Compsoneuria Eaton, 1881

Webb, J. M., Braasch, D. & Mccaferty, W. P., 2006, Reevaluation of the Genera Compsoneuria Eaton and Trichogenia Braasch & Soldán (Ephemeroptera: Heptageniidae), Zootaxa 1335, pp. 55-68 : 56-60

Compsoneuria Eaton

Compsoneuria Eaton, 1881: 23

Compsoneuriella Ulmer, 1939: 563 ; (syn.) Braasch and Soldán, 1986a: 46 Cosmetogenia Eaton, 1883: 18 (obj. syn.)

Notonurus Crass, 1947: 126 ; (syn.) Gillies, 1963: 232

Wang and McCafferty (2004) placed Compsoneuria as the sister group of Trichogenia in the Heptageniinae tribe Compsoneuriini because larvae described as C. spectabilis by Ulmer (1939) have a row of setae on the ventral surface of the maxillae, triangular glossae, no distinct dorsal apical projection on the femora, two rows of setae on the lateral margin of the mandibles, and a wide labrum. Male adults of C. spectabilis , however, possess Ecdyonurinae characters such as parallel-sided medial margins of the mesothoracic furcasternum as well as forewings with a reduced number of crossveins. Because Ulmer (1939) did not rear any larvae of C. spectablis, Jensen (1972) , Braasch and Soldán (1986a), and Edmunds and Polhemus (1990) viewed this association with caution. We found Ulmer's (1939) association to be incorrect because forewings dissected out of a mature larva similar to that described by Ulmer (1939) have numerous crossveins and are shaped differently than those of adults of C. spectabilis , and also have other Heptageniinae characters noted above. Therefore, the true larva of C. spectabilis is still unknown. The larva described by Ulmer (1939) as C. spectabilis is an unnamed species of Trichogenia which is further discussed under that genus (see below).

Based on the evidence provided above, Compsoneuria can no longer be included in the Heptageniinae and is placed in Ecdyonurinae , with the name Compsoneuriini accompanying its type genus to Ecdyonurinae . The potential relationships of Compsoneuria to other Ecdyonurinae are discussed below. The phylogenetic concept of the Heptageniinae clade defined by larval characters that Wang and McCafferty (2004) named Compsoneuriini remains the same; however, this concept is now relevant only to the genus Trichogenia . Given the constraints of a strict phylogenetic classification, we must establish a new tribe, Trichogeniini Webb & McCafferty N.TRIB., allowing each of the additional eight Heptageniinae genera to retain their generic status.

Braasch and Soldán (1986a) noted that adults of Compsoneuriella thienemanni Ulmer, 1939 , the type species of Compsoneuriella Ulmer , were congeneric with C. spectabilis adults and synonymized Compsoneuriella with Compsoneuria . Wang and McCafferty (2004) did not recognize this synonymy because they believed Compsoneuria to be a member of the Heptageniinae , and placed Compsoneuriella as a junior synonym of Thalerosphyrus instead, based on the shared possession of long, pointed supracoxal sclerites in the larvae. Examination of larvae and males of several species of Thalerosphyrus and Compsoneuriella indicates, however, that the two are distinct. The penes of Compsoneuriella are slightly expanded laterally, are only partially fused medially, have distinct dorsal sclerites both apically and laterally, and have ventral and dorsolateral spines. The penes of Thalerosphyrus are not expanded laterally, are fused medially for most of their length, do not have ventral or dorsolateral spines (small dorsolateral spines may be present in some species), and do not have as well-developed apical sclerites. Additionally, the forewings of the two genera differ in that those of Compsoneuriella tend to have a reduced number of crossveins and a slightly sigmoidal bend in the costa and subcosta, whereas those of Thalerosphyrus have numerous crossveins and a straight costa and subcosta. The legs of Compsoneuriella have numerous black spots that are not present in Thalerosphyrus . The larvae of Thalerosphyrus and Compsoneuriella also differ in many characters. Thalerosphyrus larvae, for example, have well developed posterolateral spines on the abdomen, simple setae on the inner surface of the hindtibiae, slight posterolateral extensions of pronotum, distinct thickening of the anterior margin of the head capsule, do not possess any black spotting on the head capsule or femora, and the glossae are subquadrate and laterally expanded. Compsoneuriella larvae, on the other hand, generally have shorter posterolateral spines on the abdominal segments, have fimbriate setae on the inner surface of the hindtibiae, generally lack posterolateral expansions on the pronotum, do not have the anterior margin of the head capsule thickened, usually have conspicuous black spots on the head capsule and/or femora, and the glossae are narrow and pointed apically. For these reasons, we no longer recognize any synonymy of Compsoneuriella with Thalerosphyrus and because all of the adult characters of Compsoneuriella are present in Compsoneuria , we recognize the previous synonymy of Compsoneuriella with Compsoneuria . All African species transferred to Thalerosphyrus by McCafferty (2003) are recombined with Compsoneuria .

The phylogenetic position of Compsoneuria (and Compsoneuriini) within the Ecdyonurinae is unclear. The possession of ventral spines on the penes, the general shape and the sclerotization of the penes, and the possession of black spotting on the head capsule indicate Compsoneuria may be related to members of the tribe Leucrocutini Wang & McCafferty. However, unlike other members of the Leucrocutini, the apical setae on the ventral side of the galealaciniae are simple rather than fimbriate, and the distal dentisetae on the maxillae are branched rather than simple, indicating a closer relationship to the Notacanthurini Wang & McCafferty and Atopopini Wang & McCafferty. The long supracoxal sclerites of the larvae are similar to those found in most Thalerosphyrus (a member of Atopopini) but we believe this to be a case of convergence as Compsoneuria lacks apomorphies of the Atopopini such as a thickened anterior margin of the larval head capsule. In most males of Atopopini, the first foretarsal segment is greater than 0.5X the length of the second segment; in Compsoneuria , the length varies from 0.3-1.0X the length of the second segment.

We presently consider Compsoneuria to include the following species:

Compsoneuria bequaerti (Navás) AFROTROPICAL

Adenophlebia bequaerti Navás, 1930: 316 (orig.)

Adenophlebia eatoni Navás, 1931a: 271 ; (syn.) Demoulin, 1956: 45 Adenophlebia inflexa Navás, 1932: 285 ; (syn.) Demoulin, 1956: 45 Notonurus cooperi Crass, 1947: 127 ; (syn.) Demoulin, 1970: 97 Compsoneuriella cooperi ; (comb.) Gillies, 1963: 232

Compsoneuriella bequaerti ; (comb.) Gillies, 1963: 232

Notonurus bequaerti ; (comb.) Demoulin, 1956: 45

Compsoneuria bequaerti ; (comb.) Braasch and Soldán, 1986a: 46 Thalerosphyrus bequaerti ; (comb.) McCafferty, 2003: 792

Compsoneuria cingulata (Navás) ORIENTAL

Thalerosphyrus cingulatus Navás, 1933: 18 (orig.)

Compsoneuria cingulata ; (comb.) Braasch and Soldán, 1986a: 46 Compsoneuria diehli Braasch & Soldán ORIENTAL Compsoneuria diehli Braasch & Soldán, 1986a: 48 (orig.)

Compsoneuria flowersi Braasch & Soldán ORIENTAL Compsoneuria flowersi Braasch & Soldán, 1986b: 59 (orig.)

Compsoneuria josettae (Sartori & Elouard) N.COMB. AFROTROPICAL Thalerosphyrus josettae Sartori & Elouard, 1996: 125 (orig.)

Compsoneuria lieftincki (Ulmer) N.COMB. ORIENTAL Heptagenia lieftincki Ulmer, 1939: 571 (orig.)

Compsoneuria njalensis (Kimmins) AFROTROPICAL Afronurus njalensis Kimmins, 1937: 433 (orig.)

Compsoneuriella njalensis ; (comb.) Gillies, 1963: 232

Compsoneuria njalensis ; (comb.) Braasch and Soldán, 1986a: 46 Thalerosphyrus njalensis ; (comb.) McCafferty, 2003: 792

Compsoneuria sinuosa (Navás) AFROTROPICAL Adenophlebia sinuosus Navás, 1931b: 109 (orig.)

Notonurus sinuosus ; (comb.) Demoulin, 1970: 98

Compsoneuria sinuosa ; (comb.) Braasch and Soldán, 1986a: 46 Ecdyonurus sinuosus ; (comb.) Kluge, 1988: 298

Thalerosphyrus longinosi McCafferty, 2003: 792 (unnecessary name change)

Compsoneuria spectabilis Eaton ORIENTAL Compsoneuria spectabilis Eaton, 1881: 23 (orig.)

Compsoneuria thienemanni (Ulmer) ORIENTAL

Compsoneuriella thienemanni Ulmer, 1939: 564 (orig.)

Compsoneuria thienemanni ; (comb.) Braasch and Soldán, 1986a: 46 Thalerosphyrus thienemanni ; (comb.) Wang and McCafferty, 2004: 17

Compsoneuria tortinervis (Navás) AFROTROPICAL Adenophlebia tortinervis Navás, 1930: 317 (orig.)

Notonurus tortinervis ; (comb.) Demoulin, 1956: 48

Compsoneuriella tortinervis ; (comb.) Gillies, 1963: 232

Compsoneuria tortinervis ; (comb.) Braasch and Soldán, 1986a: 46 Thalerosphyrus tortinervis ; (comb.) McCafferty, 2003: 792

Thalerosphyrus is restricted presently to T. bishopi Braasch & Soldán, 1986 a ; T. determinatus ( Walker, 1853) ; T. flowersi Venkataraman & Sivaramakrishnan, 1987 ; T. sinuosus ( Navás, 1933) ; T. sumatranus ( Ulmer, 1939) ; T. torridus ( Walker, 1853) ; and T. vietnamensis ( Dang, 1967) . While T. ethiopicus Soldán, 1977 , and T. separatus Nguyen & Bae, 2004 are likely members of Afronurus Lestage we are retaining them in Thalerosphyrus until we can examine specimens. Similarly, Thalerosphyrus melli Ulmer, 1925 , is probably a member of Epeorus Eaton , but we have not examined specimens to confirm this and we are therefore retaining it in Thalerosphyrus for the time being.

Material examined. Compsoneuria flowersi , 2ɗ, Indonesia, Java, GC Crampton [ PERC]. C. lieftincki , 2ɗ syntypes, Indonesia, West Java, 150m, Djasinga, Tjibarangbang Fluss, abends an der Lampe im Flussbett, 6-VII-1935, Lieftinck [ FAMU]. C. njalensis , 5ɗ 2Ψ, South Africa, no other data [ PERC]; 1ɗ, South Africa, Skandinawiedrift, 30-III-1965 [ PERC]. C. spp. 1ɗ, India, Kalial (Kerala), 7-I-1962, F. Schmid [ PERC]; 1ɗ, Madagascar, Majunga Prov, 25km SW Ambalanjankomby, XI-1962, ED Cashatt [ PERC]; 1ɗ 5Ψ, South Africa, Vaal R at Skandinawiedrift, 30-III-1965, 28-IV-1965, 10-IV-1969, HJ Schoonbee [ PERC]; 1ɗ, Sri Lanka, Sabaragamuwa Prov, Helihul-Oya, 1-III-1962 [ PERC]; 1ɗ, Sri Lanka, Sabaragamuwa Prov, Rakwana, 27-II-1962 [ PERC]. C. spectabilis , 3ɗ, Indonesia, W Java, Buitenzorg (now Bogor), 13-II-1932, VII-1932, Lieftinck [ PERC]. C. thienemanni , 3ɗ, Malaysia, Sabah, Mansuncun R NE of Kota Kinabalu, 18-VIII-1972, WL and JG Peters [ PERC]; 26ɗ 11Ψ, Vietnam, Gia Lai, An Khe District, Cha R 3km E Buoenloy, 25-VI-1996, B Hubley, DC Currie, J Swann ROM 961090 [ ROM]. C. tortinervis ?, 1ɗ, B Congo, 9mi NW of Bunyakiri, 8-IX-1957 [ PERC].