Bromeloecia wolverinei Yau and Marshall, 2018

Yau, Tiffany & Marshall, Stephen A., 2018, A revision of the genus Bromeloecia Spuler (Diptera: Sphaeroceridae: Limosininae), Zootaxa 4445 (1), pp. 1-115 : 74-85

Bromeloecia wolverinei Yau and Marshall sp. n.

(Figs. 15.1–15.14)

Description: Body length 0.9–1.2 mm (males), 0.9–1.4 mm (females).

Head: Interfrontal setae in 5 pairs. Length of anterior fronto-orbital seta only slightly shorter than posterior seta. Flagellomere 1 with 1 long pit on basal outer edge. Genal height 0.6x eye height.

Wing: Angle between costa and R4+5 70°. Distance between radial fork and r-m 1.7x length of r-m; dm-cu subequal to length of r-m and distance between crossveins equal to length of dm-cu. M-pseudovein slightly bent towards costa, leveling off in distal third and ending 1.5x length of dm-cu from wing margin.

WIPs: Basal angle of distal spot wide, 100°.

Legs: Fore femur with a row of 5 strong posteroventral setae on distal two thirds. Ventral surface of mid tibia with a large apical seta.

Male abdomen and terminalia: Anteroventral arm of synsternite 6+7 arched, apex triangular; posteroventral lobe simple, sclerotized prior to anteroventral edge. Subanal plate concave, appearing golden due to microtrichia. Subepandrial sclerite reduced to simple π-shape. Cerci thimble-shaped. Surstylus with 3 highly sclerotized flattened setae and a posterior process with apical seta and a blunt spur.

Hypandrium rectangular with sclerotized anterior hypandrial apodeme on anterior half; hypandrial arms slender. Ejaculatory apodeme and bulb not observed. Distiphallus WS apically spatulate with strong wide stalk; LFS strong and wide; FDS reach dorsally to meet with the lateral cheeks of WS; SDS also bends dorsally and forms a bridge by fusing medially.

Female abdomen and terminalia: T6 with setae across width of tergite. Each half of T8 ovoid, with 1 large and 4 smaller setae. Epiproct glossy at anterolateral corners. Cercus diamond-shaped. Paired spermathecae with thimble-shaped pit apically and basally, single spermatheca with 1 apical and 2 basal pits, spermathecal ducts with mid-length swelling.

Type specimens: Holotype: ♂: ECUADOR: Galápagos: Santa Cruz 2 km N Bellavista, guava thicket, agricultural area, 360 m, FIT, 14 May–13 Jul 1985, S. & J. Peck ( QCAZ) .

Paratypes: ECUADOR: Galápagos: Floreana , agriculture zone, 350 m, sweeping, 22 Mar 1989, B. J. Sinclair (2♂, 2♀, DEBU) ; Isabella Sierra Negra , 1000 m, sweeping pampa, 3–11 Feb 1989, B. J. Sinclair (1♂, 6♀, DEBU) ; same as holotype (2♂, 2♀, QCAZ, DEBU); Santa Cruz, Los Gemelos , Scalesia forest, 600 m, dung trap, 31 Jan–4 Feb 1989, B. J. Sinclair (19♂, 26♀, QCAZ, DEBU) ; Seymour , arid zone, seal dung, yellow pans, 24 Jan 1989, B. J. Sinclair (2♂, 6♀, DEBU). PANAMA : Chiriquí: Lagunas, 5 km SW Hato del Volcan , 1360 m, dung trap, 22–26 May 1977, S. Peck (14♂, 13♀, DEBU) . PUERTO RICO: Jayuya: Toro Negro National Forest Rd 143, km 2.5, human dung trap, 3–4 Jul 1977, J. Micheli & R. E. Woodruff (2♂, DEBU) ; Luquillo: National Forest Lesser Antilles El Junque Rd , field of kail, dung trap, 4–5 Jul 1977, R. E. Wooddruff (1♂, 1♀, DEBU). THE FEDERATION OF SAINT KITTS AND NEVIS : Saint Kitts: Wing Field Mountain , old field/forest, 400 m, Malaise trap, 1–31 Oct 1985, L. D. Coote (1♂, 1♀, DEBU) . TRINIDAD AND TOBAGO : Saint George: Curepe , 10°38ʹ00ʺN, 061°24ʹ00ʺW, Malaise trap, 17 Dec 1978, F. D. Bennett (3♂, 2♀, DEBU) .

Other material examined: BELIZE: Cayo: Augustine, pine savannah, 1500ʹ, dung trap, 1–4 Aug 1972, S . & J. Peck (2♂, 1♀, DEBU); Beaver Dam , 39 mi W Belize City, swamp forest, dung trap , 6–10 Aug 1972, S. & J. Peck (1♀, DEBU); Belmopan , palm forest, carrion trap , 26–30 Jul 1972, S. Peck (1♀, DEBU); Belmopan , creekside, gallery forest, dung trap , 11–16 Aug 1972, S. & J. Peck & R. Anderson (6♂, 6♀, DEBU). COSTA RICA : Alajuela: Florencia, forest, dung trap, 28 Feb 1980, H . Howden (1♀, DEBU); Puntarenas: Osa Peninsula , Rincón, 2.5 km S, 08°42ʹ01ʺN, 083°30ʹ50ʺW, secondary forest, ~ 50 m, dung pans, 11 Aug 2001, M . Buck (1♂, DEBU); DOMINICAN REPUBLIC: Alcoa Road , 600 m, FIT carrion , 17–20 Jan 1989, S. A. Marshall & J. Swann (1♂, 1♀, DEBU); Baranona: La Cienega , FIT , 11–22 Jun 1989, S. A. Marshall & J. E. Swann (1♀, DEBU); 7 km NW Paraiso , rainforest reminant, 200 m, fungus on logs , 27 Nov 1991, S. & J. Peck (5♂, 7♀, DEBU); Pedernales: 28 km N Cabo Rojo , evergreen dry forest, 760 m, carrion trap , 29 Nov–3 Dec 1991, S. & J. Peck (3♂, 2♀, DEBU). ECUADOR: Galápagos: Floreana, Black Beach, agric. zone, 370 m, carrion trap , 22–25 Feb 1989, B. J. Sinclair (1♂, DEBU); Floreana, 3 km E Black Beach , up. arid zone, 120 m, carrion traps , 22–25 Mar 1989, B. J. Sinclair (5♂, 4♀, DEBU); Floreana, 8 km E Black Beach , littoral-arid, 10 m, FIT , 21–28 Mar 1989, Peck & Sinclair (1♀, DEBU); Isabella, 12 kmNW Villamil , jaboncillo forest, 125 m, carrion trap , 9–12 Mar 1989, B. J. Sinclair (1♀, DEBU); Isabella, Villamil 12 km NW, trans. forest, 150 m, FIT, 2–15 Mar 1989, Peck & Sinclair (2♀, DEBU); San Cristóbal, 4 km E Baquerizo , trans. z., 150 m, FIT , 12–23 Feb 1989, Peck & Sinclair (1♂, 2♀, DEBU); San Cristóbal, Casetta , 560 m, pampa dung trap , 16–19 Feb 1989, B. J. Sinclair (47♂, 82♀, DEBU); San Cristóbal, Progresso 1 km E, guava ravine, 370 m, Malaise , 15–23 Feb 1989, Peck & Sinclair (1♀, DEBU); Santa Cruz, Bellavista , agriculture zone, 160 m , 27 Feb 1989, B. J. Sinclair (1♀, DEBU) Santa Cruz, Bellavista , agriculture zone, field, 160 m, FIT , 1–9 Apr 1989, Peck & Sinclair (1♂, 1♀, DEBU); same as holotype (20♂, 17♀, DEBU); Santa Cruz, 2 km N Bellavista , guava thicket, agricultural area, 360 m, FIT , 13 Jul 1985, S. & J. Peck (1♂, 1♀, DEBU); ditto , 2 Jul 1985 (9♂, 10♀, DEBU); Santa Cruz, 25 km N Bellavista, guava thicket, agricultural area, 360 m, Malaise , FIT , 13 Jul 1985, S. & J. Peck (3♂, 4♀, DEBU); Santa Cruz, 3 km S Bellavista, trans. Z., 115 m, Malaise , 1–29 Feb 1989, Peck & Sinclair (1♀, DEBU); Santa Cruz, 4 km N Bellavista, Media Luna , Miconia Zone , 620 m, FIT , 14 May–13 Jul 1985, S. & J. Peck (29♂, 39♀, DEBU); Santa Cruz, 4 km N Bellavista, Media Luna , Miconia Zone , 620 m, carrion traps , 17–21 May 1985, S. & J. Peck (4♂, 1♀, DEBU); Santa Cruz, 4 km N Bellavista, Media Luna , Miconia Zone , 620 m, dung trap , 21–31 May 1985, S. & J. Peck (3♂, 1♀, DEBU); Santa Cruz , CDRS, arid zone, tortoise dung trap, 31 Jan–9 Feb 1989, B. J. Sinclair (3♂, 3♀, DEBU); Santa Cruz , CDRS, arid zone, 50 m, MV-lights, 3 Feb 1989, B. J. Sinclair (2♂, 2♀, DEBU); Santa Cruz, Los Gemelos , Scalesia forest, 600 m, dung trap , 31 Jan –- 4 Feb 1989, B. J. Sinclair (1♂, DEBU); Santa Cruz, Los Gemelos , forest, 600 m, dung trap , 31 Jan–4 Feb 1989, B. J. Sinclair (2♂, 3♀, DEBU); Santa Cruz , Los Gemelos, 31 km N Santa Rosa , Scalesia forest, 570 m, FIT & Malaise, 28 Jun 1985, J . Peck (1♂, 2♀, DEBU); Santa Cruz , Los Gemelos , 31 km N Santa Rosa, Scalesia forest, 570 m, FIT & Malaise , 23 Jun 1985, S. & J. Peck (5♂, 6♀, DEBU); ditto , 13 Jul 1985 (1♀, DEBU); Santa Cruz, Media Luna , Miconia zone , 500 m, FIT-trough , 1–29 Feb 1989, Peck & Sinclair (1♂, 1♀, DEBU); Santa Cruz, Media Luna , Miconia Zone , 550 m, FIT , 1–30 Mar 1989, Peck & Sinclair (1♂, DEBU); Santa Cruz, 1 km N Puntudo , Scalesia , 650 m, FIT , 1–8 Apr 1989, S. Peck (1♂, DEBU); Santiago, 3 km NE Aquacate Camp , moss elfin forest, 740 m, FIT , 5–9 Jun 1991, S. Peck (1♂, DEBU); Marchena Pta. Espejo , arid zone, Bursera grove , 11–24 Mar 1992, S. Peck (1♀, DEBU); Pichincha: 23 km E Alluriquin, Chiriboga , 4600ʹ, dung trap , 19–27 Jun 1975, S. Peck (1♀, DEBU); Palenque , 01°25ʹ56ʺS, 079°45ʹ14ʺW, day 3 trap , 24–25 Mar 1976, S. Peck (1♂, DEBU); Rio Palenque , J. Glasser trap , 26 Feb 1976, S. Peck (1♂, DEBU); 47 km S Santo Domingo, Rio Palenque Station , 250 m , 17–25 Feb 1979 (1♂, DEBU); 16 km SE Santo Domingo, Tinalandia , 680 m, dung trap , 21–22 Jun 1975, S. Peck (2♀, DEBU). JAMAICA: Saint Ann: Ocho Rios , fern gully, FIT , 19 Feb–1 Mar 1989, D. H. Lindmon (1♂, 1♀, DEBU); Saint Thomas: Corn Puss Gap, 4 mi N Bath , 2100ʹ, dung trap , 3–8 Aug 1974, S. Peck (4♂, 2♀, DEBU); Trelawny: Good Hope , 8 Aug 1966, H. F. Howden (1♀, DEBU). MEXICO : Chiapas: 5 mi S Bochil, oak, 5000ʹ, human dung, 18–24 Aug 1971, A. Newton (2♂, 1♀, DEBU); Palenque , rainforest, 100 m, carrion trap , 6–9 Jul 1983, S. & J. Peck (2♂, 4♀, DEBU); Palenque , 2° veg., 80 m, FIT , 2–23 Jul 1983, S. & J. Peck (2♂, DEBU); 4 mi S Palenque , rainforest, 700ʹ, human dung , 7–15 Aug 1971, A. Newton (1♂, 2♀, DEBU); Guerrero: 9 mi NE Iguala , deciduous, 4400ʹ, human dung , 29 Aug–4 Sep 1971, A. Newton (2♀, DEBU); Oaxaca: 9 mi E El Camaron , 4300ʹ, human dung , 23 Aug–6 Sep 1973, A. Newton (1♀, DEBU); Querétaro: 45 km NE Ixtapa , 10–12 Jul 1987, B. Gill (3♂, 6♀, DEBU); San Luis Potosí: 40 km W Xilitla , pine-oak forest, 1700 m, FIT , 12 Jun–6 Aug 1983, S. & J. Peck (1♀, DEBU); Tabasco: 46 mi SE Villahermoss , km 74, 2° veg. rainforest, 150ʹ, human dung , 8–15 Aug 1971, A. Newton (16♂, 23♀, DEBU); Veracruz: 8.2 mi W Ciudad Mendoza on Mex. 150D, oak woodland, 6200ʹ, carrion , 27 Jul–3 Aug 1973, A. Newton (1♂, DEBU). PANAMA : Chiriquí: 2 km W Cerro Punta, Baldwin forest , 1760 m, dung trap , 30 May–2 Jun 1977, S. Peck (1♂, DEBU); Chirrari, Hartmann’s Finca, dung trap, 30 May 1977, S. J. Peck (1♂, DEBU); Hartmann’s Finca, 15 km NW Hato de Volcan , 1500 m, dung trap , 20–25 May 1977, S. Peck (1♂, DEBU); Hartmann's Finca, 15 km NW Hato de Volcan , 1200 m, dung trap , 20–25 May 1977, S. Peck (1♀, DEBU); Lagunas, 5 km SW Hato del Volcan , 1360 m, dung trap , 22–26 May 1977, S. Peck (1♀, DEBU); Panamá: Canal Zone, Madden Forest , carrion trap , 10–13 Jun 1977, S. Peck (1♀, DEBU). TRINIDAD AND TOBAGO : Saint George: Arima Ward, NY Zoological Society Station, pig dung trap, 11 Jun 1977, R . Wooddruff (1♂, DEBU); Curepe , 10°38ʹ00ʺN, 061°24ʹ00ʺW, Malaise trap , 29 Nov 1978, F. D. Bennett (1♂, DEBU); ditto , Aug 1978 (1♀, DEBU); Simla House, 4 mi post on Blanchisseuse Rd. N Arima, 10°41ʹ00ʺN, 061°17ʹ00ʺW, 250 m, garden Malaise , 3–10 Jun 1987, M. Pickles (1♀, DEBU); U. Santa Cruz, Gasparillo , gr/for. edge, Malaise trap , 15 Nov 1987, R. Borneo (2♂, DEBU). SAINT VINCENT: St. Andrews: Botanical Garden Kingstown , Malaise trap , 1–3 Oct 1991, R. E. Wooddruff (1♂, DEBU). VENEZUELA:

Táchira: 57 km SE San Cristóbal, El Pinal , 1500ʹ, dung trap, 19–21 May 1974, S. Peck (1♀, DEBU).

Etymology: This species is named for the group of 3 heavily sclerotized, flattened setae on the ventral surface of the surstylus, which closely resembles the claws and fist of the comic book character "Wolverine".

Comments: B. wolverinei is found throughout Central America, several of the northern countries of South America, and the Caribbean. It is the only Bromeloecia species found on the Galápagos Islands, and according to the label data, occurs on seven of the largest islands. This wide distribution suggests that B. wolverinei is a highly vagile species with wide ecological tolerances. Although B. wolverinei and B. triunguia closely resemble each other externally, B. wolverinei can be easily identified by the presence of a posterior process and the very wellsclerotized flattened setae on the surstylus; the posterolateral corners of the subanal plate also lack clusters of setae and microtrichia. See also comments below the description of B. triunguia .

Bromeloecia bromeliarum (Knab and Malloch)

(Figs. 16.1–16.10, 17.1–17.10)

Limosina bromeliarum Knab and Malloch, 1912: 413

The type series of B. bromeliarum includes specimens with two different wing forms, with the first wing form (known only from males) matching the holotype. The holotype male is redescribed below; both sexes of the second wing form are described separately under the heading “Variation”.

Redescription: Body length 0.9 mm (males).

Head: Frons uniformly brown except slightly paler at base of interfrontal setae and darker between interfrontal setae and fronto-orbital plate. Postocellar setae large and parallel. Paravertical setae small and converging. Interfrontal setae in 4 pairs. Length of anterior fronto-orbital seta subequal to posterior seta. Flagellomere 1 with 2 closely contiguous oval pits on basal outer edge. Longest aristal branches as long as length of basal aristomere. Eyes ovoid; genal height 0.4x eye height; gena with 4–5 setulae at genal margin and anteroventral corner. Clypeus height obscured on available material.

Thorax: One large prescutellar dorsocentral seta, remaining 2 postsutural dorsocentrals slightly longer than acrostichals. Katepisternum with 1 dorsal seta.

Wing: Costa with 1 costagial seta; costal section between humeral and subcostal break with 2 pairs of setae and a dorsal setula in between; costa ends at 4/5 of wing length. R2+3 straight. Angle between costa and R4+5 70°. Distance between radial fork and r-m 2x length of r-m; dm-cu subequal to length of r-m and distance between crossveins 0.5x length of dm-cu. M-pseudovein slightly bent towards costa, leveling off in distal half and ending close to wing margin. CuA1 extends slightly beyond dm-cu and continues as pseudovein towards wing margin. Distal half of A1-pseudovein straight or only slightly curved.

WIPs: Cell r2+3 generally of uniform colour with a thin compact band of colour pattern at anterior edge. Basal angle of distal spot wide, 100°; ripple pattern skewed, larger gradient length towards wing tip; posterior edge elbowed at basal one third. Distal colour incision wide.

Legs: Base of fore femur obscured on available material (basal golden hairs therefore not visible), posterodorsal half of femur with 1 strong seta; ventrally with 1 large ventrobasal seta and posteroventrally with a row of 8 strong setae along femur, basal 4 more closely spaced than distal 4. Male fore tibia anteroventrally with a comb of 4 small stout setae on distal quarter. Mid tibia with a proximal and middle anterodorsal seta and a pair of distal dorsal (anterodorsal and dorsal) setae; ventral surface of mid tibia with small middle anteroventral seta and a large apical seta. Hind tarsus simple, claw symmetrical.

Male abdomen and terminalia: Sternite 4 unmodified. S5 posteromedially covered with microtrichia. Anteroventral arm of synsternite 6+7 apically bent, dilated, covered with microtrichia and distally desclerotized to fuse with membrane; medioventral lobe long and thin, weakly sclerotized, curves anterodorsally, apically desclerotized and blending into membrane of genital pouch; posteroventral lobe long and curled inwards to abdominal cavity, with anteroventral edge sclerotized. Anteroventral process on right side of epandrium short and truncated, articulating with right lateral edge of hypandrium. Subanal plate present, with a pair of strong setae. Subepandrial sclerite with medial plate and thin lateral arms. Cerci wide at base and not contiguous. Surstylus simple, ventral surface with scattered setulae.

Hypandrium Y-shaped, lateral extension plate asymmetrical; outer margin of posteromedial hypandrial processes articulating with hypandrial arms. Phallapodeme symmetrical, distal end fan-like. Ejaculatory apodeme and bulb not observed. Epiphallus strong, chin-like. Distiphallus WS long, tube-like, not bifurcate; LFS strong, possibly fused with FDS distally; FDS distally shielded by spinulose membrane, dorsally with a lightly striated wing of membrane; SDS spatulate.

Type specimens: Holotype: ♂: “C ordoba Mex, 21 Mch ’08, from arboreal bromeliad, Fred K. Knab collector. Type No. 14913 U.S.N.M. Limosina bromeliarum Type K. & M., Limosina bromeliae Knab & Mall type” (USNM).

Paratypes: “ Cordoba Mex, 17 Mch ’08, from arboreal bromeliad, Fred K. Knab collector. Type No. 14913 U.S. N.M., Limosina bromeliarum Paratype ” (1♂, USNM).

Variation:

The type series of B. bromeliarum includes 1 male and 4 females that differ from the holotype in having a broad and staggered subcostal break. These specimens probably represent a second species, but the material at hand is inadequate to confirm this. They are therefore described below and tentatively treated as a variant of B. bromeliarum , which is otherwise known only from two males.

Body length 0.9 mm (male), 0.9–1.1 mm (females).

Head: Length of anterior fronto-orbital seta 0.8x posterior seta. Flagellomere 1 with 2 closely contiguous circular pits on basal outer edge.

Wing: Subcostal break deep; CS2 basally dipped significantly lower than CS1. R1 distally shortened; R2+3 generally straight except slightly curved basally. Distance between radial fork and r-m 2x length of r-m; dm-cu slightly longer than r-m and distance between crossveins 0.5x length of dm-cu.

Legs: Fore femur anterodorsally with 2 short golden hairs at base. Male fore femur not observed (specimen without fore legs); female fore femur ventrally with 1 large ventrobasal seta and posteroventrally with a row of 8 strong setae equally spaced along femur.

Male abdomen and terminalia: Distiphallus whip-like sclerite (WS) long, tube-like, not bifurcate, apically tipped with contracted spinulose membrane.

Female abdomen and terminalia: S7 simple with scattered setae. Each half of T8 huge, larger than S7, shielding most of epiproct, posteriorly with a neat row of 4 setae. S8 rectangular. Epiproct large, with medial stripe of microtrichia, anteromedially extended with well-sclerotized edge, base of posterior point with a pair of setae on each side, laterally followed by a V-shaped groove. Cercus quadrate, apical seta long. Each spermatheca with 1 large apical thimble-shaped pit and 10 small basal pits.

Specimens of this second wing type, all paratypes of B. bromeliarum : “ Cordoba Mex, 17 Mch ’08, from arboreal bromeliad, Fred K. Knab collector. Type No. 14913 U.S. N.M., Limosina bromeliarum Paratype ” (1♂, 4♀, USNM).

Comments: B. bromeliarum can be easily distinguished from other species by the long aristal branches (as long as the basal aristomere) and the posterior edge of the distal spot of WIP that is elbowed at basal one third. This species is only known from a few specimens collected on epiphytic bromeliads on mango trees in Mexico. As noted and described above, the type series of B. bromeliarum includes specimens with two different wing forms, one known only from males and the other from both sexes. Comparing the males of the two variants, there are only 2 distinctive morphological differences: the modification of the subcostal break and the presence and absence of the spinulose membrane at the tip of the distiphallus whip-like sclerite (the entire remaining male terminalia of the two variants are identical to one another). Since the thin morphological evidence and distributional data are inadequate to support the idea of a second species, and the number of specimens and condition of the series is not suitable for molecular work, the two variants are treated as one species, B. bromeliarum .