Lucilia Robineau-Desvoidy

Whitworth, Terry, 2014, A revision of the Neotropical species of Lucilia Robineau-Desvoidy (Diptera: Calliphoridae), Zootaxa 3810 (1), pp. 1-76 : 8-22

Genus Lucilia Robineau-Desvoidy

Lucilia Robineau-Desvoidy, 1830: 452 . Type species: Musca caesar Linnaeus, 1758 , by subsequent designation of Macquart (1834: 162).

Phaenicia Robineau-Desvoidy, 1863: 750 . Type species: Phaenicia concinna Robineau-Desvoidy, 1863 , by subsequent designation of Townsend (1916b: 8).

Bufolucilia Townsend, 1919b: 542 . Type species: Lucilia bufonivora Moniez, 1876 , by original designation.

Francilia Shannon, 1924: 74 . Type species: Francilia alaskensis Shannon, 1924 , by monotypy.

Viridinsula Shannon, 1926: 131 (as a subgenus of Lucilia Robineau-Desvoidy, 1830 ). Type species: Musca pionia Walker, 1849 , by original designation. Shannon (1926) erected the subgenus Viridinsula for L. pionia based on the anteroventral expansion of the head seen in male specimens. Curran (1934b) elevated Shannon’s subgenus to full genus level for both L. pionia and L. deceptor . James (1966) concluded Curran’s elevation of Viridinsula to full genus status was not warranted and retained it as a subgenus and applied it to all three species endemic to the Galápagos. Rognes (1991) did not recognize subgenera and listed Viridinsula as a synonym of Lucilia .

Diagnosis. The genus can be distinguished by a bare stem vein above; presence of a sclerite on the suprasquamal ridge with a conspicuous cluster of setae near the base of the scutellum and the lower calypter bare above ( Whitworth 2006, fig. 14). It also normally has a metallic, shining body color. Other families ( Muscidae , Sarcophagidae , and Tachinidae ) have species with shining body color which may be encountered in the region, see the key to families in Whitworth (2006: 693) and the discussion in Vargas & Wood (2010) to distinguish them.

Discussion. This study provides a key to the 23 Lucilia species known to occur in the Neotropical Region. See Tables 1 and 2 for a comparison of important character states used to identify these species. Twelve species were selected for detailed study, including six new species. The five species of Lucilia endemic to the West Indies, and L. cluvia will not be discussed in detail herein because they were addressed in Whitworth (2010). Three species, L. deceptor , L. pionia (Walker) , and L. setosa (James) are found in the Ecuadorean Galápagos Islands. Lucilia deceptor is also found in the Costa Rican Cocos Islands. James (1966) provided descriptions and figures of male genitalia for each of these species, but they were not studied in detail. Tantawi & Sinclair (2013) conducted a more detailed study of the Galápagos species providing illustrations of male genitalia and frons of both sexes. The cosmopolitan species L. cuprina and L. sericata are also not detailed herein since they have been described by numerous authors ( Hall 1948, Whitworth 2006). The Nearctic species L. coeruleiviridis Macquart , L. illustris (Meigen) , and L. silvarum (Meigen) which likely occur in Mexico ( James 1970) are not included in this key; these species may be keyed using Whitworth (2006: 718). Lucilia coeruleiviridis was listed as a species found in the Neotropical Region by Whitworth (2010) following Townsend (1908) who recorded it from Cuba (as L. oculata ). Whitworth (2010) listed L. coeruleiviridis in Guatemala, based on a single specimen examined. The listing by Kosmann et al. (2013) of this species in the Neotropical Region appears to be based on Whitworth (2010). Since that study, the Guatemala specimen was re-examined and it was determined to be a discolored and nontypical specimen of L. eximia . Lucilia coeruleiviridis has been excluded from the key since I have not found it in the Neotropical Region and I have been unable to verify its presence in Cuba. A search of Townsend material in USNM revealed no paratypes of L. oculata , and no USNM specimens of L. coeruleiviridis from Cuba were found (Norman Woodley, pers. comm.). The record of L. oculata , and hence L. coeruleiviridis , from Cuba is assumed to have been based on a misidentified specimen. Lucilia coeruleiviridis can be determined using the key to Lucilia in Whitworth (2006) if its presence is suspected. However, it is included in the barcode diagram to show its relationship to other Lucilia ( Fig. 161 ).

Despite the paucity of good distinguishing characters in Neotropical Lucilia , this study has revealed six new species, which are described herein. The status of three species, L. japuhybensis , L. ochricornis , and L. purpurascens has been clarified herein.

Head Characters. The ratio of frons to head width measured at the narrowest area of the frons is a useful character to help separate many species, especially for males. Six species have males with exceptionally broad frons (0.115–0.23); four have frons of medium width (0.05–0.06) and thirteen have narrow frons (0.01–0.04) (Table 1). In the species studied, the female frons is always wider than the male (0.21–0.39) frons; widths in females tend to be less distinctive than in males, but can be useful to help separate some species (Table 1). Seven species have some pale setae on the gena along with dark setae (as in Fig. 1 ). When pale setae are present on the gena, they may be limited to the posterodorsal corner of the gena or extend forward and downward to cover almost all of the gena. Species with this character state include L. albofusca , L. cluvia , L. deceptor , L. nitida , L. pulverulenta , L. rica and L. rognesi , while the remaining 16 species have only dark setae on the gena (as in Fig. 2 ). Occasional specimens in the group with dark seta on the gena may have a few pale setae that occur just above the junction of the postgena with the gena; specimens with this condition should still be considered to have dark setae on the gena.

Fourteen species have pale, weak setae below and behind the postocular row (as in Fig. 3 ), eight species have stout dark setae, the condition of these setae in L. problematica was not determined (Table 1). Some species with weak setae, behind the postocular row, like L. eximia , and L. woodi , have a few stout dark setae behind the posteroventral corner of the eye, but the setae behind the rest of the postocular row are weak. These specimens are considered to have weak setae below and behind the postocular row of setae.

Eye facet size was measured in 11 selected species to help separate poorly known species. See the methods section for an explanation of how facet diameters were measured. See Table 1, for a comparison of facet size by species and sex. Males of six species have anterior facets that are relatively large, ranging from 0.55mm–0.68mm (as in Fig. 6 ). Males of the remaining five species studied have smaller anterior facets ranging from 0.40mm–0.49mm (as in Fig. 5 ). Anterior facets in L. woodi are twice the size of posterior facets in both sexes. In L. purpurascens the anterior facets in males are twice the diameter of the posterior facets (0.64mm/0.32mm) ( Fig. 6 ), while in females the anterior and posterior facets are much smaller and more similar in size (0.45mm/0.34mm).

Thorax and Abdomen Characters. Chaetotaxy patterns on the thorax are similar in most species of Neotropical Lucilia , and this character is not mentioned unless the pattern varies. Exceptions are two cosmopolitan species of Lucilia , L. cuprina and L. sericata , which have three postsutural acrostichal setae, while the remaining 21 species have two postsutural acrostichals. In the Galápagos species L. deceptor , the presutural intra-alar is sometimes much reduced or absent, while it is usually stronger in all other species. Color of upper and lower calypter disc and rim are also important characters that vary with species and in some cases, by sex. Some species have upper and lower calypters pale in both sexes, L. cluvia , L. cuprina , L. deceptor , L. pionia , L. sericata , L. setosa , in others, calypters are pale except lower calypter faintly tan to brown in males only, L. eximia , L. ibis , L. mexicana , L. rica , L. ochricornis , and L. retroversa to both upper and lower calypters dark in both sexes, L. fayeae , L. nitida , L. pulverulenta , L. purpurascens , L. vulgata , L. woodi , L. japuhybensis , L. rognesi ( Fig. 8 ), to upper and lower dark in males and upper pale and lower dark in females, L. problematica , to upper calypter pale and lower dark in both sexes, L. albofusca ( Fig. 9 ), L. lucigerens . Basicosta color is dark brown to tan in most species ( Fig. 11 ); only L. cluvia , L. cuprina , L. deceptor , L. problematica , L. retroversa and L. sericata have whitish to orange colored basicostas ( Fig. 10 ). The presence or absence of the coxopleural streak can help distinguish species (Table 2). When present, it is a suture-like depression between the katepimeron and meron (see Rognes 1991, fig. 6 or Whitworth 2006, fig. 16). The streak is often paler in color and distinct, when there is no color difference, the suture can be hard to see. On the mainland of Central and South America, species with pale setae on the gena lack the streak, except it is present in L. cluvia , which occurs in Central America. In the Galápagos, Cocos and West Indies islands, L. deceptor (Galápagos and Cocos) has pale setae and lacks the streak, while L. rica (West Indies) has both pale setae and the streak. Again on the mainland, all species with dark setae on the gena have the streak present, except L. purpurascens and L. woodi . The island species of L. pionia and L. setosa (Galápagos) and L. retroversa (West Indies) have dark setae on the gena, and lack the streak; in L. fayeae (West Indies) the streak is variable, it is distinct in some specimens and absent in others.

Microtomentum patterns on the thorax and abdomen provide useful characters; see Table 2 for a comparison between species. The thorax may be covered with microtomentum (as in Fig. 12 ), limited to the presutural area (as in Fig. 13 ), small patches (as in Fig. 14 ) or the whole thorax may be polished (as in Fig. 15 ). In most species, the abdomen has T1+2 dark greenish or bluish black T1+2 and T3 usually have microtomentum, while some or all of segments T4 and T5 are polished. The posterior edges of T3 and T4 have black bands in the three endemic Galápagos species. Faint bands can sometimes be seen in the same area on other Lucilia species. Generally body color cannot be relied on for species distinctions, with a few exceptions. Two Neotropical species lack the bright, shining body color, the West Indies L. problematica , and the Galápagos and Cocos Islands L. deceptor (Curran) are dull colored with dense microtomentum and only faint underlying metallic blackish-green color. For L. ibis , the very distinctive violet-pink body color is a useful character ( Fig. 21 ), as is the bright aeneous coloration on T 5 in most L. lucigerens .

Males. Frons width in males tends to be very consistent within species and is an important key character for many species. See Table 1 for a comparison of frons to head ratios for each species. The shape of surstyli and cerci tends to be very similar between species, but in lateral view, they fall into a few more or less distinct groups: surstylus short broad, parallel sided (digitate), L. cuprina ( Hall 1948, figs. K, L), L. ibis , L. ochricornis , L. sericata ( Hall 1948, figs. F, G), L. vulgata ( Figs. 43, 44 ; 51, 52 , 59, 60 respectively); surstylus a little longer, narrower, parallel sided, L. woodi ( Fig. 61, 62 ); surstylus medium length, digitate, gradually expanded toward distal end of surstylus, L. nitida ( Fig. 49, 50 ); surstylus medium length, curved forward with distal end expanded, L. purpurascens ( Figs. 55, 56 ), L. lucigerens ( Whitworth 2010, figs. 46, 47), L. retroversa ( Whitworth 2010, figs. 48, 49); surstylus medium length, parallel sided, L. albofusca ( Figs. 39, 40 ), L. problematica ( Hall 1948, p. 423, A, B), L. pulverulenta ( Figs. 53, 54 ), L. rica ( Whitworth 2010, figs. 50, 51), L. rognesi ( Figs. 57, 58 ); surstylus with distal end narrower, and curved slightly forward, L. cluvia (figs. 38, 39, Whitworth 2010), L. eximia ( Figs. 41, 42 ), L. pionia ( James 1966, fig. 2; Tantawi & Sinclair 2013, figs. 3G, H); surstylus medium length, narrower at base, distal half expanded, L. fayeae ( Whitworth 2010, figs. 44, 45); surstylus in lateral view medium length, slightly curved forward, slender and parallel sided; in posterior view the cerci and surstyli are about equal in length, L. setosa ( James 1966, figs. 7, 8; Tantawi & Sinclair 2013, figs. 2E, F); surstylus longer, slender, curved forward, parallel sided, L. mexicana ( Figs. 47, 48 ) and L. japuhybensis ( Figs. 45, 46 ). Lucilia mexicana also has distinctive upside down Y-shaped cerci in the posterior view, and tip of cercus with distinct hook ( Figs. 47, 48 ). Lucilia japuhybensis surstyli and cerci are exceptionally long, and in lateral view, the tip of the cercus is hooked like L. mexicana . Lucilia deceptor cerci are long and slender, much longer than surstyli; tips of cerci diverge in posterior view, in lateral view, the tip of the surstylus is enlarged and extends posteriorly ( James 1966 figs. 4, 5; Tantawi & Sinclair 2013, figs 1E, F. Phalli of the 12 species studied in detail herein ( Figs. 63–86 ) are very similar to those illustrated in Whitworth (2010), figs. 52–63. The epiphallus is broad and cupped and the distal end curves forward, tip angling more or less sharply downward. The epiphallus originates near posterior end of basiphallus in most species; it is slightly farther forward in L. mexicana and L. japuhybensis . The venter of hypophallus is serrated; tip of paraphallus curves down and usually slightly to the rear; acrophallus similar in all species, anterior end with posterior pointing denticles. Shape of hypandria is more or less distinctive in each species although variation is minimal between species; width of distal end tends to be narrower in species such as L. japuhybensis and L. woodi , while broader in species like L. mexicana , L. ochricornis , L. purpurascens , and L. vulgata . The pre- and postgonites of all species are similar, the pregonite has 3–4 setae, including one on the tip, while the postgonite has a single long seta originating from near the base. The ejaculatory sclerites are similar, broader in L. mexicana and L. nitida , while narrower in the remaining species. See Figs. 87–122 for photos of hypandria, pre- and postgonites, and ejaculatory sclerites. The sternites are also similar, though the exact shape and size of each sternite varies between species, especially in segments ST2 and ST4–5 ( Figs. 123–134 ).

Females. Females share many characters with males, but they consistently differ in several ways. In the 23 species studied, frons width was broader than males, (0.21–0.39 of head width at narrowest) (see Table 1 for a comparison of species), with one lateroclinate and two proclinate orbital setae. Typically the frontal vitta and fronto-orbitals are much broader in females than males and the frontal setae extend the full length of the frontoorbital plate, while in most males, they do not. Normally the pattern of microtomentum on the thorax does not vary significantly between sexes, but in some species the pattern on the abdomen may vary. If it is significant, it is mentioned in the species discussion. The most extreme example is in L. mexicana where females have all but the front edge of T5 polished, while in males, only the rear half of T5 is polished. For some species, calypter color of the upper and lower calypters differs between sexes (Table 2). See Whitworth (2010) for a discussion of ovipositor characters of four species endemic to the West Indies, and L. cluvia . Lucilia problematica females were not dissected. The female genitalia of Galápagos species L. deceptor , L. pionia , and L. setosa also were not dissected in this study and their condition is unknown. In the 12 species studied in detail, female genitalia were dissected and ovipositors and spermathecae were illustrated ( Figs. 135–158 ). Tergites and sternites 6–8 of the ovipositor exhibited some variation between species. In 9 species, the cuticle on the anterior margin of T6 is weakened and translucent, in 2 species, L. mexicana and L. rognesi , only the anterolateral corners are weakened, and in one species, L. eximia , there is no weakening. ST6 of most species have lateral margins weakened and ST 6–7 has a posterior digit-like prolongation into the intersegmental membrane. This digit-like extension is found in 11 of the 12 species studied herein; only L. purpurascens lacks these extensions ( Fig. 143 ). When the ovipositors of West Indies Lucilia were studied ( Whitworth 2010) these structures were not noted, but upon re-examination the projections are present to some degree in all. Rognes (1991) also found these extensions in some species of Lucilia he studied in Fennoscandia and Denmark. The central portion of T7 is at least partially divided by weakened, translucent cuticle and/or membrane in all 12 species. The pattern of cuticular weakening extends the full length of the tergite in L. eximia , L. ibis , L. ochricornis , L. nitida , L. purpurascens , and L. woodi ; in the remaining species the division of the tergite ends just short of the posterior margin. The weakened cuticle is further separated by membrane in all species, ranging from a tiny area of membrane at the anterior edge in L. ochricornis to the anterior two-thirds of the segment in L. albofusca , L. eximia , L. nitida , L. purpurascens , L. rognesi , L. vulgata , and L. woodi . T8 is composed of two tergites broadly separated by membrane, in some species the tergites join at the posterior edge of the segment. In L. ibis , L. japuhybensis , L. purpurascens , and L. vulgata the posterior edge of the tergite is more or less connected by a narrow strip of cuticle. The shape of ST8 is somewhat variable among species. The shape of the epiproct and hypoproct are similar between species though they exhibit some variation in shape ( Figs. 135–158 )). The spermathecae of all 12 species are similar as well ( Figs. 147–158 ).

Distribution. Some species have very limited distributions so range information can be used to narrow down species identity. For example, endemic species in the West Indies and Galápagos Islands are not found on the mainland. Luclia mexicana is known only from Mexico, Guatemala, and Honduras in the Neotropics while L. ibis is known only from the eastern slope of the Andes in Argentina, Bolivia, and Peru. When distribution data helps distinguish species, it is included in the key.

DNA Analysis. Samples of specimens for selected species were submitted to the BOLD project at the University of Guelph in an effort to increase information needed to distinguish valid species. See details of this project under Methods. CO1 barcodes were generated for 14 species of Neotropical Lucilia . See the barcode diagram showing the clusters of species analyzed ( Fig. 161 ). Details of barcoding results, when available, are given under each species.

Key to the Species of Neotropical Lucilia

See Tables 1, 2 for a comparison of important key characters in each species.

1 Three postsutural acrostichal setae; basicosta orange (as in Fig. 10 ); abdomen usually with apparent mesal division in which one half is microtomentose, the other half shining when viewed from a sharp angle laterally......................... 2

- Two postsutural acrostichal setae; basicosta usually tan to dark brown ( Fig. 11 ) (except basicosta orange in L. cluvia , two West Indies species and one Galápagos species); abdomen usually uniformly metallic or microtomentose................... 3

2 Central occipital area with single seta just below inner vertical seta ( Whitworth 2006, fig. 73); abdomen dull coppery; humeral callus with 2–3 small setulae along posterior margin ( Whitworth 2006, fig. 74); metasternum bare; frons of male broad, much broader than width of parafacial at level of lunule, 0.20 (0.19–0.21/7) of head width; male genitalia (as Phaenicia pallescens ) as in Hall (1948, fig. 24, J–M); widespread in the Neotropical Region.................................. 3. L. cuprina

- Central occipital area with 2–5 setae below inner vertical seta ( Whitworth 2006, fig. 73); abdomen usually bright green, occasionally shining coppery; humeral callus with 6–8 small setulae along posterior margin ( Whitworth 2006, fig. 74); metasternum setose; frons of male narrower, about equal to width of parafacial at level of lunule, 0.13 (0.12–0.14/6) of head width; male and female genitalia as in Rognes (1991, figs. 455–465); common near many larger cities and developed areas in the Neotropical Region......................................................................... 20. L. sericata

3 Two species known only from the Galápagos Islands, and one species, L. deceptor , known from the Galápagos Islands and Cocos Island off the west coast of Costa Rica (where it coexists with some mainland species). Lucilia deceptor is distinct from the mainland species of Lucilia in having the thorax and abdomen dull blackish to blackish green with dense whitish microtomentum on thorax and all abdominal segments; frons of male very broad, 0.10 head of width or more, only male L. cluvia among species with two posterior acrostichals has such a wide frons (about 0.12 of head width)...................... 21

- Not known from the Galápagos or Cocos Islands (except L. eximia ); male frons narrower, 0.075 of head width or less..... 4

4 Body color violet-pink with aeneous highlights ( Fig. 21 ); face bright yellow to gold from above, including lower frontoorbital, parafacial, and gena; all abdominal tergites microtomentose; upper calypters white, both sexes, lower calypter light tan in male, white in female; basicosta brown; fifth tergite aeneous in some; male frons broad, at narrowest, 0.05 (0.04–0.06/5) of head width, slightly narrower than the width of first flagellomere; female frons, at narrowest, 0.26 (0.25–0.27/5) of head width, known primarily from Peru, Junin Region near Chuquisunca, San Felix, San Ramon and Cusco Province near Huadquina and Huancabamba ( Baumgartner & Greenberg 1985), also Santa Cruz, Bolivia and Rio Tapia, Tucaman Province, Argentina 7. L. ibis

- Body color combination not as above; face usually darker; usually part or all of T4 and T5 polished; male frons width variable .................................................................................................... 5

5 Known only from Jamaica, fifth abdominal tergite coppery or aeneous (not always obvious in some females); intrapostocular area golden; body color dark blue sometimes with purple highlights; upper calypter white, lower calypter tan in both sexes. See Whitworth (2010) for more information....................................................... 9. L. lucigerens

- not as above......................................................................................... 6

6 Known only from Bermuda. Body color metallic-tan; basicosta pale orange; all abdominal tergites microtomentose; known only from six specimens, probably extinct. See Whitworth (2010) for more information............... 14. L. problematica

- Body color normally shining metallic green, blue, or purple................................................... 7

7 Gena with, at least, some pale setae mixed with dark setae on the posterior edge, anterior to the postgena. In some species the pale setae extends forward to midway on the gena or beyond ( Fig. 1 )............................................ 8

- Gena with dark setae only ( Fig. 2 ) (note the postgena has some pale setae in all species). Rarely a few pale setae may be found just ahead of the postgena in specimens with dark setae, but they are not found beyond the edge of the postgena......... 13

8 Basicosta pale whitish to yellow or orange, rarely light tan; both calypters pale in both sexes; rear of T4 and all of T5 polished (fig. 37, Whitworth, 2010). Male frons exceptionally broad, 0.10–0.12 of head width, at narrowest, broader than the width of the first flagellomere; female frons, at narrowest, 0.29 (0.28–0.30/4) of head width; known from Mexico, Guatemala, Honduras, Costa Rica, and West Indies................................................................. 2. L. cluvia

- Basicosta light brown to dark brown; lower calypters dark in both sexes except for L. rica which is known only from the West Indies; condition of T4 and T5 variable; male frons narrower, 0.01-0.035 of head width; female frons narrower, 0.21–0.26 of head width........................................................................................... 9

9 Known only from the West Indies; upper calypter pale both sexes, lower calypter dark in male, pale in female. Rear edge of T3, all of T4 and T5 polished when viewed from rear (fig. 36, Whitworth, 2010); anterior edge of presutural area of thorax microtomentose....................................................................................18. L. rica

- Known only from Central and South America. Upper and lower calypters brown in both sexes (except upper pale and lower brown in both sexes of L. albofusca ); only rear one-third to two-thirds of T4 and T5 polished; microtomentum on presutural area of thorax variable from a broad band to absent.......................................................... 10

10 Upper calypter pale in both sexes (sometimes light tan, in poor specimens or in low light), lower calypter dark in both sexes ( Fig. 9 ); face pale yellow-gold from above, orange from below; anterior edge of gena often orange; dorsum of thorax all polished; male frons narrow, 0.02 (0.01–0.02/5) of head width, about equal to width of anterior ocellus; female frons narrow, 0.21 (0.20–0.22/5) of head width. Known from Brazil, Colombia, Ecuador, French Guyana, Guyana ( British Guiana), Panama, Peru, and Venezuela ( Figs. 159 , 160 )........................................................... 1. L. albofusca

- Upper calypter light tan to darker brown, lower calypter brown to dark brown (as in Fig. 8 ); face and edge of gena darker; thorax microtomentum variable; frons width variable.......................................................... 11

11 Presutural area of thorax with broad band of whitish microtomentum (as in Fig. 13 ); male, surstylus digitate, medium length, parallel sided ( Fig. 53 ); male frons very narrow, 0.018 (0.015–0.02/5) of head width at narrowest, narrower than the width of the medium ocellus; female frons, at narrowest, 0.24 (0.23–0.25/5) of head width. Known from Colombia, Costa Rica, Ecuador, Honduras and Panama ( Figs. 159 , 160 )................................................. 15. L. pulverulenta

- Presutural area of thorax all polished ( Fig. 15 ), or at most only a few small patches of microtomentum (as in Fig. 14 ); male genitalia variable; male frons 0.015–0.03 of head width; female frons 0.21–0.24 of head width...................... 12

12 Presutural area of thorax all polished (as in Fig. 15 ); male surstylus digitate, medium length, gradually expanding toward distal end, cercus about equal in length to surstylus ( Figs. 49, 50 ); known from northern South America, Brazil, Peru, and Venezuela ( Fig. 160 ); male frons narrow, 0.02, (0.015–0.03/6) of head width at narrowest, about equal to the width of median ocellus; female frons, at narrowest, 0.23 (0.21–0.24/6) of head width … 11. L. nitida

- Presutural area of thorax mostly polished with microtomentose streak between acrostichal and dorsocentral setae and a whitish patch between dorsocentral seta and humeral callus; surstylus medium length, parallel sided, cercus shorter and broader at base ( Figs. 57, 58 ) known from Central America, Costa Rica, Honduras and Panama ( Fig. 159 ); male frons, at narrowest, 0.02 (0.015 –0.025 /4) of head width, about equal to width of medium ocellus; female frons, at narrowest, 0.23 (0.23/4) of head width at narrowest............................................................................... 19. L. rognesi

13 Basicosta pale yellow to orange ( Fig. 10 ); upper calypter pale in both sexes, lower calypter tan in male, pale in female; known only from the West Indies (see Whitworth 2010 for details)........................................17. L. retroversa

- Basicosta brown (rarely a few L. ochricornis are seen with an orange basicosta); not known from the West Indies, except for L. fayeae and L. eximia ................................................................................. 14

14 Disc of upper calypter usually pale in both sexes, from whitish to yellowish, rim may be tan, clearly lighter than lower calypter in male; lower calypter pale in female, light tan to brown in male; coxopleural streak present (see this character in Whitworth 2006, fig. 16); anterior one-third to one-half of gena orange in two of three species, silvery in L. mexicana ............ 15

- Disc of upper and lower calypters light tan to dark brown in both sexes; gena normally dark silvery to brown; coxopleural streak absent in all but two species ( L. fayeae , L. vulgata )..................................................... 17

15 One or more rows of stout dark setae below and behind strong postocular row (as in Fig. 4 ); T4 with, at most, the rear edge polished to only rear half of T5 polished; male frons broad, about the width of the first flagellomere (0.05–0.07 of head width); female frons 0.26–0.28 of head width................................................................... 16

- Setae below and behind strong postocular row pale whitish or yellow and usually weak, except a few stronger setae can be found near the posteroventral corner of the eye ( Fig. 3 ); polished area larger on abdomen with rear half to one-third of T4, and all of T5 polished (in West Indies specimens most or all of T4 is polished); male frons narrow, less than the width of fifth flagellomere, 0.04 (0.03–0.05/19) of head width, at narrowest; female frons 0.25 (0.24–0.28/11); widespread from the southern United States through Central and South America.................................................. 5. L. eximia

16 Lower parafacial and anterior half or more of gena orange; usually rear edge of T4 and all of T5 polished; proepisternal depression usually with pale setae; male frons, at narrowest, 0.06 (0.05–0.075/5) of head width, about equal to the width of the parafacial; female frons, at narrowest, 0.26 (0.24–0.26/5) of head width; cerci in male genitalia nearly parallel when viewed from rear ( Fig. 52 ); first flagellomere in male is usually narrower than parafacial; rarely with an orange basicosta. Known from Argentina, Brazil, Paraguay, Peru and Uruguay ( Fig. 160 ).................................... L. ochricornis … (12)

- Parafacial and gena dark silvery; in males, only rear half T5 of abdomen polished, in females, all of T5, or all but front edge of T5 polished; proepisternal depression usually with dark setae; male cerci in the shape of an inverted Y when viewed from the rear ( Fig. 48 ); in male, first flagellomere usually wider than parafacial; male frons, at narrowest, 0.055 (0.05–0.06/8) of head width, slightly narrower than the width of the parafacial; female frons, at narrowest, 0.28 (0.26–0.30/6) of head width. Known from the southern United States south through Mexico into Guatemala and Honduras................... 10. L. mexicana

17 Known only from the West Indies. Setae below and behind strong postocular row pale and weak. Presutural area of thorax with heavy microtomentum; rear one-half to two-thirds of T4 and all of T5 polished; see Whitworth (2010) for more information............................................................................................. 6. L. fayeae

- Not as above....................................................................................... 18

18 Entire dorsum of thorax covered with heavy whitish microtomentum when viewed from rear ( Fig. 12 ); T4 and T5 usually entirely polished; calypters very dark brown; usually a strong row of brown setae below and behind the postocular row, some setae may be weaker; male frons extremely narrow, 0.01/5 of head width at narrowest, eyes almost touch, a thin line midway, narrower than width of a single adjacent eye facet; surstylus in lateral view short, curved forward, with distal end expanded ( Fig. 55 ); in male, anterior thoracic spiracle much enlarged, about equal to humeral callus; anterior eye facets much larger than posterior facets ( Fig. 6 ); female frons broad, at narrowest, 0.28 (0.27–0.29/4) of head width. Known from Argentina, Bolivia, Costa Rica, Colombia, Ecuador, Guatemala, Mexico, Panama, Peru, and Venezuela.................. 16. L. purpurascens

- Only presutural area of thorax with microtomentum ( Fig. 13 ) or whole thorax bare and shining ( Fig. 15 ); abdomen with, at least, some microtomentum on T4; calypters not dark brown; condition of setae below and behind postocular row variable; male frons wider, 0.03–0.07 of head width, in two species, similar in one ( L. woodi ); surstylus not as above; anterior thoracic spiracle smaller; anterior eye facets smaller (except similar in L. woodi ); female frons 0.23–0.25 of head width, except up to 0.28 in L. japuhybensis which is known only from southern Brazil............................................. 19

19 Setae below and behind strong postocular row pale and weaker (some strong setae may occur behind the posteroventral corner of the eye); rear one-third to two-thirds of T4 on abdomen polished; coxopleural streak absent; calypters darker brown; postgena either all pale setae or anterior quarter with dark setae; male frons usually 0.017– 0.03 of head width............. 20

- Row of stout dark setae behind and below the strong postocular row ( Fig. 4 ); anterior edge of presutural area with heavy microtomentum (as in Fig. 13 ); only rear edge of T4 polished; coxopleural streak present; calypters lighter tan; anterior half of postgena with dark setae, remainder pale; surstylus in male digitate, medium length, gradually expanding toward distal end ( Fig. 59 ); male frons wide, 0.05 (0.045–0.07/7) of head width at narrowest, slightly narrower than width of parafacial at lunule. Known from Argentina, Bolivia, Brazil, Colombia, Peru, and Venezuela ( Fig. 160 )...................... .. 22. L. vulgata

20 Presutural area of thorax usually all polished with little or no microtomentum (as in Fig. 15 ); only rear third T4 of abdomen polished; proepisternal depression usually with dark setae (rarely pale); anterior fourth of postgena with dark setae, remainder pale; surstylus long and curved to rear ( Figs. 45, 46 ); male frons broader, at narrowest, 0.03 (0.02–0.04/5) of head width, about two-thirds the width of the parafacial at lunule; female frons, at narrowest, 0.25 (0.23–0.28/5) of head width; known only from Brazil ( Fig. 160 )............................................................................................................................................................................. 8. L. japuhybensis

- Front edge of presutural area normally with a broad band of whitish microtomentum ( Fig. 13 ), some specimens with only patchy microtomentum on the anterior edge of the presutural area (as in Fig. 14 ); T4 polished except for anterolateral corners in males, all polished in females; proepisternal depression usually with pale setae; all setae on postgena pale; male surstylus short and digitate ( Figs. 61, 62 ); known from Costa Rica, Honduras and Panama ( Fig. 160 ); male frons narrower, 0.017 (0.01–0.02/5) of head width, about equal to width of median ocellus; female frons, at narrowest 0.24 (0.22–0.25/5) of head width..................................................................................... 23. L. woodi

21 Thorax and abdomen dull blackish to greenish black with heavy tan to whitish microtomentum; basicosta pale orange; gena with pale setae on posteroventral area; setae below and behind postocular row pale and weak; ST5 is deeply incised with elongate lobes, as in Tantawi & Sinclair (2013, fig. 1G); Male frons broad, 0.23 (0.21–0.24/5) of head width at narrowest, almost twice the width of the parafacial at narrowest; female frons about 0.35 (0.35/1) of head width at narrowest; genitalia are distinctive, cercus is slender and much longer than surstylus, the tips of the cerci diverge in posterior view, in lateral view, the tip of the surstylus is enlarged and projects posteriorly ( Tantawi & Sinclair 2013, figs. 1E, F)..................4. L. deceptor

- Thorax and abdomen with areas of shining blue green, or coppery cuticle with whitish or brown microtomentum; basicosta brown; gena with only dark setae; setae below and behind postocular row strong; ST5 not deeply incised as in Tantawi & Sinclair (2013, fig. 3I); male frons narrower, 0.12–0.18 of head width at narrowest; genitalia not as above; female frons narrower (0.31–0.33 of head width)............................................................................ 22

22 Lower gena and parafacial expanded anteroventrally as in Tantawi & Sinclair (2013, fig. 3C) (except see discussion of variant found on the island of Española, Tantawi & Sinclair 2013); thorax and abdomen shining bright metallic green to coppery with weak microtomentum (tends to be stronger in males than females); in females, disc of T5 without stout, black setae; presutural intra-alar seta often weak or absent (variable, seen in four of six female specimens); male frons narrower, 0.115 (0.10–0.13/2) of head width at narrowest, slightly narrower than width of parafacial at narrowest; male genitalia as in Tantawi & Sinclair (2013, figs. 3G, H); female frons 0.31 (0.30–0.32/3) of head width at narrowest.......................... 13. L. pionia

- Lower gena and parafacial not expanded anteroventrally as in Tantawi & Sinclair (2013, fig. 1C); thorax and abdomen shining green showing through heavy whitish microtomentum; disc of T5 with stout black setae; presutural seta present; male frons broader, 0.17 (0.16–0.18/3) of head width, about 1.25x the width of parafacial at narrowest; male genitalia as in Tantawi & Sinclair (2013, figs. 2E, F); female frons 0.33 (0.33–0.34/3) of head width at narrowest..................... 21. L. setosa

Species descriptions

Tables 1 and 2 provide a comparison of important morphological characters for all 23 species.

Figures

FIGURE 159. Map of Lucilia species found in Central America.

FIGURE 160. Map of Lucilia species found in South America.

FIGURE 161. A neighbor joining tree of specimens and species of Neotropical Lucilia using CO1 DNA barcode, made with K2P distance and generated in BOLD.

FIGURES 1–6. 1–2. Left lateral view of gena=g and postgena=pg showing color of setae on gena. 1. L. cluvia, setae pale. 2. L. eximia, setae dark. Left lateral view of head showing condition of setae below and behind postocular row on occiput (white arrow). 3. L. albofusca female, setae pale and weak. 4. L. ochricornis female, setae stout and dark. Left lateral view of eye facet size, see white arrows 5. L. ochricornis male, anterior and posterior facets very small. 6. L. purpurascens male, anterior facets twice the diameter of posterior facets.

FIGURES 7–15. 7–9. Upper and lower calypter color. 7. L. eximia female, both pale. 8. L. japuhybensis female both dark, both sexes. 9. L. albofusca female, upper pale, lower dark, both sexes.10–11. Basicosta color left lateral view. 10. L. cluvia female, basicosta orange. 11. L. eximia female, basicosta brown. 12–15. Pattern of microtomentum on thorax viewed from the rear. 12. L. purpurascens female, heavy on whole dorsum of thorax. 13. L. woodi female, broad band on presutural area of thorax. 14. L. rognesi female, patches on presutural area of thorax. 15. L. nitida female, microtomentum absent, presutural area of thorax all polished.

FIGURES 16–21. Photos of adults and labels. 16–20. Adult types and labels. 16. L. ochricornis (Wiedemann) (lectotype male of Musca ochricornis Wiedemann). 17. L. eximia (Wiedemann) (holotype male and paratype female of L. hirtiforceps Shannon, in copula). 18. L. ochricornis (Wiedemann) (cotype male of Lucilia mera Shannon & Del Ponte). 19. L. ochricornis (Wiedemann) (cotype male of L. primaveris Shannon & Del Ponte). 20. L. purpurascens (Walker) (holotype male of L. ocularis Shannon). 21. L. ibis, left lateral view of non-type female from San Ramon, Peru.

FIGURES 43–46. Left lateral and posterior views of male genitalia of Lucilia spp. 43, 44. L. ibis. 45, 46. L. japuhybensis.

FIGURES 51–54. Left lateral and posterior views of male genitalia of Lucilia spp. 51, 52. L. ochricornis. 53, 54. L. pulverulenta.

FIGURES 59–62. Left lateral and posterior views of male genitalia of Lucilia spp. 59, 60. L.vulgata. 61, 62. L. woodi.

FIGURES 47–50. Left lateral and posterior views of male genitalia of Lucilia spp. 47, 48. L. mexicana. 49, 50. L. nitida.

FIGURES 55–58. Left lateral and posterior views of male genitalia of Lucilia spp. 55, 56. L. purpurascens. 57, 58. L. rognesi.

FIGURES 39–42. Left lateral and posterior views of male genitalia of Lucilia spp. 39, 40. L. albofusca. 41, 42. L. eximia.

FIGURES 63–72. Phallus of male Lucilia in left lateral and dorsal views. 63, 64. L. albofusca. 65, 66. L. eximia. 67, 68. L. ibis. 69, 70. L. japuhybensis. 71, 72. L. mexicana.

FIGURES 73–82. Phallus of male Lucilia in left lateral and dorsal views. 73, 74. L. nitida. 75, 76. L. ochricornis. 77, 78. L. pulverulenta. 79, 80. L. purpurascens. 81, 82. L. rognesi.

FIGURES 83–86. Phallus of male Lucilia in left lateral and dorsal views. 83, 84. L. vulgata. 85, 86. L. woodi.

FIGURES 87–98. Hypandrium, pre- and postgonites, and ejaculatory sclerite of male Lucilia. 87–89. L. albofusca. 90–92. L. eximia. 93–95. L. ibis. 96–98. L. japuhybensis.

FIGURES 99–110. Hypandrium, pre- and postgonites, and ejaculatory sclerite of male Lucilia. 99–101. L. mexicana. 102–104. L. nitida. 105–107. L. ochricornis. 108–110. L. pulverulenta.

FIGURES 111–122. Hypandrium, pre- and postgonites, and ejaculatory sclerite of male Lucilia. 111–113. L. purpurascens. 114–116. L. rognesi. 117–119. L. vulgata. 120–122. L. woodi.

FIGURES 123–134. Male Lucilia abdominal sternites ST1–5. 123. L. albofusca. 124. L. eximia. 125. L. ibis. 126. L. japuhybensis. 127. L. mexicana 128. L. nitida (ST1 is missing).129. L. ochricornis. 130. L. pulverulenta. 131. L. purpurascens. 132. L. rognesi. 133. L. vulgata. 134. L. woodi.

FIGURES 135–138. Ovipositors of Lucilia. 135. L. albofusca. 136. L. eximia. 137. L. ibis. 138. L. japuhybensis.

FIGURES 139–142. Ovipositors of Lucilia. 139. L. mexicana. 140. L. nitida. 141. L. ochricornis. 142. L. pulverulenta.

FIGURES 143–146. Ovipositors of Lucilia. 143. L. purpurascens. 144. L. rognesi. 145. L. vulgata. 146. L. woodi.

FIGURES 147–158. Spermathecae of Lucilia. 147. L. albofusca. 148. L. eximia. 149. L. ibis. 150. L. japuhybensis. 151. L mexicana. 152. L. nitida. 153. L. ochricornis. 154. L. pulverulenta. 155. L. purpurascens. 156. L. rognesi. 157. L. vulgata. 158. L. woodi.