ARTICLE

Taxonomy of black scavenger (Diptera: ) from Luzon, Philippines

Socrates D. Letana

Institute of Biology, College of Science, University of the Philippines, Diliman, Quezon City 1101, Philippines

ourteen of black scavenger flies in four gene- (Ozerov 2005, Ozerov 2010, Iwasa 2008, Iwasa et al. 2008, Ang ra are treated taxonomically in this paper. Five spe- and Meier 2010). This discovery is largely from the Afrotropical cies belonging to the subfamily Sepsinae are record- region, also with the highest generic endemicity, with some rep- ed for the first time in the Philippines, namely: Sep- resentatives in the Neotropics, Oriental and Australasian regions. sis sepsi Ozerov, Dicranosepsis dudai Ozerov, Di- There are about 78 species and two endemic genera from the Fcranosepsis transita Ozerov, Dicranosepsis sauteri Ozerov, and Oriental region (5.4% generic endemism) (Ozerov 2005). Iwasa Dicranosepsis pseudotibialis Ozerov. A new Philippine island mainly contributed to the discovery and description of the Asian distribution record of Australosepsis niveipennis (Becker) is also and Oriental species (Iwasa 1980, Iwasa 1981, Iwasa 1982, noted. Taxonomic keys for the genera and species are presented. Iwasa 1984a, Iwasa 1984b, Iwasa 1985, Iwasa 1986, Iwasa 1995, Iwasa 1999, Iwasa 2001, Iwasa 2007, Iwasa 2008). INTRODUCTION The taxonomic literature and knowledge regarding the Phil- In the most recent world catalogue of Sepsidae (Ozerov ippine black scavenger flies are very limited. The early works of 2005), there are about 37 genera with 312 described species of Duda (1926a,b) and Zuska (1977b) included Philippine materi- black scavenger flies worldwide. Twelve genera are distributed als. Later, Baltazar’s inventory of Philippine (Baltazar in the Oriental region and six of these are from the Philippines. 1990) paved the way for gathering pertinent information about Previously, Baltazar (1990) listed 12 species in five genera of this group and there are four species added since then in the sepsid flies from the Philippines. In his catalogue, Ozerov (2005) world catalogue (Ozerov 2005). Taxonomic keys in the generic accounted for 14 species in six genera, adding Toxopoda viduata and species levels, literature citation, general description, distri- and three more species of Dicranosepsis. Iwasa (2008) described bution and illustrations are included in this study. and added Toxopoda angulata as a Philippine endemic. The Sep- sidae of the Philippines have been poorly investigated and rec- This work is limited to the collections made in Luzon, Phil- ords are scattered. This study includes 14 species, five of which ippines (Camarines Norte: Daet; Laguna: Mt. Makiling, Univer- are reported for the first time. sity of the Philippines (UP) Los Banos (UPLB), International Rice Research Institute (IRRI), UP Land Grant; Pangasinan: From 2000 to 2010, approximately an average of six sepsid Sison; Quezon: Alabat island; and Zambales) covering a variety species per year are being discovered and are new to science of ecosystems, such as ricefields, pasture and grazing land, resi- dential area, coastal area, dipterocarp forest, and bordering *Corresponding author mossy forest. Email Address: [email protected] Submitted: March 31, 2013 Accepted: April 11, 2014 KEYWORDS Published: May 10, 2014 Philippine Sepsidae, black scavenger flies, ant-mimicking flies, Guest Editor: Victor P. Gapud Dicranosepsis, , Australosepsis, Meroplius

155 Philippine Science Letters Vol. 7 | No. 1 | 2014 The Sepsidae had received little attention due to their minute lected live female specimens were kept alive and fed sugarwater size and association with filth and disgusting habitat. Larvae and solutions for rearing. As for the armature of specimens with clus- adults of this group are exclusively saprophagous with most spe- tered legs, the femoro-tibial joint was temporarily softened with cies associated with mammal feces (Meier 1996, Pont and Meier a drop of xylene and the tibia gently teased away from the femur 2002). with a fine pin. All specimens are currently in the author’s col- lection. The black scavenger flies are considered an important group of flies in the decomposition of pasture dung. There are species Method of Identification associated with both cow and human excrements (Meier 1996). This group of flies is believed to be of some public health con- The adults were identified using a light stereomicroscope up cern due to potential harm as vectors of pathogens and their to 40x magnification and good light quality to detect some of the synanthropic association (Pont and Meier 2002). They are even subtleties of bristling and pruinosity against the black back- reported to be attracted to perishable foods, or foods prepared for ground of the sepsid integument. storage, which may induce the growth of bacteria and fungi (Gagné 1987). A single-female rearing technique was employed in associ- ating collected live female sepsid flies. Males are more easily This group of flies has been a subject of studies in mating identified because of their strong dimorphic characters especially behaviors, e.g., courtship involving leg displays, female recep- in their forelegs: tubercles, number of setae, setulae, spines, spi- tivity as related to ovarian status, precopulatory guarding, and nules, and osmeterium, and their shape. Head and thoracic chae- copulation posture in understanding male foreleg morphology totaxy and wing venation are important characters in sorting to (Eberhard 2001, Eberhard 2002, Ingram et al. 2008, Puniamoor- genera and species. Female sepsids are difficult to pin down to thy et al. 2008). Breeding substrates of Sepsidae are very varied, species since outstanding leg characters and genitalic features ranging from bird and mammal droppings, vertebrate carrion, are very rare and difficult as most taxonomic keys and descrip- decaying vegetation, slime molds, dead insects or snails, to de- tions deal with male specimens. Some illustrations were used caying brown algae. There are reports of swarming of this group along with photographs of observed specimens in identifying the of flies as a hibernation phenomenon (Pont 1987) that may have collection. been the result of mass emergence (Meier 1996). The species treated in this work were identified with the aid The minute black scavenger flies (or ant-mimicking flies or of pertinent taxonomic literature. ensign flies) have been used in forensic investigations and crime scene interpretations such as determining the time of death or the The terminology follows that used by McAlpine (1981). season wherein other groups of flies could not have been present Abbreviations used in the descriptions are as follows: Head: oc, (Benecke 2001). ocellar setae; or, upper fronto-orbital setae; vti, inner vertical setae. Thorax: dc, dorsocentral setae; m, mesopleural setae; npl, METHODOLOGY notopleural setae; sa, supra-alar setae; pa, postalar setae; pprn, postpronotal setae; sc, scutellar setae. Wings: bm, basal medial Collection and Preservation cell. Length of body and length of wing were measured.

Adult black scavenger flies were collected by sweeping and RESULTS AND DISCUSSION hand-netting over substrates where adults are gathering for feed- ing, mating or oviposition. Dung baits from cow, carabao, goats, Family SEPSIDAE Walker, 1883 horse and pig were used in collecting the specimens placed in various habitats such as ricefields, pasture and grazing land, for- Sepsidae Walker, 1833: 245. Type-: Sepsis Fallén, 1810, ested area, coastal and along rivers or natural water sources. Pre- by present designation. vious or relatively old excrements in the different localities were also used in collecting the sepsids. Cow dung was used in rear- ing some collected female sepsids from the field. Newly collect- ed adults were killed in 70-95% ethyl alcohol in separate vials The Sepsidae constitute a small family of acalyptrate flies in with screw caps. They were sorted and stored in smaller vials the . They are distributed in all known zoogeo- and/or tubes with 95% ethyl alcohol for future molecular work graphic regions. Melander and Spuler (1917) separated Sepsidae and slide preparation. Specimens were provided with labels and and and recognized them as different families along codes using lead pencils. with misplaced genera. The early classification scheme of the Sepsidae was summarized by Duda (1926a,b), but Hennig Specimens were kept in 90-95% ethyl alcohol in vials. The (1937), according to Meier (1996), completed the taxonomic ornamentation of the male forelegs and genitalia are highly spe- task by including Orygma luctuosum in Sepsidae; it was previ- cies-specific and are easily seen in alcohol materials. The col- ously classified in the by using the female genitalic

Vol. 7 | No. 1 | 2014 Philippine Science Letters 156 features. Steyskal (1987) modified Duda’s (1926a,b) and Zus- one pair of dorsocentral; lower margin of face projecting; ka’s (1977b) systems probably only to reflect loosely phyloge- middle femora of male bent in middle....Toxopoda Macquart netic relationships, unlike the strict and explicitly phylogenetic intention of Hennig (1949) (see, also, Meier 1996). Body usually shining, at least on parts of pleura; microsetae present, and often macrosetae on abdomen...... 3 Two subfamilies, Orygmatinae and Sepsinae, are recognized in this study. The subfamily Orygmatinae contains only one Hol- 3. A pair of strong orbital bristles present; abdomen lacking arctic species; only Orygma luctuosum Meigen of Sepsinae is macrochaetae except at tip, not or but slightly constricted recorded in this region. behind second tergite; always only one dorsocentral; wings not spotted; postvertical bristles present; no strong acrosti- Diagnostic Character chals; humeral bristle present; only one well-developed vibrissa; wings hyaline or scarcely gray; genal bristle indis- Adult Sepsidae are relatively small, elongate, myrmecomor- tinct or lacking; male hypopygial processes simple, cruciate, phic or ant-like in appearance. The abdomen is constricted basal- with anteroventrally with four bristles or without bris- ly. Vibrissa is absent but several strong peristomal hairs are pre- tles...... Meroplius Rondani sent; palp is vestigial. It can be distinguished from other families of acalyptrate Diptera by the presence of one or more setae on Orbital bristles lacking or very small; abdomen with or the hind margin of the posterior spiracle. ♂ spiracle 6 situated in without preterminal macrochaetae, strongly constricted be- tergite 6 (spiracle 7 in tergite 7+8); ♂ sternite 7 not delimited, hind second tergite; usually two dorsocentrals; wing with or either lost or fused with sternite 8 (the resulting sclerite termed without spots...... 4 sternite 7+8); and, vein A1 shortened and not reaching wing mar- gin. 4. Humerals and postoculars lacking; no wing spots; one dorsocentral; abdomen without macrochaetae; male with Taxonomic Keys peculiar lateral processes (sternite brush) on fourth sterni- te...... Perochaeta Duda Subfamilies Key to the Subfamilies of Sepsidae [adapted from Steyskal Humerals and postoculars present; wing spot present at tip (1987) and Pont and Meier (2002)] of second vein or apically or wing unmarked or marked ba- sally only...... 5 1. Eye small and gena very deep, the face strongly receding; 1 orbital and 2 frontal setae; anepisternum, anepimeron, kate- 5. Abdomen of male always with distinct macrochaetae and pisternum and disc of scutellum densely setulose; 3 strong sternite 4 absent; wing spot, if present, only in vicinity of dorsocentral and 2 postalar setae; large bristly species with vein 2...... Sepsis Fallén robust legs and short tarsi...... subfamily ORYGMATINAE, Orygma Meigen Abdomen of both sexes without distinct macrochaetae, alt- hough sometimes with somewhat stronger hairing on tergal 2. Eye large and gena narrow, the face not receding; 0-1 orbital margins and with strong anal bristles; wing darkened along and 0 frontal seta; anepisternum and katepisternum with costa basally and sometimes with apical spot; sternopleura only a few setulae, anepimeron and disc of scutellum bare; 1 shining anteroventrally….………..……Dicranosepsis Duda -2 dorsocentral setae, or if with 4-5 (Susanomira, Zuskami- ra) then 1 postalar seta; smaller species with more delicate Genus AUSTRALOSEPSIS Malloch, 1925 legs and longer tarsi...... subfamily SEPSINAE Australosepsis Malloch, 1925: 314. Type-species: Australosepsis Genera fulvescens Malloch, 1925 [=Sepsis niveipennis Becker, 1903], Key to the Philippine Genera of Sepsidae (Diptera) [modified by original designation. from several authors] Saltelliseps Duda, 1926: 25. Type-species: Sepsis niveipennis 1. First and second basal cells of wing united; orbital bristles Becker, 1903, by designation of Hennig (1949): 63. lacking; abdomen constricted behind second tergite; ab- dominal macrochaetae present...... Australosepsis Malloch DIAGNOSIS. Head roundish or slightly flattened dorsoventrally. Distance between eyes at level of vibrissae significantly larger First and second basal cells separated...... 2 than the width of postpedicel. Occipital sclerite with several se- tae. Arista bare. Chaetotaxy: or absent, but if present (1), then 2. Thorax and abdomen dull black; abdomen with silvery prui- very short, hair-like; 1 oc, 1 poc, 1 ovt and 1 ivt. 2-3 vibrissae, nose bands; abdominal tergites lacking bristles and setae; always striking from genal setulae. humeral bristle minute; outer verticals (postoculars) lacking;

157 Philippine Science Letters Vol. 7 | No. 1 | 2014 Scutum with the following paired setae: 1 pprn, 2 npl, pal part curved, clearly concave ventrally. absent or present 1 thin, hair-like, 0+2 dc. Proepisternum without setae. Metepimeral bridge absent. Mediotergite shining under Abdomen: Constricted between second and third tergum, last scutellum. Scutellum dorsally without hairs, with a pair of well- segments with long and strong marginal macrochetae. Hypopyg- developed apical setae; basal setae absent or short, hair-like. ium small, simple in structure.

Coxa of male foreleg without osmeterium. Male femur and Length: body: 2.8 to 3.9 mm; wing: 1.6 to 2.8 mm. tibia of foreleg modified. Coxa of midleg in upper half with ver- tical row of thin setulae. Femur of midleg straight. Tibia of male Specimens examined: CAMARINES NORTE: 4 ♂. Daet, Cama- hindleg with a hardly visible osmeterium-like area. rines Norte, cow dung, May 2011, S. D. Letana (Coll. #44b); LAGUNA: 15 ♂. UPLB Cattle Farm, cow dung, 30 May 2006, Wing normal, longer than abdomen, with darkened spot near S. D. Letana (Coll. #17); PANGASINAN: 3 ♂, Sison, Pan- apex R2+3, usually almost unclear from yellowish–colored speci- gasinan, unknown dung, May 2011, Ian Marca (Coll. #46); mens. Anal lobe well-developed. Cells bm and br fused. Alula ZAMBALES: 8 ♂, Cabangan, ricefield, carabao dung, 24 June well-developed, bare. Margin of upper calypter with hair, margin 2006, S. D. Letana (#Z). of lower calypter without hairs. Distribution: Australasian/Oceanian. — Australia (Nothern Abdomen constricted after syntergite 1+2. Sternite 4 of male Territory, Western Australia), New Caledonia (New Caledonia). simple. Surstyli symmetrical, fused to epandrium. Epandrial pro- Oriental. — Bangladesh, China (Guangdong, Taiwan), India cess absent. (Andhra Pradesh, Karnataka, Kerala, Orissa, Tamil Nadu), Indo- nesia (Lombok I., Sulawesi I., Sumbawa I., Timor I.), Japan Key to the species of Australosepsis (Okinawa I.), Malaysia (Borneo I.) Nepal, Philippines (Balabac I., Busuanga I., Culion I., Luzon I., Mindanao I., Palawan I.), 1. Wing clear, without white apex; Male: fore tibia ventrally Singapore, Sri Lanka, Thailand, Vietnam. Palaearctic. — Asia: with short row of stout setulae on basal third, apical half Pakistan. curved...... Australosepsis frontalis (Walker) AUSTRALOSEPSIS NIVEIPENNIS (Becker, 1903) Wing with diffuse dark subapical spot; wing apex milk- (Figures 1 and 2) white; Male: fore tibia ventrally with longer row of setulae on basal half, followed distally by 2 bare tuber- Sepsis niveipennis Becker, 1903: 143. Type-locality: Asyȗt cles...... Australosepsis niveipennis (Becker) (EGYPT); Lectotype ♂, by designation of Zuska (1968): 474, in ZMHUB. AUSTRALOSEPSIS FRONTALIS (Walker, 1860) Sepsis flava Brunetti, 1910: 351. Type-locality: Calcutta frontalis Walker, 1860: 163 (Sepsis). Type-locality: (INDIA); Lectotype ♂, by designation of Zuska (1968): “Makessar: [now= Ujung Pandang] (Indonesia: Sulawesi I.); 474, in NHML. Lectotype ♂, by designation of Zuska (1968): 472, in NHML. Sepsis tincta Brunetti, 1910: 353. Type-locality: Allahabad (INDIA); Lectotype ♀ by designation of Zuska (1968): 474, tenella de Meijere, 1906: 183 (Sepsis). Type-locality: Singa- NHML. pore; Lectotype ♂, by Zuska (1968): 472, in MTMB. Australosepsis fulvescens Malloch, 1925: 314. Type- brevis Brunetti, 1910: 361 (Sepsis). Type-locality: Baroda locality: Sydney (AUSTRALIA: New South Wales); Holo- (India); Holotype ♂, not in NHML and presumed in ZSIC or type ♂, in AMS. destroyed. Head: Frontal plate mostly shining. Outer vertical ½ and post- lieveni Frey, 1917: 25 (Sepsis). Type-locality: Anurandha- ocellar 2/3 length of inner vertical. Gena ½ to 2/3 width of an- pura (Sri Lanka); Holotype ♂, in ZMUH. tennal flagellomere. Subvibrissal setae numerous and short.

Head: Ocellar, postvertical, inner and outer vertical bristles Thorax: Scutum, postpronotal lobes and scutellum dull, only present and well-developed. 1 to 3 dorsocentral setae. Humeral around postalar callus. Anterior notopleural much shorter than bristles developed. posterior; 1-2 dorsocentral setae; acrostichal and dorsocentral ground-setulae minute. Wing apex including veins and mi- Thorax: Wings clear with microtrichiae, without apical spot. crotrichia whitish with a diffused dark spot at the end of R2+3 Anterior femora without tubercles bearing spinules; anterior tibi- vein. Fore tibia ventrally with longer row of setulae on basal ae in proximal half with flat tubercles bearing black spines, distal half, followed distally by 2 bare tubercles; distal parts of tibiae

Vol. 7 | No. 1 | 2014 Philippine Science Letters 158 A

B

Figure 1. Australosepsis niveipennis (Becker). Male. Scale Figure 2. Australosepsis niveipennis (Becker). (After Zuska bar: 1 mm. 1968) Male: A, wing; and B, surstylus, dorsal view. Scale bar: 0.1 mm not strongly curved. Protomeroplius Ozerov, 1999: 92 [described as a subgenus of Abdomen: Subshining. Strongly constricted between syntergite Meroplius]. Gender: masculine. Type-species: Meroplius tris- 1+2 and base of tergite 3. pinifer Ozerov, 1999, by original designation.

Length: body: 3.2 mm; wing: 2.0 mm. DIAGNOSIS. Head roundish or slightly flattened dorsoventrally. Distance between eyes at level of vibrissae significantly larger Specimens examined: PANGASINAN: Sison, Pangasinan, 2 ♂, than the width of postpedicel. Occipital sclerite with several se- 11 May 2011, Ian Marca (Coll #46a). tae. Arista bare. Chaetotaxy: 1 or, 1 oc, 1 poc, 1 ivt, and 1 ovt. 1 vibrissa. Distribution: Afrotropical. — Angola, Botswana, Cameroon, Ethiopia, Ghana, Kenya, Madagascar, Malawi, Namibia, Nige- Scutum with the following paired setae: 1 pprn, 2 npl, 1 ria, Republic of the Congo, Republic of South Africa, Sierra spal, 1 pal, 0+(1-2) dc; often present rows of short setae between Leone, Sudan, Swaziland, Tanzania, Togo, Uganda, Yemen, lines ac (unpaired row) and along lines dc. Proepisternum with- Zimbabwe. Australasian/Oceanian. — Australia (Australian out setae. Metepimeral bridge absent. Mediotergite with shining Capital Territory, New South Wales, Nothern Territory, Queens- (without pollen) spot under scutellum. Scutellum dorsally con- land, Western Australia), Fiji, New Caledonia (New Caledonia), vex, without hairs, with well-developed apical setae; basal setae Papua New Guinea (Bismarck Arch., New Guinea I.), Solomon short, hair-like. Islands, Vanuatu. Oriental. — Afghanistan, Bangladesh, China (Taiwan), India (Andhra Pradesh, Karnataka, Orissa, Rajasthan, Coxa of male foreleg without osmeterium. Male femur and Tamil Nadu, Uttar Pradesh, West Bengal), Indonesia (Java I., tibia of foreleg modified, usually posteriorly with two black Lesser Sunda Is, Sulawesi I.), Japan (Okinawa I.), Malaysia spines; female fore femur usually with a deta in apical third ven- (East Malaysia), Nepal, Philippines (Jolo I.). Palaearctic. — trally. Coxa of midleg in upper half bare. Femur of midleg Asia: Iraq, Israel, Pakistan, Turkmenistan; Europe: Cyprus; straight. Male tibia of hindleg with an osmeterium-like area or North Africa: Egypt, Morocco. with a conspicuous osmeterium.

Genus MEROPLIUS Rondani, 1874 Wing normal, longer than abdomen, with anal lobe well- developed. Cells bm and br separate. Alula well-developed or Meroplius Rondani, 1874: 175. Gender: masculine. Type- moderate, entirely covered with microtrichiae. Margin of upper species: stercoraria Robineau-Desvoidy, 1830 calypter with hairs, margin of lower calypter without hairs. [=Sepsis minuta Wiedemann, 1830], by original designation. Abdomen not constricted after syntergite 1+2. Sternite 4 of Parameroplius Duda, 1926a: 37 [described as a subgenus of male modified. Surstyli symmetrical, fused to epandrium. Meroplius]. Gender: masculine. Type-species: Sepsis fasciculata Epandrial process absent or present. Brunetti, 1910, by monotypy.

159 Philippine Science Letters Vol. 7 | No. 1 | 2014 MEROPLIUS FASCICULATUS (Brunetti, 1910) Genus SEPSIS Fallén, 1810 (Figure 3) Sepsis Fallén, 1810: 17. Gender: feminine. Type-species: Musca fasciculata Brunetti, 1910: 365 (Sepsis). Type-locality: cynipsea Linnaeus, 1758, by designation of “Ceylon” [=Sri Lanka]; Holotype ♂, not in NHML and pre- Curtis (1829): Plate 245. sumed in ZSIC or destroyed. Threx Gistel, 1848: 599. Gender: masculine. Unjustified substi- plumata de Meijere, 1913b: 363 (Sepsis). Type-locality: tute name for Sepsis Fallén, 1810. Type-species: Musca cynipsea “rivier Kamp” (Irian Jaya: New Guinea I.); Lectotype ♂, by Linnaeus, 1758, automatic. designation of Ozerov (1999): 51, in ZMUA. Acrometopia Lioy, 1864: 1088. Gender: feminine. Type-species: Head: Roundish, facial carina small, face receding, gena linear. Sepsis cornuta Meigen, 1826 [=Musca cynipsea Linnaeus, Orbital bristle strong and distinct, inner vertical and outer verti- 1758], by monotypy. cal setae present. Beggiatia Lioy, 1864: 1088. Gender: feminine. Type-species: Thorax: Wings with a distinct gray tinge. Distal modified bristle Sepsis barbipes Meigen, 1826 [=Musca cynipsea Linnaeus, on ventral side of fore femur small, club-like, anterior bristle in 1758], by monotypy. basal fourth of femur long and strong, row of posteroventral set- ulae in basal half not differentiated from normal pilosity; osme- Sepsidimorpha Frey, 1908: 578, 584. Gender: feminine. Type- terium about two-third as long as hind tibia. species: Sepsis loewi Hendel, 1902 [=Sepsis duplicata Haliday, 1838], by monotypy. Abdomen: Hypopygium with a long, slender, apically widened surstylus, and with a sharp anteroventral process. Nicarao Silva, 1995: 203. Gender: masculine. Type-species: Specimens examined: METRO MANILA: 1♂, Quezon Ave., Nicarao rarus Silva, 1995, by original designation. Quezon City, 4 March 2007, S. D. Letana; QUEZON: 1♂, Del Pilar, Quezon, Alabat Island, goat dung, 12 Dec. 2006, S. D. Allosepsis Ozerov, 1992: 44. Gender: feminine. Type-species: Letana (Coll. #40). Sepsis indica Wiedemann, 1824, by original designation.

Length: body: 4 to 4.3 mm; wing: 2.6 to 3 mm. DIAGNOSIS. Head roundish or slightly flattened dorsoventrally. Distance between eyes at level of vibrissae larger than width of Distribution: Australasian/Oceanian. — Papua New Guinea postpedicel. Occipital sclerite with several setae. Arista bare. (New Guinea I.). Oriental. — Bangladesh, China (Guangdong, Chaetotaxy: absent or present, 1 very short hair-like; 1 oc, 1 pos, Taiwan), India (Karnataka, Kerala, Madhya Pradesh, West Ben- 1ovt and 1 ivt. 2-3 vibrissae, always striking from genal setulae. gal), Indonesia (Java I., Sulawesi I.), Kalimantan I., Malaysia (East Malaysia), Nepal, Philippines (Luzon I., Jolo I.), Sri Scutum with the following paired setae: 1 pprn, 1-2 npl, 1 Lanka, Thailand. Palaearctic. — Asia: China (Sichuan), Japan pal (usually short, hair-like), 0+ (1-2) dc; sometimes with a row (Honshu I., Kyushu I., Shikoku I.). of thin and short setulae along each ac, dc, and ial line. Proe- pisternum without setae. Metepimeral bridge absent. Medioterg- ite shining under scutellum. Scutellum dorsally without hairs. With a pair of well-developed apical setae; basal setae absent or shot, hair-like.

Coxa of male foreleg without osmeterium. Male femur and tibia of foreleg modified. Coxa of midleg in upper half with ver- tical row of thin setulae. Femur of midleg straight. Tibia of male hindleg with a hardly visible osmeterium-like area.

Wing normal, longer than abdomen, with dark spot near apex. R2+3 or without spot, with well-developed anal lobe. Cells bm and br separate. Alula well-developed to narrow, entirely covered with microtrichiae. Margin of upper calypter with hairs, margin of lower calypter without hairs.

Abdomen constricted after syntergite 1+2. Sternite 4 of male Figure 3. Meroplius fasciculatus (Brunetti). Male habitus. simple. Surstyli symmetrical, fused to epandrium. Epandrial pro- Scale bar: 1 mm. cess absent.

Vol. 7 | No. 1 | 2014 Philippine Science Letters 160 Key to the species of Sepsis SEPSIS DISSIMILIS Brunetti, 1910

1. Wing without a dark spot at the end of R2+3vein...... 2 Sepsis dissimilis Brunetti, 1910: 355. Type-localities: “Shashthancottah, 12 miles N. N. E. of Quilon”, Rajmahal, Wing with a dark spot at the end of R2+3 vein; sternopleura and “Gathwal district, Western Himalayas” (INDIA); Syn- posterodorsally pruinose; fore femur of male with antero- types 4 ♂♂ and 1 ♀, not in NHML and presumed in ZSIC basal patch of long hair...... S. dissimilis Brunetti or destroyed.

2. Sternopleura dorsally and posteriorly pruinose...... Sepsis albolimbata de Meijere, 1913: 115. Type-locality: ...... S. lateralis Wiedemann “Tainan” (CHINA: Taiwan); Lectotype ♂, by designation of Ozerov (1997): 478, in ZMHUB. Sternopleura wholly pruinose; abdomen of the male al- ways, and often the female also, with distinct macrochaetae Sepsis albopunctata Lamb, 1914: 323. Type-locality: “Mahé, ...... 3 Cascade Estate, 800 feet or over; marshes on coastal plain of Anse aux Pins and Anse Royale” (SEYCHELLES); Syntype 3. Pteropleura pruinose...... S. indica Wiedemann s ♂♂, in NHML (2 ♂♂) and MCZC (1 ♂).

Pteropleura shining...... 4 Sepsis hirtifemur Malloch, 1925: 314. Type-locality: Mosman (AUSTRALIA: New South Wales); Holotype ♂, in 4. Fore femur of male with large ventromedial bump...... NHML...... S. sepsi Ozerov Sepsis acroleucoptera Duda, 1926: 41 [as a variety of al- Fore femur of male without bump...S. coprophila de Meijere bopunctata]. Type-locality: “Anping” (CHINA: Taiwan); Lectotype ♂, by designation of Ozerov (1997): 478, in SEPSIS COPROPHILA de Meijere, 1906 ZMHUB.

Sepsis coprophila de Meijere, 1906: 176. Type-locality: Sepsis natalensis Brunetti, 1929: 27. Type-locality: Weenen SINGAPORE; Lectotype ♂, by designation of Ozerov (REPUBLIC OF SOUTH AFRICA: Natal); Holotype ♂, in (1997): 479, in ZMHUB. NHML.

Body color of males generally reddish; sternopleura wholly Fore femur of male with anterobasal patch of long hair; pruinose; wing without a dark spot at the end of R2+3 vein; postalar bristles weak; a dark spot at the end of R2+3 vein com- Surstylus (hypopygial process) long and stout; male fore femur paratively weak; sternopleura posterodorsally pruinose; hypo- without such distinct median tubercle. Postalar bristles strong. pygial processes short and slender.

Wing apex never whitish. Fore metatarsus ventrally, near Specimens examined: CAMARINES NORTE: 7 ♂, Camambugan, base, with 2 strong, black, sinuous bristles, clearly differentiated Daet, pasture area, cow dung, May 2011, SD Letana (#44a, 44c from other ventral hairs. Male: Fore femur ventrally without and 44d); LAGUNA: 2 ♂, Science, UPLB, horse dung, tubercle. But with a row of spines; fore tibia with a posteroven- 23 April 2008, SD Letana (#27a); 8 ♂, IRRI, Laguna, pitfall cow tral tubercle; surstylus longer, wider. flatter. with long, ventrally dung trap, 11 June 2010, SD Letana (#41a and 41c); QUEZON: directed, basal bristle posteriorly. 1 ♂, Del Pilar, Alabat Island, pig dung, 30 December 2006, SD Letana (#36a); ZAMBALES: 2 ♂, Cabangan, ricefield, carabao Specimens examined: LAGUNA: 1 ♂, UP Land Grant, pasture, dung, 24 June 2006, SD Letana (#Zd). carabao dung, 23 March 2010, SD Letana (Coll. #38c); QUE- ZON: 5 ♂, Del Pilar, Alabat Island, horse dung, 29 December Length: body: 2.6 to 3 mm; wing: 1.5 to 1.8 mm. 2006, SD Letana (Coll. #35b). Distribution: Afrotropical —Democratic Republic of the Con- Length: body: 3.6 to 3.8 mm; wing: 2.5 mm. go, Ethiopia, Kenya, Madagascar, Namibia, Nigeria, Republic of South Africa, Seychelles, Swaziland, Uganda, Zimbabwe. Aus- Distribution: Oriental. — Bangladesh, China (Guangdong, Tai- tralasian/Oceanian. — Australia (Queensland, New South wan), India (Assam, Karnataka, Kerala, Maharashtra, Tamil Na- Wales), New Fiji, Hebrides, Papua New Guinea (New Guinea du, West Bengal), Indonesia (Java I., Sulawesi I., Sumatra I.), I.). Oriental — China (Taiwan), India (Andhra Pradesh, Assam, Japan (Okinawa I.), Malaysia (East Malaysia), Nepal, Philip- Bihar, Karnataka, Kerala, Uttar Pradesh), Indonesia (Lesser Sun- pines (Luzon I., Mindanao I., Palawan I.), Singapore, Sri Lanka, da Is, Sulawesi I.), Japan (Okinawa I.), Malaysia (East Malay- Thailand, Vietnam. Palaearctic. — Asia: Japan (Kyushu I.). sia), Nepal, Pakistan, Philippines (Busuanga I., Culion I., Jolo I., Luzon I., Mindanao I., Negros I., Palawan I.), Thailand, Vi-

161 Philippine Science Letters Vol. 7 | No. 1 | 2014 etnam. Palaearctic. — Asia: Japan (Honshu I., Kyushu I.). immaculata Macquart, 1843: 391 (Sepsis). Type-locality: “De l’ile Bourbon” [=Réunion]; Holotype ♂, in MNHNP. SEPSIS INDICA Wiedemann, 1824 hyalipennis Macquart, 1851: 269 (Sepsis). Type-locality: indica Wiedemann, 1824: 57 (Sepsis). Type-locality: “India EGYPT; Lectotype ♂, by designation of Pont in Pont and orient”; Lectotype♀, by designation of Ozerov (2004): 00, Meier (2002): 167, in UMO. in ZMUC. rufa Macquart, 1851: 269 (Sepsis). Type-locality: Cairo decipiens de Meijere, 1906: 177 (Sepsis). Type-locality: (EGYPT); Holotype ♀, in UMO. Stephansort, Astrolabe-Bai (Papua New Guinea: New Guin- ea I.); Holotype ♂, in MNMB. melitensis Rondani, 1874: 176 (Meroplius). Type-locality: MALTA; Syntypes ♂♂, probably in MZLSF. fusciventris Brunetti, 1910: 357 (Nemopoda). Unavailable name; citation of a Bigot MS name, as nomen nudum is syn- schembrii Rondani, 1874: 176 (Meroplius). Type-locality: onymy with Sepsis indica Wiedemann. MALTA; Holotype ♂, probably in MZSLF.

The males generally reddish or reddish-brown. Sternopleura senegalensis Bigot, 1886: 389 (Nemopoda). Type-locality: dorsally and posteriorly pruinose; postalar setae strong; wing SIERRA LEONE; Holotype ♀, in UMO. without a dark spot at the end of R2+3. The males have strongly modified forelegs with large femoral protrusions and spines. fragilis Becker, 1903: 145 (Sepsis). Type-locality: Lake Bir- Hypopygial process with long bristles. ket-el-Karȗn (EGYPT); Syntypes ♂♂, in ZMHUB

Specimens examined: LAGUNA: 3 ♂, UPCO, UPLB, cow dung, astutis Adams, 1905: 174 (Sepsis). Type-locality: 14-18 March 2008, S. D. Letana (Coll. #29a, 30a and 31a); “Salisbury” [now=Harare] (ZIMBABWE); Lectotype ♂, by QUEZON: 1 ♂, Del Pilar, Quezon, Alabat Island, horse dung, 29 designation of Ozerov (1998a): 85, in SEMK. Dec. 2006, S. D. Letana (Coll. #35a); 1 ♂, Del Pilar, Quezon, Alabat Island, pig dung, 30 Dec. 2006, S. D. Letana (Coll. #36a). lutea Duda, 1926: 51 (Sepsis). Unavailable name; citation of a Wiedemann name, in synonymy with Sepsis lateralis Length: body: 5.6 to 6.2 mm; wing: 4 to 4.8 mm. Wiedemann.

Distribution: Australasian/Oceanian. — Papua New Guinea unicoloripes Brunetti, 1929: 27 (Sepsis). Type-locality: (New Guinea I.). Oriental. — Bangladesh, China (Taiwan), India “Aburi” (GHANA); Lectotype ♂, by designation of Ozerov (Karnataka, Orissa, Tamil Nadu), Japan (Okinawa I.), Nepal, (1998): 87, in NHML. Philippines (Luzon I.),Thailand, Vietnam. Palaearctic. — Asia: Japan (Hokkaido I., Honshu I., Kyushu I.), Korea, Russia definita Brunetti, 1929: 29 (Sepsis). Type-locality: Weenen (Khabarovskiy Kray, Primorskiy Kray). (REPUBLIC OF SOUTH AFRICA: Natal); Lectotype ♂, by designation of Ozerov (1998a): 85, in NHML. SEPSIS LATERALIS Wiedemann, 1830 kwanzaensis Vanschuytbroeck, 1963a: 31 (Sepsis). Type- lateralis Wiedemann, 1830: 468 (Sepsis). Type-locality: locality: Virunga National Parc: “riv. Kombo, affl. Riv. Ru- China; Lectotype ♂, by designation of Pont in Pont and anoli, 1550m” (DEMOCRATIC REPUBLIC OF THE Meier (2002): 167, in ZMUC. CONGO); Holotype ♂, in MRAC.

complicata Wiedemann, 1830: 468 (Sepsis). Type-locality: bombokaensis Vanschuytbroeck, 1963: 50 (Sepsis). Type- China; Holotype ♂, in ZMUC. locality: Virunga National Parc: “Bomboka, prés Kyando- lire, 1650m” (DEMOCRATIC REPUBLIC OF THE CON- inpunctata Macquart, 1839: 118 (Sepsis). Type-locality: not GO); Holotype ♂, in MRAC. stated [from title: Canary Is.]; Holotype ♂, in MNHNP. migeriensis Vanschuytbroeck 1963: 71 (Sepsis). Type- algira Macquart, 1843: 389 (Nemopoda). Type-locality: locality: Virunga national Parc: “Kiribata (Migeri), Moy- Algiers (ALGERIA); Holotype ♂, probably in MNHL. enne Lume, 1760 m” (DEMOCRATIC REPUBLIC OF THE CONGO); Holotype ♂, in MRAC. lateralis Macquart, 1843: 390 (Nemopoda). Junior second- ary homonym, preoccupied by Sepsis lateralis Wiedemann, curiosa Ozerov, 1996: 144 (Sepsis). Substitute name for 1830. Type-locality: “Du Brésil ou du Chili” [probably from Nemopoda lateralis Macquart. Africa, not South America]; Holotype ♀, in MNHNP.

Vol. 7 | No. 1 | 2014 Philippine Science Letters 162 “astuta”, incorrect subsequent spelling of astutis Adams Length: body: 3.8 mm; wing: 2.5 mm. [Bezzi, 1908: 169; Vanchuytbroeck, 1961: 78]. Distribution: Oriental. — Indonesia (Sumbawa I.), Vietnam (Ha “definita Duda” , error for definita Brunetti Tay). Philippines (Luzon; New record). [Vanschuytbroeck (1962): 459].

Head: dark, with only genal area brown to yellow; to mostly yellow, with only fronto-orbital plates and median occipital scle- A C rite brown. Antenna brown to almost wholly reddish-yellow.

Thorax: wholly dark with brown postpronotal lobes and proe- pisternal area; to entirely yellow and a median line on post- notum. Scutellum black to reddish-yellow. Legs wholly yellow; to mid and hind coxae basally; fore femur mid and hind femora B except at base, mid tibia except on apical quarter, hind tibia wholly, and tarsomeres 4-5 dark.

Abdomen: wholly dark; to mainly dark with basal part and api- Figure 4. Sepsis sepsi Ozerov. (After Ang and Meier 2010) cal part reddish to yellow. Male: A, fore femur, posterior; B, fore tibia, posterior view; and C, hypopygium, dorsal view. Scale bar: 0.5 mm Specimens examined: 1♂ LAGUNA: Los Banos, Umali Subdivi- sion, attracted to TV, 28 May 2011, S. D. Letana (Coll. #47); 2 ♂, UP Land Grant trail, carabao dung, 23 March 2010, SD Letana (Coll. #42).

Length: body: 3.6 mm; wing: 2.5 mm.

Distribution: Afrotropical. — Angola, Botswana, Cameroon, Democratic Republic of the Congo, Ethiopia, Ghana, Kenya, Madagascar, Malawi, Mauritius, Namibia, Nigeria, Republic of South Africa, Republic of the Congo, Réunion, Seychelles, Sier- ra Leone, Swaziland, Tanzania, Uganda, Zambia, Zimbabwe; Asia: Yemen. Australasian/Oceanian. — USA (Hawaiian Is.), Papua New Guinea (New Guinea I.). Palaearctic. — Asia: Af- ghanistan, China (Hebei), Iraq, Israel, Japan (Kyushu I.), Syria; Europe: Cyprus, Greece, Italy, Malta, Spain (incl. Balearic Is), Turkey; North Africa: Algeria, Azores, Canary Is, Egypt, Libya, Madeira Is, Morocco, Tunisia. Oriental. — Bangladesh, China (Guangdong, Taiwan), India (Andhra Pradesh, Assam, Bihar, Himachal Pradesh, Karnataka, Kerala, Meghalaya, North Ben- gal, Tamil Nadu, Uttar Pradesh, West Bengal), Japan (Okinawa I.), Malaysia (East Malaysia), Myanmar, Nepal, Pakistan, Philip- pines (Luzon I., Negros I.), Sri Lanka, Thailand.

SEPSIS SEPSI Ozerov, 2003 (Figures 4 and 5)

Sepsis sepsi, Ozerov 2003: 1276. Type-locality: Wawó, 450m (INDONESIA: Sumbawa I.); Holotype ♂, in DEI.

Sepsis sepsi can be distinguished from other Sepsis by the row of four large spines on a large rounded ventromedial bump of the fore femur (Fig.16a). The fore tibia (Fig.16b) lacks a rounded lobe, and the surstylus differs in structure.

Specimens examined: ZAMBALES: 3 ♂, Cabangan, ricefield, Figure 5. Sepsis sepsi Ozerov. (After Sepsidnet: sepsidnet- carabao dung, 24 June 2006, SD Letana (# Zb). rmbr.nus.edu.sg) Male habitus. Scale bar: 1 mm

163 Philippine Science Letters Vol. 7 | No. 1 | 2014 Genus DICRANOSEPSIS Duda, 1926 tween av setae in the basal fifth of fore femur less than length of basal seta and apical end of distal av seta does not Dicranosepsis Duda, 1926a: 43 [described as a subgenus of reach the half-way point of fore femur...... Sepsis]. Gender: feminine. Type-species: Sepsis bicolor ...... D. revocans (Walker) Wiedemann, 1830, by original designation. Fore femur without av seta basally; mid-tibia without brown DIAGNOSIS. Head round or slightly flattened dorsoventrally. ring at the distal end...... D. javanica (de Meijere) Distance between eyes at level of vibrissae significantly larger than the width of postpedicel. Occipital sclerite with several se- DICRANOSEPSIS DUDAI Ozerov, 2003 tae. Arista bare. Chaetotaxy: or absent, 1 oc, 1 poc, 1 ovt, and 1 (Figure 6) ivt. 2-3 vibrissae, always striking from genal setulae. Dicranosepsis dudai Ozerov, 2003: 89. Type-locality: Scutum with the following paired setae: 1 pprn, 2 nol, 0-1 “Batoe Doelang: (Indonesia: Sumbawa I.); Holotype ♂, in pal, 0+2 dc. Proepisternum without setae. Metepimeral bridge DEI. present. Mediotergite shining under scutellum. Scutellum dorsal- ly without hairs, with a pair of well-developed apical setae; basal Head: Frons dark or brownish near apex. Face yellow to brown, setae absent or short, hair-like. but antennal grooves usually blackish. Gena brown to black. Antenna brownish. Occipital sclerite with several setulae. Gena Coxa of male foreleg without osmeterium. Male femur and with a row of setulae along lower margin. Postgena with seta, 2- tibia of foreleg modified. Coxa of midleg in upper half bare. 3 vibrissae. Femur of midleg straight. Tibia of male hindleg with a hardly visible osmeterium-like are. Thorax: Black. Legs yellow but femur of midleg and hindleg in apical half and tibia of midleg and hindleg in basal half usually Wing normal, longer than abdomen, with moderate anal darkened; tibia of midleg blackish near apex. Tarsomeres 4 and 5 lobe. Cells bm and br separate. Alula narrow, entirely coverd of all legs black. Wing clear, with brownish veins; basal-costal with microtrichiae. Margin of upper calypter with hairs, margin cell basally and basal cell completely blackish. Upper calypter of lower calypter without hairs. and margins white, lower calypter and margins darkened. Halter yellowish. Abdomen constricted after syntergite 1+2. Sternite 4 of male simple. Surstyli symmetrical, fused to epandrium. Epandrial pro- Abdomen: Black. Constricted after syntergite 1+2. Syntergite cess present. 1+2 at sides with 2-4 thin setae. Tergites 3-5 each with a row of thin marginal setae. Surstyli symmetrical, fused to epandrium. Key to the species of Dicranosepsis Specimens examined: LAGUNA: 2 ♂, Hortorium, UPLB, ex cul- 1. Anepimeron completely pruinose...... 2 ture from cow dung, 10 March 2007, SD Letana (#10). Length: body: 4.1 mm; wing: 2.2 mm. Anepimeron shining in anterior half or with shining spot anteriorly...... 5 Distribution: Oriental. — Indonesia (Flores I., Sumbawa I.). Philippines (Luzon; New record). 2. Fore trochanter strongly or somewhat extended ventrally ....4 DICRANOSEPSIS JAVANICA (de Meijere, 1904) Fore trochanter not extended ventrally...... 3 Sepsis javanica de Meijere, 1904: 107: Type-locality: Tosari 3. Hind trochanter posteriorly covered with hair-like setae...... (INDONESIA: Java I.); Lectotype ♂, by designation of ...... D. dudai Ozerov Ozerov (1997): 147, in ZMUA.

Hind trochanter without hair-like setae .....D. transita Ozerov “javana”, incorrect subsequent spelling of javanica de Mei- jere [Hennig (1949): 61] 4. Ventral process of fore trochanter same length as body of fore trochanter...... D. sauteri Ozerov Color of body black. Legs yellow. Wing without dark spot near apex. Height of gena+subgena approximately 10 times Ventral process of fore trochanter shorter than body of fore shorter than vertical diameter of eye . First flagellomere approxi- trochanter; fore femur with 1–2 anterobasal setae...... mately 1.4 times as long as wide Proepisternum completely ...... D. pseudotibialis Ozerov greyish pruinose. Anepimeron shining in anterior half and prui- nose in posterior half. 5. Fore femur with 2 av setae in the basal third; distance be-

Vol. 7 | No. 1 | 2014 Philippine Science Letters 164 Fore trochanter simple. Fore femur without av in basal third. Holotype ♀, in NHML. Midtibia with 1 av in apical third. Length of av and pv on first tarsomere of mid leg less than the height of tarsomere 4 of mid Sepsis acuta de Meijere, 1913a: 118 [as forma of bicolor]. leg. Length of av on first tarsomere of hind leg less than the Type-locality: “Tainan” (China: Taiwan); Lectotype ♂, by length of tarsomere 4 of hind leg. designation of Ozerov (1997): 154, in ZMHUB.

Anal vein approximately 6 times as long as the width of bm Sepsis bipilosa Duda, 1926a: 48 [as variety of bicolor]. cell. Alula approximately 1.5 times as wide as bm cell. Type-locality: “Macuyam” (China: Taiwan); Lectotype ♂, by designation of Ozerov (1997): 155, in DEI. Specimens examined: QUEZON: 1 ♂, Del Pilar, Alabat Island, carabao dung, 30 December 2006, SD Letana (#37b). Sepsis bipilosiformis Duda, 1926a: 48 [as a subvariety of bicolor variety of bipilosa]. Type-locality: Colombo (Sri Length: body: 3.7 mm; wing: 2.1 mm. Lanka; lectotype ♂, by designation of Ozerov (1997): 155, in MTMB. Distribution: Oriental. — China (Guangdong, Taiwan), India, Indonesia (Java I.), Myanmar, Nepal, Pakistan, Philippines parabipilosa Duda, 1926b: 55. Error for bipilosiformis Du- (Luzon), Sri Lanka, Thailand, Vietnam. da.

DICRANOSEPSIS PSEUDOTIBIALIS Ozerov, 2003 Head: Color of body from yellow to black. Height of (Figure 7) gena+subgena approximately 10 times shorter than vertical di- ameter of eye. First flagellomere approximately 1.5 times as long Dicranosepsis pseudotibialis Ozerov, 2003: 88. Type- as wide. locality: “Batoe Doelang” (INDONESIA: Sumbawa I.); Holotype ♂, in DEI. Thorax: Proepisternum completely grayish pruinose, anepimer- on shining in anterior half and pruinose in posterior half. Legs Head: slightly flattened dorsoventrally in lateral view. Postpedi- yellow. Fore trochanter simple. Mid tibia with 1 av in apical cel in profile long-oval approximately 1.5 time longer than wide. third. Length of av and pv on first tarsomere of mid leg less than 1 oc, 1 poc, 1 ivt, 1 ovt; or absent. Occipital sclerite with several the length of tarsomere 4 of mid leg. Length of av on first tarso- setulae. Gena with a row of setulae along lower margin. Post mere of hind leg less than the length of tarsomere 4 of hind leg. gena with 1 seta. 2-3 vibrissae. Wing without dark spot near apex.

Thorax: Scutellum with well-developed apical setae; basal setae Specimens examined: LAGUNA: 6 ♂, Makiling Forest Reserve short, hair-like. Scutum: 1 pprn, 2 npl, 1 spal, 1 pal, 0+2 dc. Fe- Trail, approx 500m asl, Mt. Makiling Laguna, old horse dung, 15 mur of foreleg with 1 av basally. Coxa of midleg bare in upper September 2008, SD Letana (#19); 2 ♂, Makiling Forest Reserve half. Femur of midleg with a row of short a. tibia of midleg with Trail, approx 500m asl, Mt. Makiling Laguna, cow dung, 15 1 v in apical third and with apical av and pv. Femur and tibia of September 2008, SD Letana (#20a and b). hindleg without striking setae. Length: body: 3 to 3.5 mm; wing: 2.1 to 2.2 mm. Abdomen: constricted after syntergite 1+2. Syntergite 1+2 at sides with 2-4 thin setae. Tergites 3-5 each with a row of thin Distribution: Australasian/Oceanian. — Australia (Queens- marginal setae. Surstyli symmetrical, fused to epandrium. land), Solomon Islands. Oriental. — China (Guangdong, Tai- wan), India (Assam), Indonesia (Flores I., Sulawesi I., Sumbawa Specimens examined: LAGUNA: 1 ♂, UPCO, UPLB, from cow I.), Japan (Okinawa I.), Malaysia, Myanmar, Philippines dung, 14 March 2008, SD Letana (#29b); 3 ♂, Tayabak Camp (Luzon), Sri Lanka, Thailand, Vietnam. Site, Mt. Makiling, fresh horse dung, 19 September 2008, SD Letana (#23a). DICRANOSEPSIS SAUTERI Ozerov, 2003 (Figure 8) Length: body: 3 to 3.3 mm; wing: 2.1 to 2.2 mm. Dicranosepsis sauteri Ozerov, 2003: 87. Type-locality: Distribution: Oriental. — Indonesia (Sumbawa I.). Philippines “Taihoku” (China: Taiwan); Holotype ♂, in DEI. (Luzon; New record). Head: Frons black, but brownish near apex. Face and gena red- DICRANOSEPSIS REVOCANS (Walker, 1869) dish yellow; antennal grooves slightly darkened. Subgena, clype- us and postcranium black. Antenna reddish yellow. 1 oc, 1 poc, 1 Sepsis revocans Walker, 1869: 163. Type-locality: ivt, 1 ovt; or absent. Occipital sclerite with several setulae. Gena “Makessar” [now=Ujung Pandang] (Indonesia: Sulawesi I.); with a row of setulae along lower margin. Postgena with 1 seta.

165 Philippine Science Letters Vol. 7 | No. 1 | 2014 2-3 vibrissae. Length: body: 3.6 mm; wing: 2.2 mm.

Thorax: Dark. Legs yellow, tibia of midleg blackish near apex. Distribution: Oriental — China (Taiwan), Philippines (Luzon; Wing clear, with brownish veins; basal-costal cell and basal cell New record). completely blackish. Upper calypter and margin white, lower calypter and margin blackish. Halter yellowish. Scutum with the DICRANOSEPSIS TRANSITA Ozerov, 1997 following paired setae: 1pprn, 2 npl, 1 spal, 1 pal, 0+2 dc. Ane- (Figure 9) pisternum in posterior half bearing scattered hairs and with long seta near posterior margin. Scutellum with well-developed apical Sepsis gracilis Duda, 1926a: 48 [as variety of bicolor]. Jun- setae; basal setae short, hair-like. ior primary homonym, preoccupied by Sepsis gracilis Zet- terstedt, 1847. Type-locality: “Chosokei” (China: Taiwan); Abdomen: Black. Constricted after syntergite 1+2. Syntergite Lectotype , by designation of Ozerov (1997): 157, in DEI. 1+2 at sides with 2-4 thin setae. Tergites 3-5 each with a row of thin marginal setae. Surstyli symmetrical, fused to epandrium. Dicranosepsis transita Ozerov, 1997: 156. Substitute name for Sepsis gracilis Duda. Specimens examined: LAGUNA: 2 ♂, UP Land Grant trail, La- guna, carabao dung, 23 March 2010, SD Letana (#39); 1 ♂, IR- Head: Dark brown to black. Height of gena + subgena approxi- RI, Laguna, cow dung pit fall trap, 11 June 2010, SD Letana mately 12-16 times shorter than vertical diameter of eye. First (#41d). flagellomere approximately 1.4 times as long as wide.

A B C A B C

D D E

Figure 6. Dicranosepsis dudai Ozerov. (After Ozerov 2003) Figure 7. Dicranosepsis pseudotibialis Ozerov. (After Ozerov Male: A, fore tibia, posterior view; B, same, anterior view; C, fore 2003) Male: A, fore tibia (left), posterior view; B, same, anterior femur (left), posterior view; D, same, anterior view; and E, tro- view; C, fore femur (left), posterior view; and D, same, anterior chanter of hindleg. view.

A B C A B C

D D E

Figure 8. Dicranosepsis sauteri Ozerov. (After Ozerov 2003) Figure 9. Dicranosepsis transita Ozerov. (After Ozerov 2003) Male: A, fore tibia (left), posterior view; B, same, anterior view; C, Male: A, fore tibia (left), posterior view; B, same, anterior view; C, fore femur (left), posterior view; and D, same, anterior view for femur (left) posterior view; D, same, anterior view; and E, frag- ment of tibia.

Vol. 7 | No. 1 | 2014 Philippine Science Letters 166 Thorax: Proepisternum and anepimeron completely grayish pru- REFERENCES: inose. Legs yellow; mid tibia completely yellow, or darkened on anterior, ventral, and posterior sides near apex, or with black ring Adams CF. Diptera Africana, I. Kans Univ Sci Bull 1905; 3: 149 near apex. Short av on first tarsomere of male hind leg. Wing -208. without spot near apex. Ang Y, Meier R. Five additions to the list of Sepsidae (Diptera) for Vietnam: Perochaeta cuirassa sp. n., Perochaeta lobo Specimens examined: LAGUNA: 8 ♂, DTRI, UPLB, cow dung, sp. n., Sepsis spura sp. n., Sepsis sepsi Ozerov, 2003 and 10 March 2007, SD Letana (#11 and 12); 1 ♂, Hortorium, Sepsis monostigma Thompson, 1869. ZooKeys 2010; 70: UPLB, carabao dung, 12 March 2008, SD Letana (#34b). 41-56. Baltazar CR. An Inventory of Philippine Insects I. Orders Neu- Length: body: 3 mm; wing: 2 mm. roptera, Strepsiptera, Trichoptera & Diptera. University of the Philippines Los Baños. 1990. Distribution: Oriental. — China (Taiwan), Philippines (Luzon; Becker Th. Ägyptische Dipteren (Fortsetzung und Schluss). Mit- New record). teilungen aus dem Zoologischen Museum in Berlin. 1903; 2: 67-195. SUMMARY Benecke M. Murder most putrid. Nature 2001; 410: 752. Bezzi M. Ditteri iritrei raccolti dal Dott. Andreini e dal Prof. This study of Philippine black scavenger flies covers the Tellini. Parte seconda. Diptera Cyclorrapha. Bulletino subfamily Sepsinae and the genera Australosepsis, Meroplius, della Societrà Entomologica Italiana. 1908; 39(1907): 3- Sepsis and Dicranosepsis, which include 14 species, four of 199. which are new records for the Philippines. The putative oriental Bigot JMF. Diptères nouveaux ou peu connus. 29e partie (suite) Australosepsis niveipennis has new Philippine distribution rec- XXXVII 2e. Essai d’une classification synoptique du ord which was previously recorded in Jolo island. groupe des Tanypezidi (mihi) et description de genres et d’espèces inèdits. Annales de la Société Entomologique Each species discussed was provided with literature citation, de France. 1886; 6: 369-392. general description, distribution, and habitat/substrate record. Brunetti E. New Oriental Sepsinae. Records of the Indian Muse- Keys and diagnoses were also provided for each category when- um. 1910; 3(1909): 343-372. ever possible. Brunetti E. New African Diptera. Annals and Magazine of Natu- ral History, series 10. 1929; 4: 1-35. ACKNOWLEDGMENTS Colless DH. Biogeography of Australian Sepsidae (Diptera). Aust J Zool 1980; 28: 65-78. This study would not have been possible without the support and Curtis J. British entomology; being illustrations and descriptions patience of my MS adviser Dr. Clare R. Baltazar, along with Dr. of the genera of insects found in Great Britain and Ire- Venus J. Calilung and Dr. Antonio J. Alcantara. Their guidance, land: containing coloured figures from nature of the most patience, and encouragement contributed immensely to this out- rare and beautiful species, and in many instances of the come. I am grateful to my UPLB professors Dr. Victor P. Gapud, plants upon which they are found. Vol. VI. Published by Dr. Jessamyn Adorada, and the late Dr. Stephen G. Reyes for the the author. London. Plates 242-289. 1829. discussions and insights given. I thank Professor Rudolf Meier Duda O. Monographie der Sepsiden (Dipt.). I. Annalen des (National University of Singapore) for the hospitality in accom- Naturhistorischen Museums Wien. 1926a; 39: 1-153. modating me for a short but meaningful stint in his laboratory; to Duda O. Monographie der Sepsiden (Dipt.). II. Annalen des Dr. Andrey L. Ozerov (Zoological Museum, Moscow State Uni- Naturhistorischen Museums Wien. 1926b; 40: 1-110. versity) who has been gracious to all my queries, for reprints and Eberhard WG. Species-specific genitalic copulatory courtship in permission to use his excellent photographs; Dr. Ireneo Lit, Jr. sepsid flies (Diptera: Sepsidae, Microsepsis) and theories (UPLB Museum of Natural History) for allowing me to study the of genitalic evolution. Evolution 2001; 55: 93-102. museum sepsid collection; Dr. Jose Sargento (Makiling Center Eberhard WG. Physical restraint or stimulation? The function(s) for Mountain Ecosystems) for issuance of GP; Dr. Finbarr G. of the modified front legs of male Archisepsis diversi- Horgan (IRRI) for allowing me to work on insects associated formis (Diptera, Sepsidae). Journal of Behavior with dung and decomposing snails both in his laboratory and 2002; 15(6): 831-850. field sites; to my fieldwork company in some sites Dr. Stephen Fallén CF. Specimen Entomologicum novam Diptera disponendi Marshall (University of Guelph), Almon Merep, Jouhannes Faid- methodum exhibens. Litteris Berlingianis, Lundae. 1810: iban, and Ian Marca. 26 pp. Feng-Yi Su K, Kutty SN, Meier R. Morphology versus mole- CONFLICTS OF INTEREST cules: the phylogenetic relationships of Sepsidae (Diptera: Cyclorrhapha) based on morphology and DNA sequence None. data from ten genes. Cladistics 2008; 24: 902-916. Floate KD, Wardhaugh KG, Boxall ABA, Sherratt TN. Fecal

167 Philippine Science Letters Vol. 7 | No. 1 | 2014 residues of veterinary parasiticides: Nontarget effects in Iwasa M. Review of the flies (Diptera) associated with animal the pasture environment. Annual Review of Entomology dung and human feces in Japan. Med Entomol Zool 2007; 2005; 50: 153-179. 58(3): 155-166. Frey R. Über die in Finnland gefundenen Arten des Formen- Iwasa M. Revisional notes on the genus Toxopoda Macquart kreises der Gattung Sepsis Fall. (Dipt.). Deutsche Ento- (Diptera, Sepsidae) in the Oriental and Australasian re- mologische Zeitschrift 1908; 577-588. gions. Entomological Science 2008; 11: 449-463. Frey R. Ein Beitrag zur Kenntnis der Depterenfauna Ceylons. Iwasa M, Kanmiya K, Lee CE. Notes on the Sepsidae (Diptera) Öfversigt af Finska Vetenskaps-Societetens Förhandlin- from South Korea. Esakia 1994; 34: 203-208. gar, 59 A. 1917; 20: 1-36. Iwasa M, Maruyama M, Nakamura E, Yamashita N, Watanabe Gagné RJ. Flies (Diptera). In: Gorham JR, ed. Insect and Mite A. 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Vol. 7 | No. 1 | 2014 Philippine Science Letters 168 Hamm, Germany:Schulz-Wundermann, 1826: 1-412. Puniamoorthy N, Kotrba M, Meier R. Unlocking the “Black Meijere JCH de. Über einige indo-australische Dipteren des Un- box”: internal female genitalia in Sepsidae (Diptera) garischen National-Musuems, bez. des Naturhistorischen evolve fast and are species-specific. BMC Evolutionary Museums zu Genua. Annales Historico-Naturales Musei Biology 2010; 10: 275. Nationalis Hungarici 1906; 4: 165-196. Robineau-Desvoidy AJB. Essai sur les Myodaires. Mémoires Meijere JCH de. H. Sauter’s Formosa Ausbeute. Sepsinae. presents par divers Savans à l’Académie Royale des Sci- (Dipt.). Annales Historico-Naturales Musei Nationalis ences de l’Institut de France 1830; 2: 1-813. Hungarici 1913a; 11: 114-124. Rondani C. Species italicae ordinis Dipterorum (Muscaria Meijere JCH de. Diptera. I. Résultats de l’Expédition scien- Rndn.). Stirps XXI. Tanipezinae Rndn. 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