Beebeomyia taccarivora Wendt and Maia, 2018

Wendt, Lisiane Dilli, Gonçalves, Eduardo Gomes & Maia, Artur Campos Dália, 2018, A new species of Beebeomyia Curran (Diptera: Richardiidae) from Brazil, with description of immature stages and notes on their association with Taccarum ulei (Araceae), Zootaxa 4369 (4), pp. 587-599 : 589-595

Beebeomyia taccarivora Wendt and Maia sp. nov.

( Figs 1–52 )

Diagnosis and comments. Beebeomyia taccarivora is quite different from all other known speCies of the genus Beebeomyia . It is the only known speCies within the genus exhibiting an entirely yellowish orange thorax with a Central brown stripe on the sCutum, and a brownish orange abdomen with a Central brown stripe on tergites 1–4. The other CongeneriCs exhibit blue to blaCk-Colored abdomens, and entirely or parts of the thorax with reddish to blaCk Coloration (see first step of identifiCation key in Hernández-Ortiz and Aguirre 2015). Similarly to B. senilis (Hennig) , B. taccarivora has only one pair of dorsoCentral setae, but the two speCies differ from one another in the general Coloration of the thorax and abdomen – whiCh are blaCk in B. senilis . Besides, differently from all other Beebeomyia speCies, B. taccarivora exhibits the following Combination of CharaCters: fore femur with one preapiCal anteroventral spine-like seta and a posteroventral row of spine-like setae on the apiCal half; legs entirely yellowish orange; postgenal seta absent; proepimeral seta robust; and abdominal tergites densely Covered with whitish setulae.

The only other speCies of Beebeomyia for whiCh the male and female terminalia have been desCribed and illustrated is B. tuxtlaensis Hernández-Ortiz and Aguirre. The female terminalia of B. taccarivora and B. tuxtlaensis are Considerably different. Beebeomyia tuxtlaensis has subtriangulate CerCi, with a very narrow and rounded and tip, whereas in B. taccarivora the CerCi are subreCtangular, with a straight lateral margin and subapiCal region narrowing gradually, but with a distinCtly straight and trunCated tip. On the other hand, the male terminalia of the two speCies are more similar, speCially the epandrium and CerCi. DifferenCes are that the medial surstylus in B. tuxtlaensis has a ventral triangular protuberanCe at Ca. mid-length, the lateral surstylus has a mid-dorsal toothlike projeCtion, and the apex of the distiphallus has two opposite rows of modified setae; in B. taccarivora , the medial surstylus has a ventral preapiCal protuberanCe, the lateral surstylus does not possess a tooth-like projeCtion, and the apex of the distiphallus has only one row of modified setae.

Adult description. Holotype male. Body length: 6 mm; wing length 4.9 mm; abdomen length 2.5. Head ( Figs 5–7 ). Yellowish orange, exCept for two dark brown spots on faCe; frons weakly and sparsely whitish, pollinose. Contour of oCelli ConspiCuously marked in blaCk. Orbital seta shorter than oCellar seta and inserted at the same level. SCape and pediCel yellowish orange. First flagellomere 4.25 times longer than high; brown-Colored with yellowish orange dorsal, ventral and proximal margins. Arista dark brown. Clypeus, palpus and prementum yellowish orange. PostoCellar seta very weak. Genal seta blaCk-Colored and strong. Postgenal seta absent. PostCranium with a band of whitish pollinosity on Contour of eye; sparsely setulose with short, yellowish setae, and with three stout and blaCk setulae at median region. Thorax ( Figs 1, 6 ). Yellowish orange with a Central brownish stripe on sCutum; Covered with whitish and thin pollinosity, distinCtly denser on Central stripe, postpronotal, notopleuron and katatergite; sCutum yellowish, setulose. Halter yellowish white; Calypter small, white-Colored with a brownish Contour. Proepimeron setulose. Proepimeral seta blaCk and strong; presutural supra-alar seta absent; one dorsoCentral seta; two sCutellar setae; intra-alar seta distinCtly weak and shorter than post-alar seta; katepisternal seta very weak. Legs ( Fig. 1 ). Femora thiCker than tibiae; yellow-Colored. Fore Coxa with yellowish setae and setulae. Fore femur with one preapiCal spine-like seta on anteroventral surfaCe and a row of spine-like setae on apiCal half of posteroventral surfaCe; posterodorsal surfaCe with a row of three blaCk setae on apiCal third. Mid Coxa with one robust and blaCk anteromedial seta. Mid and hind femora with row of spine-like antero- and posteroventral setae on apiCal half. Hind Coxa with strong and blaCk setae on anterior surfaCe. Hind femur with two preapiCal anterodorsal setae. Wing ( Fig. 3 , photo of paratype female). Entirely hyaline, exCept for very faint and small brown mark at apex. Entirely miCrotriChose, exCept Cells bm and bCu. Crossvein r-m at middle of Cell dm. Abdomen ( Fig. 1 ). Brownish orange, with Central brown stripe on tergites 1–4. Tergites densely Covered with whitish setulae, and with row of long and blaCk setae at apiCal margins of tergites 4 and 5. Syntergite 1+2 with three blaCk laterodorsal preapiCal setae; apex 2.4 times wider than base; 1.8 times longer than tergite 3. Tergites 3–4 gradually narrower than apex of syntergite 1+2. Sternites observed and illustrated using a paratype ( Figs 8–11 ): sternite 1 absent, sternite 2 divided into two parts: anterior part very reduCed and slightly sClerotized, and posterior part elongated, “Y-shaped” ( Fig. 8 ) and relatively sClerotized; sternites 3 and 4 shortly and weakly setulose, exCept for a row of strong and long setulae on apiCal margins ( Figs 9–11 ); sternite 3 longer than wide, with base narrower than apex ( Fig. 9 ); sternite 4 reCtangular, longer than wide ( Fig. 10 ); sternite 5 subquadrangular, slightly wider than long ( Fig. 11 ). Two ConspiCuous spiraCles: on tergite 6 and on syntergosternite 7+ 8 ( Fig. 14 ); spiraCle vestigial, very reduCed on sternite 6 ( Fig. 14 , spiraCle indiCated by arrow). Tergite 6 with sparse setae on ventral half, different in size ( Fig. 14 ). Sternite 6 reduCed and slightly sClerotized ( Fig. 14 ). Syntergosternite 7+8 densely pollinose, exCept on apiCal and right lateral fourth, bare; with sparse setae on apiCal half, similar in length and thiCkness ( Fig. 14 ). Male terminalia (observed using a paratype, Figs 12–16 ). Epandrium, in posterior view, slightly wider than high ( Fig. 12 ). Lateral surstylus slightly longer than medial surstylus, developed, wide, Covering outer lateral portion of medial surstylus ( Figs 12–14, 16 ). Medial surstylus Curved, without projeCtion on anterior basal, Ciliated ( Figs 12, 14, 16 ). Prensiseta robust and apiCally bifurCated, inserted at apex of medial surstylus ( Figs 12, 13, 14, 16 ). CerCi developed and protruded, slightly sClerotized, rounded and Covered with setae of different lengths ( Figs 12–14 ). Distiphallus long and flexible, with two sClerotized ribbons ConneCted by a membrane ( Fig. 14 ), setulose on basal third and bare on apiCal two-thirds; apex with a row of modified setae, similar to dentiCles, and a sClerotized proCess plate ( Figs 14, 15 ).

Variation, paratypes male. No morphologiCal variation among the paratype was found.

Paratypes female. Body length 5.5–5.7 mm (n=2); ( Fig. 2 ). As male, exCept for the following features: Abdomen ( Figs 17–22 ). Sternite 1 present, divided into two small and weakly sClerotized plates, bare ( Fig. 17 ); sternite 2 divided into two parts: anterior very reduCed and slightly sClerotized, and posterior elongated, “Yshaped” ( Fig. 18 ) and relatively sClerotized; sternites 3–5 shortly and weakly setulose, exCept for a row of strong and long setulae on subapiCal margin ( Figs 19–21 ); sternites 3 and 4 longer than wide, base narrower than apex ( Figs 19, 20 ); sternite 5 subreCtangular, distinCtly wider than long ( Fig. 21 ); sternite 6I hemispheriCal, distinCtly wider than long ( Fig. 22 ). Female terminalia ( Figs 23–26 ). OvisCape 0.8 mm long (n=1), brownish yellow, with a Central, wide brown stripe on apiCal two-thirds; apiCal margin straight ( Fig. 23 ). Taenia brown, short, one-half as long as ovisCape; weakly sClerotized, distinCtly less sClerotized than ovisCape ( Fig. 23 ). Eversible membrane subhyaline ( Fig. 23 ), weakly Covered with dentiCles on basal two-thirds. Sternite 8 0.5 mm long (n=1), weakly sClerotized, Ciliated on apiCal three-fifths, shorter than tergite 8 ( Fig. 25 ). CerCi 0.21 mm long (n=1); straight at lateral margin, with subapiCal region narrowing gradually; tip 0.042 mm wide (n=1), trunCated ( Fig. 26 ): CerCus with six setae ( Fig. 26 ): two long medial setae: lateral shorter setae, two subapiCal setae similar in length, one slightly longer preapiCal seta, and one very small apiCal seta. Two spheriCal and very sClerotized spermatheCae, similar in length ( Fig. 24 ).

Immature description. Third-instar larvae ( Figs 27–37 ). Body length: 7.96–11.20 mm; width at the sixth abdominal segment: 1.59–1.79 mm (n=5). General shape and color ( Figs 27, 32 ). Elongate, CylindriCal, distinCtly tapered at thoraCiC segments, and trunCate at Caudal end. All segments similar in length. Color ranging from Creamy white to Creamy yellow ( Fig. 27 ), anterior spiraCle yellow-Colored ( Fig. 29 ), posterior spiraCle brownish ( Figs 33, 34 ); perianal pad yellow and portly ( Fig. 32 ). Spinules present only on ventral surfaCe of abdominal segments, slightly extending to lateroventral surfaCe, in disContinuous rows of variable length. Cephalopharyngeal skeleton ( Figs 36, 37 ). Total length from tip of mandible to end of ventral Cornu 1.15–1.20 mm (not inCluding the extension of the ventral pharyngeal ridge) (n=4); strongly sClerotized (but with different degrees of sClerotization depending on the struCture), with general shape as in Figure 36 (ventral view) and Figure 37 (lateral view), Consisting of three main parts: the anterior, mandible; the median, hypopharyngeal sClerite (or intermediate sClerite); and the posterior, pharyngeal sClerite. Two mandibles, in lateral view, total length 0.18–0.21 mm, height 0.063–0.068 mm (n=3); entirely well-sClerotized, anterior portion siCkle-shaped, posterior margin ConCave; tooth on ventral margin absent. Dental sClerite very reduCed, square-shaped in dorsal view, not visible in lateral view. Hypopharyngeal sClerite lying between mandibles and pharyngeal sClerite, Consisting of two longitudinal sClerotized rods, 0.7 times as long as mandibles, wider at anterior half, ConneCted by a weakly sClerotized bridge medially; labial sClerite not visualized (possibly absent). Parasotomal bar, total length 2.12–2.20 mm (n=2) ( Fig. 37 ); inserted dorsomedially and parallel to intermediate sClerite, paired, very slender, and moderately sClerotized. Pharyngeal sClerite, distanCe between base of dorsal bridge and anterior margin of dorsal Cornu 1.45–1.47 mm (n=3) Consisting of pair of reClining, “U-shaped” sClerites – dorsal and ventral Cornua; dorsal Cornua anteriorly ConneCted by distinCtly sClerotized dorsal bridge, arChed anterodorsally; in lateral view, ventral margin straight, and dorsal margin ConCave, narrow at base, wider gradually; dorsal Cornu anterior, posterior and ventrally slightly more sClerotized than median region; membranous and Convex dorsally; ventral Cornua ConneCted by pharyngeal sClerite; pharyngeal sClerite with distinCt and relatively well sClerotized pharyngeal filter, Composed of 8 to 9 longitudinal Cibarial ridges, straight anterior and posteriorly, and sinuous medially, or Completely straight; distinCtly spaCed from one another on anterior third and then gradually Closer to one another, ConstriCted at apex of ventral Cornu, and distanCed again forming an extension beyond the apex of ventral Cornu, similar to a “tail” ( Figs 36, 37 ). Anterior spiracle ( Figs 29, 30 ). Inserted laterodorsally, at posterior margin of prothoraCiC segment; papillae projeCted from a Common base, similar in size, between 12 and 16 in number (n=3). Abdominal segments ( Figs 27 , 38–45 ). Eight segments Clearly visible, anterior margin of eaCh one with ventral transversal oval Creeping welts (CW I–VIII), Consisting of swollen ridges bearing several and variable number of rows of ConiCal and pointed spinules, different in size and arrangement ( Figs 38–45 ). CW I anteroposteriorly narrower than all other CWs, with five disContinuous and irregular rows of similar sized spinules ( Fig. 38 ). CW II–VIII with medial, almost Continuous (sometimes with few interrupted, speCially CW VIII) and regular row of similar sized spinules; and posteriorly to this row there is a gap without any spinules until the next row ( Figs 39–45 ); general distribution of spinules and rows as in Figures 37–43 . Anteriorly to perianal pad, one Complete and regular row and one disContinuous and irregular row of differently sized spinules ( Fig. 32 ). Perianal pad bifurCated ( Fig. 32 ). Variation in rows and spinules on CW: a small variation was observed in the number of rows in eaCh CW in different larvae (n=4), but this never exCeeds more than two rows; the shape and length of a row and the arrangement and size of its spinules Can vary slightly, however, the Central and regular rows on CW II–VIII are always present. Posterior spiracles ( Figs 31, 33–35 ). Inserted on a

semispheriCal yellowish protuberanCe, slightly sClerotized, dorsally to the midline of the posterior surfaCe of the larva ( Fig. 31 ); perimeter of spiraCular plate with irregular inner margin, extending between the spiraCular openings; eCdysial sCar positioned at dorsomedial margin of perimeter; two larger dorsal groups of spiraCular hairs, with simple or mostly bifurCated hairs ( Figs 31, 34, 35 ), and one lateroventral very reduCed group, with simple hairs ( Fig. 31 ); spiraCular openings Curved, almost generally parallel ( Figs 31, 33–35 ), lined by a sClerotized rimae and traversed by several narrow sClerotized trabeCulae. Puparium ( Figs 46–52 ). Total length: 5.7–6.1 mm; width at the third abdominal segment: 2.1–2.3 mm (n=5). Brown to brownish orange Coloration. Notable CutiCle shrinkage of thoraCiC segment. In lateral view, flattened ventrally, and gradually higher towards abdominal segments, and abruptly lower at segment VIII, forming an angle of approximately 45º with the anterior line of the segment ( Fig 46 ); in dorsal view, metathorax distinCtly wider than pro- and mesothorax ( Fig. 47 ). Anterior spiracles. Elongated in lateral view, dark brown, laterally inserted, prominent and protruding, Covering at least one third of the prothoraCiC surfaCe area. Abdominal segments. Creeping welts limited to ventral surfaCe, not extending to lateral margin ( Figs 46, 48 ), distribution of spinules and rows of CW similar to third larval instar, exCept for shorter spaCe between rows ( Figs 48–50, 52 ). Posterior spiraCles very similar to third larval instar, dark brown, with evident round aperture ( Fig. 51 ). Perianal pad brown-Colored, flattened, trapezoidal, with irregular posterior margin, and evident anal opening, with 1–2 irregular rows of spinules bordering the anterior margin ( Fig. 52 ).

Type material, adult. HOLOTYPE ♂ BRAZIL. “Brasil, PernambuCo, Igarassu\Estrada BR101 Norte\7º47’S 34º55’W 80 m \ 17–31.v.2013 \A.C.D. Maia Col.” “Planta hospedeira: Taccarum \ ulei (AraCeae) . InfloresCÊnCias\ imaturas” (DZUP). PARATYPES, DZUP. Same data as holotype: 3♀, 4♂ (pinned and dry), 2♀ (in alCohol).

Immatures examined, BRAZIL (DZUP). Igarassu, PernambuCo, Estrada BR101 Norte, 7º47’S 34º55’W 80 m, A.C.D. Maia Col., host plant: Taccarum ulei (AraCeae) , immature infloresCenCes: 9 third instar larvae (3 disseCted), 9 puparia (1 disseCted).

Distribution and ecology. Both adults and immatures of this speCies have only been ColleCted in the northern Coastal region of the state of PernambuCo, in Close assoCiation with the infloresCenCes of Taccarum ulei . As it is assumed that B. taccarivora is a host speCialist (Maia et al. 2013), its distribution range Could overlap with that of T. ulei , naturally found from Coastal AtlantiC Forest fragments in northeastern Brazil to the xeriC shrublands and thorn forests of the Continental Caatinga.

Etymology. The speCifiC epithet “ taccarivora ” alludes to the host plant speCifiCity of the florivorous larvae of this speCies; from Taccarum plus vora (vorare), to eat or devour.

Figures

FIGURES 1–7. Beebeomyia taccarivora sp. nov.: 1. HOLOTYPE MALE, HABITUS, LATERAL VIEW; 2–3. PARATYPE FEMALE: 2. HABITUS, LATERAL VIEW;3. WING;4. PARATYPE MALE PRESERVEDIN ALCOHOL,DORSAL VIEW;5–7. HOLOTYPE MALE, HEAD:5. FRONTAL VIEW;6. DORSAL VIEW;7. LATERAL VIEW.SCALES LINES:FIGS 1–4: 1.0 MM; FIGS 5–7: 0.5 MM.

FIGURES 8–16. Beebeomyia taccarivora sp. nov., PARATYPE MALE:8–11. DRAWINGS OF STERNITES:8. STERNITE 2; 9. STERNITE 3; 10. STERNITE 4; 11. STERNITE 5; 12–13. DRAWING OF EPANDRIUM (SETAE NOT REPRESENTED), LATERAL AND MEDIAL SURSTYLI, AND CERCI: 12. POSTERIOR VIEW; 13 LATERAL VIEW; 14. LATEROPOSTERIOR GENERAL VIEW: STERNITE 6, SYNTERGOSTERNITE 7 + 8, SPIRACLES 6 AND 7, EPANDRIUM, LATERAL AND MEDIAL SURSTYLI, CERCI, BASIPHALLUS AND DISTIPHALLUS (BLACK ARROW INDICATES THE VESTIGIAL SPIRACLE ON STERNITEVI); 15.DISTIPHALLUS,APEX INDETAIL;16.MEDIAL SURTYLUS INDETAIL.SCALESLINES:FIGS8–14: 0.2MM;FIGS 15,16:0.1 MM.

FIGURES17–26. Beebeomyia taccarivora sp. nov., PARATYPE FEMALE:17–22. DRAWINGS OF STERNITES:17. STERNITE 1; 18. STERNITE 2; 19. STERNITE 3; 20. STERNITE 4; 21. STERNITE 5; 22. STERNITE 6; 23. TERMINALIA, GENERAL VIEW: OVISCAPE, TAENIA, EVERSIBLE MEMBRANE,STERNITE 8, CERCI;24. SPERMATHECAE AND SPERMATHECAL DUCT;25. STERNITE 8 AND CERCI;26. CERCI IN DETAIL.SCALES LINES: 0.2 MM.

FIGURES 27–35. Beebeomyia taccarivora sp. nov., THIRD-INSTAR LARVA:27. GENERAL VIEW, LATERAL;28. ANTERIOR REGION,VENTRAL; 29, 30: ANTERIOR SPIRACLE (NOT CLAREAD AND CLEARED, RESPECTIVELLY); 31. DRAWING LINE OF POSTERIOR SPIRACLE; 32. POSTERIOR REGION, VENTRAL; 33 – 35: POSTERIOR SPIRACLES: 33. GENERAL VIEW; 34. IN DETAIL, NOT CLAREAD; 35. IN DETAIL, CLAREAD. ABBREVIATIONS: An: ANTENNA;ES: ECDYSIAL SCAR;MSO: MAXILLARY SENSE ORGAN; OA: ORAL AREA;P: PERITREME;SH: SPIRACULAR HAIRS;SO: SPIRACULAR OPENING.SCALES LINES:FIG.27: 2.0MM;FIGS 28, 29, 32–34:0.2 MM; FIG.30 = 0.1 MM.

FIGURES 36, 37. Beebeomyia taccarivora sp. nov., THIRD-INSTAR LARVA, CEPHALOPHARYNGEAL SKELETON: 36. VENTRAL VIEW; 37. LATERAL VIEW.SCALES LINES:0.1MM.

FIGURES 38–45. Beebeomyia taccarivora sp. nov., THIRD-INSTAR LARVA, ABDOMINAL SEGMENT, LOCOMOTORY STRUCTURES, VENTRAL VIEW: 38. CREEPING WELT I (CW I); 39. CREEPING WELT II (BLACK BAR INDICATES THE EVIDENT DISTANCE BETWEEN THE MEDIAL ROW AND NEXT ROW); 40. CREEPING WELT III; 41. CREEPING WELT IV; 42. CREEPING WELT V;43. CREEPING WELT VI; 44. CREEPING WELT VII; 45. CREEPING WELT VIII. ABBREVIATIONS:CW: CREEPING WELT.BLACK ARROWS INDICATE THE MEDIAL,ALMOST CONTINUOUS AND REGULAR ROW. SCALE LINES:0.1MM.

FIGURES 46–52. Beebeomyia taccarivora sp. nov., PUPARIUM:46. LATERAL VIEW;47. DORSAL VIEW;48. VENTRAL VIEW, PUPARIUM DISSECTED AND CLARIFIED,49–52: ABDOMINAL SEGMENTS: 49. CREEPING WELT I; 50. CREEPING WELT V; 51. SPIRACULAR PLATE, POSTERIOR SPIRACLE;52. PERIANAL PAD AND CREEPING WELT VIII. ABBREVIATIONS:CW:CREEPING WELT;T1, 2, 3:THORAXIC SEGMENT 1, 2, 3. SCALE LINES:FIGS 46–48: 1.0 MM;FIGS 49–52 =0.2MM.