Eucelatoria gladiatrix ( Townsend, 1917 )

Burington, Zelia L., 2022, A taxonomic revision of the Eucelatoria ferox species group (Diptera: Tachinidae), Zootaxa 5143 (1), pp. 1-104 : 33-35

Eucelatoria gladiatrix ( Townsend, 1917)

( Figs 1G , 12 , 35 , 57 , 80 , 101 , 116 , 130 , 146 , 163 , 174 )

Xiphomyia gladiatrix Townsend, 1917: 126 . Holotype, Panama, Cabima (USNM). Other references: Guimarães (1971: 151).

Proroglutea pilligera Townsend, 1919: 574 . Holotype, Costa Rica, San Carlos (USNM). Other references: Guimarães (1971: 146). Syn. nov.

Eucelatoria gladiatrix: Wood (1985: 40) .

Type material examined. Holotype ♀, labeled “CabimaPan/ May 17.11/ AugustBusck”, “Type No./ 20799/ U.S. N.M.”, “ Xiphomyia / gladiatrix/ ♀ / DetCHT T.” ( USNM).

Holotype ♂ of “ Proroglutea pilligera ”, labeled “SanCarlos/ CostaRica”, “CollSchild/ & Bingdorf”, “Type No./ 22252/ U.S. N.M.”, “ Proroglutea / ♂ pilligera/ DetCHT T.” ( USNM).

Other material examined. 13 ♀♀ and 13 ♂♂. Costa Rica — Guanacaste : 1 ♂, “81-SRNP-280”, “ Santa Rosa National Park / Guanacaste Province / Costa Rica, D.J. Janzen ” ( CNC) . 1 ♂, “LEGS AWAY/ FOR DNA”, “DHJPAR0035706”, “Voucher: D.H. Janzen & W. Hallwachs / DB: http://janzen.sas.upen.edu/ Area de Conservacion Guanacaste,/ COSTA RICA./ 09 -SRNP- 41488”, “ZLB_E.Ferox 00363” ( CNC) . 5 ♂♂ and 7 ♀♀, as previous except DHJPAR numbers “DHJPAR0017869”, “DHJPAR0035982”, “DHJPAR0035824”, “DHJPAR0035730”, “DHJPAR0046689”, “DHJPAR0034414”, “DHJPAR0035743”, “DHJPAR0014981”, “DHJPAR0035800”, “DHJPAR0024451”, “DHJPAR0017870”, and “DHJPAR0014980”, SRNP numbers “05-SRNP-41790”, “09- SRNP-70832”, “09-SRNP-69469”, “09-SRNP-41491”, “11-SRNP-66139”, “09-SRNP-41155”, “09-SRNP-41780”, “06-SRNP-41826”, “09-SRNP-69389”, “08-SRNP-70391”, “05-SRNP-24270”, and “06-SRNP-41925”, and specimen IDs “00364” [♂], “00365”[♂], “00367”[♂], “00360”[♂], “00378”[♀], “00377”[♀], “00379”[♀], “00380”[♀], “00381”[♀], “00382”[♀], “00375”[♂], “00376”[♀] [final two with terminalia in vial under specimen] ( CNC) . 1 ♀, “ COSTA RICA: Guanacaste Prov. / Diria N.P. 933 m / 10°08.34′ S [sic; N] 85°37.50′W / 5–6.I.2015 / J.O. Stireman, A. Eckhardt ”, “JOS115.30”, [pink label], “ Eucelatoria / sp. ( Xiphomyia grp.)”, “ZLB_E.Ferox 00328” ( JOSC). Ecuador —Orellana : 1 ♂, “ ECUADOR: Orellana Prov. / Shiripuno Lodge/ 01°06′ 282″ S 076°43′ 896″W/ 28–29- xii - 2012 / J.O. Stireman III”, “JOS1212.87”, [blank pink label], “ Eucelatoria sp. / Det. J.O. Stireman III”, “ZLB_E.Ferox 00370” ( JOSC; to be deposited in MECN). Mexico — Oaxaca : 1 ♀, “ MEXICO Oax 1 km N/ Portillo del Rayon / ca Km 188 1400m / 25.VII.92 D.M.Wood ”, “ZLB_E.Ferox 00285” ( DMW). Veracruz : 1 ♀, “ MEX: Veracruz, UNAM/ Los Tuxtlas Estac./ Biol., N Catemaco/ 16–19 Sept. 1989 / E Barrera, TJ Henry,/ & IM Kerzhner colls.”, “ Eucelatoria / gladiatrix/ (Townsend)/ det. Woodley 1989”, “USNM ENT 00039945”, “ZLB_E.Ferox 00385” ( USNM). PANAMA—PM : 1 ♂, “Pantilla Pt./ Canal Zone/ 15.I.1929 ”, “Collector/ C. H. Curran ”, “ Proroglutea / pilligera [label folded]”, “Reinhard/ Collection”, “ Proroglutea / pilligera/ R Tns.”, “ZLB_E.Ferox 00369” ( CNC). Peru — Loreto : 1 ♀, “ 24VIII 1965 San Antonio/ LORETO, PERU 69/ Coll.J.C. Hitchcock,Jr. ”, “USNM ENT 00875971”, “ZLB_ E.Ferox 00356” ( USNM). Madre de Dios : 1 ♀, “Avispas, Madre/ de Dios, PERU / 10–20.IX.1962 / L. Pena. 400m. ”, “ZLB_E.Ferox 00383” ( CNC). San Martín : 1 ♀, “Tabalosas[sic]/ 8.XI Peru”, “ Xiphomyia / gladiatrix/ TT ♀ / DetCHTT”, “USNM ENT 00039941”, “ZLB_E.Ferox 00384” ( USNM). Venezuela — Aragua : 4 ♂, “ VENEZUELA Aragua / 11 km n Rancho/ Grande 25.II.1971 / G. & M. Wood ”, the specimen IDs “ZLB_E.Ferox 00371”, “00372”, “00373”, and “00374” [final three with genitalia in vial under specimen] ( DMW) .

Recognition. This species is similar to E. falcata sp. nov., E. yanayacu sp. nov., and E. charapensis in its large body size, V-shaped thoracic vittae, and minute ocellar setae. It differs in that the thin thoracic vittae do not merge on the postsutural area, the abdominal setulae are entirely black, and the abdominal tomentum bands are uninterrupted by a median vitta. Males of E. gladiatrix are distinct among all E. ferox group species in that the abdominal T5 bears a ventrolateral pair of patches bearing strong, oily black setulae (“sex patches”). The female piercer is the longest in the E. ferox group, the aculeate lobe usually 12 times or more the height of the seventh segment base. The piercer tip is usually held against the postmetacoxal membrane, making the aculeate lobe appear as a curved rod between the abdominal apex and hind coxae in lateral view. Females are nearly identical to E. luctuosa , which also has uninterrupted abdominal bands, but differ in having strong ocellar setae, a narrower parafacial, and a shorter piercer.

Redescription. [Redescribed from 11 ♀♀ and 13 ♂♂. Head measurements are from 3 ♀♀ and 3 ♂♂.] Length 6.4–10.1 mm (mean = 8.2 mm).

Head. Parafacial, face, fronto-orbital plate, vertex, post-ocular plate, and vibrissal angle tomentum yellow, shining gold in the light, often with the ventral half of the post-ocular area shining gray. Genal dilation tomentum yellow-gray. Postgena and occiput tomentum ash-gray. Pale occipital setulae yellow to yellow-gray. Ocellar setae minute, barely differentiated from surrounding setulae of ocellar triangle, or apparently absent. Gena with 5–6 setae, subvibrissal ridge with 2 setae. Facial ridge with setulae on lower one-third or less. Postocellar setae two-thirds to three-fourths length posterior reclinate orbital seta. Paravertical seta subequal to length of postocellar setae. Outer vertical seta usually indistinguishable from post-ocular row of setae. Reclinate orbital setae 2–3. Frontal setae 6–9, usually less in females, base of last frontal seta level with arista. Eye apparently bare. Eye height to head height ratio 0.85–0.90 (mean = 0.88). Postpedicel length 0.34–0.37 (mean = 0.36) height of head. Facial ridge length 0.45–0.50 (mean = 0.48) height of head. Parafacial width 0.07–0.13 (mean = 0.09) lateral length of head. Pedicel 0.28–0.38 (mean = 0.32) length of postpedicel. Postpedicel 1.11–2.67 (mean = 2.13) times width of parafacial in lateral view. Vertex 0.15–0.25 (mean = 0.22) width of head in dorsal view. Palpus brown, subcylindrical; slightly flattened and inflated at the apex, more so in females; with short dense setulae dorsoapically, longer setulae mediolaterally, and numerous fine setulae ventrally from apex nearly to base.

Thorax. Dorsomedial length 1.3–1.5 times width of thorax. Lateral tomentum ash-gray on katepisternum, usually yellow-gray on anepisternum. Dorsal tomentum yellow, less dense and grayer between inner and outer vittae. Presutural outer vitta triangular, usually disconnected from anterior margin. Postsutural outer vitta elliptical, pointed posteriorly, distinct from inner vittae but often seems connected due to less dense intermediate tomentum. Inner thoracic vittae linear, bar shaped, slightly divergent, distinct from outer vittae, extending on postsutural area to level of second dorsocentral seta. Scutellar dorsal tomentum yellow to yellow-gray, covering three-fourths of surface. Postpronotum with 3–4 setae. Presutural area with 2 supra-alar setae. Postsutural area with 4 dorsocentral setae. Scutellum with 1 pair discal setae. Fore tibia with 2–3 posterodorsal setae. Wing hyaline to lightly infuscated around costa and radial sector [in Venezuelan specimens, infuscation darker and extending to membranes along primary wing veins in anterior half of wing]. Calypters tan to cream colored.

Abdomen. Cuticle and setulae entirely black. Dorsal tomentum bands ash-gray, gold tinted near anterior margin; continuous, without a median vitta; covering one-third to one half length of T3–5. Ventral tomentum bands ashgray to yellow-tinged gray, covering one-half to two-thirds length of T3–5; in male, T5 with ventral patch of dense, dark oily setulae on shiny, slightly concave cuticle devoid of tomentum.

Male terminalia ( Figs 130 , 146 , 163 ). Sternite 5 basal plate with median teeth squared off, apparently separated 0.6 width of basal notch; apical lobes 2 times length of basal plate. Postgonite strongly hooked, rounded on posterior margin, with anterior emargination 2 times width postgonite at mid length. Surstylus slightly longer than cercus, anterior and posterior margins subparallel on apical three-fourths, blunt at apex. Cercus in lateral view expanded at base, gradually narrowed at mid length, apical lobe with subparallel margins, cylindrical; in caudal view upper lobe rectangular, subparallel at median section, apical lobe gradually narrowed to blunt apex. Upper lobe 0.35 length of cercus. Median section 0.35 length of cercus. Apical cleft parallel sided, 0.3 length of cercus. Syncercus apex in caudal view 0.5 width of syncercus base.

Female terminalia ( Figs 101 , 116 ). Piercer generally extending past base of abdomen, in lateral view gradually curved to a slightly bent apex; in posterior view parallel sided on nearly entire length, gradually narrowed to apex on apical fifth. Aculeate lobe 6 times height of segment 7 base. End tergite narrow, length ten times width, narrowed near apex to blunt point. Cercus with 4–5 setulae; ventral extension no longer than width of main cercus body. Postgenital plate with 14 setulae.

Hosts. Thirty-eight individuals have been reared from species of Crambidae at Area de Conservación Guanacaste, Costa Rica. Five of these were reared from Desmia Westwood species [including D. benealis Schaus and D. tages (Cramer) ], one was reared from Palpusia plumipes (Dognin) , eight were reared from a Leucochromodes Amsel species, one was reared from a Pilocrocis Lederer species, one was reared from Phostria latiapicalis (Schaus) , and the remainder were reared from Piletosoma thialis Dyar and another Piletosoma Hampson species. Hosts were found feeding on several species of understory trees in the families Dilleniaceae , Rubiaceae , and Actinidiaceae , including Hamelia patens Jacq. , Uncaria tomentosa (Willd. ex Schult.) DC. , Chimarrhis parviflora Standl. , Sauraria montana Seem. , and Dolicocarpus multiflorus Standl. ( Janzen & Hallwachs 2008; see Table 2 ).

Geographic extent and seasonal occurrence. Eucelatoria gladiatrix has been collected in lowland tropical rainforest and seasonal forests from southern Mexico to Northern Peru, with intermediate records from Costa Rica, Panama, Venezuela, and Ecuador ( Fig. 174 ). Reared individuals in Costa Rica emerged throughout the year, with the largest emergences in June and July ( Janzen & Hallwachs 2008).

Discussion. The presence of male sexual patches is an uncommon condition in Eucelatoria , and it is unknown in any other species of the E. ferox group. The E. gladiatrix patches are found only on the 5th abdominal tergite, a condition rare in Tachinidae , and are composed of thick, apparently oily setulae with a round depression-like annulus at each setal base set in shiny brown cuticle. This appears to match either Type I or II in the sexual patch classification of Cerretti et al. (2015), but electron photomicrographs would be needed to further classify them. The authors suggest the position on T5 rather than the typical T4 is due to “rearward shift of the focus of the activating or permissive signal” ( Cerretti et al. 2015). As all known males from closely related species apparently lack sexual patches, it is not clear why males of E. gladiatrix uniquely possess them.

The E. gladiatrix piercer is the longest known in any Tachinidae , perhaps any Diptera barring some Tephritidae . It is somewhat flexible, such that the female can direct the apex against the postmetacoxal area when not in use. The disparity of male and female sexual characters led Townsend to name them originally as two separate genera, Proroglutea and Xiphomyia , but genetic evidence ( Fig. 1 ) supports their synonymy. Males and females are otherwise identical in color patterns and other external characters.

With close to forty reared specimens, E. gladiatrix has one of the best records of host use of any species of the E. ferox group. The species seems to specialize on leaf-rolling Crambidae caterpillars feeding on broad-leaved tropical understory trees. Its use of five host genera would suggest it has a reasonably broad range of host use, a characteristic that may explain its wide distribution in the Neotropical Region.

The Mexican species E. rivalis and E. luctuosa resemble E. gladiatrix in coloration and vittae patterns, but in both cases the ocellar setae are distinct rather than minute or apparently absent. Genetic data weakly support E. gladiatrix as sister to a clade of several species, including North American E. ferox group members ( Fig. 1 ). More interesting is the apparent genetic structure with at least three distinct and well supported genetic clusters ( Fig. 1 ). As there are few morphological differences among the examined specimens, and as the genetic clusters are not divided by locality, it is not clear whether these represent intraspecific genetic variation or a cryptic species complex.

Figures

FIGURE 1. CO1-based neighbor-joining tree of 12 species of the Eucelatoria ferox species group, with inset illustrations of mesonota. The outgroup is Eucelatoria armigera (Coquillett). Node labels indicate support values over 60. Scale bar = 2% genetic difference. The E. charapensis (Townsend) sequence with the axial break is of poor quality but is included to show unity of El Bigal specimens with the rest of the metapopulation. Mesonota are illustrated with setal annuli only for clarity. A. E. gustavogutierrezi sp. nov. B. E. kopis sp. nov. C. E. ritavargasae sp. nov. D. E. ferox (Townsend). E. E. huitepecensis sp. nov. F. E. auriceps (Aldrich). G. E. gladiatrix (Townsend). H. E. borealis sp. nov. I. E. yanayacu sp. nov.

FIGURES 10–13. Female habitus photographs of Eucelatoria ferox group spp., dorsal view. 10. E. falcata sp. nov., allotype (ZLB_E.Ferox 00276) (DMW). 11. E. fordlandia sp. nov., allotype (ZLB_E.Ferox 00314) (USNM). 12. E. gladiatrix (Townsend), Guanacaste Province (ZLB_E.Ferox 00380) (CNC). 13. E. jorgecortesi sp. nov., holotype (ZLB_E.Ferox 00401) (CNC).

FIGURES 34–37. Male habitus photographs of Eucelatoria ferox group spp., dorsal view. 34. E. fordlandia sp. nov., holotype (ZLB_E.Ferox 00299) (USNM). 35. E. gladiatrix (Townsend), Guanacaste Province (ZLB_E.Ferox 00365) (CNC). 36. E. sabroskyi sp. nov., holotype (ZLB_E.Ferox 00070) (USNM). 37. E. strigata (Wulp), Trinidad (ZLB_E.Ferox 00141) (CNC).

FIGURES 56–57. Female habitus photographs of Eucelatoria ferox group spp., lateral view. 56. E. fordlandia sp. nov., allotype (ZLB_E.Ferox 00314) (USNM). 57. E. gladiatrix (Townsend), Guanacaste Province (ZLB_E.Ferox 00380) (CNC).

FIGURES 80–81. Male habitus photographs of Eucelatoria ferox group spp., lateral view. 80. E. gladiatrix (Townsend), Guanacaste Province (ZLB_E.Ferox 00365) (CNC). 81. E. sabroskyi sp. nov., holotype (ZLB_E.Ferox 00070) (USNM).

FIGURES 94–108. Female seventh abdominal segment of Eucelatoria ferox group spp., lateral view. Numbers are “ZLB_ E.Ferox” specimen IDs. 94. E. ferox (Townsend), Santa Cruz (00398) (USNM). 95. E. aurata (Townsend), Nova Teutonia (00250) (DMW). 96. E. auriceps (Aldrich), Crane Hollow (00410) (JOSC). 97. E. borealis sp. nov., Sheffeld (00041) (DMW). 98. E. charapensis (Townsend), Chapare Province (00393) (DMW). 99. E. falcata sp. nov., Nova Teutonia (00274) (CNC). 100. E. fordlandia sp. nov., Fordlandia (00311) (USNM). 101. E. gladiatrix (Townsend), Guanacaste Province (00376) (CNC). 102. E. sabroskyi sp. nov., Herb Martyr Campground (00084) (CNC). 103. E. strigata (Wulp), Trinidad (00138) (CNC). 104. E. texana (Reinhard), Madera Greaterville Rd. (00048) (JOSC). 105. E. yanayacu sp. nov., Yanayacu (no “E.Ferox” number, labeled “DIP328”) (USNM). 106. E. gustavogutierrezi sp. nov., Guanacaste Province (00007) (CNC). 107. E. kopis sp. nov., Avispas (00217) (CNC). 108. E. sica sp. nov., Yanayacu (00211) (CNC).

FIGURES 109–123. Female abdominal segments eight through ten of Eucelatoria ferox group spp., dorsal view. Numbers are “ZLB_E.Ferox” specimen IDs. 119. E. ferox (Townsend), Santa Cruz (00398) (USNM). 110. E. aurata (Townsend), Nova Teutonia (00250) (DMW). 111. E. auriceps (Aldrich), Crane Hollow (00410) (JOSC). 112. E. borealis sp. nov., Sheffeld (00041) (DMW). 113. E. charapensis (Townsend), Chapare Province (00393) (DMW). 114. E. falcata sp. nov., Nova Teutonia (00274) (CNC). 115. E. fordlandia sp. nov., Fordlandia (00311) (USNM). 116. E. gladiatrix (Townsend), Guanacaste Province (ZLB_ E.Ferox 00379) (CNC). 117. E. sabroskyi sp. nov., Herb Martyr Campground (00084) (CNC). 118. E. strigata (Wulp), Trinidad (00138) (CNC). 119. E. texana (Reinhard), Madera Greaterville Rd. (00048) (JOSC). 120. E. yanayacu sp. nov., Yanayacu (no “E.Ferox” number, labeled “DIP328”) (USNM). 121. E. gustavogutierrezi sp. nov., Guanacaste Province (00007) (CNC). 122. E. kopis sp. nov., Avispas (00217) (CNC). 123. E. sica sp. nov., Yanayacu (00211) (CNC).

FIGURES 124–135. Male terminalia of Eucelatoria ferox group spp., left lateral view. Numbers are “ZLB_E.Ferox” specimen ID numbers. 124. E. aurata (Townsend), Nova Teutonia (00100) (CNC). 125. E. auriceps (Aldrich), Springfield (00018) (JOSC). 126. E. borealis sp. nov., St. Lawrence Island (00032) (CNC). 127. E. charapensis (Townsend), Chiapas (00202) (DMW). 128. E. falcata sp. nov., Nova Teutonia (00271) (CNC). 129. E. fordlandia sp. nov., Fordlandia (00361) (USNM). 130. E. gladiatrix (Townsend), Guanacaste Province (00375) (CNC). 131. E. sabroskyi sp. nov., Greaterville Rd. (00071) (USNM). 132. E. strigata (Wulp), Trinidad (00147) (CNC). 133. E. texana (Reinhard), College Station (00048) (CNC). 134. E. woodorum sp. nov., Xochipala (00201) (DMW). 135. E. yanayacu sp. nov., Yanayacu (00189) (USNM).

FIGURES 136–155. Male terminalia of Eucelatoria ferox group spp., left lateral view, and caudal view of cerci and surstyli. Numbers are “ZLB_E.Ferox” specimen ID numbers. Specimen information for caudal views is identical to that for Figs 124– 139. 136. E. gustavogutierrezi sp. nov., lateral view, Guanacaste Province (00008) (CNC). 137. E. hafelei sp. nov., lateral view, Chapare Province (00386) (DMW). 138. E. ritavargasae sp. nov., lateral view, Narupa (00215) (USNM). 139. E. sica sp. nov., lateral view, Yanayacu (00213) (CNC). 140. E. aurata (Townsend), caudal view. 141. E. auriceps (Aldrich), caudal view. 142. E. borealis sp. nov., caudal view. 143. E. charapensis (Townsend), caudal view. 144. E. falcata sp. nov., caudal view. 145. E. fordlandia sp. nov., caudal view. 146. E. gladiatrix (Townsend), caudal view. 147. E. sabroskyi sp. nov., caudal view. 148. E. strigata (Wulp), caudal view. 149. E. texana (Reinhard), caudal view. 150. E. woodorum sp. nov., caudal view. 151. E. yanayacu sp. nov., caudal view. 152. E. gustavogutierrezi sp. nov., caudal view. 153. E. hafelei sp. nov., caudal view. 154. E. ritavargasae sp. nov., caudal view. 155. E. sica sp. nov., caudal view.

FIGURES 156–173. Male terminalia and fifth abdominal sternite of Eucelatoria ferox group spp. Specimen information for fifth sternites is identical to that for Figs 124–139. 156. Generic male terminalia, left lateral view (left) and caudal view (right), setae and membrane omitted.Abbreviations: aed, aedeagal apodeme; bas, basal lobe of surstylus; C, depth of postgonite anterior emargination; cer, cercus; epa, epandrium; hyp, hypandrium; pha, phallus; pos, postgonite; pre, pregonite; sur, surstylus. 157. E. aurata (Townsend). 158. E. auriceps (Aldrich). 159. E. borealis sp. nov. 160. E. charapensis (Townsend). 161. E. falcata sp. nov. 162. E. fordlandia sp. nov. 163. E. gladiatrix (Townsend). 164. E. sabroskyi sp. nov. 165. E. strigata (Wulp). 166. E. texana (Reinhard). 167. E. woodorum sp. nov. 168. E. yanayacu sp. nov. 169. E. gustavogutierrezi sp. nov. 170. E. hafelei sp. nov. 171. E. ritavargasae sp. nov. 172. E. sica sp. nov.

FIGURES 174–179. Distribution maps of Eucelatoria spp. 174. E. yanayacu sp. nov., E. gladiatrix (Townsend), E. falcata sp. nov., E. charapensis (Townsend), and E. fordlandia sp. nov. 175. E. strigata (Wulp) and E. aurata (Townsend). 176. E. hafelei sp. nov., E. gustavogutierrezi sp. nov., and E. ritavargasae sp. nov. 177. E. ferox (Townsend), E. huitepecensis sp. nov., and E. inclani sp. nov. 178. E. kopis sp. nov., E. sica sp. nov., E. makhaira sp. nov., and E. tenebrionis sp. nov. 179. E. rivalis (Reinhard), E. luctuosa (Wulp), E. woodorum sp. nov., E. crambivora sp. nov., and E. jorgecortesi sp. nov.