Triplocania Roesler (Psocodea: ‘Psocoptera’: Ptiloneuridae): review of the internal classification, new species, and new records for the state of Bahia, Brazil

Four species of Triplocania collected in the Brazilian state of Bahia, are described and illustrated, based on male specimens, namely: T. annyae sp. nov., T. diamantina sp. nov., T. lauzie sp. nov. and T. miltoni sp. nov. They differ from the other species in the genus, in which the males are known by hypandrium and phallosome structures. New records of T. capixaba Silva-Neto, García Aldrete & Rafael, 2016 and T. manueli Silva-Neto, García Aldrete & Rafael, 2016 for the state of Bahia are presented. In addition, an update on the Triplocania internal classification is proposed.


INTRODUCTION
Triplocania Roesler, 1940 is one of 12 genera in the psocopteran family Ptiloneuridae. It is the most speciose genus of the family, presently including 98 described species, with 42 species known only from males, 23 species known only from females, and 33 species known from both sexes (González et al., 2017;. Brazil includes 26 states and a Federal District. A total of 30 species of Triplocania have been recorded in 11 of these states ( Table 1). The most diverse state for Triplocania species is the state of Amazonas, in the North region of Brazil (Table 1). Among the Brazilian regions, the Northeast region has the lowest number of Triplocania species records, with only three species records, one species in the state of Ceará (Tiplocania manueli Silva-Neto, García Aldrete & Rafael) and two species in the state of Bahia (Triplocania ariasi New and Triplocania calori Silva-Neto, García Aldrete & Rafael) (Silva-Neto & García Aldrete, 2020).
Recently, two of us (AMSN and DML) have found specimens of four Triplocania undescribed species and specimens of T. capixaba Rafael, 2016 andT. manueli Silva-Neto, García Aldrete &Rafael, 2016 in the state of Bahia. This study aims, based on male specimens, to describe and illustrate these species to include new Triplocania species records in the state of Bahia, and to update the Triplocania species list in Brazil. In addition, we propose an update on the Triplocania internal classification.

MATERIAL AND METHODS
Thirteen male specimens and three female specimens were available for study. They were dissected in 80% ethanol, and their parts were mounted on 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 fore-and hind-wing ISSN On-Line: 1807-0205 ISSN Printed: 0031-1049 ISNI: 0000-0004-0384-1825 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 specimens studied were stored in CD boxes, as described by Silva-Neto et al. (2016a).
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). All specimens used in this study will be deposited in the Entomological collection Prof. Johann Becker of the Museu de Zoologia of the Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil (MZFS).

Triplocania annyae sp. nov. Male (Figs. 1-7)
Diagnosis: Forewings with a pale brown spot darkening the pathways through which it passes, around the vein that joins R-R s , passing through the junctions between M and R s , M and Cu, and part of Cu; a pale brown band distally in the pterostigma; a small brown spot distally on R₄₊₅ and M₁ (Fig. 2). Hypandrium of one sclerite, glabrous, with sides deeply concave in the middle, anteriorly almost trapeziform, heavily sclerotized, posteriorly elliptic, less sclerotized, with irregular posterior border (Fig. 5); Phallosome ( Fig. 6) anteriorly membranous, scaly, organized like reptile skin; three pairs of endophallic sclerites, an anterior chain-shaped pair, behind the side struts; an antero-lateral pair small, narrow, anteriorly rounded, posteriorly acuminate; posterior pair stout, elongate, anteriorly heavily sclerotized, almost rectangular, narrowing in the middle and dilated posteriorly.
Morphology: Head vertex slightly concave in the middle, below the level of the compound eyes' upper border, these without interommatidial setae (Fig. 8). Outer cusp of lacinial tips broad, with six denticles (Fig. 11). Forewing pterostigma wider in the middle, narrow anteriorly; areola postica wide, slanted posteriorly, apically round; M stem slightly concave, M₁ almost straight, M₂ slightly sinuous, M₃ proximally convex, then almost straight to the end, R s stem convex, R₂₊₃ proximally straight then distally concave, R₄₊₅ slightly concave proximally, then almost straight ( Fig. 9). Hindwing R s stem straight, R₂₊₃ straight, and R₄₊₅ slightly concave proximally, then almost straight, M stem sinuous, proximally slightly convex (Fig. 10). Hypandrium ( Fig. 12) of three sclerites with central piece anteriorly with two less sclerotized circular parts and bristles distributed over the surface, a small projection in the middle, also round. Phallosome (Fig. 13). Epiproct concave in connection with the clunium, obtuse on the sides, almost straight at the apex, with three mesal macrosetae (Fig. 14). Paraprocts broad, broadly triangular, setose as illustrated; sensory fields with 33 trichobothria on basal rosettes (Fig. 14). Diagnosis: Forewings with a brown marginal pigmented band, from R₄₊₅ to wing base, with hyaline fenestrae at wing margin as illustrated; pterostigma with a small proximal brown band, and a large, distal brown band, with a central, hyaline area (Fig. 16). Hypandrium of one sclerite, anteriorly straight, with mesal lateral processes, posteriorly with sides converging to broad apex, with an almost triangular, acuminate process on each side, directed outward, leaving a wide concavity between them (Fig. 19). Phallosome (Fig. 20) with side struts independent, inverted Y-shaped, not fused posteriorly to external parameres; anterior pair of endophallic sclerites broadly elliptic, with irregular outer borders, the left one with three short discrete posterior projections and a rounded apex; a mesal pair, small, strongly sclerotized, almost touching in the middle; posterior pair scythe-shaped.
Three pairs of endophallic sclerites, an antero-mesal pair slender, sinuous, subdivided into pieces V-shaped organized in chain (catenulate), a lateral-pair slender, straight anteriorly, slightly curved inwards from three quarters of its length to the apex, this rounded; a posterior pair, stout, elongated, basally wide, with a medial concavity, which give rise almost triangular projections, on each side antero-lateral, narrowing distally, with apex almost triangular (Fig. 27). Epiproct almost trapezoidal, posteriorly slightly concave in the middle, anteriorly rounded with some bristles, other setae as illustrated (Fig. 28). Paraprocts broadly elliptic, sensory fields with 32 trichobothria on basal rosettes, setae as illustrated (Fig. 28).

DISCUSSION
The species here described raise to 102 the number of species in the genus and raise to 34 the species recorded in Brazil (33% of the total) (Table 1). Silva-Neto et al. (2015) recognized two species groups in Triplocania, as follows: -MPB group: characterized by having forewing venation caeciliusid, that is, with R s of two branches, and M with only three primary branches, and without secondary branches. -MSB group: characterized by having forewing M with three primary and secondary branches, this group is here referred as MSB group and it is divided in two subgroups: -Subgroup MSB1: characterized by having more than one M vein with secondary branches, the branches originating closer to the wing margin than to the main M. -Subgroup MSB2: characterized by having only one secondary branch, in M₃, resulting in M₃ a and M₃ b , and with branches originating closer to the main M than to the wing margin.
In a monograph on Colombian Triplocania, González et al. (2017), recognized two species groups in the genus, but the authors forgot that there was already a classification previously made by Silva-Neto et al., 2015, abovementioned. Later Silva-Neto & García Aldrete (2019) created the subgroup magnifica within the group MPB, characterized by having forewing with a U-shaped band from the areola postica's apex to the basal and distal pterostigma part. Forewing M deeply concave before its first bifurcation, areola postica low, very wide, side struts proximally expanded forming a shield; v1 stout, wider in the middle.
Here we propose a modification of the subgroups' classification present in the MSB and MPB groups proposed by Silva-Neto et al. (2015). In addition, we also propose some 'Sub subgroups' to homogenize the Triplocania internal classification, as follows: -MSB group (Equivalent to the group II of González et al. (2017): characterized by having forewing M with three primary and secondary branches, this group is divided in two subgroups: -Subgroup MSB1: characterized by having more than one M vein with secondary branches or more than two secondary branches in M₃.  González et al. (2017)): characterized by having forewing with M with only three primary branches, and without secondary branches.
MPB group includes 79 species, 23 species these species are known only to females. Here we propose two new subgroups only for the species with known males, except for Triplocania marginepicta Roesler and Triplocania chulumanensis (Williner), the types of which are lost, the original publications have poor descriptions or do not have illustrations and we did not get additional specimens. In the future with the unknown males' description, the species only with known females could be included in their respective subgroups whose diagnoses are based on male genital characteristics: -Subgroup MPB1 (Equivalent to the subgroup IA. of González et al. (2017)): characterized by having hypandrium of of one sclerite and side struts not expanded forming a shield. differs from all other abovementioned species, by having the unique shape of central piece of the hypandrium, in particular by the shape and details of its posterior projections and by having only a pair of endophallic sclerite and in other details described in its diagnosis. The recognition of species groups and subgroups is an important step to facilitate their identification and to understand relationships among these taxa, especially in taxons with many species, as in Triplocania. Future phylogenetic studies, using morphological and molecular data will be necessary to test the monophyly of these groups and subgroups and to understand the Triplocania evolution. However as mentioned by Yoshizawa (2002) and reminded by Silva-Neto et al. (2016b), the highly modified male genital structures present in the genera Ptiloneuridae could be generating significant synapomorphies for them, but unfortunately, they are extremely variable within the family and thus, it is difficult to decide on their homologies. This fact, along with the loss of the thorax or its destruction in part during the specimens' dissection, weakens phylogenetic analyzes in Ptiloneuridae using only morphological characteristics. Based on the abovementioned facts, species groups and subgroups in Triplocania are important in facilitating this genus' taxonomy, even without the support of a current phylogenetic analysis for the time being.