Bugilliesia Lugo-Ortiz & McCafferty 1996

Kluge, Nikita J., 2015, First description of winged stages of Thraulobaetodes Elouard & Hideux 1991 and reclassification of Rhithrocloeoninae (Ephemeroptera, Baetidae), Zootaxa 3949 (4), pp. 491-514 : 506-508

Plesiomorphon Bugilliesia Lugo-Ortiz & McCafferty 1996

( Figs 50–55 )

Rank-free hierarchical name: Bugilliesia /g(1).

Type species: Afroptilum guineense Gillies 1990a .

References: Gattolliat et al. 2009: larva, imago; Kluge 2012: larva, imago.

Composition. Seven described species belong without doubt in Bugilliesia , being known as male imagoes, which have characteristic genital structure and well developed hind wings; these are: biloba Gattolliat 2006 [ Bugilliesia ]; cavalliensis Gattolliat 2006 [ Bugilliesia ]; griseum Gillies 1990 [ Afroptilum ]; guineense Gillies 1990 [ Afroptilum ]; notabile Kimmins 1956 [ Centroptilum ]; sudanense Ulmer 1916 [ Centroptilum ]; truncata Gattolliat 2006 [ Bugilliesia ]. Among them, only griseum [ Afroptilum ], guineense [ Afroptilum ] and sudanense [ Centroptilum ] larvae have been associated with the imago by rearing ( Gillies 1990a). Besides this, two species, margaretae Gattolliat & Barber-James 2009 [ Bugilliesia ] and mirandei Lugo-Ortiz & McCafferty 1997 [ Cheleocloeon ] are known as larvae only, and their relation with the species described as imagoes is unknown ( Gattolliat et al. 2009). Placement of the species nitidum Ulmer 1916 [ Centroptilum ] (= nigroalbum Navas 1932 [ Cloeon ] = bredoanum Navas 1933 [ Cloeon ]) to Bugilliesia is not grounded ( Kluge 2012). Besides these formally described species, some undescribed species belong to Bugilliesia (see below).

Material examined: Bugilliesia notabilis : UGANDA, Jinja, F.I.R.R.I., 6–10.VII.2007, coll. N. Kluge: 8 I ♀; Bujagali Falls 6.VII.2007, coll. N. Kluge: 1 S♂, 12 I ♀. ZAMBIA, Mwinilunga District: River Mudanyama in Mwinilunga, 14–17.VIII.2014, coll. N. Kluge & L. Sheyko: 1 I ♀; River Lwakela, 22 km N Mwinilunga, 18– 21.VIII.2014, coll. N. Kluge and L. Sheyko: 1 I ♀.

Bugilliesia sudanensis : SUDAN, White Nile, 26.I.1964, coll. A.V. Monakov: 4 larvae; Ed-Dueim, 19.XI.1963, coll. A.V. Monakov: 1 male larva.

Bugilliesia grisea : UGANDA: Victoria Nile at Bujagali Falls, 7.VII.2007, coll. N. Kluge: 1 female larva; Kasese District, River Nyamagasan near Kiburara, 8–13.VIII.2007, coll. N. Kluge: 1 mature female larva with well-developed eggs.

Bugilliesia biloba : MALI, Bafing bei Tinko, 1.X.1991, coll. D. Tobias: 1 I ♂.

Bugilliesia sp. K: Kluge 2012: British East Africa, Kahavati, 20.I.1911, coll. Svatosh: 1 I ♂.

Bugilliesia sp. N: Kluge 2012: UGANDA, Jinja, F.I.R.R.I., 6–10.VII.2007, coll. N. Kluge: 5 I ♂, 10 I ♀; Bujagali Falls 6.VII.2007, coll. N. Kluge: 1 S-I♂.

Bugilliesia sp. NZ: ZAMBIA, River Zambezi near Victoria Falls 25–31.VIII.2014, coll. N. Kluge & L. Sheyko: 1 gynandromorph imago. This specimen has one left well-developed gonostylus of the same structure as in Bugilliesia sp. N, with the same numerous spines on inner sides of 2nd segment ( Kluge 2012: Figs 40–42 ); other structure as in female, with well developed eggs ( Fig. 53 ). Differs from Bugilliesia sp. N by presence of reddish maculae on abdomen, which are situated asymmetrically (possibly, individual feature of gynandromorph).

Plesiomorphies. Hind wings developed, with hooked costal process (in contrast to Mutelocloeon and Rhithrocloeon , whose hind wing is reduced or lost). Larval claw retains both rows of denticles ( Fig. 55 ) (in contrast to Mutelocloeon , whose larval claw lack denticles, and Rhithrocloeon , whose larval claw has one row of denticles).

Other characters of larva. Described by Gattolliat et al. (2009) and Kluge (2012).

Egg structure ( Figs 50–54 ). Chorion with characteristic structure in form of round impressions, some interconnecting, some separated by partitions with flat surfaces; while all impressions usually have similar size and shape, partitions are variable in thickness and shape, and absent in some places ( Figs 50, 51 ). This distinguishes Bugilliesia from many other mayflies, whose eggs have a net-like relief with closed cells, where partitions have equal thickness, while cells can have unequal size and shape ( Figs 47, 48 ). Very similar structure on egg chorion is found in the examined species of Bugilliesia — B. notabilis ( Fig. 51 ), B. grisea ( Fig. 54 ) and Bugilliesia sp. N ( Fig. 51 ). In Bugilliesia sp. NZ relief is often smoothed out; if present, cells are somewhat smaller and less regular than in other species, and partitions between them are thick and integral ( Fig. 53 ).

Just the same relief of egg chorion as in Bugilliesia , is found in Mutelocloeon (see below and Figs 56–59 ), while in Rhithrocloeon relief of egg chorion is quite different (see below and Figs 65–70 ).

Comments. Based on reared material, Gillies (1990a) described larvae of Afroptilum sudanense , A. griseum and A. guineense (recently attributed to Bugilliesia ) without detailed figures of their claws; he only reported that "tarsal claws toothed and without a seta before apex" ( Gillies 1990a: 114) and gave drawings of total legs of A. griseum , where claws are too small to show their teeth (ibid., Figs 79–81). There are no other descriptions of Bugilliesia which would be based on larval exuviae associated with reared imagoes. Lugo-Ortiz & McCafferty (1996) established the new genus Bugilliesia without examination of any material. Based on literature data only, they stated that in Bugilliesia "tarsal claws poorly dentate" ( Lugo-Ortiz & McCafferty 1996: 184) and that "larvae of ... Bugilliesia ... lack the 2 rows of denticles on the tarsal claw" ( Lugo-Ortiz & McCafferty 1998: 2). Gattolliat et al. (2009: 169) correctly state that "there are always two rows of abundant teeth in Bugilliesia " and gave illustration of the claw of B. margaretae (ibid., Fig. 32 ). Larvae of B. grisea and B. sudanensis , examined by me, have two equal and symmetrically arranged rows of teeth, each with several (2–5) most distal teeth much larger than more proximal ones ( Fig. 55 ).

Figures

FIGURES 50 – 55. Bugilliesia spp.: 50 – 54, eggs: 50, 51, Bugilliesia notabilis (extracted from female imago); 52, Bugilliesia sp. N (extracted from female imago); 53, Bugilliesia sp. NZ (extracted from gynandromorph imago); 54, Bugilliesia grisea (extracted from female larva). 55, Bugilliesia sudanensis, larval claw.

FIGURES 38 – 44. Thraulobaetodes cumminsorum. 38, apex of abdomen of male imago, lateral view; 39, genitals of male subimago, ventral view; 40, subimaginal exuviae of genitals; 41, genitals of male imago of the same individual, ventral view; 42, female imago; 43, male imago (arrows show protuberances—remnants of larval spines on abdominal terga); 44, mesonotum, lateral-dorsal view.

FIGURES 45 – 48. Thraulobaetodes cumminsorum. 45, 46, subimaginal exuviae spread on slide (on the same scale): 45, left half of mesonotum; 46, part of mesopleuron and metapleuron, with fragments of fore and middle legs. 47, 48, egg and its enlarged fragment (extracted from female imago). Abbreviation: psa, postsubalar sclerite of mesothorax.

FIGURES 56 – 60. Mutelocloeon bihoumi. 55 – 59, eggs (extracted from female imago); 60, male imago.

FIGURES 65 – 70. Rhithrocloeon spp., eggs. 65, 66, Rhithrocloeon insuetum (extracted from female imago); 67, 68 Rhithrocloeon munyagae (poorly developed, extracted from female larva); 69, 70, Rhithrocloeon elgonensis (extracted from female larva).

FIGURES 31 – 37. Thraulobaetodes cumminsorum. 31, fore and hind wing on the same scale; 32, hind wing, enlarged; 33, apex of fore wing of another individual; 34, head of female imago; 35, outline of abdominal segment IX of mature male larva with developing subimaginal genitals inside (in left half muscles shown by interrupted lines); 36, exuviae of left gonostylus of male subimago; 37, genitals of male imago, the same individual (muscles shown by interrupted lines; in left half gonostylar muscle not shown, internal sclerotized part of penis shown by interrupted line and dotted; in right half cuticular pigmentation shown by dots). Abbreviations: gs 1, gs 2, gs 3, segments of gonostylus; m. gs, gonostylar muscle; m. gv 1, m. gv 2, muscles of gonovectis; m. s, sterno-styligeral muscle; us, unistyliger.