New species of Triplocania Roesler (Psocodea, ‘Psocoptera’, Ptiloneuridae), from Brazil and Ecuador

Abstract Four species of Triplocania, three with M3 simple, based on male specimens and one with forewing M3 forked, based on male and female specimens, are here described and illustrated, namely: Triplocania bravoi sp. n. (Napo: Ecuador), Triplocania erwini sp. n. (Napo: Ecuador), Triplocania trifida sp. n. (Mato Grosso and Rondônia: Brazil) and Triplocania lamasoides sp. n. (Rondônia: Brazil). They differ from all the other species in the genus, in which the males are known, by the hypandrium and phallosome structures. The female is first described for the M3 forked group. The identification key for males of the M3 forked group is updated.


Introduction
Triplocania Roesler (1940) is one of 12 genera in the psocopteran family Ptiloneuridae; it is the most species rich genus of this family. It presently includes 32 described species that, according to forewing venation, can be separated in two groups: a large one with 25 species, characterized by having forewing venation caeciliusid, that is, with Rs of two branches, and with M of three branches, this group is here referred as MPB group (M with only primary branches); and a smaller group with 7 species, characterized by having M with three primary and secondary branches, this group is here referred as MSB group (M with secondary branches); this group is divided in two subgroups: the first (MSB1) is represented by Triplocania palaciosi García Aldrete & Casasola González (2012), it is characterized by having more than one M vein with secondary branches, the branches originating closer to the wing margin than to the main M. The second subgroup (MSB2) is represented by 6 species; it is characterized by having only one secondary branch, in M 3 , resulting in M 3a and M 3b, and with branches originating closer to the main M than to the wing margin. The purpose of this work is to describe and illustrate three new species of Triplocania belonging in group MPB, based on males, and to describe a new species belonging in subgroup MSB2 mentioned above, based on males and females.

Material and methods
Ten specimens were available for study; they were dissected in 80% ethanol; their parts (head, right legs and wings, and genitals) were mounted in Canada balsam. Before dissecting, whole specimens were placed in 80% ethanol under a dissecting microscope, illuminated with cold, white light, and observed at 50× to record color. 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 length, 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 dorsal view of head, D and d: antero-posterior and transverse diameter, respectively, of right compound eye in dorsal view of head, PO: d/D. The types of the Brazilian species will be deposited in the Invertebrate Collection of the Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Amazonas, Brazil. The types of the Ecuadorian species will be deposited in the Sección de Entomología, Instituto de Ciencias Biológicas, Escuela Politécnica Nacional, in Quito, Ecuador (EPN). Etymology. This species is dedicated to the Ecuadorian entomologist Freddy Rubén Bravo Quijano, of the Universidade Estadual de Feira de Santana, Bahia, Brazil, in recognition to his important contributions in the taxonomy of Neotropical Psychodidae (Diptera), also for the support to AMSN, to pursue a career studying Psocodea, 'Psocoptera'.
Diagnosis. Differing from the known species of Triplocania, in having the hypandrium with side sclerites fused proximally to the central piece, and having two forked posterior projections, horn shaped; also by having a U-shaped phallosome, a phallobase with lateral extensions covering partly the anterior pairs of endophallic sclerites, and in having ornamented areas on the endophallus.
Male. Color. Compound eyes black, ocelli hyaline, with ochre centripetal crescents; head pattern ( Fig. 1). Scape and pedicel pale yellow, f 1 -f 3 pale yellow, with apex white. Mx4 pale brown. Femora pale yellow; tibiae yellow, distally pale brown; tarsomere 1 pale yellow, tarsomeres 2-3 pale brown. Forewing with a brown marginal band from R 4+5 to almost CuA 2 ; a brown, almost rectangular band, well pigmented proximally and distally, weakly pigmented in the middle, from the apex of the areola postica to posterior end of the pterostigma; pterostigma with proximal and distal brown bands. Proximal third of forewing dark brown, limited posteriorly by level of crossvein Rs-M, as illustrated (Fig. 2). Veins brown. Hindwing with area below CuP dark brown, and a marginal pale brown band from R 4+5 to almost CuA (Fig. 3).
Morphology. As in diagnosis, plus the following: compound eyes without interommatidial setae. Outer cusp of lacinial tip broad, with five denticles (Fig. 4). Forewing pterostigma long, widest in the middle. Areola postica very wide basally, slanted posteriorly, tall proximally with apex round and narrow, distally sinuous and low; R 2+3 and R 4+5 sinuous, M stem slightly concave proximally, M 1, M 2 and M 3 sinuous. Hindwing Rs almost straight, R 2+3 straight, R 4+5 almost straight, M sinuous. Hypandrium of three sclerites, central piece anteriorly convex, with side projections almost parallel to side sclerites, forked posterior projections densely setose, other setae as illustrated (Fig. 5). Phallosome U-shaped, side struts independent, V shaped, fused posteriorly to external parameres, these stout, bearing a field of pores distally, each with an small projection heavily sclerotized distally; three pairs of endophallic sclerites, a posterior pair, curved outwards distally, close to the ends of the external parameres, a mesal pair, originating from behind the side struts, crossing behind the external parameres, distally dilated, with a small, acuminate projection apically; anterior pair stout, curved inwards, apically pointed and proximally rounded; phallobase with anterior border concave; lateral extensions covering partly the anterior pair of endophallic sclerites; endophallus membranous, with areas thickened and ornamented as illustrated (Fig. 6). Paraprocts broad, a field of setae along inner margin, other setae on apex; sensory fields with 27-28 trichobothria on basal rosettes (Fig. 7). Epiproct wide based, posteriorly rounded, with three large mesal setae, next to anterior margin, other setae as illustrated (Fig. 7).
Measurements ( Etymology. This species is dedicated to Dr. Terry L. Erwin, of the Smithsonian Institution, Washington, D. C., USA., in recognition to his seminal studies in biodiversity, in estimating the number of arthropods on this planet, in systematics and biology of the Carabidae, and for making available for study to ANGA, the psocid specimens collected by his team in Napo, Ecuador, by canopy fogging.
Diagnosis. Differing from the known species of Triplocania, in having the hypandrium with side sclerites and central piece similar in size; central piece with two short, lateral posterior projections, and two short, blunt ended, median posterior projections, leaving between them a small concavity, in having the external parameres with a distinct lobe apically on the inner side and, in having two pairs of endophallic sclerites.
Male. Color. Body pale brown, with ochre spots as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents; head pattern ( Fig. 8). Scape brown and pedicel yellow, f 1 -f 3 yellow, with apices white. Mx4 brown. Tergal lobes of meso and methathorax brown, pleura with ochre spots above the level of the coxae; dark brown bands on proximal and distal ends of coxae, femora yellow with three brown equidistant bands, a middle one, and one on each end of the femur; tibiae pale brown, tarsomeres 1-3 yellow. Forewing with an irregular, submarginal brown band from R 2+3 to areola postica, this with a small brown spot proximally, and a dark brown spot between its apex and M, a dark brown spot below the proximal end of CuA, and a brown spot at confluence of CuP-1A; a pale brown spot between proximal ends of R 4+5 -M 1 ; pterostigma with brown bands anteriorly and posteriorly; veins brown, with brown spots at wing margin (Fig. 9). Hindwing almost hyaline, veins brown, with a pale brown spot at confluence of CuP and wing margin (Fig. 10).
Morphology. As in diagnosis plus the following: compound eyes without interommatidial setae. Outer cusp of lacinial tip broad, with five denticles (Fig. 11). Fore- wing pterostigma basally narrow, wider in the middle; areola postica wide basally, slightly slanted posteriorly; R 2+3 and R 4+5 sinuous, M stem slightly concave proximally, M 1, M 2 and M 3 sinuous. Hindwing Rs almost straight, R 2+3 and R 4+5 straight, M sinuous. Hypandrium with two large setae posteriorly between each lateral-median posterior projections (Fig. 12). Phallosome with side struts independent, V shaped, fused posteriorly to external parameres, these stout, with pores posteriorly; the distal lobe of each heavily sclerotized; anterior pair of endophallic sclerites elongate, almost touching anteriorly, inserted on a membranous, V shaped, thickened area, lying distally behind the external parameres, ending in a rounded apex; posterior pair of endophallic sclerites triangular, each anteriorly concave, close to the inner border of the external parameres, with apices sinuous.
Measurements ( Etymology. The specific name refers to the proximity of this species to T. lamasi Silva-Neto, Rafael & García Aldrete. Diagnosis. Differing from T. lamasi in having the posterior sclerite of the hypandrium thicker in the middle, with the posterior projection more than twice as long; sickle-shaped lateral projections distal to the anterior sclerite barely reaching the inner margins of the lateral sclerites. Male. Color. Body yellowish brown, with dark brown spots as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents; head pattern ( Fig. 15). Scape and pedicel pale brown; flagellomeres pale yellow. Mx4 pale yellow. Tergal lobes of meso-and metathorax reddish brown; episternum of mesothorax ochre. Coxae, trochanters and femora creamy white, tibiae and tarsomeres pale yellow. Forewings hyaline, as illustrated (Fig. 16); veins brown. Hindwing (Fig. 17), hyaline throughout, veins brown.
Morphology. As in diagnosis, plus the following: compound eyes with interommatidial setae. Outer cusp of lacinial tip broad, with six denticles (Fig. 18). Forewing pterostigma long, widest in the middle. Areola postica wide basally, slightly slanted posteriorly, apex round, narrow. R 2+3 and R 4+5 sinuous, M stem concave, M 1 almost straight, M 2 sinuous, M 3 branched, the branching point closer to M than to the wing  margin. Hindwing Rs almost straight. Hypandrium (Fig. 19) of four sclerites, anterior piece broad, setose, bearing distally two sickle-shaped lateral projections, heavily sclerotized at both ends, and having also a well defined, setose sclerotized area in the middle; posterior sclerite concave anteriorly, with a long, slender posterior projection in the middle, flanked by two large, broadly triangular lateral sclerites. Phallo-some ( Fig. 20) with side struts independent, V shaped, fused posteriorly to external parameres, these stout, each with an elongate projection on inner margin, with field of pores; three pairs of endophallic sclerites; anterior pair long, slender and curved, mesal pair wide proximally, narrowing distally, pointed, and posterior pair parallel to the inner margin of the external parameres, with three acuminate projections distally. Paraprocts broad, wide proximally, narrowing to round apex; with a field of short setae along inner margin, other setae as illustrated; sensory fields with 30-31 trichobothria on basal rosettes (Fig. 21). Epiproct mesally with an almost elliptic protuberance, with a field of setae posteriorly, and three large mesal setae next to anterior margin (Fig. 21).
Measurements ( Etymology. The specific name refers to the characteristic of the hypandrium, having three posterior projections.
Diagnosis. Differing from the known species of Triplocania, in having the central piece of the hypandrium with three posterior projections, a middle one, pointed, setose, flanked by lateral, strongly sclerotized, glabrous acuminate projections. Phallo- T. trifida and T. lamasoides increase the diversity of Triplocania in Brazil to 16 species, this country being the most species rich so far for described species of Triplocania.
The transverse bridge in the phallosome of T. trifida is a character that distinguishes it from other species of Triplocania; this character also appears in some species of Loneura (L. jinotegaensis García Aldrete, L. mirandaensis García Aldrete, L. tuluaen-sis García Aldrete, Mendivil & González, and L. andina García Aldrete, Mendivil & González. The structure the phallosome of T. trifida is also very similar, except for the bridge, to the phallosome of L. gorgonaensis. The pattern of pigmentation and wing venation in T. trifida is similar to T. erwini with small differences, but the hypandrium and phallosome structures of the two species are different.
The remarkable similarities of phallosome and hypandrium in species of Triplocania and Loneura may indicate that the two genera are closer than previously thought.
T. lamasoides and T. lamasi constitute a pair of sister species within Triplocania. The morphological structure that separate them, are the lenght of the posterior projections of the anterior and posterior sclerite of the hypandrium. With more knowledge on the diversity of Triplocania, perhaps new cases of species complexes will be found, possibly confirming that the hypandrium is the most variable structure in Triplocania. The pair of species T. lamasoides-T. lamasi alerts also on the difficulty of association between males and females in Triplocania. The female of T. lamasoides is the first female described for the subgroup MSB2, it was associated with the male because they were collected in the same place and date, but the wings and patterns of body pigmentation are also identical to T. lamasi.