males and one female syntypes with subimaginal exuviae, Indonesia, West Java, Djasinga, Tjibarangbang River, 150 m., caught at light in the evening above the river, 6. VII. 1935, Lieftinck leg [ZMH].	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433B4A14FF7AEABA95D35ACB.taxon	description	Description. Male imago. (Note: Only significant characters are mentioned here; for a complete description see Ulmer 1939). All specimens faded. Forewings translucent, except costal and subcostal fields milky. Hindwings entirely colored in light brown, color presumably more pronounced on fresh material. Foreleg similar to C. spectabilis, with first segment of tarsus very short (Ulmer, 1939, fig. 160). Forewing (Fig. 22) with 12 crossveins in costal and subcostal fields; row of crossveins readily apparent in middle of radial-median field, crossveins on distal part of radial field less distinct. Overall number of crossveins greater than number in other species (> 90). Gonopods, styliger plate and genitalia characteristic of genus (Fig. 23). Penis lobes (Fig. 24) with small apical spine at base of well-developed apical sclerite, protruding outwards slightly, giving apex of lobe more concave appearance than found in other species; subapical spine and titillators slender; basal sclerite not easily observed, with only acute and long spine visible; lateral sclerite broad and quadrate. Nymphal stage unknown.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433B4A14FF7AEABA95D35ACB.taxon	discussion	Discussion. Ulmer (1939) described this species in the genus Heptagenia Walsh, 1863, because the first segment of the hind tarsus is not longer than the second. As illustrated by Ulmer (1939, fig. 161), this first segment is subequal in length to the second one in H. lieftincki. The presence of a transverse suture on the mesonotum, together with the median depression of the furcasterum of mesothorax subparallel clearly indicates that C. lieftincki belongs to the subfamily Ecdyonurinae and therefore cannot be a member of the Heptageniinae where belongs the genus Heptagenia. Consequently, the combination proposed by Webb et al. (2006) is followed here. Compsoneuria lieftincki possesses genitalia in accordance with the actual concept of Compsoneuria, but the forewing venation is intriguing since it possesses many more crossveins than in C. spectabilis. Compsoneuria lieftincki exhibits few crossveins in the costal and subcostal fields, as in C. spectabilis, together with rows of crossveins in the radial and median fields, together with the reduced first tarsal segment of the forelegs. The female of C. lieftincki described by Ulmer (1939, p. 573) does not belong to the same species or even genus as the male mentioned here, because of the wing venation with numerous crossveins in the costal and subcostal fields and the hind leg proportions, the latter being already mentioned by Ulmer (1939, p. 574) as something strange (“ auffällig ”).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433D4A12FF7AEBEE95C65D40.taxon	diagnosis	Diagnosis. Imago. Overall number of crossveins in the forewing reduced in number compared to other Heptageniidae, except other members of the tribe Compsoneuriini (Compsoneuria and Notonurus). Cross veins in the costal and subcostal fields, sometimes also in the radial and median fields, trimmed with violet or brown. Costal and subcostal fields with numerous crossveins (> 15), no rows of crossveins visible in the radial-median fields, sometimes present at apex of the radial field (Fig. 25). Male foreleg with first tarsal segment slightly shorter than the second one (Figs 26 – 27). Styliger plate faintly convex (Fig. 28). Last segment of the gonopods generally smaller than the previous one. Male genitalia with median titillators, penis lobes with apical, subapical and basal spines (Fig. 29). Lateral sclerite narrow, and basal sclerite well-developed. Nymph. Head not thickened, dark with whitish spots near the anterior margin (Figs 81 – 82). Distal dentisetae on maxillae with inner and outer one simple (Figs 48 – 51). Labial glossae conical with concave outer margin apically (Figs 35, 68, 72, 75). Scattered setae on the maxillae generally fimbriate (Figs 52 – 53). Supracoxal spurs well developed, sharply acute and often pointed (Figs 81 – 82). Spines on the upper face of femora pointed, or at least with strongly convergent margins (Figs 37, 66, 69). Inner margin of femora with long and thin setae (Figs 36, 65). Gills generally whitish with well visible tracheation (Figs 38 – 39). All gills with fibrillous and plate like structures, except gill VII only plate-like. Hind tibiae with one row of fine setae on the outer margin (Figs 36, 65). Tarsal claw with 0 - 2 subapical teeth. Caudal filaments with whorls of spine like setae on each segment, without long and thin setae. Eggs. Chorion covered by flat, fibrillose and rounded structures; on each pole, KCT´s around two times the size of the others; micropyle with single margin (Figs 41 – 42; Boonsoong & Braasch 2013, fig. 7 A – B).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433D4A12FF7AEBEE95C65D40.taxon	discussion	Species included: Compsoneuriella thienemanni Ulmer, 1939 (imagos, nymphs): Sumatra, Java Compsoneuriella sp. 1 (imagos, nymphs): Vietnam, Thailand Compsoneuriella langensis (Braasch & Boonsoong, 2010) (imagos, nymphs): Thailand comb. nov. Compsoneuriella tagbanua (Braasch & Freitag, 2008) (imagos, nymphs): Philippines (Palawan) comb. nov. Discussion. As already mentioned, the genus Compsoneuriella corresponds to the concept of Compsoneuria by Braasch & Soldán (1986 b) and subsequent authors. At the nymphal stage, Compsoneuriella is distinguished from its relatives by the contrasted color pattern of the cephalic capsule, with at least 4 whitish dots close to the anterior margin, the acute supracoxal spurs, the presence of long and thin setae on the inner margin of the femora, the bristles on the upper face of femora pointed or at least with margins clearly convergent, the outer distal dentiseta of the maxillae entire, and by the presence of a single row of setae on the hind tibiae. At the imaginal stage, the combination of the number of forewing crossveins in the costal and subcostal fields, the male foretarsi proportions and genitalia will distinguish the genus from Compsoneuria, but differences with Notonurus were not found. The eggs of Compsoneuriella differ from those of Compsoneuria by the arrangement of the chorion and the margin of the micropyle. To our knowledge, no eggs of Notonurus are presently described. Within the tribe Compsoneuriini, Compsoneuriella share several features with the Afrotropical genus Notonurus in the nymphal stage, including the overall color pattern of the head and the pointed supracoxal spurs. Compsoneuriella, however, typically has fimbriate scattered setae of the maxillae together with entire distal dentisetae (Figs. 48 – 53) whereas those of Notonurus have simple scattered setae on the maxillae and numerous distal dentisetae (Figs 55 – 58).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433C4A06FF7AEB9A95C75923.taxon	materials_examined	Material examined. Specimens in ethanol. 1 female imago, Sumatra, Wai Lima, Lampongs, XI – XII. 1921. Karny leg. [ZMH] 2 male imagos (fore- and hind legs mounted on slide), 1 male subimago, Sumatra, Lake Singkarak, III. 1929, Lampe, Prof. Thienemann leg. [ZMH] 1 male subimago (with corresponding nymphal exuvia on slide), Sumatra, shore of the Lake Singkarak, 23. II. 1929, F 3 e, Prof. Thienemann leg [ZMH] 4 nymphs (one mounted on slide), Indonesia, West Java, Fish ponds south of the Puntjak pass, 13. IX. 1928, B 13, Prof. Thienemann leg [ZMH, MZL] 1 nymph, Indonesia, West-Java, Buitenzorg, lateral branch of the Tjiliwung River, IX. 1928, B 3, Prof. Thienemann leg [ZMH] 3 nymphs (two mounted on slides), Indonesia, Sumatra Barat, Harau Canyon, stream, 540 m, 0 ° 04.428 ’ S 100 ° 38.002 ’ E, 27. IX. 2009, SUM 09 / UN 11, M. Balke & D. Amran leg. [ZMH, MZL] 1 nymph, Indonesia, Sumatra Barat, Sawahlunto, Sikalang, 200 m, 0 ° 37.678 ’ S 100 ° 46.314 ’ E, 21. IX. 2009, SUM 01, M. Balke & D. Amran leg. [LIPI] Pinned specimen. 1 female imago, E. Jacobson, Toentang Java, Oct. 1910 [ZMH] (sub. nom C. spectabilis) Lectotype designation. Ulmer did not designate a holotype for Compsoneuriella thienemanni, although he mentioned that the type was a male imago; therefore all the original material constitutes the type series and consequently all specimens are syntypes. In the ZMH collections catalogue, under this species name, appears the mention of a holotype male imago. This information is also available in Weidner (1964). This terminology cannot be accepted because the “ holotype ” has not been designated by Ulmer and cannot be ascertained by the presence of a single specimen (see also Recommendation 73 F. Avoidance of assumption of holotype, ICZN, 1999). Weidner (1962; 1964) also used the term “ paratypoides ” which should correspond to our actual concept of “ paratypes ”. Finally Ulmer himself brought some confusion; when describing the nymph of C. thienemanni, he mentioned in the material examined the four nymphs belonging to the species C. spectabilis (see above), with the following comment in bracket “ darunter das typische Stück ”, which means “ among which the typical one ”. This cannot be taken into account for the designation of a type specimen, and this is not a nomenclatorial act according to the Code. As obviously all the specimens examined by Ulmer under the name Compsoneuriella thienemanni encompass more than one species, a lectotype is designated among them, which is the best preserved male imago, with dissected genitalia, and which bears the following labels: i) Zool Mus. Hamburg — Coll. G. Ulmer — Eing Nr. 6 – 1963 ii) Compsoneuriella Thienemanni Ulm. iii) Singkarak, Lampe III. 29; iii) Compsoneuriella thienemanni Ulmer, 1939 LECTOTYPUS ♂, M. Sartori des., 2013 The type locality is therefore restricted to Singkarak, Sumatra, Indonesia (articles 73.2.3 and 76.2, ICZN, 1999).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433C4A06FF7AEB9A95C75923.taxon	description	Description. Male imago (lectotype). Coloration faded, legs yellowish, thorax and compound eyes brownishorange, wings translucent, costal and subcostal fields milky in distal part, abdomen translucent, except segments VII – X yellowish, with ventral nerve cord whitish, gonopods yellowish, cerci whitish, some junctions between segments lightly tinted in orange in proximal part, suggesting cerci originally banded with darker color on ca. onethird of their length. Measurements: Body length: ca. 6.6 mm; Forewing length: 6.3 mm; Gonopods length: 0.8 mm; Cerci length: ca. 10 mm but probably broken at apex. Head: Compound eyes large, meeting in middle at distance superior to distance between lateral ocelli; lateral ocelli two times larger than median ocellus. Frons translucent with well-marked carina. Thorax: Transversal suture on mesothorax present; medial depression of furcasternum of mesothorax subparallel. Forewing (Fig. 25) with numerous crossveins, those of costal and subcostal fields tinted with orange (blackish originally?); no rows of crossveins visible. Foreleg (Fig. 26) with tibia slightly longer than femur, and tarsi 1.75 x longer than tibia; tarsal segment 2 and 3 subequal in length, segment 1 shorter, ca. 0.7 x length of segment 2, segment 4 0.55 x length of segment 3 and segment 5 0.5 x length of segment 4; tarsal claws dissimilar, one obtuse, one hooked. Hind leg with tibia shorter than femur, tarsi subequal length of tibia, tarsal segments 1 longest, segments 2, 3 and 5 subequal, only slightly shorter than segment 1, segment 4 shortest. Abdomen (genitalia): styliger plate (Fig. 28) slightly convex; basal segment rounded, 2 x longer than wide, with inner corner well-marked; segment 1 short and broad, segment 4 slightly shorter than segment 3, together between ½ and ⅓ length of segment 2. Penis lobes stocky. In ventral view (Fig. 29, left), apical sclerite reaching half of lobe width, with distinct basal spine; well-developed subapical spine present below apical sclerite; on inner face of lobes, two stout and pointed titillators. In dorsal view (Fig. 29, right), lateral sclerite narrow and constricted at apex, basal sclerite strongly developed and bearing on outer margin one small and acute spine directed inwards. Nymph. Size: body length up to 7 mm and 8 mm in male and female nymphs respectively. Coloration (Fig. 81): overall medium brown with yellowish brown markings. Head (Fig. 30) with lateral margin yellowish and 4 rounded spots near anterior margin, median spots apparently extend backward in yellowish band through antennae and to anterior margin of compound eyes. Femora also contrasted, yellowish with two brownish maculae longitudinal to outer margin and two to inner margin, base of stout bristles on upper face strongly colored in dark brown; tibiae whitish with three ringed brownish maculae, largest macula in middle of tibia, others present in proximal and distal positions. Prothorax greyish brown with two diagonal elongated yellowish bands on each side, mesothorax with two blackish maculae situated medially and close to prothorax. Abdominal tergites with much contrasted patterns: tergite III with two median and longitudinal black thick lines, tergite IV with triangular black lines not reaching posterior margin of segment, tergite V with small triangular black lines on proximal fourth of segment, tergite VI with blackish lines faded in background, but with two rounded yellowish anterior dots, tergite VII broadly similar to tergite IV, tergites VIII and IX dark brown with two small yellowish dots in middle of segment, tergite X medium brown with two large yellowish maculae in middle of segment, reaching anterior margin. Cerci light brown, banded with dark brown segments. All patterns slightly variable from one individual to another, with respect to size and intensity of maculae, black lines and spots. Head: Labrum (Fig. 31) greatly extended laterally, with narrow tips; anterior margin almost straight, covered by numerous long and simple setae; dorsal surface with long row of small stout setae; ventral surface with scattered long and simple setae directed toward middle of labrum and concentrated near anterior margin. Mandibles strongly arched, both with lateral margin covered with several rows of long and thin setae; right mandible (Fig. 32) with outer incisor saw-like on both margins (those on outer margin often difficult to see on microscopic slides, Fig. 32 c), composed of ca. 6 teeth, inner incisor trifid (Fig. 32 b); presence of 4 fimbriate setae below incisors and 4 long and simple setae below mola (Fig. 32 a); left mandible with outer incisor saw-like on its inner margin, composed of ca. 6 teeth, and with subapical tooth in ventral position; inner incisor slender and pointed, bearing 2 setae-like structures in apical position (Fig. 33 b); 3 fimbriate setae present below incisors, followed by tuft of short setae above mola, and 5 long and simple setae below mola (Fig. 33 a). Maxillae with row of ca. 12 comb-shaped setae on anterior margin, medial ones bearing 7 – 9 teeth (Fig. 54); ventral surface of galea covered by scattered long fimbriate setae (Figs 52 – 53); proximal dentiseta bifid and fimbriate, with two distal dentisetae simple (Figs 48 – 51). Hypopharynx (Fig. 34) with well-developed superlinguae, densely covered with long and simple setae on lateral margin, ending below rounded apex of lobes; lingua without or with few short setae arranged in tuft. Labium (Fig. 35) with glossae somewhat slender and with inner margin slightly concave before apex, inner margin covered with dense simple setae, outer margin with fimbriate setae, dorsal face with single pointed bristle; paraglossae large and extending laterally, two times longer than wide. Thorax: Leg ornamentation rather similar between all three pairs, except forefemora lack row of stout spines on inner margin. Supracoxal spurs strongly developed, especially on mid- and hindlegs, acute apically. Hindleg (Figs 36, 64) with femur bearing row of long and simple setae on its outer margin, together with stout spines on distal half; inner margin with row of stout spines directed toward apex, row of long and very thin setae and incomplete submarginal row of stout spines (Fig. 36); bristles on upper face of femora elongated, with lateral margins clearly convergent, with rounded apex (Figs 37, 66); tibiae (Fig. 65) each with one row of long and very thin setae on outer margin; inner margin with only 5 – 6 short spines regularly arranged. Tarsal claw (Fig. 67) hooked, with no subapical denticles. Abdomen: Posterolateral projections weakly developed on segments I – III, more pronounced on segments IV – IX. Gills (Fig. 38) broadly oval or rounded and asymmetrical, fibrillate part well developed on gills I – VI, absent on gill VII; plate gills with tracheation apparent, gill VII (Fig. 39) more elongated than others, with inner margin concave near apex. Posterior margin of tergites (Fig. 40) with row of large and pointed spines, together with intercalary small and medium ones; several rows of microdenticles apparent. Cerci with whorl of stout and pointed setae on each segment. Eggs (from mature nymph and female): Size: 110 – 125 µm x 85 – 100 µm. Chorion covered with flat, fibrillose and rounded structures (Figs 41 a, 42 a), some bearing KCTs; on each pole, KCTs around two times size of KCTs located elsewhere (Figs 41 b, 42 b); micropyle with single margin (Figs 41 c, 42 c).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0433C4A06FF7AEB9A95C75923.taxon	discussion	Discussion. The lectotype description fits the diagnosis proposed by Ulmer (1939), but there are some intriguing points. The first one concerns the forewing venation. When describing its new genus Compsoneuriella, Ulmer stated that the wing venation was very close to the one of Compsoneuria (… ” ganz ähnlich wie bei Compsoneuria Etn ”), a statement not correct when comparing our Figs 1 and 8, with Ulmer´s (1939) fig. 143 and Ulmer´s (1924) fig. 55. In C. thienemanni, the arrangement and number of crossveins are quite different than in C. spectabilis. The “ rows ” of crossveins are absent, and the crossveins are more numerous than in the previous species, especially in the costal and subcostal fields. An answer to this “ paradox ” can be found in his 1924 paper. When redescribing C. spectabilis, he illustrated the forewing (Ulmer 1924, fig. 54) and the abdomen in lateral view (Ulmer 1924, fig. 53 B) of a pinned female imago (coll. Jacobson), mentioning that this female possesses different abdominal color patterns as well as different wing venation, but Ulmer considered them as intraspecific variation. When making the diagnosis of the genus Compsoneuriella (Ulmer 1939, p. 563), he wrote that the wing venation of the new genus is similar to the one of this female from 1924 (“ wie bei der von mir in Treubia, 6, 1924, f. 53 b, 54 abgebildeten dunkleren Form (Tuntang, Java) der C. spectabilis Etn ”). This female possesses numerous crossveins, more than 15 in the costal field and more than 12 in the subcostal one, and the general pattern is similar to the lectotype of Compsoneuriella thienemanni. One can therefore conclude that this female was misidentified by Ulmer, and rather than being a variation of C. spectabilis, constitute a female imago of C. thienemanni. The foreleg proportions illustrated in my Fig. 26 are a little bit different from the drawing made by Ulmer (1939, fig. 140) as well as his description, and this can be due to some distortion in the drawing since it has been done with the leg attached to the body, and therefore all pieces not always in the same plane. Fig. 27 presents the leg of a specimen on Ulmer’s slide, which can belong to the lectotype or to the other male imago, and which kept its original coloration. In this figure, the leg proportions are more in accordance with Ulmer’s description, except that tarsal segment 3 is longer than the first one, whereas Ulmer stated that they are of the same length. The coloration of the leg is as follow: on the dorsal face of the femur, the same small blackish maculae as in C. spectabilis can be found; on the ventral side there are three large dark spots, one apical, one subapical and one in the middle of the femur; tibia is ringed by three maculae, one apical, one in the middle and one subproximal; distal part of the tarsal segments tinted in blackish, the coloration fading towards the apex of the tarsus to become light brown on segment 4. Major differences have to be found on the genitalia. Ulmer’s drawings (1939, figs. 145 – 149) did not mention any spines or sclerotized structures, but the general shape is in accordance with the drawings. These genital structures are difficult to represent because the penis is thick due to a very developed basal sclerite on dorsal side, and the structures not easy to see. Anyway, penis lobes bear a spine of the apical sclerite together with subapical spine below the apical sclerite, an inner spine on the basal sclerite, and titillators. Although broadly similar, genitalia of C. thienemanni differ from those of C. spectabilis in several respects; the basal sclerite is much more developed and the basal spine is small, the lateral sclerite is narrower, the apical spine is less developed and the titillators are much more developed than in C. spectabilis. One misinterpretation by Ulmer needs to be corrected. When describing the maxillae, he mentioned that the crown of the galea bears two rows of comb-shape setae (Ulmer 1939, fig. 453 e); in fact the mounted nymph was about to molt, and what Ulmer interpreted as a second row is in fact the row of the next molt. The nymph of C. thienemanni described here is very closely related to the one illustrated by Braasch & Soldán (1986 a) under the name Compsoneuria thienemanni (?) from Vietnam, and reported later on also from Thailand (Braasch & Boonsoong 2010). In peculiar the abdominal patterns are almost identical between insular and continental populations (see Braasch & Soldán 1986 a, fig. 14.5 and Boonsoong & Braasch 2013, fig. 3 I). Anyway, both are distinct species, as demonstrated below. Compsoneuriella thienemanni occurs in Java and Sumatra; there is no evidence at the moment that the species inhabits other islands. The pinned specimens reported by Ulmer (1939, p. 566) from Borneo (Nanga Serawei, 12 - 18. XI. 1924, Prof. Winkler leg) could not be related to C. thienemanni, although probably belonging to the genus Compsoneuriella (see below).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043284A05FF7AEFA4971A5E19.taxon	materials_examined	Material examined. 2 nymphs, Thailand, Chiangmai Province, near Hot, Mae Nam Chaem, 200 m., 18 ° 12 ’ 07.54 ’’ N, 98 ° 36 ’ 32.85 ’’ E, IV. 2003, D. Braasch leg. [MZL]	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A043284A05FF7AEFA4971A5E19.taxon	discussion	Discussion. The Thai material differs from C. thienemanni by the following characters: maxilla with 8 – 9 comb-shape setae on the crown, instead of 12 in C. thienemanni; glossae (Fig. 68) of the labium more conical, dorsal face with two spines instead of one; bristles on the upper face of femora clearly pointed (Fig. 69), posterior margin of the abdominal tergites without rows of microdenticles (Fig. 70), apex of gill VII clearly pointed instead of rounded. As in C. thienemanni, the tarsal claw bears no subapical teeth. The Vietnamese material present similar characters, but in the absence of material in hand, it cannot be ascertained that both populations belong to the same species. Anyway, this Thai species needs to be redescribed and named on the basis of a much more abundant material than covered here (Boonsoong & Sartori, in prep.).	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0432B4A05FF7AE8A290875C07.taxon	materials_examined	Material examined. 2 nymphs, Thailand, Mae Hong Son Province, Soppong, Nam Lang River, 605 m., 19 ° 34.447 ’ N, 98 ° 16.727 ’ E, 3 – 25. IV. 2003, D. Braasch leg. [MZL]	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0432B4A05FF7AE8A290875C07.taxon	discussion	Discussion. The species has been adequately described by Braasch & Boonsoong (2010), and differs from the two previous ones by its labrum less expended laterally (Fig. 71), by the shape of the glossae, with inner margin regularly convex and outer margin slightly concave (Fig. 72), the inner margin of the fore femora with a row of spines and fine and long setae, the presence of microdenticles on the posterior margin of the abdominal tergites, and by the presence of 2 subapical teeth on the tarsal claw. Contrary to what has been previously illustrated (Braasch & Boonsoong 2010: Fig. 45), gill VII is not obtusely rounded but is apically narrowly pointed (Fig. 73). Finally the overall body coloration and pattern is not as contrasted as in the previous species.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0432B4A05FF7AEAD991BA58F0.taxon	materials_examined	Material examined. 2 nymphs, Philippines, Palawan, Mun. Narra, Estrella Falls, 7 km N. Narra, Taritien River, in forest, 100 – 200 m., 9 ° 18 ’ N, 118 ° 23 ’ E, 5. IV. 1992, M. Zerning leg. [MZL] The nymph of this species has been incompletely described, and some important characters have not been mentioned or drawn. A complementary description is given below.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0432B4A05FF7AEAD991BA58F0.taxon	description	Complementary description. Coloration: see Fig. 82. Head: Labrum (Fig. 74) moderately extended laterally, with narrow tips; anterior margin slightly concave, covered by numerous long and simple setae. Mandibles with 10 – 11 fimbriate or simple setae below inner incisor (Fig. 76); right mandible with 5 long and simple setae below mola; left mandible with inner incisor slender and bifid, with some minute teeth (Fig. 76); 6 long and simple setae below mola. Maxillae each with row of ca. 9 combshaped setae on anterior margin, medial comb-shaped setae with 11 – 12 teeth each; ventral surface of galea covered by scattered long and simple setae (Fig. 77); proximal dentiseta bifid and fimbriate, two distal dentisetae simple. Hypopharynx with well-developed superlinguae, densely covered with long and simple setae on lateral margins, ending at tip of rounded apex of each lobe. Labium (Fig. 75) with glossae with inner margin regularly convex and outer margin slightly concave. Thorax: Hindleg with femur similar to other species, but tibia with row of submarginal bristles close to inner margin. Tarsal claw with 2 subapical teeth. Abdomen: Posterior margin of tergites with row of large and pointed spines, together with intercalary small ones; several rows of microdenticles apparent.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
03A087A0432B4A05FF7AEAD991BA58F0.taxon	discussion	Discussion. The nymph of C. tagbanua fits the present concept of Compsoneuriella with one major exception: the scattered setae on the galea are not fimbriate but simple, such as those found in the Afrotropical genus Notonurus. At the moment, it is considered as a case of convergence. Besides this, C. tagbanua does possess the characteristic dentisetae typical for Compsoneuriella, and all other characters. In addition to the scattered setae of the galea, this species differs from all others by the high number of setae below the inner incisor of the mandibles.	en	Sartori, Michel (2014): The concept of Compsoneuria Eaton, 1881 revisited in light of historical and new material from the Sunda Islands (Ephemeroptera: Heptageniidae: Ecdyonurinae). Zootaxa 3835 (1): 1-32, DOI: 10.11646/zootaxa.3835.1.1
