Speonemadus, Jeannel, 1922

Fresneda, Javier, Faille, Arnaud, Fery, Hans & Ribera, Ignacio, 2019, A molecular phylogeny of Speonemadus Jeannel, 1922 with description of two new species from Morocco (Coleoptera: Leiodidae: Cholevinae: Anemadini), Zootaxa 4543 (1), pp. 1-36 : 26-27

Phylogeny of Speonemadus

After the description of the two new species and the reinstatement of S. gracilis and S. pulchellus , the genus Speonemadus adds to 16 species, of which our molecular phylogeny includes ten ( Tables 1, 2). Other than the newly described S. comasi sp. n., S. transversostriatus is a missing species with an uncertain phylogenetic position, the only member of its species group sensu Giachino & Vailati (1993). These authors did not include S. transversostriatus in the escalerai -group due to some differences in the sexual dimorphism of males: they have simple protibiae, without the characteristic keel of the species of the escalerai -group ( Reboleira et al. 2017). However, all other characters agree with the typical morphology of the escalerai -group: elytral sculpture of " type c", median lobe of the aedeagus bottle-shaped, parameres of the aedeagus only shortly longer than the median lobe ( Giachino & Vailati 1993), and females with elytral apex more acute than males (strongly acute and with a cleavage in the females of the escalerai -group, Blas 1984). It seems thus likely that S. transversostriatus is either sister or within the clade formed by the species of the S. escalerai -group. The other four missing species ( S. escalerai , S. algarvensis , S. orchesioides and S. subcostatus ) are morphologically very close to some species included in the phylogeny, as already noted.

Of the species groups defined in Giachino & Vailati (1993), we recovered as monophyletic the studied species of the escalerai- and vandalitiae -groups. However, according to the characters used to define the vandalitiae -group the newly described S. brusteli sp. n. should have been included in it, but we recovered it as sister to S. maroccanus , which belongs to the clathratus -group. Some of the characters used to define species groups were reconstructed as having been independently gained in different species. The thick and blunt setae (the "spine") on the ventral side of the apex of the parameres ( Figs 19–26 ) is a derived state of clade A, present in S. maroccanus ( Fig. 26 ), in the species of the S. escalerai -group ( Figs 19–23 ) including S. transversostriatus ( Fig. 24 ) and in S. clathratus ( Fig. 25 ). According to our reconstruction the "spine" was secondarily lost in S. brusteli sp. n. and its likely sister S. orchesioides , and it is primarily absent in clade B and in Anemadus ( Fig. 28 ).

The shortening of the parameres from the apparent plesiomorphic condition of being much longer than the median lobe has occurred independently in S. maroccanus and the S. escalerai and S. clathratus -groups. Other characters used by Giachino & Vailati (1993), such as the presence of more or less developed striae on the elytral disc, are difficult to define precisely. Despite their usefulness in identifying the species of the genus, our results suggest that the species groups defined within Speonemadus are in need of revision.

Figures

FIGURES 13–28. Apex of the parameres of: (13) Speonemadus orchesioides (Ammi-Moussa, Algeria); (14) holotype of S. brusteli sp. n. (Talembote, Morocco); (15) S. gracilis (Hornos, Spain); (16) S. tenuipes (Ifri Semedane, Algeria); (17) S. vandalitiae (Villanúa, Spain); (18) S. pulchellus (Ficuzza, Sicily); (19) holotype of S. algarvensis (Gruta do Vale Telheiro, Portugal); (20) S. angusticollis (Rute, Spain); (21) S. bolivari (Ardales, Spain); (22) S. breuili (Motillas, Spain); (23) S. escalerai (Jumilla, Spain); (24) S. transversostriatus (La Bastida, Spain); (25) S. clathratus (Linarejos, Spain); (26) S. maroccanus (Taza, Morocco); (27) holotype of S. comasi sp. n. (Tagelft, Morocco); (28) Anemadus hajeki (Yunnan, China).