A new genus of Ptiloneuridae, its position within the family, and descriptions of five species (Psocodea, ‘Psocoptera’)

Abstract Upon examination of ptiloneurid specimens recently collected in forests of natural areas and Natural National Parks in Colombia, ten males and seven females were found that represent five species of an undescribed ptiloneurid genus. It differs from the other genera in the family by characters of the wings, hypandrium, phallosome, epiproct, female gonapophyses, and sternum IX. An identification key to the males of the genera of Ptiloneuridae, and a key to the species of the new genus are presented. A test on the validity and monophyly of the new genus, and its position within the family was also conducted.

Within the framework of the project "Revisión Taxonómica y Endemismo de los Psócidos (Insecta: Psocodea: 'Psocoptera') de Areas Protegidas de Colombia", 17 specimens of Ptiloneuridae, corresponding to five species that could not be assigned to any of the known ptiloneurid genera were found. These specimens were collected in localities of Valle del Cauca, Meremberg Nature Reserve and La Candelaria (Huila), Planes de San Rafael Nature Reserve (Risaralda), National Natural Park Tamá (Norte de Santander), and Chicaque Nature Reserve (Cundinamarca) (Figure 48). Here we describe and illustrate these new species, erect a new genus for them, and discuss its position within the family.

Materials and methods
Seventeen specimens were available for study, ten males and seven females. Nine specimens were dissected in 80% ethanol, and their parts (head, right legs and wings, and genitals), were mounted in Canada balsam, following standard procedures. Before dissecting, whole specimens were placed in 80% ethanol and observed at 50× to record color. Standard measurements were taken with a filar micrometer, and are given in mm. Abbreviations of parts measured are as follows: FW and HW: right fore-and 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, Mx4: length of fourth segment of right maxillary palp, IO: minimum distance between compound eyes, D and d: antero-posterior and transverse diameter, respectively, of right compound eye, all in dorsal view of head. PO: d/D, H: head length, MxW: maximum head width, Ratio head length (H)/(D), L/W: forewing length/forewing width, lp/wp: pterostigma length/pterostigma width, al/ah: areola postica length/tall, l/w: hindwing length/hindwing width. The specimens studied are deposited in the Entomological Museum, Universidad del Valle (MUSENUV), Santiago de Cali, Colombia.
To test the validity and the monophyly of the new genus, and to establish its position within the family, a cladistic analysis was performed, using the free Software TNT 1.1 (Goloboff et al. 2008a) under implied weight schemes, between K = 1-9, determines how strongly homoplasious characters are downweighted, lower K values indicate that homoplastic characters are more strongly penalized or given less weight (Goloboff 1993) and given the concavity constant value K should be calculated as a function of N, which is the ratio of a single extra step at the cost of the most homoplastic character (Goloboff et al. 2008b), we obtained K values through a TNT script (setk.run) written by Salvador Arias to calculate an appropriate value for K. The script returned a value of K = 2 for our data set, which was then employed. Given the purpose of the analysis, only one outgroup, Spurostigma Eertmoed (Spurostigmatidae) was used (see Yoshizawa 2002;García Aldrete 2005;Casasola González 2006;Silva-Neto et al. 2016). The characters were treated as nonadditive and the topologies were evaluated with an implicit weight (IW) analysis, using Traditional Search with the algorithm TBR (100 repl/10 TperRp), with1000 replicas and Traditional Search with Bootstrap and Symmetric Resampling (Goloboff et al. 2003). The support of the clades was calculated with ACCTRAN and DELTRAN optimization in WINCLADA ver 1.00.08 (Nixon 2002). The resulting strict consensus tree was exported and the names were edited in CorelDraw X7. Neto et al. (2016b), only one species was used to represent each genus, except for Triplocania (two species) and Colocania gen. n., in which the males (four species) of each species were included in the analysis. The 17 species included were: Brasineura diamantina Silva Neto & García Aldrete, Colocania candelaria sp. n., C. chicaque sp. n., C. occidentalis sp. n., C. pilosa sp. n., Euplocania cerata New, Loneura jinotegaensis García Aldrete, Loneuroides colombianus García Aldrete, González  20. Costal margin, between base and nodus: (0) straight (1) convex (Figures 1,7,13,19,25,31,37). 21. Shape of proximal area of pterostigma (at around 1/3 of its length): (0) not petiolated ( Figure 43), (1) little petiolated ( Figure 44), (2) strongly petiolated (Figures 1,7,13,19,25,31,37). 22. Anterior region of hypandrium: (0) without ringed, short digitiform process, (1) with vestigial or short ringed digitiform process (Figures 4,10,22,34). 23. Oval cuticular depression on central-median area of clunium: (0) absent, (1) present (Figures 45, 46).   From 88 most parsimonious cladograms, obtained for each K value analyzed (K = 1-9), 17 consensus topologies were retained (L = 75-76, CI = 48-49, RI = 61-62). In all of them the clade formed by the species of Colocania was maintained, although between K = 5-9 the relations of this with the rest of the genera is unstable. This however, remains constant between K = 1-4 (L = 76, CI = 48, RI = 61), which includes the value calculated as appropriate (K = 2) through the use TNT script (setk.run) on the data set. In this way, the strict consensus topology obtained in the cladistic analyses with different optimality criteria of parsimony, showed that Colocania is a monophyletic group, supported by high symmetric resampling (88%) and Bootstrap (82%) ( Figure 47). This monophyly is supported by two unambiguous synapomorphies: costal margin, between base and nodus, convex (char. 20:1), and clunium with oval cuticular depression on central-median area (char. 23:1), present in both males and females (Figures 45,46). This genus is also supported by the homoplasic condition: pterostigma proximally petiolate for around 1/3 of its length (char. 21:2), external parameres stout (char. 17:0); and absence of a mesal transverse endophallic sclerite (char. 18:0). It is related to Ptiloneuropsis diamantina, forming a clade supported by one unambiguous synapomorphy: anterior region of hypandrium with vestigial or short ringed digitiform process (char. 22:1) and by a homoplasic condition: marginal area of the forewing from R4+5 to Cu1a hyaline (char. 10:0). The clade above is related to a clade conformed by Loneura, Loneuroides and Ptiloneura, forming a clade supported by one unambiguous synapomorphy: hindwing with four primary branches in vein M, which could change in future analyzes if we consider all the possible variations that can be observed in the species of these three genera. Furthermore, this clade, as well as the related to the other genera, showed unstable arrangements when the characters tend to have the same weight (K = 5-9) (cladograms not shown Diagnosis. Forewings hyaline, without marginal bands as in C. chicaque sp. n., and C. occidentalis sp. n., differing from them by wing characters, phallosome, and details of hypandrium and epiproct. Unlike the above species, the pterostigma has large pigmented bands proximally and distally (Figure 1); phallosome with mesal endophallic sclerite widened, apically narrow and with two elongated teeth, one of them curved outwards ( Figure 6).

As in Silva
Description. Color (in 80% ethanol). Body pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex with three dark brown areas, a central one and two lateral ones between the compound eyes. Front with brown area between ocellar group and epistomal sulcus, as illustrated ( Figure 3). Postclypeus brown, with diagonal dark brown stripes. Genae, anteclypeus, labrum, mandibles, maxillae, and labium brown to pale brown. Antennae brown, scape pale brown, flagellomeres distally cream. Maxillary palps pale brown, Mx4 dark brown distally. Tergal lobes of meso-and metathorax brown. Thoracic pleura pale brown. Mesopleura with dark brown spots. Legs: coxae, trochanter, and femora pale brown, tibiae and tarsi brown. Wings hyaline, forewing pterostigma with a large dark brown band proximally and distally. Abdomen cream, with subcuticular brown spots; clunium, hypandrium, and phallosome dark brown; epiproct and paraprocts brown.  Figure 4), with three pigmented areas, two antero-lateral, curved, backwards and a central, posterior one, wide and narrow, setose as illustrated; phallosome with side struts independent, with two separate basal stems anteriorly wide, narrowing distally and curved outwards, basally articulated to a mesal process that projects to the hypandrium; external parameres laminar, dilated distally, apex rounded, bearing pores ( Figure 6); anterior pair of endophallic sclerites oval, antero-lateral pair curved as illustrated, rounded distally and overlapping with the basal part of the external parameres ( Figure 6). Paraprocts ( Figure 5) broad, elliptic, with distal setal field as illustrated, sensory fields oval, with 34 trichobothria on basal rosettes. Epiproct ( Figure 5) broadly trapeziform, with a group of three mesal macrosetae and a setal field distally on each side; posterior border with a field of microsetae and a row of four-five setae.
Measurements Diagnosis. Forewings hyaline, without marginal bands as in C. candelaria sp. n., and C. occidentalis sp. n., differing from them by characters of the forewing, phallosome and details of hypandrium and epiproct. Unlike C. candelaria the pterostigma has only a distal pigmented band, sometimes with a median, little defined, small pigmented area (Figure 7); phallosome with narrow mesal endophallic sclerite, with distal processes curved as illustrated. Unlike C. occidentalis, the forewing has M fivebranched, with M5 distally forked (Figure 7).
Measurements Diagnosis. Related to C. pilosa sp. n. in having the forewing with similarly pigmented pattern and shape of the areola postica. It differs from the female of C. pilosa by having the hindwing M widely forked, by the shape of sternum IX, and by the shape and pilosity of the epiproct, without distally curved setae.
Male. Color (in 80% ethanol). Body cream to pale brown, with pigmented dark brown areas as indicated below. Compound eyes black, ocelli hyaline, with ochre centripetal crescents. Vertex with dark brown spots, central and lateral between com-  Genae brown. Postgena pale cream. Antennae brown, flagellomeres cream apically. Maxillary palps pale brown, Mx4 dark brown distally. Tergal lobes of meso-and metathorax dark brown; thoracic pleura dark brown with pale spots. Legs pale brown, coxae with brown spots basally, femora with gray-brown ring basally and gray-brown spots widely distributed; tibiae with gray-brown spot, tarsi 1 brown apically; tarsi 2-3 brown. Wings hyaline, forewing pterostigma with one distal large dark brown band, some with additional proximal spot (not illustrated). Abdomen pale cream, with subcuticular bands brown; clunium and hypandrium brown; clunium with pale central area, hypandrium anteriorly cream; phallosome brown, with sclerites dark brown; epiproct and paraprocts brown.
Morphology. As in diagnosis, plus the following: Head elongate ( Figure  Hypandrium with three pigmented, setose areas, the two anterior widely separated and weakly connected to the posterior area, the latter narrow and extended laterally as a boomerang of little curvature ( Figure 22). Phallosome ( Figure 24) with laminar external parameres, wide, with apically rounded lobe bearing pores, curved inwards; mesal sclerite process with curved outwards apical teeth as illustrated. Side struts Vshaped, basally with an attached complex structure that projects over the hypandrium, distally curved outwards. Anterior endophallic sclerites laminar and oval, with serrate anterior margin, lateral sclerites curved as illustrated. Paraprocts ( Figure 23) almost elliptic, with a distal setal field; sensory fields with 24 trichobothria on basal rosettes. Epiproct (Figure 23 Female. Color (in 80% ethanol). Body, head, legs, epiproct, paraprocts, and wings as in the male, plus the following: Subgenital plate with pigmented area V-shaped, arms wider proximally. Clunium, epiproct and paraprots brown. Gonapophyses dark brown. Sternum IX cream yellowish, darker on the edges.
Measurements. FW: 4350, HW: 2925, F: 1300, T: 2050, t1: 940, t2: 104, t3: 124, Mx4: 330, ctt1: 35, f1: 1130, f2: 1100 Etymology. The specific name refers to the distribution of the species in localities of the western Andean Cordillera in Colombia. Diagnosis. It is related to C. norsantanderina sp. n., described above (see diagnosis of the latter). Both species seem to have a sister-group relationship, to be confirmed when the male of the latter be found. The ninth sterna and gonapophyses in both species are quite similar, but the epiprocts are distinct. The male presents side struts with arms independent, proximally paralell, with processes directed outwards, unique among the species of the genus, distally curved outwards and not fused to the external parameres ( Figure 36).
Colocania is, so far, endemic to Colombia, its species have been collected between 1687 and 2450 meters of altitude, in Andean and Subandean areas (see Rangel and Aguilar 1995) of Central and East-western mountain ranges ( Figure 48). The genus might also be found in Venezuela, as C. norsantanderina was found in a montane area near the Merida mountain range in Venezuela. On basis of the forewing pigmentation pattern, two species groups are recognized, group A, with the forewings having a marginal pigmented band, from R4+5 to distal end of Cu2, including C. norsantanderina sp. n. and C. pilosa sp. n., and group B, with forewings hyaline, including C. candelaria sp. n., C. chicaque sp. n. and C. occidentalis sp. n. However, C. candelaria and C. chicaque are related only on the basis of one homoplasic character, the forewings with a ratio of crossvein Rs-M and the portion of vein M before it of 1:2 (char. 12) (Figure 47). ultad de Ciencias Naturales y Exactas of Universidad del Valle for sponsoring this publication. We thank Dimitri Forero for the loan of the holotype of C. chicaque. We also thank Svanhild Büch and Rubén Luna of the Meremberg Nature Reserve, Ovidio Ledesma, from Planes de San Rafael (Risaralda), Libardo Suárez Fonseca and Orlando Benítez of the National Natural Park Tamá. We thank Oscar Saenz (Universidad del Valle, Santiago de Cali, Colombia) for the phylogenetic analysis. RGO, NSC and JMN thank Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad del Valle, Santiago de Cali, Colombia, for research support. ANGA thanks Instituto de Biología, Universidad Nacional Autónoma de México, for continuous research support.