Teloganodes Eaton 1882

Kluge, Nikita, 2023, Reclassification of Teloganodes Eaton 1882 (including subgenera Dudgeodes Sartori 2008 and Derlethina Sartori 2008) with a new species from Sulawesi (Ephemeroptera, Teloganodidae), Zootaxa 5244 (6), pp. 527-552 : 529

Genus Teloganodes Eaton 1882

( Figs 1–101 )

Genus Teloganodes Eaton 1882: 208 (imago); Eaton 1883 –1888 (imago); Ulmer 1939: 512 (imago), 627 (larva). Teloganodes/fg1: Kluge 2004: 320 (larva, subimago, imago).

Macafertiella Wang in Wang & McCafferty 1996: 15 (synonymized by Sartori, Peters & Hubbard 2008).

Dudgeodes Sartori in Sartori, Peters & Hubbard 2008: 25 , syn. n.

Derlethina Sartori in Sartori, Peters & Hubbard 2008: 44 , syn. n.

Type species: Cloe tristis Hagen 1858 .

Systematic position. Teloganodes belongs to the taxon Ephemerella/fg1, or superfamily Ephemerelloidea , which is characterized by peculiar structure of tergalii with lamellate dorsal lobe and two-branched ventral lobe bearing processes on outer sides of its branches (character 7 of Ephemerella/fg 1 in Kluge 2004: 294) ( Figs 1 , 50–57 ), presence of crossvein cua-cup near base of fore wing (character 6 of Ephemerella/fg 1 in Kluge 2004: 293) ( Fig. 94 ), certain reductions in larval maxillae and labium (characters 1–3 of Ephemerella/fg 1 in Kluge 2004: 293), peculiarities of larval thorax (characters 4–5 of Ephemerella/fg 1 in Kluge 2004: 293), loss of 4th segment of gonostylus (character 8 of Ephemerella/fg 1 in Kluge 2004: 294) ( Figs 42 , 97 ), and some other characters (characters 9–19 of Ephemerella/fg 1 in Kluge 2004: 294–295).

The holophyletic taxon Ephemerella/fg1 is divided into two holophyletic taxa, Ephemerella/fg2 (or family Ephemerellidae ) and Pantricorythi Kluge 2014, or Tricorythus /fg1 (usually treated as two or more families).

The taxon Pantricorythi is characterized by retention of fully developed tergalii on the 2nd abdominal segment ( Figs 1 , 62 ), long 1st segment of the 3-segmented gonostyli ( Figs 42 , 97 ) and peculiar arrangement of setal rows on anterior (dorsal) side of larval fore femur, which have a regular row crossing the femur near its middle and continuing distally along the outer margin of the femur ( Figs 19–20 ). Among Teloganodes s. l., this arrangement of setal rows occurs in most species; in the known species of subgenus Teloganodes s. str., the transverse setal row of fore femur is either secondarily lost ( Sartori et al. 2008: fig. 59–60) or detached from the row on outer margin ( Selvakumar et al. 2014: fig. 50).

The taxon Pantricorythi contains Vietnamella Tshernova 1972 , Austremerella Riek 1963 , Ephemerellina Lestage 1924 , Lithogloea Barnard 1932 , Lestagella Demoulin 1970 , Nadinetella McCafferty & Wang 1998 , Manohyphella Allen 1973 , Indoganodes Selvakumar et al. 2014 , Melanemerella Ulmer 1920 , Teloganella Ulmer 1939 , Teloganodes Eaton 1882 , Ephemerythus Gillies 1960 , Dicercomyzon Demoulin 1954 , Machadorythus Demoulin 1959 , the taxon with the non-typified circumscriptional name Tricorygnatha Kluge 2004 and the large taxon with the rank-free hierarchical name Leptohyphes/fg1 ( Kluge 2004).

Various authors either arrange these taxa among the families Tricorythidae Lestage 1942 and Ephemerellidae Klapálek 1909 , or accept greater number of families. In the past, the genus Teloganodes was placed in the family Ephemerellidae , which was paraphyletic in such sense. Currently, it is used to place Teloganodes in a family Teloganodidae Allen 1965 , which has no generally accepted circumscription and diagnosis.

Figures

FIGURE 1. Tergalii figured by Kluge (2004: figs 96E–H) and two classifications of Teloganodes. E, tergalii of T. (Teloganodes) kodai; F, tergalii of T. (Dudgeodes) romani (= Teloganodes/fg1 sp. T4: Kluge 2004); G, tergalii of T. (Dudgeodes) jacobusi (= Teloganodes/fg1 sp. T2: Kluge 2004); H, tergalii of T. (Derlethina) eloisae (= Teloganodes/fg1 sp. T3: Kluge 2004).

FIGURES 2–7. Details of Teloganodes s.l. 2–4, Teloganodes (Derlethina) eloisae, mature larvae (Malaysia, Sabah, Kindssang, Ranau, Negeri, 9.IX.1994, coll. S.C. Kang): 2, apical part of fore protopteron; 3, hind protopteron; 4, hind leg (si, outer margin of subimaginal femur developing inside). 5, egg of T. (Teloganodes) insignis (Sri Lanka, south-eastern slope of Sri Pada, river Battulu oya, Fishing Hut, 10.II.2020, coll. N. Kluge & L. Sheyko). 6–7, T. (Dudgeodes) celebensis, dorsal cuticle from labrum: 6, focus on proximal part; 7, focus on distal part.

FIGURES 8–11. Teloganodes (Dudgeodes) celebensis. 8, young larva in dry condition, to show relief; 9–11, fore, middle and hind legs of young larva to show black hypodermal coloration.

FIGURES 12–18. Teloganodes (Dudgeodes) celebensis, larval cuticle. 12, latero-posterior margin of head; 13, left half of pronotum and mesonotum; 14–18, left half of abdomen and its enlarged fragments.

FIGURES 19–26. Teloganodes (Dudgeodes) celebensis, exuviae of larval legs. 19–20, fore leg and its fragment; 21, middle leg; 22–26, hind leg and its fragments.

FIGURES 27–30. Teloganodes (Dudgeodes) celebensis. 27–28, left half of subimaginal exuviae of mesonotum; 29–30, hind wing in falling and transmitted light. Abbreviations: acz, achromozone; cz.a, anterior chromozone; cz.l-p, latero-posterior chromozone; cz.s, chromozonal suture; LPs, lateroparapsidal suture; MPs, medioparapsidal suture; MS, medioscutum; SMS, submedioscutum.

FIGURES 31–39. Teloganodes (Dudgeodes) celebensis. 31, male imago; 32, abdomen of male imago; 33, head and thorax of female imago; 34, fore leg of female imago; 35–36, fore legs of male imagines; 37, middle leg of male imago; 38–39, exuviae of fore and middle legs of male subimago (32–39 with equal magnification).

FIGURES 40–45. Teloganodes (Dudgeodes) celebensis. 40–41, penis of male imago, dorsal view; 42, genitalia, ventral view; 43, cuticle of abdominal sternum IX and protopenis of male larva; 44, developing subimaginal penis extracted from the same larva; 45, position of developing penis in larval cuticle of protopenis.

FIGURES 46–49. Teloganodes (Dudgeodes) celebensis, eggs.

FIGURES 50–60. Tergalii. 50–53, Teloganodes (Dudgeodes) celebensis, tergalii II–V; 54–57, Teloganodes (Dudgeodes) rasus sp. n., tergalii II–V; 58–60, tergalius V of two individuals (59–60, holotype).

FIGURES 61–62. Teloganodes (Dudgeodes) rasus sp. n. 61, male larva of last instar; 62, younger female larva.

FIGURES 63–68. Teloganodes (Dudgeodes) rasus sp. n. 63–64, left and right mandibles; 65, maxilla, 66, apex of maxilla; 67, hypopharynx and superlingua; 68, labium. Abbreviations: in, incisor; kd, kinetodontium; m, mola; prs, prostheca.

FIGURES 69–74. Teloganodes (Dudgeodes) rasus sp. n., larval exuviae (holotype). 69–70, parts of head; 71, half of pronotum and mesonotum; 72, abdomen; 73, fragment of abdominal segment I; 74, fragment of abdominal segment VIII.

FIGURES 75–83. Teloganodes (Dudgeodes) rasus sp. n., larval exuviae of legs (holotype). 75–76, fore leg and its fragment; 77, middle leg; 78–82, hind leg and its fragments; 83, claw.

FIGURES 84–92. Teloganodes (Dudgeodes) rasus sp. n., male (holotype). 84, head and thorax of imago; 85, subimaginal exuviae of abdomen; 86, imaginal abdomen; 87–88, two portions of imaginal cercus; 89–90, subimaginal exuviae of fore and hind legs; 91–92, imaginal fore and hind legs.

FIGURES 93–94. Teloganodes (Dudgeodes) rasus sp. n., wings of male imago (holotype). 93, in falling light; 94, in transmitted light (arrow shows constant crossvein cua-cup).

FIGURES 95–101. Teloganodes (Dudgeodes) rasus sp. n., male (holotype). 95–96, penis, dorsal view; 97, genitalia, ventral view; 98–99, halves of subimaginal exuviae of mesonotum, in Canadian balsam and in dry condition; 100–101, hind wing, in falling and transmitted light.