A revision of Lycinella Gorham, 1884 with the description of six new species (Coleoptera, Lycidae, Calopterini)

Abstract The Neotropical genus Lycinella Gorham, 1884 is revised. Lycinellaopaca Gorham, 1884 and Lycinellaparvula Gorham, 1884 are redescribed and illustrated. Six new species are described for the genus: Lycinellaadamantissp. n., L.hansonisp. n., L.millerisp. n., L.cidaoisp. n., L.marshallisp. n. and L.pugliesaesp. n.. Lycinellahumeralis Pic, 1933 is moved to Ceratoprionhumerale (Pic, 1933), comb. n. A key to the species of Lycinella, illustrations and a distribution map is provided.


Introduction
While searching for cantharoid beetles in Malaise traps samples generated by the Costa Rican Malaise Trap Network project, we found a remarkable number of tiny Lycidae specimens of a unique form. These Costa Rican specimens have eight pronotal stemmata ( Figure 11), an apparently unpublished character discovered by Richard S. Miller (1991) in Leptolycus Leng & Mutchler, 1922 (Leptolycini). However, they did not seem to belong to the Leptolycini, based on current diagnoses (Ferreira andIvie 2016, Ferreira et al. 2018).
In discussions with Miller, he pointed us to Lycinella Gorham, 1884, a poorly known genus of Calopterini with three named species from Central America (Bocáková 2003(Bocáková , 2005) that exhibit these unreported structures. Following this lead, we found the new specimens belong to Lycinella, and represent several new species. In this study we take the opportunity to rediagnose and redescribe the genus and the two original species described by Gorham (Lycinella opaca Gorham, 1884 andLycinella parvula Gorham, 1884), describe six new species, and provide illustrations, distribution maps and a key to all the species. One last problem was the species Lycinella humeralis Pic, 1933, known in the literature only from a short description. Based on examination of the type, it is moved to Ceratoprion Gorham, 1884 (Leptolycini).

Materials and methods
The specimens were examined under a Leica Wild M3C stereoscopic microscope with magnification up to 40×. Photos were taken using a JVC (DC Ky-F75U) digital camera mounted on a Leica MS5 stereoscope, a Visionary Digital Passport II imaging system, equipped with a Canon 6D DSLR (http://www.duninc.com), and a Canon T3i DSLR with a MP-E 65 mm lens and stacked using the software Zerene Stacker version 1.04. Enhancements to digital images were made in Adobe PhotoShop CC 2018. Drawings were prepared based on photographs using Adobe Illustrator CC 2018. The distribution map technique follows Ferreira (2016): the map was generated using the software Google Earth and Quantum GIS 2.18.9, using the maps available on the website http://www.naturalearthdata.com, a free public database of maps.
Morphological terminology follows Crowson (1944), Bocáková (1990), Miller (1991), Kazantsev (2003) and Lawrence et al. (2011). Of particular note is the term "stemma" (pl. stemmata) for a unique form of structure on Leptolycini and related Calopterini, including Lycinella. These structures were first noted by Miller (1991), who was the first to use stemmata in this way. His usage has been subsequently followed by Ferreira and Ivie (2016) and Ferreira et al. (2018). Pronotal stemmata are tiny hemispherical white objects that occur on the pronotum, coxae and antenna of adult males (Figs 11,12). The number and placement of these structures are diagnostic at the generic and species level. Their function and homology is unknown.
Male genitalia were dissected after the entire specimen was soaked in hot water. For disarticulation and clearing processes the specimens were left overnight in a warm solution of KOH after which they were dissected and left in cold KOH for approximately 2 hours, time enough for the musculature to detach from inner structures. Transcription of label data from specimens follows Ivie (1985): the end of each line on a label is indicated by a ";" (semicolon); the individual labels are separated by a "/" (slash).
The majority of specimens treated here were taken in the Costa Rican Malaise Trap Network project headed by Paul Hansen. Further data on this project and the localities and methods is available at Hansen (1992).
Material examined is deposited in the following collections (respective curators are indicated in parentheses):

MAIC
Michael A

Results
Examination of Pic's type showed that his L. humeralis belongs in the Leptolycini genus Ceratoprion Gorham, 1884. Therefore, we are moving it, in anticipation of a revision of the Leptolycini in progress by VF.
Description. General dorsal coloration dark brown to black, with pronotum black, yellow-brown or yellow in some species bearing dark macula in discal portion or with complete longitudinal medial region (Figs 1-9). Body densely setose, dorsal pubescence long and erect, remainder of body with fine yellow pubescence throughout (Figs 1-9).

Distribution.
Lycinella is known to occur in Panama, Guatemala, and Costa Rica ( Figure 37).
Biology and immature. Females are unknown and presumably neotenic. Although information about the ecology and biology of Lycinella is unknown we can infer from the fact they were virtually all taken in Malaise traps that males of Lycinella species are flight active species.
Taxonomic placement of Lycinella. The initial tribal placement of Lycinella was difficult because it is among the genera that, like Cephalolycus Pic, 1926 and Aporrhipis Pascoe, 1887, shares features of both Calopterini and Leptolycini (see Miller 1991;Bocáková 2003Bocáková , 2005Ferreira and Ivie 2016;Ferreira et al. 2018). Bocák and Bocáková (1990) placed the genus in the Leptolycini, but based on examination of Lycinella humeralis, here moved to the Leptolycine genus Ceratoprion.

Key to the species of Lycinella
Diagnosis. Both the completely black body (Figure 4) and the unique male genitalia with median lobe tapered apically ( Figure 29) will distinguish L. adamantis from all other Lycinella species.
Pronotum trapezoidal, with posterolateral angles pronounced and acute, divergent, with weakly visible longitudinal carina in anterior portion of pronotum, bifurcate posteriorly forming an areola. Prosternum V-shaped; posterior margin rounded; laterally reaching hypomeron.
Elytra 9× longer than pronotum; costae II and IV visible, I and III weakly visible. Humeral region rounded in dorsal view. Legs slender, elongate (Figure 4). Pro-and mesocoxae with stemmata absent. Aedeagus with median lobe tapered apically, Etymology. The species was named after VSF's friend, Felipe Francisco Barbosa, a.k.a. Cidão, for his priceless advice and discussions on beetle taxonomy and systematics.
Diagnosis. The weak black discal macula that does not reach any margin is unique in this species, as most other species with bicolored pronota have the strong discal macula reaching the hind margin. The exception is L. opaca, which has a strongly demarcated discal macula. They are further distinguished by a black antennomere XI (white in L. cidaoi). The unique male genitalia have a stout median lobe which is 1.3× longer than parameres and a rounded apex (Figure 32).
Pronotum trapezoidal, not constricted medially, with posterior margin straight, anterolateral angles rounded, with posterolateral angles and pronounced and acute, divergent, with weakly visible longitudinal carina in anterior portion of pronotum, bifurcate posteriorly forming an areola, hardly visible. Prosternum V-shaped; posterior margin rounded; laterally reaching hypomeron. Elytra approx. 11× longer than pronotum, slightly expanded in 2/3 portion; costae I, II and IV strongly visible. Humeral region rounded, non-pronounced (Figure 7). Legs slender, elongate. Pro-and mesocoxae bearing stemmata. Aedeagus with median lobe stout, apex rounded, 1.3× longer than parameres; parameres 1.  Diagnosis. The unique genitalia of this species, characterized by the subquadrate apex of the short median lobe, which is 0.6× the length of the parameres, will distinguish this otherwise rather generalized species from all other Lycinella species. Among the species with a bicolored pronotum, only L. opaca also lacks stemmata on the pro-and mesocoxae. These two species are easily distinguished by the anteriorly rounded pronotum and strong elytral costae of L. opaca (Figure 1), in contrast to the angulate anterior angles and weakly costate elytra of L. hansoni ( Figure 5).
Distribution. Costa Rica: San José ( Figure 37). Diagnosis. The stout antennae and rounded anterior margin of the pronotum are unique to this species (Figure 1). The strong pronotal macula that does not reach the base of the pronotum is also diagnostic. The scaliform setae on antennomeres IV-XI are likewise unique in the genus (Figure 18). The male genitalia (Figure 34) can be used to confirm the identification.