taxonID	type	description	language	source
275ECD603464FFB53EF4FBD06AAD35B1.taxon	description	Kinzelbach, 1971: 149; Miyamoto and Kifune, 1984: 143; Kathirithamby, 1989: 71; 1990: 471; Pohl et al., 1996: 108; Pohl and Beutel; 2005: 359; Cook and Tribull, 2013: 314	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603464FFB53EF4FB2D6AB13713.taxon	description	Males: Tarsi five-segmented, strongly reduced tarsal claw often present; antennae seven-segmented with segments III and IV laterally flabellate; antennal segment V as long as or longer than segments VI and VII together. Abdominal segment IX ventrally elongated. Aedeagus long, straight or sinuous, but not hooked. Known hosts of the family are species of Heteroptera: Cydnidae.	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603464FFB33EF4F9426D3E3471.taxon	description	(Figures 1 A – B; 2 A – G; 3 A – C; 4 C; 5 A, H; 6 A)	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603464FFB33EF4F9426D3E3471.taxon	materials_examined	Holotype (Fig. 1 A): Male, collected in light trap at Coari, Rio Urucu, Amazonas, Brazil (4 ° 51 ’ 56 ” S – 65 ° 04 ’ 56 ” W); Paulo Bührheim et al., col.; 25 Jan. - 10 Feb. 1995. Specimen preserved whole in glycerin. Paratype (Fig. 1 B) — Male, same collection data as holotype. Paratype specimen received with head separated from body; wing, head, one antenna, and thorax + abdomen mounted on 4 slides. Type material deposited in the Strepsiptera Collection, INPA, Manaus, Brazil. Host and female are unknown.	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603464FFB33EF4F9426D3E3471.taxon	description	Description. Length from tip of frons to distal end of 9 th sternite 3.47 mm. Head (Figs. 2 A, 4 C) — 1.08 mm wide (between edges of eyes); eyes with 14 visible, large, round eyelets occupying ca. 2 / 3 of the head’s width dorsally; inter-eyelet ridges with dense but very short pubescence (microtrichia) (Figs. 4 C). Antefrons in shape of broadly-based isosceles triangle, with rounded angles (Figs. 4 C); vertex plates (epicraneal plates of Kogan, 1958 and Brailovsky and Márquez, 1974) widely separated frontally, oblique, sclerotized, sub-rectangular, sharply narrowing posteriorly, following curvature of eye base; integument smooth, not dimpled. Antennae (Figs. 2 A, 2 C) seven-segmented; segments I and II very short, segments III and IV extended with slender lateral flabella; antennal segments III-VII and flabella covered with sensorial pits of variable shapes and areas; length of antennal segments (in µm): I ≈ II = 50 each; III = 60 (body of segment), flabellum = 810; IV- base + flabellum = 870; V = 500; VI = 230; VII = 280. Mandibles absent. Maxilla (Figs. 2 B, 5 H) club shaped; width at base and tip sub-equal; maxillary palp covered with sensorial trichomes, inserted at proximal end of sub-apical, wide, oval pit, extending by half its length beyond tip of maxilla; length of maxilla 330 µm; palp 200 µm. Thorax (Figs. 2 D-E, 5 A) — Pronotum and mesonotum (Fig. 2 D) very short; mesonotum slightly protruded anteriorly in center, expanding laterally to connect to metanotum. Metanotum (Fig. 2 E) 1.88 mm long, being 6.75 times longer than pro- and mesonotum combined; praescutum broadly rounded anteriorly, protruding from scutum; scutellum in shape of isosceles triangle, with sinuous base; postlumbium lip-shaped, narrow, sinuous; postnotum nearly as long as the other metanotal components together, narrowly rounded posteriorly. Hind wings (Fig. 6 A) with span of 2.64 mm, from base to middle of radial sector, 2.60 mm long from base through edge of sub-costal margin; R 1 almost fused to Sc along middle 1 / 3 of its length; R 2 very short, proximal end slightly overlapping with distal end of Sc; R 3 branching off distal 1 / 3 of R 4; R 4 distal end fading before reaching edge of wing; R 5 about as long as R 3, reaching edge of wing; MA 1 very short, MA 2 extending full length of medial sector; CuA 1 stout at base and most of its length, but fading before edge of wing; CuA 2 well developed, approaching edge of wing; CuP short but evident; illustration (Fig. 6 A) based on slide-mounted wing of paratype that shows vein-like lines caused by folds of anal sector. Legs (Figs. 2 F – G) with hind coxae very short; trochanters of front and middle legs nearly as long as femora; trochanter of hind legs short and stubby, with small lobe projecting beyond insertion of femora (Fig. 2 G); femora and tibiae long and slender; tarsi five-segmented, 1 st tarsomere slender, almost as long as tibiae; tarsomeres 2 - 4 each about half the length of previous one; tarsomeres 1 - 3 with round sensorial pits; tarsomere 5 slim and short, claws barely discernible. Abdomen (Figs. 3 A – C) — Abdominal segments uniformly sclerotized. Terminalia (Figs. 3 A, B, C) with abdominal segment IX elongated, sub-oval, extending ventrally to twice the depth of segment VIII; aedeagus (Fig. 3 C) 0.78 mm long, bi-sinuous, very slender, sharply pointed at tip; segment X (proctiger) reduced to a subrectangular flap slightly curved laterally and truncated (straight edged) distally.	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603464FFB33EF4F9426D3E3471.taxon	discussion	Discussion. T. buehrheimi sp. n. is considerably larger than any of the known congeneric, Neotropical and Nearctic species (Table 2). The most distinctive characters of T. buehrheimi are found in the morphology of the head (Figs. 4 A – C) including: a) the length and density of the inter-eyelet pubescence which is very short and dense in T. buehrheimi, very long in T. paulistana, and intermediate in length and less dense in T. mexicana; b) shape and integumental texture of vertex plates; c) shape of the antefrons, which is similar to that of T. paulistana, and apparently also that of T. vernalis, but clearly different from the antefrons of T. mexicana and T. tecpanensis; and d) shape of the maxilla and point of insertion of the maxillary palp. Maxillae and maxillary palps (Figs. 5 F – I) of T. mexicana and T. tecpanensis are similar, with the palp attaching nearly apically to the body of the maxilla; in both T. buehrheimi and T. vernalis, palps are sub-apically attached to the body of the maxillae; the maxillary palp of T. buehrheimi is much longer than the palps of all other species (Figs. 2 B & 5 H) (structure of T. paulistana maxilla has not been described). Additional distinctive characters in T. buehrheimi are the hind trochanter that projects beyond insertion of the femur (Fig. 2 G); and the aedeagus long, sinuous, and much more slender than that of T. paulistana or T. mexicana (Figs. 3 A, D, E). Presence of a short, but discernible M 1, long CuA 1, and well defined CuP veins in the hind wing further distinguishes T. buehrheim i from the other species. Size variability as a function of host size has been well documented in Strepsiptera (Kathirithamby and Johnston 1992, Cook 2000). Intra-specific size variability of known hosts of Triozocera (Heteroptera: Cydnidae) is likely to occur and would have an influence on the size of the parasites (see below). If this is the case, it follows that body size still may be of significant diagnostic value in this genus, as pointed out by Kifune and Brailovsky (1987), but it must be considered cautiously. Triozocera buehrheimi, T. paulistana, T. mexicana, T. tecpanensis, and T. vernalis form a complex of species clearly differentiated by key morphological features. With only type specimens available for T. buehrheimi, T. paulistana, and T. tecpanensis neither molecular studies nor scanning electron micrograph images can be obtained at this time. Should additional collections be made in the respective type locations or elsewhere in the zoological region, future studies, including molecular analyses, may be possible and will help clarify the affinities of Triozocera spp. in the Neotropics. Such possibility was demonstrated by the identification of a cohort of T. vernalis collected in South Carolina and Georgia, USA (Reeves and Cook (2005). Dedication: The species is dedicated to Paulo Bührheim † former Professor of Biology, Universidade Federal do Amazonas, Manaus, State of Amazonas, Brazil, outstanding entomologist and collector, and a colleague during the time MK was a biologist on the staff of the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil, in early 1960 s.	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603462FFBF3EF4FA626D3C3097.taxon	description	(Figures 3 E, 4 B, 5 C, & 6 I)	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
275ECD603462FFBF3EF4FA626D3C3097.taxon	discussion	T. paulistana was the first species of Strepsiptera originally described from Brazil. A single specimen was collected in a primitive light trap and kept dry for over five years before it was hot KOH treated and mounted in Canada balsam for study. Although the type specimen was damaged, all major diagnostic characters were intact. The species was listed by Kinzelbach (1971) as synonymous of T. mexicana, without the criteria used for the synonymy. Since then, Kifune and Brailovsky (1987) argued against the synonymy based on the difference in size and disjunct geographic distribution of T. mexicana and T. paulistana. They questioned, as well, the synonymy of T. mexicana and T. texana Pierce, 1911, based on an assessment by Luna de Carvalho (1967). Kathirithamby (1990) retained the synonymy of T. mexicana and T. texana, but listed T. paulistana as a separate species; she later accepted that synonymy (Kathirithamby, 1993). Triozocera paulistana distinguishing characters from T. mexicana in the original description were confirmed and expanded in this study. Those differences are reiterated here based on a reassessment of the holotype and Pierce’s original descriptions; examination of 4 specimens identified as T. mexicana from Puerto Rico, Cuba, and Southern USA (Gainesville, Mississippi) (see Table 1); as well as photos posted on the Web of representatives of a large cohort of male Triozocera, likely to be T. mexicana, collected in light traps near Austin, Texas (Quinn, 2008). The list of those diagnostic characters follows and a key to the species of Nearctic and Neotropical species is included at the end of the paper: 1. Vertex (epicraneal) plates narrower in T. paulistana (Fig. 4 B) and widely separated anteriorly (Kogan, 1958; Oliveira and Kogan, 1959) while the plates are wider and nearly touch each other frontally, as seen in all illustrations of T. mexicana (see Pierce, 1913, Plate 1 Fig. 1; 1918, Plate 54 fig. 10 (T. texana), Plate 55 fig. 1 (T. mexicana )); observed in the four specimens used in this comparative study (Fig. 4 A); and seen in photos published in the Web (e. g., Quinn, 2009). 2. Antefrons (frontal tubercle) in T. paulistana with 2 lateral- and one medial lobe projecting anteriorly, in shape of a triangle with rounded angles; postfrons extending posteriorly separating the vertex plates; antefrons in T. mexicana rounded anteriorly (Figs. 4 A & B). 3. Profuse, long inter-eyelet trichomes in T. paulistana (Fig. 4 B); profuse but much shorter trichomes in T. mexicana (Fig. 4 A). 4. Although the wings of the T. paulistana holotype were damaged, it was possible to reconstitute the key elements of the venation. It seems that R 4 does not fork out of the extension of R 5, a character apparently shared with T. tecpanensis that differentiates them from the other species. 5. Subtle differences were also observed in the terminalia (9 th abdominal sternite, 10 th tergite), and base of aedeagus in dorsal view (Figs. 3 D & E). 6. The only known host of T. mexicana is Pangaeus bilineatus Say (Heteroptera: Cydnidae). The species does not seem to occur in Brazil (Lis et al., 2000), but the genus is represented by two other species P. aethiops (Fabr.) and P. neogeus Froeschner (ESALQ-USP, 2004). Although host specificity in Triozocera has not been documented, the absence of the T. mexicana host in Brazil, may offer an additional argument for the separation of the two species. As for the arguments raised by Kifune and Brailowsky (1987), the following considerations are in order. Differences in size (T. paulistana much larger than T. mexicana) resulted from artifact of the mounting procedure of T. paulistana. Boiling the specimen in 70 % KOH caused the abdomen to expand. Using the illustration in Kogan (1958), Fig. 1, the area corresponding to the inter-segmental membranes of the abdomen was deleted and the length of the specimen reassessed. The length was reduced from the originally reported 3.08 mm, to 2.89 mm, well within the range of T. mexicana, T. vernalis and T. tecpanensis, but smaller than T. buehrheimi (Table 2). The importance of body size in Strepsiptera should be considered in light of the size variation of the host, as demonstrated by Kathirithamby and Johnston (1992) and Cook (2000) for Caenocholax fenyesi Pierce, the males of which are parasitic on ants. Pangaeus bilineatus, the host of T. mexicana, is a ubiquitous cydnid in North America, and body length variability has been observed but not measured (Lis et al., 2000). Intraspecific size variability in Heteroptera has been studied in a few species, but the range of variability seems to be considerably less than that observed in Formicidae. For instance, body length variability in Pyrrhocoris apterus L. (Heteroptera: Pyrrhocoridae), as affected by rearing conditions, was under 10 % (Honĕk, 1987). In Nezara viridula L. (Heteroptera: Pentatomidae), in a family that contains Strepsiptera hosts, male size variability may reach 20 % (McLain, 1985). Length variability in Formicidae, however, is much greater. Cushman et al. (1993) reported a 52 % difference in body size among ants in a species of Camponotus. We infer from this that body size variability is expected in Triozocera, but the range of that variability probably is significantly smaller than that found in Caenocholax. Unless the various body proportions show tendency to gigantism, as in T. buehrheimi, smaller differences, particularly in total body length, probably are less significant. Finally, the argument of the disjunct geographic distribution of the two species needs to be reconsidered. Caenocholax fenyesi Pierce, 1909 is an example of introgression of a Neotropical Strepsiptera species both ways into North and South America. C. fenyesi was originally described from males collected in Cordoba, Mexico. Its recorded range has since been extended north to nine Southern USA States, and south to 10 Central American countries and the Antilles, and to three and possibly five countries in South America (Cook et al., 1997; Kathirithamby and Hughes, 2002). This expansive distribution may, perhaps, involve a complex of cryptic lineages the nature of which is beginning to be unraveled (Hayward et al., 2011). What seems certain, however, is that Strepsiptera distribution depends on suitable host availability (Kathirithamby, 2009). We conclude that T. paulistana differs from T. mexicana in various morphological characters (apomorphies), but we deemphasize the importance of differences in body size and disjunct distribution of the two species. The key differential characters are: a) profuse and long inter-eyelets pubescence; b) shape and structure of the vertex plates with wide separation of the plates anteriorly; c) shape of the antefrons (frontal tubercle); d) apparent lack of an R 4 vein off of the R 5; and, e) subtle differences in the terminalia. The absence in South America of Pangaeus bilineatus, host of T. mexicana, may also be of significance in support of the revalidation of T. paulistana.	en	Kogan, Marcos, Cook, Jerry L. (2014): The genus Triozocera Pierce, 1909 (Insecta: Strepsiptera: Corioxenidae) in South America. Zootaxa 3779 (1): 48-60, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.7
