On Loneura crenata Navás and Loneura ocotensis García Aldrete (Psocodea, ‘Psocoptera’, Ptiloneuridae)

Here it is shown that Loneura crenata Navás, 1927 and Loneura ocotensis García Aldrete are distinct species, so the previously proposed synonymy of the latter with the first is not valid. Illustrations of the L. crenata holotype, deposited in the Hamburg University Zoological Museum, are here presented for the first time.


INTRODUCTION
The Costa Rican species Loneura crenata Navás, 1927 is the type species of Loneura Navás. The description, in Latin, refers to the body's color and length, as well as the fore-and hind-wings' length and venation. The fore-and hind-wings are illustrated, the first shows a submarginal pigmented band from vein R₄₊₅ to the areola postica, which is tall, broadly triangular, and slightly slanted posteriorly. The pterostigma is subtriangular, semioval, with vein R₁ well pigmented, the stems of veins R s and M are straight, and vein M is six-branched ( Fig. 1). The hindwing is unpigmented, with vein M four-branched (Fig. 2). Although is not indicated in the description, the holotype is a male, dry mounted, i.e., it is deposited in the Centrum für Naturkunde, of the Hamburg University Zoological Museum (M. Huseman, 2020, in litt.) . Later, New (1976) reports that there are also three male paratypes in the same collection and, based on the most complete paratype, redescribed and illustrated the species (Figs. 1-2, 3-6). Here we highlight that these three specimens of L. crenata, considered as paratypes by New (1976) do not have the status of paratypes because they are not mentioned in the original description by Navás (1927) and are therefore non-type specimen from the type locality.
Loneura crenata Navás belongs to the species group I, subgroup IA, defined by García Aldrete  (1995)]. L. crenata is also related to the Nicaraguan L. mombachensis García Aldrete, 2003, and despite having the same six M branches on the forewing, differs by lacking a pigmented band from R₄₊₅ to the areola postica in the forewings, from the middle of Cu₁A to the wing margin, by lacking distal dilated sides in the central sclerite of the hypandrium, and by the phallosome's endophallic sclerites [see Figs. 11 (wings), 16 (hypandrium), and 17 (phallosome) in García Aldrete (2003)].
Loneura crenata and L. ocotensis García Aldrete are remarkably similar in having fore-and hindwings, with the same M terminal branches (6 and 4, respectively), the head's same color pattern, hypandrium with central projection distally acuminated, phallosome with the same shape on the side strut and external parameres, and the endophallic sclerites' same shape and number. Based on the abovementioned, Castro (2007), in an un-published master's thesis conducted at the Instituto Nacional de Pesquisas da Amazônia, in Manaus, Brazil, established that L. ocotensis was a synonym of L. crenata. Then, this was mentioned by González Obando et al.
(2020). One of the most important facts that explained and suggested these species' synonym is that both species have six M terminal branches in the forewing, but in L. crenata all six branches are primary, while in L. ocotensis only four branches are primary and the fifth is bifurcated.
Previous taxonomic works related to Loneura, like those of Navás (1927) and New (1976), were based on characters such as the number of branches with venation, which was considered invariable and only the terminal branches were considered disregarding the primary branches. The wing color pattern, head color, and morphology of the genitalia's structures was simply described and little studied. Now, with the increase in the number of species and specimens, we know that the head color and the number of veins, due to having great intraspecific variation, are no longer safe characters to separate Loneura species. The genitalia morphology is more informative, it has very well-defined patterns and it has been shown to be the best character to safely separate this genus' species.
The purpose of this paper is to present details on genitalia morphology and illustration of the L. ocotensis male holotype to investigate the synonym with L. cren-   New (1976). Scales in mm (Fig. 3 without scale). ata, which, for being the type species of the genus and being inserted in this dubious synonym, was bringing uncertainty to the taxonomy of the genus.

MATERIAL AND METHODS
Three L. ocotensis males were available on loan from García Aldrete (UNAM) for study. They were dissected in 80% ethanol, and their parts were mounted on glass slides in Canada balsam. Standard measurements (in μm) were taken with a filar micrometer. Abbreviations of parts measured are as follows: FW and HW: right foreand hind-wing lengths; F, T, t1, t2, and t3: lengths of femur, tibia, and tarsomeres 1, 2, and 3 of right hind leg; f1…fn: lengths of flagellomeres 1…n of right antenna; Mx4: length of fourth segment of right maxillary palpus; IO: minimum distance between compound eyes in head dorsal view; D and d: antero-posterior and transverse diameter, respectively, of right compound eye in head dorsal view; PO: d/D. The final storage of the specimens was in CD boxes, as described by Silva-Neto et al. (2016).
Photographs of the parts mounted were taken with a Leica DFC500 digital camera attached to a Leica M205C stereomicroscope, connected to a computer with the Leica Application Suite LAS V3.6 software, which includes an Auto-Montage module (Syncroscopy software).

RESULTS
The decision to synonymize L. ocotensis with L. crenata is wrong. After analyzing more specimens, we have found that the two species, although related, are distinct and differ as follows: -In the posterior process of the central sclerite of the hypandrium of L. ocotensis, the sides are parallel; distally, the process has a pointed projection on each side, and the apex is obtusely convex (Fig. 15). In L. crenata, the posterior process of the central sclerite of the hypandrium has sides not parallel, slightly converging to a slightly obtusely concave apex, it has, subapically, a triangular flap on each side of the longitudinal midline (Fig. 4). -In L. ocotensis, the distal fourth of the posterior endophallic sclerites of the phallosome is distinctly narrowed, while in L. crenata the distal fourth of the posterior endophallic sclerites is continuous with the rest of the sclerite. Likewise, and as far as it can be observed, the anterior endophallic sclerites are quite different (compare the phallosome (Fig. 5) illustrated by New (1976) with the phallosome (Fig. 16) of L. ocotensis in this paper).
The observations above suggest that L. crenata Navás and L. ocotensis García Aldrete are different species, therefore, the synonymy proposed by Castro (2007) and reported by González Obando et al. (2020) has to be rejected.