Some Scuttle Flies (Diptera: Phoridae) of the Galápagos Islands

R. Henry L. Disney & Bradley J. Sinclair

Twelve species of Phoridae are reported from the Galápagos Islands. At least nine of these were evidently introduced by humans (Chonocephalus dominicanus Borgmeier, C. heymonsi Stobbe, Dohrniphora cornuta (Bigot), Gymnoptera simplex (Brues), Megaselia scalaris (Loew), M. seticauda (Malloch), Puliciphora borinquenensis Wheeler, P. occidentalis (Melander & Brues), P. sobria Borgmeier). Megaselia edenensis Disney, sp. n., M. peckorum Disney, sp. n. and Puliciphora species A (not named until its female has been procured) are described. The majority of the specimens of M. scalaris are darker than the typical form and many are smaller than usual, especially those from upland localities, but intermediates also occur. A key to Galápagos species is provided. The use of more varied collecting methods is likely to increase this list in the future. R. Henry L. Disney *, Cambridge University Museum of Zoology, Downing Street, Cambridge CB2 3EJ, England. [email protected] Bradley J. Sinclair, Entomology, Ontario Plant Laboratories, Canadian Food Inspection Agency, 960 Carling Ave., K.W. Neatby Bldg, C.E.F., Ottawa, ON, K1A 0C6, Canada. [email protected]

Introduction species, all classified as introduced invasive species The first scuttle fly (Diptera: Phoridae) recorded from (Causton et al. 2006). the Galápagos Archipelago (Megaselia scalaris (Loew, We report here primarily on the collections of scuttle 1866)) was collected during the Williams Expedi- flies made by S. & J. Peck and BJS and identified by tion (1923) (Johnson 1924) and a second species RHLD. (Dohrniphora cornuta (Bigot, 1857)) was collected during the Norwegian Zoological Expedition (1925) (Curran 1932). Additional material of the latter spe- Material and methods cies was collected on the Templeton Crocker Expedi- The methods of collection included direct searching tion of 1932 (Van Duzee 1933; Curran 1934). In of litter, carrion baited traps (CT), dung traps (DT), 1985, Stewart Peck (Carleton University, Ottawa) Malaise traps (MT), and yellow pan traps (ypans). started the first of five collecting expeditions to the The collectors and locality are abbreviated as fol- Galápagos, with the aim of generating an invertebrate lows: Charles Darwin Research Station – CDRS; inventory of the island fauna. There is little doubt S.B. Peck – SBP; S. & J. Peck – SJP; B.J. Sinclair that these surveys have had a tremendous impact on – BJS. The following abbreviations are used in the the knowledge of the terrestrial invertebrates, pro- descriptions: supra-antennal seta – SAs; subcutane- ducing numerous publications in many and several ous pit sensilla – SPS. Terms primarily follow those major monographs (Peck 2001, 2006). BJS partici- of Disney (1994). pated in the second Peck expedition of 1989. Vast Most specimens were preserved in alcohol and many numbers of specimens were collected on these two were mounted by RHLD on slides in so-called surveys, which resulted in the listing of four phorid Berlese Fluid (Disney 2001). A few pinned

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1 A from the rear of the dorsal shield-like epan- c drium embracing the penis complex (which may be withdrawn (Fig. 31) or extruded b (Fig. 21). The smaller ventral shield, the p hypandrium, typically bears projecting lobes s E from its rear margin. The hypopygium may a be asymmetric, so that the hypandrium is displaced to one side ...... males – 2 g l H – With or without wings. Tip of abdomen simpler (Figs 7 and 22). The cerci clearly separated ...... females – 13 2. Mid tibia with a pair of bristles, one dorsal 2 and one anterodorsal, in basal half. A single, conspicuously feathered bristle on axillary ridge ...... 3 – Without this combination ...... 4 3. Palpus single-segmented. Vein 3 unforked. Mesopleuron bare. Hypopygium as Figure 28. Abdominal tergite 6 entirely brown. Hind femur brown and base as Figure 27. Hind tibia without a dorsal hair palisade . . Figs 1–2. Chonocephalus males. – 1, hypopygium, ...... Gymnoptera simplex diagrammatic ventral view of a generalised species. – Palpus two-segmented. Vein 3 is forked. A = anal tube (with median hypoproct and paired cerci); E = epandrium; H = hypandrium; X = penis complex Mesopleuron with hairs in upper part. Hy- (simplified). a, p = anterior and posterior processes (prob- popygium as Figure 21. At least the anterior ably = subepandrial sclerites) of epandrium; l = lobe(s) half of abdominal T6 largely yellow. Hind fe- of hypandrium; g = gonopod; b = bridge (probably = mur yellow apart from being narrowly brown sternite 10); c = left surstylus (clasper); s = spine (modi- along dorsal edge and at tip; with a patch of fied bristle) of epandrium; 2, C. heymonsi, right wing. scattered hairs in a posterodorsal position at base and spinules as Figure 20. Hind tibia with a dorsal hair palisade ...... Dohrniphora cornuta specimens were remounted on slides after treat- 4. Vein 3 unforked (Figs 2, 47). Hind tibia with ment with Barber’s Fluid. Voucher material has been spaced hairs but without a dorsal, longitu- deposited in the Canadian National Collection of dinal, hair palisade ...... 5 Insects, Ottawa (CNC) and the Cambridge Univer- – Vein 3 forked (Fig. 37). Hind tibia with a sity Museum of Zoology (CUMZ). dorsal hair palisade ...... 10 5. Base of vein 4 missing and vein 3 ending in small clear vesicle (Fig. 2). Anal tube clearly Key to the species of Phoridae recorded shorter than midline length of dorsal face of from the Galápagos Islands epandrium (Figs 1, 5–6) ...... 6 The identifications should be checked against all the – Wing as Figure 47. Anal tube at least longer figures provided. If not agreeing in all the details a than midline length of dorsal face of epan- specimen should be checked out in the literature drium (Figs 48, 53, 56) or if shorter the cerci cited below for the genus in question. If not run- are clearly longer than wide (Fig. 57) . . . . . 7 ning out to a genus a specimen can be run down 6. Hypopygium as Figures 3–6, with the right in the most recent key to world genera (Disney gonopod narrowest at tip...... 1994)...... Chonocephalus dominicanus – Hypopygium as Figures 9–10, with the right 1. Wings always present. The abdomen ends in gonopod broadest at tip ...... the complex hypopygium, comprising the ...... Chonocephalus heymonsi epandrium, hypandrium, penis complex and 7. Anal tube relatively long (Figs 48, 56) . . . . 8 anal tube. The cerci are closely approximated – Anal tube relatively short (Fig. 53 and 57) . . . 9 and lie along the dorsolateral regions of the 8. Epandrium higher than long (Fig. 48) . . . proctiger to form anal tube, which emerges ...... Puliciphora borinquenensis

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– Epandrium longer than high (Fig. 56) . . . – With more numerous hairs at rear of abdom- ...... Puliciphora sobria inal tergites (Fig. 11). Spermatheca larger (its 9. Hypopygium as Figure 53, with the dorsal diameter being clearly greater than that of face of the epandrium being shortened . . . the stirrup-like loop of sternite 8 – Fig. 13) ...... Puliciphora occidentalis ...... Chonocephalus heymonsi – Hypopygium as Figure 57, with the dorsal 18. Mid tibia with a pair of bristles, one dorsal face of the epandrium not shortened . . . . and one anterodorsal, in basal half. A single, ...... Puliciphora species A conspicuously feathered, bristle on axillary 10. Mesopleuron bare (e.g. Fig. 41) ...... 11 ridge ...... 19 – Posterior part of mesopleuron with at least a – Without this combination ...... 20 dozen hairs. (Hypopygium as Fig. 30. Basal 19. Hind tibia without a dorsal hair palisade. half of hind femur with a ventral row of ro- Palpus single-segmented. Vein 3 unforked. bust hairs – Fig. 31) . . . . . Megaselia edenensis Mesopleuron bare. Proboscis as Figure 29 . . 11. Anal tube long (Figs 38 and 46). Noto- ...... Gymnoptera simplex pleuron with only two bristles. Haltere knob – Hind tibia with a dorsal hair palisade. Palpus largely pale yellow ...... 12 two-segmented. Vein 3 is forked. Mesopleu- – Anal tube short (Fig. 34–36). Notopleuron ron with hairs in upper part. Proboscis elon- with three bristles. Haltere knob brown . . . gated (Fig. 24). (Abdominal venter obviously ...... Megaselia peckorum hairy – Fig. 22) ...... Dohrniphora cornuta 12. With a single strong bristle on left side of 20. Mesopleuron bare (e.g. Fig. 41) ...... 21 epandrium (Fig. 38) . . . . . Megaselia scalaris – Posterior part of mesopleuron with at least a – With several strong bristles on left side of dozen hairs ...... Megaselia edenensis epandrium (Fig. 46) . . . . . Megaselia seticauda 21. Abdominal tergite 6 clearly narrower than 13. Wings and halteres absent ...... 14 T5 (e.g. Fig. 32) ...... 22 – Wings and halteres present ...... 18 – T6 clearly broader (and shorter) than T5 14. Ocelli present. Abdominal tergite 5 (T5) (Fig. 39) ...... Megaselia scalaris divided into an anterior semicircular flap 22. T5-T7 as Figure 32. With three bristles on (hinged along its front margin) and a larger notopleuron. Haltere knob brown . . . . . posterior portion (Figs 49, 52 and 54) (Note: ...... Megaselia peckorum frequently T1 is lost or fused to T2) . . . . 15 – T6 and T7 relatively long and narrow. With – Ocelli absent and T5 undivided (Figs 7 and only two bristles on notopleuron. Haltere 11) ...... 17 knob pale yellow ...... Megaselia seticauda 15. The anterior flap of T5 relatively broad (Figs 49 and 55). Wing represented by a bristle, which at most is situated on a small protuberance ...... 16 Taxonomy – The flap of T5 relatively small (Figs 52 and In the following list of species, only those synonyms 54). The alar bristle is situated on a small pertinent to the Galápagos fauna are listed. Full wing rudiment (Fig. 51) ...... synonyms for these species can be found in various ...... Puliciphora occidentalis revisions by the senior author. Note: The female of Puliciphora species A is either unknown or its male has not been pre- Chonocephalus Wandolleck, 1898 viously described. 16. The anterior flap of T5 a little shorter than Type species. Chonocephalus dorsalis Wandolleck, 1898 by midline length of the rest of the tergite monotypy. (Fig. 55) ...... Puliciphora sobria – The flap of T5 is longer than the midline The marked sexual dimorphism, along with the length of the rest of the tergite (Fig. 49) . . . naming of species based on the description of one sex ...... Puliciphora borinquenensis only, has caused much confusion. Species recogni- 17. With fewer hairs at rear margins (arising tion is now based on the male hypopygia in the first from pale spots embraced by the dark bands) instance. The then known species were reviewed by of abdominal tergites (Fig. 7). Spermatheca Disney (2002), including a key to the species of the smaller (subequal in diameter to the stirrup- Nearctic and Palaearctic Regions. Most of the latter like loop of sternite 8 – Fig. 8) ...... are tramp species transported around the world by ...... Chonocephalus dominicanus humans.

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3 5

Figs 5–6. Chonocephalus dominicanus male, hypopygium. – 5, left face; 6, right face tilted a little dorsally. Scale bar = 0.1 mm. Figs 3–4. Chonocephalus dominicanus male, details of hypopygium from below. – 3, right gonopod; 4, left side of hypopygium and epandrial processes of right side. Scale bar = 0.1 mm. evidently been transported across the world by humans.

Chonocephalus dominicanus Borgmeier Chonocephalus heymonsi Stobbe Figs 3–8 Figs 2, 9–18 Chonocephalus dominicanus Borgmeier, 1967: 206. Type Chonocephalus heymonsi Stobbe, 1913: 131 (female only). locality: Dominica. Type locality: Surinam.

Material examined. Santa Cruz: 1?, 2 km N Bellavista, Material examined. Isabela: 3?, Sto Thomas, 350 m, 360 m, guava thicket, agricultural area, MT, 2.vii.1985, CT, 6–9.vii.1985, SJP. Santa Cruz: 4?, 1/, Acad- SJP; 2?, 13.vii.1985. 1?, Los Gemelos, 3 kmN Santa emy Bay, CDRS, 30 m, arid zone thorn scrub, MT, Rosa, 570 m, Scalesia forest, MT, 13.vi.–15.vii.1985, SJP; 10.v.-3.vi.1985, SJP; 7?, 1/, 10.v.-14.vii.1985, SJP; 13?, 2?, 13.vii.1985. 2/, 14.vii.1985, SJP; 21?, 1/, 10.v.-14.vii.1985, SJP; 17?, 62/, Academy Bay, CDRS, 8 kmN Grieta w/water, Natural history. This species has been reared from Cueva de Iguana, bait traps, 10.v.1985, SJP; 14?, 1/, 4 kmN the rotting spadices of Carludovica palmata Ruiz & Bellavista, Media Luna, 620 m, 2.vii.1985, SJP; 2?, Pav. (Cyclanthacaeae) and Heliconia mariae Hook.f. Miconia zone, MT, 14.v.-13.vii.1985, SJP; 10?, 2 kmN (Heliconiaceae) (Disney 1980); it has also been Bellavista 360 m, guava thicket, agricultural area, MT, reared from bread fruit (Artocarpus altilis (Parkinson) 2.vii.1985, SJP; 16?, 3/, 13.vii.1985, SJP; 6?, Bel- lavista, field, 160 m, agricultural zone, 1–9.vi.1989, SBP Fosberg, Moraceae) (Disney 1994). & BJS; 9?, 8/, 1 kmN Bellavista, agricultural zone, Remarks. This is a Neotropical species that has 150 m, fruit-baited trap, 31.iii.-8.iv.1989, BJS. Downloaded from Brill.com09/25/2021 12:33:17AM via free access Disney & Sinclair: Phoridae of the Galápagos Islands 119

7 8 11 12 13

Figs 11–13. Chonocephalus heymonsi female. – 11, whole fly from above; 12, tergite 8; 13, sternite 8, furca and spermatheca. Scale bars = 0.1 mm.

14 Figs 7–8. Chonocephalus dominicanus female. – 7, whole fly from above; 8, sternite 8, furca and spermatheca (S = spermatheca, F = furca). Scale bars = 0.1 mm. 15 9

Figs 14–15. Chonocephalus heymonsi larvae. – 14, first instar; 15, third instar. (After Borgmeier 1935.)

16 17

Figs 16–18. Chonocephalus heymonsi pupae. – 16, eclo- Figs 9–10. Chonocephalus heymonsi male, hypopygium. sion plate of puparium; 17, anterior cap of puparium; – 9, from below; 10, left face. Scale bars = 0.1 mm. 18, left face of pupa. (After Borgmeier 1935.)

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Natural history. The larvae (Figs 14–15) and pupar- 19 ium (Figs 16–18) were described by Borgmeier (1935). The respiratory horns of the latter are unu- sual in this genus in being two branched. It has been reared from bread fruit (Artocarpus altilis (Parkinson) Fosberg, Moraceae). A female caught on the edible paddy straw mushroom Volvariella (Plutaceae) was probably ovipositing on an over ripe sporophore, as was the case with the females recorded on rotting Termitomyces (Amanitaceae) on a termite mound (Disney 1994). Remarks. This primarily warm-climate species has been transported across the world by humans (Dis- ney 2002), but is seemingly most abundant in Africa 20 south of the Sahara (Disney 2005b).

Dohrniphora Dahl, 1898 Type species. Dohrniphora dohrni Dahl, 1898, by monotypy.

More than one hundred species are known, with most only identifiable in the male sex in our present state of knowledge. The literature for the identification of the Neotropical species is summarised else- where (Disney 1994), supplemented by Kung & Brown (2005, 2006). Figs 19–20. Dohrniphora cornuta male. – 19, hind coxal lobe; 20, posterior face of base of hind femur. Scale bars = 0.1 mm. Dohrniphora cornuta (Bigot) Figs 19–26 Phora cornuta Bigot, 1857: 348. Aquacate, 600 m, FIT, humid forest, 4–9.vi.1991, SBP. Phora venusta Coquillett, 1895: 107; Curran 1932: 353; Curran 1934: 152. Natural history. The larvae feed on a wide range of Dohrniphora crockeri Van Duzee, 1933: 70. Dohrniphora willowsi Van Duzee, 1933: 70. decaying organic materials ranging from rotten potatoes, onions, beans, rice bran, chick peas, dead insects, dead snails, moribund turtle eggs, sour milk, Material examined. Floreana: 6?, 9/, Black Beach, lit- human faeces, general garbage, dead mice and other toral, CT, 21–26.iii.1989, BJS; 9?, 12/, 5 kmE Black small mammals, etc. (Disney 1994). They are also Beach, 250 m, CT, 22–25.iii.1989, BJS; 5?, 25/, occasional facultative predators. For example lar- 5.5 kmE Black Beach, 350 m, Scalesia forest, CT, 22–25. vae feeding on the sewage film of micro-organisms iii.1989, BJS; 7/, 3 kmE Black Beach, 120 m, upper arid zone, CT, 22–25.iii.1989, BJS; 3?, 11/, 8 kmE Black in trickling filter sewage beds, when confronted by Beach, 370 m, agricultural zone, CT, 22–25.iii.1989, competition for this resource, may prey on the eggs, BJS; 2?, 2/, 3 kmE Black Beach, 100 m, arid zone, DT, larvae and pupae of Psychodidae (Kloter et al. 1977). 24–28.iii.1989, BJS. Isabela: 7/, Sto Thomas, 350 m, In Thailand adults were reported to be the dominant CT, 6–9.vii.1985, SJP; 8?, 19/, 300–350 m, agricul- species of insect visiting the flowers of Aristolochia tural zone, CT, 9–12.iii.1989, BJS; 5?, 2/, Sierra Negra, baenzigeri B. Hansen & L. Phuphathanaphong and 1000 m, pampa, CT, 9–12.iii.1989, BJS. San Cristóbal: 1?, 1 kmE El Junco, 550 m, Miconia stream valley, CT, 18–21. to be a significant carrier of its pollen (Bänziger & iii.1989, BJS. Santa Cruz: 3/, Academy Bay, CDRS, 1 m, Disney 2006). supra-littoral zone, CT in shrubs, 10–13.v.1985, SJP; 3/, Remarks. This primarily warm-climate species CDRS, sweeping insect carrion, 29.i.1989, BJS; 3?, 4/, has been transported across the world by humans Cueva Bellavista, 160 m, cheese baited traps, iii-iv.1989, (Disney 1994). Records indicate that D. cornuta SBP; 47?, 480/, 21 larvae, Media Luna, 600 m, deep soil is found from the littoral to pampa zones on the / traps, ii.1989, SBP & BJS; 1 , Sierra Crocker, hill-top, Galápagos. 8.ii.1989, sweeping, 860 m, BJS. Santiago: 3?, 1 kmE

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T9 T10

S10 24

Figs 23–24. Dohrniphora cornuta female. – 23, left face of rear of abdominal segment 6 and of segment 7; PC 24, dorsal face of distiproboscis. Scale bars = 0.1 mm.

H 25 26

Fig. 21. Dohrniphora cornuta male, left face of hypopygium (T9 = tergite 9, C = cercus, H = hypandrium, P = proctiger, PC = penis complex, T10 + S10 = fused tergite 10 and sternite 10 to form a ring). Scale bar = 0.1 mm.

Fig. 22. Dohrniphora cornuta female, left face of termi- Figs 25–26. Dohrniphora cornuta. – 25, third instar nal segments of abdomen from rear of segment 5 on- larva from above; 26, respiratory horn of pupa. Scale wards. Scale bar = 0.1 mm. bars = 0.1 mm.

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Fig. 29. Gymnoptera simplex female, proboscis from above but focussed down onto hypopharynx.

Gymnoptera simplex (Brues) Figs 27–29 Conicera simplex Brues, 1905: 553 (female only).

Material examined. Santa Cruz: 2/, CDRS, arid zone, at insect carrion, 29.i.1989, BJS.

Natural history. This species has been previously reared from dead molluscs, dead beetles and a dead rat.

Figs 27–28. Gymnoptera simplex male. – 27, posterior Megaselia Rondani, 1856 face of base of hind femur; 28, left face of hypopygium. Type species. Megaselia crassineura Rondani, 1856 (a Scale bars = 0.1 mm. junior synonym of M. costalis (von Roser, 1840)) by monotypy.

Gymnoptera Lioy, 1864 This giant genus includes more than 1400 described species and several times this number awaiting Type species. Phora vitripennis Meigen, 1830, by description. A now dated guide to the literature is monotypy. given by Disney (1994), as well as keys to Neotropi- cal species being by Borgmeier (1962). The three known species are keyed by Disney (2003b), where two are Palaearctic and the third, Megaselia edenensis Disney sp. n. which is recorded below, is a tropical tramp species. Figs 30–31 The latter species has been recorded from all other Biogeographic Regions except the Nearctic, but Material examined. Holotype?, Santa Cruz: Pto Ayora, o o under four different names, three of which have been El Eden, 00 44’43.9’’S, 090 18’70.3”W, Malaise trap, synonymized since (loc. cit.). T.Vacio, 30 October 2006, P. Lincango & E. Lomas (INV. ZU 025) (CNC). Paratypes. 5?, 2/, same data as holotype except 2? and 1/ (CUMZ).

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30 tibia extends about 0.6 of length. Hairs below basal half of hind femur more robust (Fig. 31) and longer than those of anteroventral row of outer half. Hind tibia with some dozen differentiated posterodorsal hairs, the last not being differentiated from the rest. Spinules of apical combs simple. Wings 0.9–1.1 mm long. Costal index 0.40–0.43. Costal ratios 3.5–4.6 : 1.0–1.2 : 1. Costal cilia (of section 3) 0.05–0.06 mm long. Hair at base of vein 3 about as long as costal cilium at level of humeral cross vein. Two axillary bristles, outer slightly longer than costal cilia of section 3. Sc not reaching R1. Thick veins yellowish 31 grey, thin veins 4–6 grey and 7 very pale. Membrane almost colourless. Haltere brown. Female. Head similar to male but labrum slightly darker and at least 1.2as wide as postpedicel. Tho- rax as in male. Abdominal tergites brown and well developed. T2-T6 being progressively slightly narrower that preceding one. T6 about 0.27 mm wide and 0.13 mm long, with small median emargination Figs 30–31. Megaselia edenensis male. – 30, left face of on front margin (about 0.07 mm wide) and slightly hypopygium; 31, posterior face of hind femur. Scale concave hind margin. T7 subrectangular and about bars= 0.1 mm. 1.2as long as wide and with unpigmented median stripe extending for much of its length. Venter brown, with hairs below segments 3–6. Sternite 7 reduced to Etymology. Named after the type locality El Eden. a narrow band of pigment linking bases of four hairs on posterior margin. Posterolateral lobes at rear of Male. Frons brown, slightly broader than long, sternum 8 short, pale and with 3 long hairs near hind with 94–120 hairs and dense but fine microsetae. margin and 3–4 shorter hairs further forward. Cer- SAs robust but lower pair slightly shorter. Bristle cus pale and only about twice as long as wide. Two arrangement very similar to Figure 33. Gena with rectal papillae present. Furca not evident. Dufour’s 4–6 bristles and jowl with 2–3 longer ones. Postpedi- crop mechanism rounded behind and 0.6–0.17 mm cel subglobose, brown but not dark, without SPS long and about 0.07 mm wide. Legs similar to male vesicles. Palpus pale straw yellow, with 3 long and but basitarsus and segment 2 of front legs not so 1–3 short bristels and 5–9 hairs. Labrum coloured as thickened and hairs below basal half of hind femur palpus and about 0.7–0.8as wide as postpedicel. not obviously robust and subequal in length to those Labella also pale and with only several short spinules of anteroventral row of outer half. Wing and haltere below. Thorax brown, being paler on sides. Meso- as in male, except length 1.1–1.2 mm. Costal index pleuron with 14–24 hairs. Three notopleural bristles 0.41–0.42. Costal ratios 3.8–4.6 : 1.1–1.4 : 1. Costal and no cleft in front of these. Scutellum with ante- cilia 0.06–0.07 mm long. rior pair of fine hairs subequal to those in middle Affinities. In the keys of Borgmeier (1962) this spe- of scutum and posterior pair of bristles. Abdominal cies runs out on page 304 in Abteilung V to couplet tergites brown, with short fine hairs but those at rear 2, lead 2, to M. paupercula Borgmeier. However the of T6 longer than others. Venter brown, with hairs male is immediately distinguished by the differenti- below segments 3–6. Hypopygium brown, with ated bristles on the epandrium. In some specimens paler brown anal tube, and as in Fig. 30. Both lobes 1–3 hairs at the rear margin of the mesopleuron of hypandrium vestigial. Two rectal papillae present. are slightly differentiated from the rest, but still Legs with brown hind femur and mid coxa; other <0.06 mm long. These might then be run down in femora, mid and hind tibiae partly pale brown and Abteilung III to couplet 4, on page 303. However, partly pale straw yellow; remaining leg segments pale the wing colour fits lead 2 but the structure of the straw yellow. Fore tarsus with posterodorsal hair pali- epandrium fits lead 1. Furthermore in both the spe- sade on segments 1–5 and segment 5 slightly longer cies of this couplet the single differentiated bristle on than 4; basitarsus and segment 2 slightly thickened, the mesopleuron is >0.08 mm long. the former being 0.8–0.9as wide as tibia and only 2.8–2.9as long as wide. Dorsal hair palisade of mid

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Megaselia peckorum Disney sp. n. 33 Figs 32–36 Type material. Holotype?, Española: Bahia Man- zanilla, littoral Cryptocarpus and Prosopis, Malaise trap, 5–10 June 1985, S. & J. Peck (CNC). Para- types. Española: 1/, Prosopsis grove behind beach, CT, 5–10.vi.1985, SJP; 1/, sand beach, CT, 7–10. vi.1985, SJP (CNC and CUMZ). Santa Cruz: 1/, Academy Bay, CDRS, 1 m, supra-littoral zone, CT in shrubs, 10–13.v.1985, SJP (CNC); 1?, 1/, CDRS, littoral zone, ypans, 19.i.-9.ii.1989, BJS 32 34 (CNC); 1?, Pto. Ayora, Pelican Bay, 0.74272oS, 90.331211oW, pitfall 14c Muyuyo, 17.iii.2006, A. Velastegui (CUMZ).

Etymology. Named after the collectors S. & J. Peck.

Male. Frons brown, with 112–142 hairs and dense but very fine microsetae, and with bristles arranged as in Figure 33. SAs robust and about equal, lower pair being slightly less robust. Gena with 2 bristles and jowl with two 2–3 longer ones. Postpedicel subglobose, brown, and densely packed with SPS vesicles most of which are smaller than diameters of sockets of lower SAs. Palpus two-segmented, with Figs 32–34. Megaselia peckorum small basal segment brown and second segment . – 32, female, abdomi- inal tergites 5–7; 33, female, frons with bristles repre- straw yellow lightly tinged brown, with 6–7 bristles sented by their basal sockets only, 34, male, left face of (most apical being about as long as lower SAs) and hypopygium. Scale bars 0.1 mm. 4–9 hairs. Labrum coloured as in palpus and about 0.75as wide as postpedicel. Labellum pale, narrow and with only some short spinules below but with some strong, pale teeth on their inner edges. Thorax 35 36 brown, being slightly paler on sides. Mesopleuron bare. Three notopleural bristles and no cleft in front of these. Scutellum with anterior pair of hairs subequal to those at rear of scutum) and posterior pair of bristles. Abdominal tergites brown with hairs at rear of T6 slightly longer than rest (Fig. 34). Venter brown, with hairs below segments 3–6. Hypopygium pale brown, with almost colourless anal tube and left lobe of hypandrium, and as Figs 34–36. Right Figs 35–36. Megaselia peckorum male hypopygium with phallus complex withdrawn. – 35, right face; lobe of hypandrium shortened. Legs light brown, 36, left face. Scale bars 0.1 mm. the front pair being palest. Fore tarsus with posterodorsal hair palisade on segments 1–3 and segment 5 1.2–1.3as long as 4. Dorsal hair palisade of mid tibia extends almost 0.7 its length. Hairs below Thick veins yellowish grey, thin veins 4–6 grey and basal half of hind femur 1.1–1.2 as long as those 7 very pale. Membrane very lightly tinged grey. Hal- of anteroventral row of outer half. Hind tibia with tere brown, but not dark. 8 differentiated posterodorsal hairs. Spinules of api- Female. Frons brown, with 154–162 hairs and dense cal combs simple. Wings 0.9 mm long. Costal index but very fine microsetae, and with bristles arranged 0.43–0.44. Costal ratios 3.6–3.7 : 1.8–1.9 : 1. Cos- as in Figure 33. SAs robust and about equal, the tal cilia (of section 3) 0.04–0.05 mm long. No hair lower pair being slightly less robust. Gena with two at base of vein 3. Two axillary bristles, outer being bristles and jowl also with two. Postpedicel subglo- longer than costal cilia. Sc pale and not reaching R1. bose, brown, with at least a dozen SPS vesicles most

Downloaded from Brill.com09/25/2021 12:33:17AM via free access Disney & Sinclair: Phoridae of the Galápagos Islands 125 of which smaller than diameters of sockets of lower agricultural zone, CT, 22–25.iii.1989, BJS; 4/, 3 kmE SAs. Palpus two-segmented, with small basal segment Black Beach,100 m, arid zone, DT, 24–28.iii.1989, BJS. brown and second segment pale straw yellow to very Isabela: 8/, Sto Thomas, 350 m, CT, 6–9.vii.1985, SJP; 38?, 32/, pampa zone, 700 m, ypans, 3–12.iii.1989, lightly tinged brown, with 6–7 bristles (most api- BJS; 1?, 21/, 300–350 m, agricultural zone, CT, 9–12. cal being about as long as lower SAs) and 4–9 hairs. iii.1989, BJS; 122?, 346/, Sierra Negra, 1000 m, pampa, Labrum light brown but clearly darker than palpus CT, 9–12.iii.1989, BJS; 1/ (dark form) V. Wolf, 6 km and about as wide as postpedicel. Labellum pale, nar- E. P. Bravo, 750 m, humid forest, pit trap, SBP (96–193). row, with only some short spinules below but with San Cristóbal: 2/, 2 kmNE of Baquerizo, arid zone, CT, some strong, pale teeth on their inner edges. Thorax 15–18.ii.1989, BJS; 13/, pampa zone, 560–700 m, CT, / as in male. Abdominal tergites brown. T5-T7 as Fig- 18–21.ii.1989, BJS; 19 , 1 kmE Progresso, 320 m, agricultural zone, CT, 18–21.iii.1989, BJS; 61/, 1 kmE ure 32. Venter greyish brown, and with hairs below El Junco, 550 m, Miconia stream valley, CT, 18–21. segments 3–6. Sternite 7 an isosceles triangle, as long iii.1989, BJS. Santa Cruz: 5?, 2/, Academy Bay, CDRS, as T7 and rear margin as wide, bearing four longer 30 m, arid zone thorn scrub, MT, 10.v.-3.vi.1985, SJP; hairs on rear margin and 7–12 smaller ones further 1?, 10.v.-14.vii.1985, SJP; 1?, 3/, 10.v.-14.vii.1985, forward. Posterolateral lobes at rear of sternum 8 very SJP; 1/, 1 m, sublittoral zone, CT on litter, 16.v.1985, short, but wide, and with three longer hairs at rear SJP; 7?, 14/, Academy Bay, CDRS, 8 kmN Grieta w/ water, Cueva de Iguana, bait traps, 10.v.1985, SJP; 233/, margin and four smaller ones further forward. Cer- Academy Bay, CDRS, 1 m, supra-littoral zone, CT in cus pale brown and about 4as long as broad. Two shrubs, 10–13.v.1985, SJP; 3/, 4 kmN Bellavista, Media rectal papillae present. Furca not apparent. Dufour’s Luna, 620 m, Miconia zone, CT, 17–21.v.1985, SJP; 1?, crop mechanism rounded behind and measuring 14.v.–13.vii.1985, SJP; 1?, Media Luna, 600 m, deep soil 0.16 mm long and 0.11 mm greatest breadth. Legs traps, ii.1989, SBP & BJS. 1/ (typical form), CDRS, at similar to male but with at least a dozen differen- window of lab, arid zone, SBP (96–141). Santa Fé: 1/, tiated posterodorsal hairs on hind tibia. Wings and littoral, beach, MT, 4–6.iv.1989, SBP & BJS. haltere as in male, except 1.0–1.4 mm long. Costal Natural history. The egg as illustrated in Figure 45. index 0.45–0.47. Costal ratios 2.9–3.5 : 1.4- 1.7 : 1. The larvae (Figs 42–44) exploit a wider range of Costal cilia 0.05–0.07 mm long. decaying organic materials than any other known Natural history. All records of this new species are insect. The resources exploited range from decaying from the littoral zone, collected in Malaise, carrion plants to human corpses. They also occur as facul- and yellow pan traps. tative parasites of vertebrates, especially humans, in Affinities. In the keys of Borgmeier (1962) this spe- cases of wound, intestinal, urogenital, etc., myiasis. cies runs to in Abteilung VII or Abteilung VIII They invade laboratory cultures of insects and other depending on whether the costal index is long (0.44 organisms and pose a hygiene risk when exploiting or longer) or short (0.43 or less). In Abteilung VII human foods. Adults are important visitors to flow- it keys out on page 310 at couplet 55, to M. label- ers of Aristolochia species (Aristolochiaceae) and have lata Borgmeier, but its costal index is at least 0.5 and also been reported visiting Barbarea vulgaris R. Br. the hypopygium is different. The male hypopygium, (Cruciferae). The widely scattered literature on the especially its very short anal tube, eliminates some biology of this species has recently been reviewed subsequently described species that run to the same (Disney 2008). couplets. In the supplementary keys (listed by Disney Remarks. This primarily warm-climate species has 1994) it fails to key out. In Abteilung VIII it runs to been transported across the world by humans (Dis- couplet 3, lead 2, to M. morula Borgmeier. The lat- ney 2008). However, the Neotropical and Nearctic ter, however, has paler wings, longer hairs below the M. imitatrix Borgmeier, 1969 is apparently the most basal half of the hind femur and a brown anal tube. closely related species, whose larvae are aquatic in container habitats (Hanson & Disney 2008). Megaselia scalaris (Loew) The species is somewhat variable in size, and related to this the anterior scutellar bristles are often shorter Figs 37–45 than allowed for in most keys. In Borgmeier’s (1962) Phora scalaris Loew, 1866: 53. primary keys to Neotropical species such specimens will run to couplet 58, lead 2, on page 310, to Material examined. Española: 1?, 1/, Bahia Manza- M. pruinosifrons Borgmeier, 1962. The male of the nilla, littoral Cryptocarpus and Prosopis, MT, 5–10.vi.1985, latter has a short anal tube and the anterior scutel- SJP; 4/, Prosopis grove behind beach, CT, 5–10.vi.1985, lars are minute hairs, instead of bristles. The female / SJP. Floreana: 1 , 5.5 kmE Black Beach, 350 m, Scalesia abdominal tergite 6 is not broader than T5, as is the forest, CT, 22–25.iii.1989, BJS; 3/, Black Beach, littoral, CT, 21–26.iii.1989, BJS; 5/, 8 kmE Black Beach, 370 m, case in M. scalaris (and M. imitatrix). Many, if not most, specimens in samples from the

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41 N

Fig. 37. Megaselia scalaris male. (Drawn by J. W. Rod- ford.) Figs 40–41. Megaselia scalaris female. – 40, furca (internalised sternite 9); 41, left face of part of thorax 38 to show mesopleuron and adjacent features (M = mid- mesopleural ridge, P = post-propleural ridge, N = posterior notopleural bristle, S = anterior spiracle). Scale bars 0.1 mm. 42

43 44

Figs 42–45. Megaselia scalaris pre-adult stages. – 42, larval stage III, left face of mandibles and ce- Figs 38–39. Megaselia scalaris. – 38, left face of male phalopharyngeal skeleton; 43, left mandible of larva; hypopygium; 39, female abdominal tergites 5–7. Scale 44, median tooth of larva; 45, dorsal view of egg. Scale bars 0.1 mm. bars 0.1 mm.

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Puliciphora Dahl, 1897 Type species. Puliciphora lucifera Dahl, 1897, by monotypy.

Females of the Neotropical species are keyed by Disney (2003) supplemented by Disney & Kistner (2003) and Disney (2005a, c.)

Puliciphora borinquenensis Wheeler Figs 47–50 Figs 46. Megaselia seticauda left face of male hypopygium. Scale bar 0.1 mm. Puliciphora borinquenensis Wheeler 1906: 269.

Material examined. Floreana: 3?, 5/, 5.5 kmE Black Beach, 350 m, Scalesia forest, CT, 22–25.iii.1989, BJS. Galápagos islands are nearer the small end of the Isabela: 5?, 12/, Sto Thomas, 350 m, CT, 6–9.vii.1985, / range of variation and have somewhat shorter than SJP; 1 , pampa zone, 700 m, ypans, 3–12.iii.1989, BJS; 1/, 3 kmE Black Beach, 100 m, arid zone, DT, usual anterior scutellar bristles. In addition they tend 24–28.iii.1989, BJS; 1?, Sierra Negra, 1000 m, pampa, to be darker. Thus the postpedicels and labrum are CT, 9–12.iii.1989, BJS. Santa Cruz: 15?, 97/, Acad- more yellowish brown than yellow, the legs may be emy Bay, CDRS, 1 m, supra-littoral zone, CT in shrubs, brownish yellow and the normally yellow parts of the 10–13.v.1985, SJP; 59/, 1 m, sublittoral zone, CT on abdominal tergites are light brown or even as dark litter, 16.v.1985, SJP; 1/, Academy Bay, CDRS, 30 m, as the rest of the tergites. The abdominal tergite 7 arid zone, thorn scrub, MT, 10.v.-14.vii.1985, SJP; 1/, 14.vii.1985, SJP; 31/, 2 kmN Bellavista, 360 m, avo- of the dark-form females tends to have smaller ante- cado grove, leaf and fruit litter, 14.v.1985, SJP; 2?, 4/, rolateral processes. Typical forms tend to predomi- 360 m, guava thicket, agricultural area, MT, 2.vii.1985, nate in coastal samples but the darker forms tend to SJP; 3/, 14.v.-13.vii.1985, SJP; 4?, MT, 13.vii.1985, dominate samples from montane localities. In both SJP; 6/, 4 kmN Bellavista, Media Luna, 600 m, sifted situations a few intermediates may be encountered. Sphagnum, 21.v.1985, SJP; 115/, Sphagnum bog, CT, It is probable that this species reached the Galápagos 27.v.1985, SJP; 2?, Media Luna, 620 m, 2.vii.1985, SJP; ? / ? a long time ago and has tended to diverge from the 66 , 206 , Miconia zone, CT, 17–21.v.1985, SJP; 72 , 60/, 620 m, Miconia zone, MT, 14.v.-13.vii.1985, SJP; original mainland form. However, repeated subse- 1?, 620 m, Miconia zone, MT, 19.vi.1985, SJP; 1?, 1/, quent introductions by modern shipping may have Media Luna, 600 m, deep soil traps, ii.1989, SBP & BJS; prevented this divergence proceeding to a separate 1/, Cerro Banderas, 4 kmNE Santa Rosa, 620 m, sink subspecies or even a distinct sibling species. The pro- hole Miconia litter; 1/, CDRS, ex Rattus rattus, 15.v.1992, curement of molecular signatures for samples from R.L. Palma & E.M. Inca; 12?, 2/, Los Gemelos, 3 kmN different populations could be rewarding. Santa Rosa, 570 m, Scalesia forest, MT, 13.vi.-15.vii.1985, SJP; 1/, in Scalesia litter, 13.vi.1985; 2?, Scalesia forest, MT, 28.vi.1985; 2?, 13.vii.1985; 1?, Los Gemelos, 600 Megaselia seticauda (Malloch) m, Scalesia zone, DT, 31.i.-4.ii.1989, BJS. Fig. 46 Aphiochaeta seticauda Malloch, 1914: 32. Natural history. The egg is illustrated as Figure 50. The larvae exploit a wide range of decaying organic Material examined. San Cristóbal: 6?, El Progreso, materials, from ripe fruits to vertebrate carrion S 00.9083, W 089.56154, 20.i.2005, J. Loayza. (Disney 1994). The flightless female is transported by the male and deposited on suitable pabulum for Natural history. Not known apart from cryptic data oviposition (Miller 1979). The majority of collec- on specimen labels quoted by Borgmeier (1962) thus tions for this species are from the humid zones. “rd corn” and “in green corn car”. Remarks. This primarily warm-climate species has Remarks. This species has been reported from main- been transported around the world by humans land Ecuador, Costa Rica, Mexico, USA (Texas), (Disney 2003a). and the West Indies (Dominica) (Borgmeier 1966, 1969a).

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47 51

Figs 47–48. Puliciphora borinquenensis male. – 47, right wing; 48, left face of hypopygium. Scale bars 0.1 mm.

Figs 51–53. Puliciphora occidentalis. – 51, female, right wing rudiment; 52, female abdominal tergite 5; 53, male, left face of hypopygium. Scale bars 0.1 mm.

Figs 49–50. Puliciphora borinquenensis. – 49, female, Fig. 54. Puliciphora occidentalis female, abdominal abdominal tergites 4–6; 50, egg, with anterior end to tergites 4–6. Scale bar 0.1 mm. right. Scale bars 0.1 mm.

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(Melander & Brues) Puliciphora occidentalis 55 Figs 51–54 Stethopathus occidentalis Melander & Brues, 1903: 17. 56

Material examined. Santa Cruz: 1/, Academy Bay, CDRS, 30 m, arid zone, thorn scrub, MT, 14.vii.1985, SJP (CNC).

Remarks. This is a Nearctic species. In the keys to Neotropical females (Disney 2003a) it runs to couplet 27, where the species of neither lead fits. With the subsequently proposed synonymy of Pulicimyia Borgmeier, 1960 with Puliciphora (Disney 2005c) two of its species will run to the same couplet. Puliciphora pauxilla (Borgmeier, 1960) has the small anterior flap of abdominal tergite 5 clearly broader than its midline length. Puliciphora triangularis Figs 55–56. Puliciphora sobria. – 55, female, abdominal (Borgmeier, 1960) has this flap distinctly more tri- tergites 4–6; 56, male, left face of hypopygium. Scale bares 0.1 mm. angular, and like P. occidentalis it has a small wing rudiment but it is larger than that of the latter species (although that of P. occidentalis is a little variable in size). In the keys to the Nearctic species (Disney 2005c) the specimen runs out as P. occidentalis. The posterior part of abdominal tergite 5 is sometimes incomplete (cf. Figs 52 and 54).

Puliciphora sobria Borgmeier Figs 55–56 Puliciphora sobria Borgmeier, 1960: 317.

Material examined. Floreana: 1/, 5.5 kmE Black Beach, 350 m, Scalesia forest, CT, 22–25.iii.1989, BJS. San Cristóbal: 7?, 9/, pampa zone, 560–700 m, CT, 18–21. ii.1989, BJS; 3?, 28/, 1 kmE Progresso, 320 m, agric. zone, CT, 18–21.iii.1989, BJS; 15?, 42/, 1 kmE El Junco, Fig. 57. Puliciphora species A, male, left face of hypopy- 550 m, Miconia stream valley, CT, 18–21.iii.1989, BJS. gium. Scale bar 0.1 mm.

Natural history. Numbers of males were reported in light traps in caves in Trinidad (Disney vertex. Postpedicels brown, subglobose, with about 1995). 10 SPS vesicles, largest being about same diameter Remarks. This species has been previously recorded as basal sockets of lower SAs. Palpus light brown from Costa Rica, Dominica, Mexico, Trinidad and with 6 bristles and as many hairs. Labrum slightly the USA (Alabama). paler than palpus but labella pale. Thorax brown but paler on sides. Three bristles on notopleuron. Puliciphora species A Scutellum with anterior pair of hairs and posterior pair of bristles. Abdomen with brown tergites and Fig. 57 brownish grey venter. The latter with hairs absent Material examined. Santa Cruz: 1?, Media Luna, Mico- from segments 1–5. Hypopygium straw yellow with nia zone, 14.xi.2001, ex sweeping, BJS (CNC). some brown on epandrium and anal tube and as Figure 57. Legs with light brown hind femur and Male. Frons brown with four supra-antennal bris- tibia, partly brown partly straw yellow front and mid tles, a pair of antial bristles, each situated midway femora and tibiae and otherwise straw yellow. Front between upper SAs and anterolaterals that are slightly basitarsus with apical ventral extension below basal higher on frons; and 6 bristles (including ocellars) on third of segment 2. Hairs below basal half of hind

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Table 1. Summary of numbers of Galápagos phorids caught on the different islands

Species Islands

Española Floreana Isabela San Cristóbal Santa Cruz Santiago Santa Fé

C. dominicanus males - - - - 6 females - - - - - C. heymonsi males - - 3 - 119 females - - - - 79 D. cornuta males - 25 13 1 50 3 females - 66 28 - 491 G. simplex males - females 2 M. edenensis males 6 females 2 M. peckorum males 1 - - - 2 females 2 - - - 2 M. scalaris males 1 - 161 - 16 females 5 13 408 95 257 1 M. seticauda males - - - 6 - females - - - - - P. borinquenensis males - 3 6 - 180 females - 5 14 - 588 P. occidentalis males - - - - - females - - - - 1 P. sobria males - - - 25 - females - 1 - 79 - Puliciphora sp. A males 1 females -

No. Species (12) 2 4 4 4 10 1 1 No. Flies (2767) 9 113 633 206 1802 3 1

femur not differentiated from those on anterior face. associated with its female. Of the described males it Spinules of transverse combs of hind basitarsus all most closely resembles P. parvula Borgmeier (1969b) simple. Wing length 1.30–1.35 mm. Costal index in that it has an unusually short anal tube. However, 0.52. Costal ratios 0.6–0.7 : 1. Costal cilia 0.04 mm Species A has a slightly shorter costal index (0.52 long. The costa is narrower in the basal two-thirds as opposed to 0.56), a slightly longer costal section of section 1. Thick veins light brown, thin veins 4–6 1 (its costal ratios being 0.6–0.7 : 1 as opposed to grey and 7 paler. Membrane only very lightly tinged 0.5–0.6 : 1) and furthermore it differs in that the grey but colourless between vein 1 and the unforked costa is narrower in the basal two-thirds of section 1 vein 3. The anal lobe slightly more pronounced than and the anal lobe is more pronounced than that of in many species of genus. P. parvula (cf. Fig. 136 in Borgmeier, 1969b). Remarks. This species cannot be named until

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Discussion on Isabela. More intensive collecting on the other A total of 2767 flies were collected representing islands listed in Table 1 and on other islands would twelve species (Table 1), but 65% were from Santa be likely to yield further species. Cruz. Furthermore, the methods of collecting were limited. The larval habits of scuttle flies are more diverse than those of any other family of insects Acknowledgements (Disney 1994). However, the prevalence of tramp RHLD’s studies of Phoridae are currently supported species on remote islands has tended to give rise to an by a grant from the Balfour-Browne Trust Fund atypical preponderance of species breeding in decay- (University of Cambridge). We thank the authorities ing organic materials. The above list of recorded spe- of the Charles Darwin Research Station and those cies reflects this even more than was to be expected, of the Galápagos National Park for allowing field probably because the majority of the collections were work and for logistical support. BJS wishes to thank made with carrion baited traps or direct from carrion Stewart Peck (Carleton University) who funded his baits. participation in the 1989 expedition and has contin- It is of interest to note that half the species recorded ued to support his studies. Bernard Landry (Museum (the genera Chonocephalus and Puliciphora) have d’Histoire Naturelle, Genève) and Charlotte Causton flightless females, which supports the supposition (Charles Darwin Research Station, Galápagos) have that they were introduced to these islands by humans. encouraged BJS to continue research on Galápagos Their spread within each island, however, was prob- Diptera. ably mainly undertaken unaided by humans. In support of this supposition are the observations made by Wilton (1961), who examined the colonisation References of 68 dustbins by flies in Honolulu. Puliciphora Bänziger, H. & R.H.L. Disney, 2006. Scuttle flies (Dip- borinquenensis successfully colonised 50.0% of these tera: Phoridae) imprisoned by Aristolochia baenzigeri containers, and was thus no less successful at colonis- (Aristolochiaceae) in Thailand. – Mitteilungen der ing these bins than Megaselia scalaris, whose females Schweizerischen Entomologischen Gesellschaft 79: are fully winged. 29–61. Another consequence of the majority of the collec- Bigot, J.M.F., 1857. Dipteros. – In: R. de la Sagra, Historia fisica, politica y natural de la Isla de Cuba. Segunde tions being made with carrion baited traps, or direct parte. Historia natural. Vol. 7: 328–349, Paris. from carrion or dung baits, is a preponderance of Borgmeier, T., 1935. Sobre o cyclo evolutivo de Chono- females. Thus some sex ratios (the number of females cephalus Wandolleck e uma nova espécie de Melaloncha caught for every male) were as follows: 5.7 : 1 (n = 868) Brues, endoparasita de abelhas. – Archivos do Instituto for M. scalaris at carrion and dung traps, compared de Biologia Vegetal, Rio de Janeiro 2: 255–265. with 0.8 : 1 (n = 80) in yellow pan traps plus Malaise Borgmeier, T., 1960. Gefluegelte und ungefluegelte Phori- traps. Likewise the ratios for P. borinquenensis were den aus der neotropischen Region, nebst Beschreibung 5.4 : 1 (n = 578) in carrion and dung traps, com- von sieben neuen Gattungen (Diptera, Phoridae). pared with 0.7 : 1 (n = 160) in Malaise traps. – Studia Entomologica, Petropolis 3: 257–374. It is axiomatic, but frequently ignored, that every Borgmeier, T., 1962. Versuch einer Uebersicht ueber die collecting method is selective with regard to the neotropischen Megaselia-Arten, sowie neue oder list of species obtained at a particular locality (e.g., wenig bekannte Phoriden verschiedener Gattungen Disney et al. 1982, Disney 2004). In the above report, (Dipt. Phoridae). – Studia Entomologica, Petropolis while a range of collecting methods was employed, 5: 289–488. Borgmeier, T., 1966. Revision of the North American it is evident that typically only a few methods were phorid flies. Part III. The species of the genus Megas- employed at each locality sampled. The above list is elia, subgenus Megaselia (Diptera, Phoridae). – Studia therefore unlikely to represent a complete list of the Entomologica, Petropolis 8 [1965]: 1–160. species of scuttle fly to be found on these islands. Borgmeier, T., 1967. New or little-known phorid flies, Future collecting using different methods more with the description of two new genera (Diptera, extensively would be likely to turn up other species: Phoridae). – Anais da Academie Brasileira de Ciencias for example the more widespread use of light traps 39: 195–210. and pitfall traps; or direct collecting, such as those Borgmeier, T., 1969a. Bredin-Archbold-Smithsonian visiting flowers, honeydew or ripe fruits; or rearing Biological Survey of Dominica: The Phoridae of from fungi or invertebrate hosts of parasitoid spe- Dominica (Diptera). – Smithsonian Contributions cies; or catching specimens observed attacking ants, Zoology 23: 1–69. beetle pupae, etc. Furthermore the collecting was Borgmeier, T. 1969b. New or little-known Phorid flies, concentrated on Santa Cruz, and to a lesser extent mainly of the Neotropical Region (Diptera, Phoridae). – Studia Entomologica, Petropolis 12: 33–132. 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Brues, C.T., 1905. Phoridae from the Indo-Australian Disney, R.H.L. & D.H. Kistner, 2003. New species and region. – Annales Musei Nationalis Hungarici 3: new host records of scuttle flies (Diptera: Phoridae) 541–555. associated with army ants and termites (Hymenoptera: Causton, C.E., S.B. Peck, B.J. Sinclair, L. Roque- Formicidae; Isoptera: Termitidae). – Sociobiology 42: Albelo, C.J. Hodgson & B. Landry, 2006. Alien 503–518. insects: threats and implications for conservation of Hanson, H. & R.H.L. Disney, 2008. An example of an Galápagos Islands. – Annals of the Entomological aquatic scuttle fly (Diptera: Phoridae) – a need for care Society of America 99: 121–143. in identification. – Antenna 32: (In press). Coquillett, D.W., 1895. A synopsis of the dipterous genus Johnson, C.W., 1924. Diptera of the Williams Galapagos Phora. – The Canadian Entomologist 27: 103–107. Expedition. – Zoologica 5: 85–92. Curran, C.H., 1932. [The Norwegian Zoological Expe- Kloter, K.O., L.R. Penner, & W.J. Widmer, 1977. Interac- dition to the Galapagos Islands 1925, conducted by tions between the larvae of Psychoda alternata (Dipt., Alf Wollebaek. IV.] Diptera (excl. of Tipulidae and Psychodidae) and Dohrniphora cornuta (Dipt., Phori- Culicidae). – Nyt Magazin Naturvidenskaberne 71: dae) in a trickling filter sewage bed, with descriptions 347–366. of the immature stages of the latter. – Annals of the Curran, C.H., 1934. The Templeton Crocker Expedition Entomological Society of America 70: 775–781. of the California Academy of Sciences, 1932. no.13. Kung G. & B.V. Brown, 2005. New species of Dohrni- Diptera. – Proceedings of the California Academy of phora related to D. longirostrata (Enderlein) (Diptera: Sciences, Series 4, 21: 147–172. Phoridae). – Annals of the Entomological Society of Disney, R.H.L., 1980. A new species of Chonocephalus America 98: 55–62. (Diptera: Phoridae) from Panama. – Entomologica Kung, G. & B.V. Brown, 2006. Review of the Caribbean Scandinavica 11: 363–365. species of Dohrniphora Dahl (Diptera: Phoridae). Disney, R.H.L., 1994. Scuttle Flies: The Phoridae. Chap- – Journal of Natural History 40: 1931–1945. man and Hall, London. xii + 467 pp. Loew, H., 1866. Diptera Americae septentrionalis indi- Disney, R.H.L., 1995. Cave Phoridae (Diptera) of Trini- gena. Centuria septima. – Berliner Entomologische dad. – Giornale Italiano di Entomologia 6 [1993]: Zeitschrift 1866: 1–54. 417–436. Malloch, J.R., 1914. Costa Rican Diptera. Paper 1. A par- Disney, R.H.L., 2001. The preservation of small Diptera. tial report on the Borboridae, Phoridae and Agromyzi- – Entomologist’s Monthly Magazine 137: 155–159. dae. – Transactions of the American Entomological Disney, R.H.L., 2002. Revisionary notes on Chonocephalus Society 41: 8–36. Wandolleck (Diptera: Phoridae) with keys to species of Melander, A.L. & C.T. Brues, 1903. Guest and parasites the Holarctic Region. – Zootaxa 60: 1–36. of the burrowing bee Halictus. – Biological Bulletin of Disney, R.H.L., 2003a. Seven new species of New World the Marine Laboratory Woods Hole 5: 1–27. Puliciphora Dahl (Diptera: Phoridae) with a new key Miller, P.L., 1979. Intraspecific phorid phoresy. – Ento- to the Neotropical species. – Zootaxa 162: 1–22. mologist’s Monthly Magazine 114 [1978]: 211–214. Disney, R. H. L., 2003b. Revision of Gymnoptera Lioy Peck, S.B., 2001. Smaller orders of insects of the Galápagos (Diptera, Phoridae). – Fragmenta Faunistica 46: Islands, Ecuador: Evolution, Ecology and Diversity. 203–208. – NRC Research Press, Ottawa. xii + 278 pp. Disney, R.H.L., 2004. Species preferences for white versus Peck, S.B., 2006. The beetles of the Galápagos Islands, yellow water traps for scuttle flies (Dipt., Phoridae). Ecuador: Evolution, Ecology and Diversity (Insecta: – Entomologist’s Monthly Magazine 140: 31–35. Coleoptera). – NRC Research Press, Ottawa. xiv + Disney, R.H.L., 2005a. A new species of Puliciphora Dahl 313 pp. (Dipt., Phoridae) from Ecuador. – Entomologist’s Stobbe, R., 1913. Chonocephalus heymonsi, eine neue Monthly Magazine 141: 65–67. Art aus Südamerika. – Deutsche Entomologische Disney, R.H.L., 2005b. Revision of Afrotropical Chono- Zeitschrift 1913: 191–192. cephalus Wandolleck (Diptera: Phoridae). – Journal of Van Duzee, M.C., 1933. The Templeton Crocker Expedi- Natural History 39: 393–430. tion of the California Academy of Sciences, 1932. no. Disney, R.H.L., 2005c. Rudimentary halteres of some 7. Dolichopodidae and Phoridae. – Proceedings of the flightless female Phoridae (Diptera) reassessed, and California Academy of Sciences, Series 4, 21: 65–74. some taxonomic implications. – Zootaxa 1042: Wheeler, W.M., 1906. A new wingless fly (Puliciphora 39–54. borinquenensis) from Porto Rico. – Bulletin of the Disney, R.H.L., 2008. Natural history of the scuttle fly, American Museum of Natural History 22: 267–271. Megaselia scalaris. – Annual Review of Entomology Wilton, D.P., 1961. Refuse containers as a source of flies in 53:39–60. Honolulu and nearby communities. – Proceedings of Disney, R.H.L., Y.Z. Erzinclioglu, D. J. de C. Henshaw, the Hawaiian Entomological Society 17: 477–481. D. Howse, D.M. Unwin, P. Withers & A. Woods, 1982. Collecting methods and the adequacy of at- tempted fauna surveys, with reference to the Diptera. Received: 17 October 2007 – Field Studies 5(4): 607–621. Accepted: 28 February 2008

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