Eupotemus limicola ( Delève, 1967 ), Differential

Fikáček, Martin, Matsumoto, Keita, Perkins, Philip, Prokin, Alexander, Sazhnev, Alexey, Litovkin, Stanislav & Jäch, Manfred A., 2021, The family Epimetopidae (Coleoptera: Hydrophiloidea): review of current knowledge, genus-level phylogeny, and taxonomic revision of Eupotemus, Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 61 (1), pp. 1-34 : 18-22

Eupotemus limicola ( Delève, 1967)

( Figs 8M–Q , 9G–I )

Material examined. Hඈඅඈඍඒඉൾ: J ( MRAC): ʽHOLOTYPUS // Biot. No 14 / banks de / vase // I. R. S. A. C. – MUS. CONGO / Kivu: Kitutu, terr. Mwenga / riv. Lubushwa 650 m / B. 14 N. Leleup 12-IV-58 // TYPE // J. Delève det. 1966 / Eumetopus / limicola n. sp. // Aedeagus /drawn by / P. D. Perkinsʼ. P*ò*©*òඍඒඉൾඌ: 9 ♀♀ ( MRAC), 1 J 1 ♀ ( IRSNB): same locality data as the holotype.

Differential diagnosis. Very similar to the other species of the E. limicola species group from which it can be reliably distinguished by the male genitalia only. The aedeagus ( Figs 8M–Q ) differs in rounded apex from E. bilobatus and in the widely expanded parameres in lateral view from all species except E. smithi . From E. smithi it may be distinguished by the much shorter phallobase, more deeply excised ventral fork of the median lobe, and the complete ridge on elytral interval 3.

Redescription. Body 2.5–3.1 mm long (holotype 2.7 mm) and 1.4–1.7 mm wide (holotype 1.5 mm). Dorsal surface brown to dark brown. Habitus and sculpture as in Figs 9G–I ; ridge on elytral interval 3 not interrupted; ridge on interval 5 interrupted anteriorly and sometimes also throughout its length or just before its posterior end; ridge on interval 7 complete throughout or interrupted in posterior 0.2–0.3 of elytral length. Elytral punctures connected by low elevated line. Aedeagus ( Figs 8M–Q ): 0.90 mm long. Parameres ca. 1.8× longer than phallobase, moderately bisinuate on outer face, strongly widen- ed apically in lateral view. Median lobe with ventral impression narrow in lateral view; apical disc ca. 1.6× longer than wide, concave in lateral view, its apex very weakly sinuate. Phallobase basally with narrow, slightly asymmetrical manubrium.

Biology. No data available.

Distribution. The species was originally described from the Democratic Republic of Congo (locality of most specimens examined) and Côte d’Ivoire (based on a single female). The female from Côte d’Ivoire was not found in the collections. However, the newly collected specimens from Côte d’Ivoire all belong to a very similar but different species ( E. smithi sp. nov.) and we suppose the same applies for the female paratype examined by Dൾඅජඏൾ (1967). We hence exclude Côte d’Ivoire from the distribution range of E. limicola .

Eupotemus ophioglossus sp. nov. view. From E. limicola it differs in a shallowly excised

( Figs 8G–I , 10D–F ) ventral fork. Eupotemus smithi differs from all species of

Material examined. Hඈඅඈඍඒඉൾ: J ( NHMW): ʽGABON / Bissok the group externally in the largely interrupted keels on the (Oyem) / 3.- 10.2.1991 / leg. Bilardo // Eupotemus limicola (Del.) / det elytral intervals 3, 5 and 7. Jäch 1998ʼ. P*ò*©*òඍඒඉൾ J: 1( NHMW): ʽTOGO: Plateux / Pref. Kloto, Description. Body 2.6–3.1 mm long (holotype 2.8 mm) ca. 5 km from / Konda (village), 9.II.2006 / leg. Komarek & Houngue and 1.4–1.7 mm wide (holotype 1.5 mm). Dorsal surface (28) // 06°58’05.3”N 00°34’18.2”E, ca. 510 m a.s.l / small stream in prim. forestʼ. brown to black. Habitus and sculpture as in Figs 10A–C ; ridge on elytral interval 3 interrupted posteriorly; ridge on

Differential diagnosis. Very similar to the other species of interval 5 interrupted in posterior half to fourth; ridge on the E. limicola species group from which it can be reliably interval 7 interrupted in posterior half to third of elytral distinguished by the male genitalia only. The aedeagus length. Elytral punctures connected by low elevated line. is unique in the shape of the ventral fork of the median Aedeagus ( Figs 8J–L, X–Y ): 0.90 mm long. Parameres lobe which is very deeply excised ( Fig. 8V ), otherwise ca. 1.8× longer than phallobase, strongly sinuate on outer it resembles that of E. cameroonensis by the moderately face, strongly widened apically in lateral view. Median widened apex of the paramere in lateral view ( Figs 8H, W ) lobe with narrow ventral impression in lateral view; apical and in the narrow elongate apical disc of the median lobe disc ca. 1.5× longer than wide, concave in lateral view, its ( Fig. 8G ). Externally, it can be only distinguished from E. apex rounded. Phallobase basally with narrow, slightly smithi in the complete ridge on elytral interval 3 (see under asymmetrical manubrium. E. smithi for details). Variation. The single male from Tai National Park is Description. Body 2.6–2.7 mm long (holotype 2.6 mm) brachypterous, smaller than the remaining specimens exa- and 1.3–1.4 mm wide (holotype 1.3 mm). Dorsal surface mined (2.6 mm long), and the ridges on its elytral intervals brown to black. Habitus and sculpture as in Figs 10D–F ); 3, 5 and 7 are completely subdivided into a series of elonridge on elytral interval 3 not interrupted; ridge on interval gate tubercles. Yet, it corresponds with the macropterous 5 interrupted anteriorly and posteriorly; ridge on interval specimens from Yéale village, with which it also agrees in 7 interrupted in posterior 0.2–0.4 of elytral length. Elytral all details of the aedeagus morphology. We hence consider punctures connected by low elevated line. Aedeagus (Figs this specimen conspecific to the holotype and hypothesize 8G–I, V–W): 0.80 mm long. Parameres ca. 2.2× longer that the differences may correlate to the brachyptery. than phallobase, strongly sinuate on outer face subapically, Etymology. This species is named after Richard E. L. moderately widened apically in lateral view. Median lobe Smith, who is the founder of the African Natural History with ventral impression narrowly rounded in lateral view; Research Trust (ANHRT). apical disc ca. twice as long as wide, weakly concave in Biology. Specimens from Yéalé village were all collected lateral view, its apex rounded. Phallobase basally with at light in the middle of the village which is surrounded narrow, slightly asymmetrical manubrium. by a belt of secondary forests and plantations followed by Etymology. The latinised Greek noun ophioglossus means intact forest. The brachypterous specimen in the Tai NP was ‘a snake tongueʼ, in reference to the unique shape of the collected by sifting and washing plant debris accumulated ventral fork of the median lobe in this species. after a flood (M. Geiser & E. Ruzzier, pers. comm.). Biology. The paratype was collected at the small stream Distribution. The species was collected in two lowland in a primary forest. localities in western Côte d’Ivoire close to the border to Distribution. Known from two rather distant localities, one Liberia, situated ca. 220 km apart ( Fig. 13A ). in southern Togo and one in northern Gabon ( Fig. 13A ). Unidentified specimens

Eupotemus smithi sp. nov. of the Eupotemus limicola group

( Figs 2A, C, E–F, H ; 3A–B, J, Q ; 4C, K–L, Q ; 5A, D, J, M ; 6A–F ; 8J–L, X–Y ; 10A–C ) Material examined. CAMEROON: 1 ♀ (NMPC):Mamengole, iv.1949,

lgt. Tesárek. GABON: 1 ♀ (NHMW): Batéké Plateau National Park,

Material examined. Hඈඅඈඍඒඉൾ: J (macropterous) ( BMNH): ʽCÔ- camp Ntsa, ʽforêt denseʼ [= dense forest], 8–13.ix.2008, A. Bilardo lgt. TE D’IVOIRE, 380m, / Yeale Village, Mt. Nimba / 07°31’35.3”N 08°25’20.1”W, / 18-29. IV.2016 Light Trap,//Aristophanous,M., / Geiser, M., Moretto, P., leg., / BMNH(E) 2016-109, / Trip Ref. CI-003 (ANHRT Eupotemus carinaticollis group 17)ʼ. P*ò*©*òඍඒඉൾඌ: 14 spec. (incl. DNA voucher MF2207.W) ( BMNH, NMPC):same data as the holotype.; 1 spec. ( BMNH):ʽCÔTE D’IVOIRE, Eupotemus carinaticollis ( Basilewsky, 1956) 380m, / Yéalé Village, Mt. Nimba / 07°31’35.3”N 08°25’20.1”W, / ( Figs 11A–C , 12A–C ) 8.V.2016 //Aristophanous, M., / Geiser, M., Moretto, P., leg., / BMNH(E) Material examined. Hඈඅඈඍඒඉൾ: J ( MRAC): ʽHOLOTYPUS // COLL. 2016-109,Trip / Ref.:CI-003(ANHRT 17)ʼ; 1 J (brachypterous) ( BMNH): MUS. CONGO / Urundi: Rumonge 800 m / 7-III-1953 / P. Basilewsky

ʽCÔTE D’IVOIRE, 174m, / Tai NP,Tai Research Station, / 05°49’59.8”N, // Georyssus / carinaticollis / n. sp. Type / P. Basilewsky det., 19 // J.

07°20’32.0”W, / 14-23.xi.2015 // Leaf litter by river bank /Aristophanous, Delève det. 1966 / Eumetopus / carinaticollis / Basilewsky // Aedeagus

M., / Moretto, P., Ruzzier, E. leg., / BMNH(E) 2015-177ʼ. / drawn by / P. D. Perkinsʼ.

Differential diagnosis. Very similar to the other species of Additional specimens examined: DEMOCRATIC REPUBLIC the E. limicola species group from which it can be reliably OF THE CONGO: 1 J 1 ♀ (MRAC): Kivu: Uvira, vest. de forêt sclerophylle [= remnants of a sclerophyll forest] / I-1958 N. Leleup.

distinguished by the male genitalia only. The aedeagus ( Figs 8J–L, X–Y ) differs from all species except E. limicola Differential diagnosis. Very similar to the other species by the largely widened apex of the parameres in lateral of the E. carinaticollis species group from which it can be reliably distinguished by the male genitalia only. The aedeagus differs from E. uluguru sp. nov. in the apically widened and subapically constricted parameres ( Fig. 11A ). This character is shared with E. taianus sp. nov. from which E. carinaticollis differs in the narrower and more elongated median lobe.

Redescription. Body 2.9–3.4 mm long (holotype 2.9 mm) and 1.6–1.8 mm wide (holotype 1.7 mm). Dorsal surface black. Habitus and sculpture as in Figs 12A–C ; ridge on elytral interval 3 and 5 complete throughout; ridge on interval 7 interrupted posteriorly. Elytral punctures connected by low elevated line. Aedeagus ( Figs 11A–C ): 0.85–0.90 mm long. Parameres ca. 1.8× longer than phallobase, constricted subapically, widened at apex. Median lobe 3.4× longer than wide, apical part narrowing in apical fourth, sides of median lobe narrowing to apex in a straight line. Phallobase basally with narrow symmetrical manubrium.

Biology. Unknown. The labels of the Burundi specimens indicate that they were collected in remnants of a sclerophyll forest (i.e. in a dry forest with hard-leaved trees).

Distribution. The species is known from two localities situated around the northern part of Lake Tanganyika, one in the Democratic Republic of Congo, the other in Burundi ( Fig. 13B ).

Figures

Fig. 8. Male genitalia of the species of the Eupotemus limicola species group, holotypes.A–C, R–S – E. bilobatus sp. nov.; D–F, T–U – E. cameroonensis sp. nov.; G–I, V–W – E. ophioglossus sp. nov.; J–L, X–Y – E. smithi sp. nov.; M–Q – E. limicola (Delève, 1967). A, D, G, J, M – dorsal view; B, E, H, K, N – lateral view; C, F, I, L – ventral view; O – apex of the median lobe, dorsal view; P, R, T, V, X – fork of the median lobe; Q, S, U, W, Y – apex of parameres in lateral view.

Fig. 9. Habitus photographs of the species of the Eupotemus limicola species group, holotypes: A–C – E. bilobatus sp. nov.; D–F – E. cameroonensis sp. nov.; G–I – E. limicola (Delève, 1967).

Fig. 10. Habitus photographs of the species of the Eupotemus limicola species group, holotypes (A–F) and differences between species groups in pronotal morphology (G–H). A–C – E. smithi sp. nov.; D–F – E. ophioglossus sp. nov.; G–I – pronotal sculpture: G – E. carinaticollis species group; H – E. limicola species group.

Fig. 13. Known distribution of Eupotemus species.A – species of E. limicola species group; B – species of E. carinaticollis species group.

Fig. 2. Morphology of mouthparts and head appendages of the genera of Epimetopidae.A–I – Eupotemus Ji & Jäch, 1998: A, C, E, F, H – E. smithi sp. nov., B, D, G – E. limicola (Delève, 1967), I – E. cameroonensis sp. nov. J–Q – Eumetopus Balfour-Browne, 1949: J, L, M, N, P – E. schuelkei Jäch, 2002, Q – E. bullatus (Sharp, 1875), K, L*, O – E. sp. from Sri Lanka. R–Z, a–l – Epimetopus Lacordaire, 1854: R–W – E. mendeli Fikáček et al., 2011, X–Z, a–c – E. costaricensis Perkins, 1979, g, j – E. thermarum Schwarz & Barber, 1917, e, k – E. trogoides (Sharp, 1874), h – E. costatus group, f, i, l – E. punctipennis Perkins, 1979. Body parts:A–B, J–K, R, X, g–i – labrum; C, L, S, Y – mandibles; D, L*, e–f – mandibular apex; E, M, T, Z – maxilla; U – detail of basal part of ultimate palpomere; F–G, N–O, V, a–b, j–l – mentum and prementum; H–I, P–Q, W, c – antenna. Not to scale.

Fig. 3. Thoracic morphology of the genera of Epimetopidae. A–B, J, Q – Eupotemus smithi sp. nov.; C–D, L–O – Eumetopus schuelkei Jäch, 2002; E–F – Epimetopus mendeli Fikáček, Barclay & Perkins, 2012; G–I, P – Epimetopus costaricensis Perkins, 1979. A–I – prothorax in ventral view (B, D, F, H – detail of ventral sculpture of the pronotal hood; I – detail of closed procoxal cavity. J–L – meso- and metathorax in ventral view. M–N – elytron in ventral view (M – general view; N – detail of the ventral ridge, notice the spiny surface on inner face of the ridge (in) and on the elytral plectrum posteriorly of it (pl)). O – mesotrochanter. P–Q – details of pretarsus with the leaf-like empodial seta. Not to scale.

Fig. 4. Head and leg morphology of the genera of Epimetopidae. A–B, H, L – Eumetopus acutimontis Ji & Jäch, 1998; D, M – Eumetopus schuelkei Jäch, 2002; C, K–L, Q – Eupotemus smithi sp. nov.; F–G – Epimetopus punctipennis Perkins, 1979 (adopted from Pൾ*©κංඇඌ 2012); E, I–J, R – Epimetopus mendeli Fikáček, Barclay & Perkins, 2011; N–O – Epimetopus costaricensis Perkins, 1979. A – head in dorsolateral view (can – clypeal canthus causing the eye emargination); B–C – head in dorsal view (arrow – the ridge dividing anterior declined part of clypeus); D – mentum and maxilla; E – head in ventral view; F–G – occipital part of the head with median raised tubercle, dorsolateral view (arrows: parallel impressions corresponding to ventral ridges of pronotal hood); H–I – antenna, scapus largely omitted; J – labial palp; K – apical tibial armature, ventral view, mesothoracic leg; L – apical tibial armature and tarsus, ventral view, metathoracic leg; M – mesotibia; N–O – tarsus (N – posterior; O – anterior); P–R – metatibia, dorsal view. Not to scale.

Fig. 5. Morphology of Epimetopidae. A–B – ventral morphology: A – Eupotemus smithi sp. nov.; B – Epimetopus mendeli Fikáček, Perkins & Barclay, 2011. C–F – hind wings: C – Eumetopus schulkei Jäch, 2002; D – Eupotemus smithi sp. nov.; E – Epimetopus mendeli, adopted from Fංκගඹൾκ et al. (2011); F – Epimetopus costaricensis Perkins, 1979. G – dorsal part of the metathorax. H – scutellum. I–K – abdominal ventrites (I – Eumetopus schuelkei; J – Eupotemus smithi sp. nov.; K – Epimetopus mendeli). L–M – ovipositor (L – Eumetopus schuelkei; M – Eupotemus smithi). O – elytral punctation, slide-mounted elytra of Epimetopus costaricensis. P – female of Eumetopus acutimontis in ventral view, with the egg cases carried under the abdomen. Not to scale.

Fig. 6. Male genitalia and associated structures of Epimetopidae. A–H – Eupotemus Ji & Jäch, 1998 (A–F – E. smithi sp. nov.; G – E. cameroonensis sp. nov.; H – E. carinaticollis (Basilewsky, 1956)). I–R – Epimetopus Lacordaire, 1854 (I–N – E. mendeli Fikáček et al., 2011; O – E. cf. burruyacu Oliva, 1986; P – E. multiporus Perkins, 2012; Q – E. clandestinus Perkins, 2012; R – E. thermarum Schwarz & Barber, 1917). S–Z – Eumetopus Balfour-Browne, 1949 (S–Y – E. acutimontis Ji & Jäch, 1998; Z – E. bullatus (Sharp, 1875)).A, I, S – sternite VIII; B, J, T – tergite VIII; C, K, U – sternite IX; D, L, V – aedeagus dorsally; E, M, W – aedeagus laterally; F, N, X – aedeagus ventrally (basal part omitted in X); Y – sperm pump. Color coding: green – ventral projections of the median lobe; red – median lobe; blue – paramere (pale blue – dorsal lobe; dark blue – ventral lobe). Not to scale.

Fig. 11. Male genitalia of the Eupotemus carinaticollis species group, holotypes.A–C – E. carinaticollis (Basilewsky, 1956); D–F – E. taianus sp. nov.; G–I – E. uluguru sp. nov. A, D, G – dorsal view; B, E, H – lateral view; C, F, I – ventral view.

Fig. 12. Habitus photographs of the species of the Eupotemus carinaticollis species group, holotypes: A–C – E. carinaticollis (Basilewsky, 1956); D–F – E. uluguru sp. nov.; G–I – E. taianus sp. nov.