taxonID	type	description	language	source
0D42878FFF98FF9E20116323AA7BF9E2.taxon	description	In the classification of Ephydrinae, three groups have been described: the group Coenia plus related genera (Mathis 1979 b) as the sister group of Scatella and related genera (Mathis 1979 b), and a group with Paracoenia and related genera (group II; Mathis 1979 b), Austrocoenia and Ephydrini. Olafsson (1991) presented a phylogeny of Ephydrinae based on genera from the western Palearctic Region. In his study, Olafsson proposed a monophyletic group without postpronotal setae and named it as the tribe Scatellini, thus moving Coenia to Ephydrini. The other genera comprising the group, Paracoenia, Austrocoenia and Coenia, were later moved to Ephydrini, leaving 13 genera in the tribe (Zatwarnicki 1992). Through the years, the taxonomy, phylogeny, and classification of the tribe Ephydrini have undergone many modifications. These studies were usually based on limited groups of species or restricted geographic areas, never on a comprehensive, global basis, making the relationships among these genera less well understood. To clarify these relationships, we have undertaken a comprehensive phylogenetic analysis to test the monophyly of tribe Ephydrini and the tribe’s included genera and subgenera, based on morphological characters from adult males and females. A hypothesis of relationships among taxa within Ephydrini and the other tribes of Ephydrinae are discussed.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF93FF97227D6357AA5FFA00.taxon	description	Figs 3 – 9	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF93FF97227D6357AA5FFA00.taxon	diagnosis	Diagnosis. Austrocoenia is distinguished from other genera of Ephydrini by the following combination of characters: Medium-sized to large shore flies, body length 3.65 – 5.10 mm; coloration generally gray; wing mostly hyaline; setae generally reduced. Head: Mesofrons conspicuously setulose, completely microtomentose, dull, lacking large setae, cruciate or otherwise, parafrons more sparsely microtomentose; lateroclinate fronto-orbital setae 2; medial and lateral vertical setae both well developed; paravertical setae either reduced or lacking; antenna short; basal flagellomere lacking a lateral seta; basal flagellomere subequal in length to that of pedicel from dorsal view; arista short, only slightly longer than length of basal flagellomere, basal 2 / 3 thickened; antennal grooves deeply impressed; face long, distinctly protuberant anteriorly; facial setae small, setula-like; eye subspherical, slightly higher than wide; gena mostly bare, lacking a prominent seta, high, gena-to-eye ratio at least 0.60; maxillary palpus well developed. Thorax: Acrostichal setae in two rows, these extended posteriorly to base of scutellum, a well-developed prescutellar pair, these inserted slightly laterad of alignment of anterior setulae, rows of setulae slightly more divergent posteriorly; dorsocentral setae 5 (1: 4), 1 well-developed dorsocentral seta inserted near base of scutellum, with 3 – 4 larger setae along dorsocentral tract, anterior setae more weakly developed; postpronotal setae 1 – 2, these subequal in length to presutural supra-alar seta; postsutural supra-alar seta reduced or lacking; disc of scutellum bare; lateral scutellar setae 2; prosternum bare. Wing normally developed, mostly hyaline; costal margin with short spinelike setae between 2 nd costal break and the apex of vein R 2 + 3; costal vein long, extended to vein M 1; R stem vein bare dorsally; costal vein ratio 0.18, M 1 vein ratio 0.78. Pulvilli well developed; tarsal claws short and distinctly curved; hindfemur of male not differing markedly from fore- or midfemur, lacking stout, closely set setae; hindtibia of male lacking tuft of setulae; hindtarsi of male evenly cylindrical, normal. Abdomen: Tergites generally setose, setae along margins larger. Males with five visible segments, with tergite 5 longest, trapezoidal, broadly truncate posteriorly; females with 7 segments, sternites narrow, bearing robust and long setae on subanal plate. Male terminalia: Cercus of male terminalia very elongate, length equal to width of epandrium at midheight, fused ventrolaterally with epandrium; surstylus evidently fused with ventral margin of epandrium; gonite plate-like; aedeagus greatly reduced. Female terminalia: Female ventral receptacle with operculum subtrapezoidal, asymmetrical, extended process more or less C-shaped. Distribution. Neotropical: Austrocoenia is endemic to southern South America on the Atlantic Ocean side (Patagonia, between 49 ° – 52 ° S). Remarks. Austrocoenia is a monotypic genus and is somewhat of an anomaly due to the autapomorphic condition of several of its characters. In our morphological studies (Wirth 1970, Mathis 1980) and in this phylogenetic analysis, Austrocoenia was recovered at the base of the cladogram of Ephydrini. We further highlight that the next node in Ephydrini after Austrocoenia is the lineage giving rise to Notiocoenia, a genus that is likewise found only in the southern latitudes of South America (Chile). Wirth (1970: 6 – 7) was perplexed by his new genus, noting that although Austrocoenia is clearly a member of the subfamily Ephydrinae, “ ... it is not closely allied with any known genus. It appears to be closest to Coenia Robineau-Desvoidy, a Holarctic genus, which it resembles to a slight extent in the presence of four pairs of dorsocentral setae and curved tarsal claws. Otherwise, its similarities are diverse and rather remote. Austrocoenia is doubtless an annectant form surviving and modified from a very early offshoot of the Ephydrinae. ” The single included species, A. aczeli Wirth, is only known from the southern temperate regions of the Neotropics (Argentina, Chile). Immature stages are unknown.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF92FF91227862EBAC95F92B.taxon	description	Figs 10 – 15	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF92FF91227862EBAC95F92B.taxon	diagnosis	Diagnosis. Specimens of Calocoenia closely resemble those of Paracoenia but may be distinguished from the latter and other genera of Ephydrini by the following combination of character states: Generally dark colored. Head: Mesofrons subshiny to shiny, inconspicuously setulose; 2 large, lateroclinate fronto-orbital setae; medial and lateral vertical setae both well developed; paravertical seta well developed, about half length of cruciate vertical seta. Arista pectinate, dorsally branching rays subequal to half the length of cruciate vertical seta; arista pectinate, dorsally branching rays subequal to half width of basal flagellomere. Face with interfoveal carina, but dorsal crease not as distinct as in specimens of Paracoenia. Eye subspherical to sub-elliptical, oriented at slight oblique angle to general plane of head. Thorax: Acrostichal setulae uniformly small, in 2 rows that extend to base of scutellum; 5 (1 + 4) well-developed dorsocentral setae; postpronotum with 2 – 3 larger setae; postsutural supra-alar seta well developed, subequal to anterolateral postalar seta; scutellar disc almost flat; prosternum bare. Costal margin with evenly spaced spine-like setae distinct from remaining setae; R stem vein bare. Hindcoxa bare posteriorly along ventral margin; with dorsum subplanate; pulvilli well developed; tarsal claws short and distinctly curved. Abdomen: Male tergite 5 slightly wider than long; tergites generally unicolorous. Male terminalia: Surstyli as angulate processes extended from ventral margin of epandrium, contiguous medially; aedeagus well developed, acutely pointed apically. Female abdomen: Female ventral receptacle with small operculum, conformation subtrapezoidal in lateral view, extended process much longer than width of operculum. Distribution. See under each subgenus. Remarks. Although first described as a subgenus within Paracoenia (Mathis 1975), this taxon was later accorded generic status, as it was recovered as the sister group to the combined Paracoenia / Thiomyia lineage and not as a lineage from within Paracoenia (Mathis 1980: 7 – 8).	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF92FF91227862EBAC95F92B.taxon	description	Zatwarnicki 1995: 235 [world catalog]. Diagnosis. Although similar to Paracoenia s. str. and Leptocoenia, Calocoenia may be distinguished from either as follows: Body length 3.4 – 4.0 mm; subshiny to shiny, metallic brown to greenish-brown; microtomentose, gray ventrally. Head: Mesofrons shining, bronze-gold metallic reflections; pectinate branches of arista not more than twice aristal width at base; pruinose face tan; interfoveal hump not as prominent as Paracoenia s. str., dorsally sloping; eye large, subcircular, width in profile double the length of projecting face in profile; gena-to-eye ratio 0.25; width-to-height ratio 0.66; height-to-length ratio 0.90. Chaetotaxy of head and thorax like Paracoenia s. str. except acrostichal hairs. Thorax: Acrostichal setulae in 2 rows; dorsum microtomentum to subshiny; 5 (1 + 4) well-developed dorsocentral setae; scutellum flattened; pleural areas concolorous with mesonotum centrally, becoming microtomentum, grayed marginally, halters yellow. Wings with costal setae on dorsal and ventral margins; costal vein ratio 0.20; M 1 vein ratio less than 0.80. Male midfemur without comb of setulae. Abdomen: Generally uniform, concolorous; male tergites more noticeably narrowed apically; subshiny to shiny, brown metallic reflections; tergite 5 of male more or less truncate, without anteroventral process; sternite 5 with three posteriorly oriented prongs. Male terminalia: Symmetrical; epandrium subelliptical with closely fused surstyli ventrally and a medial groove; surstylus broadly attached to ventral margin of epandrium, approximate medially, shallowly bifurcate apically with small medial process and larger lateral process; aedeagus elongate, narrow, tapered to acutely pointed, curved apex; phallapodeme robustly crescent shaped in lateral view; gonite with apical half similar to apex of aedeagus in lateral view, tapered, curved, acutely pointed. Female abdomen: Female ventral receptacle with operculum wider than high, extended process considerably longer than operculum. Distribution. Nearctic: Temperate western Nearctic Region from Alberta in Canada south through California and New Mexico. Remarks. Calocoenia s. str. is a monotypic subgenus that only occurs in western North America within temperate zones. We know nothing about the natural history of this species or its immature stages.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF92FF91227862EBAC95F92B.taxon	diagnosis	Diagnosis. Specimens of Leptocoenia closely resemble those of Calocoenia s. str. and Paracoenia but may be distinguished from the latter and other genera of Ephydrini by the following combination of character states: Moderately small shore flies, body length 2.1 – 2.6 mm; generally dark brown, microtomentose. Head: Fronto-orbital areas, mesofrons nearly concolorous, the later subshiny; pectinate aristal branches at most 2.5 X aristal width at base; interfoveal hump not prominent, without pronounced dorsal indentation; microtomentose face light tan; longest setae along ventral margin of face approximately 3 / 4 length of interfoveal hump height; genal seta weak, subequal to humeral setae. Gena-to-eye ratio 0.175; width-to-height ratio 1.00 – 0.65; height-to-length ratio 0.93; eye-width-to-face-length ratio 0.30. Thorax: Mesonotum sparsely microtomentose; scutellum partially flat; acrostichal setae in two rows; 4 (1 + 3) well-developed dorsocentral setae; postpronotal setae present; halteres yellowish-brown to brown. Costal setae weak, developed only on dorsal margin; costal vein ratio 0.18 – 0.20; M 1 vein ratio 0.63 – 0.65. Posteroventral surface of midfemur not bearing a row of comb-like setae. Abdomen: Ventral margin of male tergite 5 not produced as a lobe. Male terminalia: Surstylus well separated apically with a small medial triangular process between surstyli comparable to structure in Paracoenia s. str. (triangular process in Paracoenia s. str. better developed in comparison with surstyli); aedeagus generally L-shaped, robust, apically tapered to acute point; phallapodeme in lateral view hemispherical; gonite narrow, with a small, shallow, subapical projection, apex curved anteriorly, point- ed. Female abdomen: Female ventral receptacle with small operculum, helmet-like, extended process three times longer than opercular height. Distribution. The only known species, C. paurosoma, has a disjunct Holarctic distribution, occurring in the western and temperate Nearctic Region (Alberta in Canada south through Wyoming and Colorado) but also in Finland and Sweden in the western Palearctic Region. We suggest that this distribution may be the result of sampling error, i. e., a lack of sampling of potential habitats between these vastly disjunct areas. Remarks. Leptocoenia, like the subgenus Calocoenia, is a monotypic subgenus, and it too would be an excellent candidate for molecular analysis to respond better to questions such as: (1) Are the two disjunct areas genetically as well as morphologically related? (2) Is the current status of the subgenus within Calocoenia validated?	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF97FF92226067D4AB89FB54.taxon	description	Figs 16 – 23	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF97FF92226067D4AB89FB54.taxon	diagnosis	Diagnosis. Cirrula is distinguished from other genera of Ephydrini by the following characters: Moderately large to large shore flies, body length 4.83 – 8.52 mm. Head: Cruciate, interfrontal setae 1 – 2 pairs, size generally subequal to fronto-orbital setae (weakly developed in C. gigantea, especially females); lateroclinate, fronto-orbital setae either 2 or 3 – 4 pairs, slightly divergent, if two, then dorsocentral setae (1 + 4), if 3 – 4, then dorsocentral setae (2 + 4); antenna simple, lacking secondary seta inserted on lateral surface just below arista; arista bare to macropubescent; face uniformly setose with marginal setae larger, declinate, one species with patches of long setae above middle height of face on anterior surface of interfoveal hump. Thorax: Prescutellar, acrostichal setae variable; dorsocentral setae 5 – 6 pairs (5 (1 + 4) in specimens with two fronto-orbital setae, 6 (2 + 4) in specimens with 3 – 4 fronto-orbital setae), well developed in Neotropical species; supra-alar seta present; presutural supra-alar seta variable; intrapostalar seta well developed. Costal vein ratio 0.18 – 0.19; M 1 vein ratio 0.95 – 1.10. Legs sexually dimorphic; hindtibia lacking apical, long seta; pulvilli greatly reduced or essentially absent; tarsal claws long and nearly straight. Abdomen: Male terminalia symmetrical; surstyli complex, situated at ventral apex of epandrium, covering other internal structures in repose, fused medially; phallapodeme more or less C-shaped with a dorsal lobe extended far into epandrial cavity; aedeagus generally simple except in males of C. austrina, where trilobate process arises at anterior base of aedeagus. Female ventral receptacle with large operculum, generally as high as wide; extended process J-shaped, length about as long as height of operculum; conformation of receptacle in females of D. spinosa exceptional, operculum trapezoidal and much smaller, extended process three times as long as operculum length. Third-instar larva: Prolegs distinct, better developed than those of Ephydra species; segment 3 of third-instar larvae with distinctive transverse band on venter. Natural History: Mathis and Simpson (1981) published a detailed report on the natural history of C. austrina (Coquillett). They successfully reared specimens from several localities along the coast of Virginia and noted that immatures and adults reached their highest densities where the habitat was partially dried, leaving algal mats on firm ground. Adults were collected commonly by sweeping over these mats. Unlike the larvae of Dimecoenia spinosa, those of C. austrina have eight pairs of well-developed prolegs and the third-instar larva also has a dark transverse strap at the anteroventral margin of segment three. Aldrich (1912) and Hutchinson (1937) provided excellent and detailed accounts of the biology of the so-called “ alkali-fly, ” C. hians (Say). The larvae are usually found on the bottom of the water in pools of water where they feed and do not come to the surface. The puparia attach to any available object. In storms, the puparia detach and form wind rows along the shore. Distribution. New World. Primarily the temperate western region of the Nearctic Region but with a southward extension into the northern Neotropical Region (Belize (Stann Creek District )). Remarks. Cirrula now comprises four species (C. austrina (Coquillett), C. currani (Wirth), C. gigantea Cresson, and C. hians (Say )) and is similar and closely related to the genus Dimecoenia.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF97FF8C20906386ACB4FA94.taxon	description	Figs 24 – 26	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF97FF8C20906386ACB4FA94.taxon	diagnosis	Diagnosis. Coenia is distinguished from other genera of Ephydrini by the following combination of characters: Moderately small shore flies, body length 2.20 – 2.80 mm. Head: Frons much wider than high, only moderately to sparsely microtomentose, especially subshiny mesofrons; no cruciate interfrontal setae; two well developed fronto-orbital setae; paravertical setae weakly developed, not subequal to vertical setae. Arista bearing long rays dorsally on basal 3 / 4, length of longest rays subequal to height of basal flagellomere. Face with lateral margins and oral margin bearing long setae, otherwise setulose, although medial area with setae shorter than marginal setae. Gena short, about half height of basal flagellomere; genal seta well developed. Thorax: Mesonotum generally dark colored, blackish brown; scutellum triangular, posterior angle rounded; no well-developed prescutellar acrostichal pair of setae; 4 (1 + 3) well-developed dorsocentral setae; postpronotal seta (e) either weak, at most 1 / 4 length of posterior notopleural seta, or lacking; 1 presutural supra-alar seta; 1 supra-alar seta; 1 postalar seta; prosternum bare; 2 scutellar setae; 2 notopleural setae; 1 large anepisternal seta; 1 well developed katepisternal seta. Costal vein ratio 0.21 – 0.22; M 1 vein ratio 0.66 – 0.69. Hindcoxa bare posteriorly; pulvilli well developed; tarsal claws short and distinctly curved. Abdomen: Male with five visible tergites dorsally, sternite 5 longer than wide, triangular. Male terminalia: Surstylar bases adjacent, separated by a narrow groove, or fused, apically separated as surstylar arms, arms pointed or digitiform, sometimes irregularly; gonite moderately well developed, elongate, regularly to irregularly tapered to acute apex; aedeagus elongate, moderately thin to very thin, shallowly to conspicuously curved, tapered from base to apex, apex acutely pointed; phallapodeme short to elongate, shallowly to conspicuously curved, thin to moderately thick medially, if thick tapered toward apices, distinct keel not evident. Female abdomen: Female ventral receptacle with small operculum, helmet-like, extended process three times longer than opercular height, C-shaped with short dorsal extension into operculum. Third-instar larva (based on Foote 1990): Small, body length 7.60 mm; elongate, nearly cylindrical, legless but with creeping welts; integument bearing slightly pigmented scales of similar shape, scales not forming patterns on dorsal surface; with circles of pigmented scales on thoracic segments, abdominal segments fully covered in scales; two types of sensilla: rayed and rosette-like (peg-like); pseudocephalon bilobed; antennae two-segmented; oral papillae comb-like; mouth brushes a long, fluffy comb at distal end; tips of oral hooks highly dentate; anterior spiracles with 4 – 6 finger-like projections; retractable respiratory tube branched at middle with two fleshy protrusions; anal opening smooth, transversely elongated. Natural history: Krivosheina (2001: 1367) wrote that larvae feed on detritus and that adults are found at sites with stagnant water during the summer. Foote (1990) suggested that there can be as many as 9 – 12 generations during the summer season. Distribution. The composite distribution for species of Coenia is Holarctic with six of seven species occurring in the Palearctic Region. Most species have relatively small distributions geographically with five species (C. caucasica Krivosheina, C. deserta Krivosheina, C. elbergi Dahl, C. palustris (Fallén), C. vulgata Krivosheina) being found thus far only in the Palearctic Region, and one species (C. alpina Mathis) being exclusive to the Nearctic Region. Only C. curvicauda (Meigen) is more widespread, having a Holarctic distribution. Most species occur at higher latitudes with only C. palustris ranging southward to the Azores and Canary Islands in the Old World. Remarks. The most recent revisions are Mathis (1975) for the Nearctic species and Krivosheina (2001) for the Palearctic species. Like some other genera of Ephydrini, specimens of Coenia are not commonly collected and are thus poorly represented in collections.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF89FF8D20F66247ACE9FD04.taxon	description	Figs 27 – 30	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF89FF8D20F66247ACE9FD04.taxon	description	Zatwarnicki 1995: 238 – 240 [world catalog]. Diagnosis. Dimecoenia is distinguished from other genera of the tribe Ephydrini by the following combination of characters: Moderately large to large shore flies, body length 4.25 – 6.15 mm; mostly dull, olivaceous brown to grayish brown, dorsum with some subshiny to shiny areas dorsally. Head: Frons with shiny, submetallic mesofrons, parafrons dark but not shiny. one pair of well-developed cruciate interfrontal setae; two well-developed lateroclinate fronto-orbital setae, slightly divergent; paravertical setae small; both medial and lateral vertical setae well developed. Basal flagellomere simple, lacking secondary seta inserted laterally just below arista; arista tapered gradually from thickened base to style-like apex, approximately basal 2 / 3 with dorsal, hair-like rays, thereafter bare, aristal rays nearly as long as width of pedicel. Facial hump poorly developed, shallowly projected, dorsal portion of hump with some shiny metallic coloration; ventral margin of antennal grooves nearly horizontal, not sloping ventrally at conspicuous angle; facial setae best developed along lateral and oral margins; one large genal seta. Gena relatively short, height about equal to height of basal flagellomere. Thorax: Chaetotaxy: Acrostichal area with setulae only, no well-developed prescutellar pair; 5 (1 + 4) dorsocentral setae, posterior seta displaced laterally; 1 presutural supra-alar seta; 1 supra-alar seta; postpronotum with 1 large seta and a smaller seta; 1 postalar seta; 2 lateral scutellar setae; prosternal setulae sparse on at least posterior portion; anepisternum with 1 long seta; katepisternum with 1 long, dorsoclinate seta. Wing generally hyaline; dorsal costagial seta subequal in length to anteroventral costagial seta; costa with numerous, conspicuous, spine-like setulae; legs of both sexes similar; costal vein ratio 0.24 – 0.26; M 1 vein ratio 0.81 – 0.85. Tarsal claws long and nearly straight; pulvilli greatly reduced or essentially absent. Abdomen: Generally subshiny; anterior portion of each tergite dark brown posterior portion lighter, grayish green; male tergite 5 as long as wide, longer than tergite 4. Male terminalia: Epandrium more or less oval in posterior view, anteroventral margin evenly rounded; surstyli with large medial flange and posterolateral, slender processes; gonite 4 X longer than wide, anteroventral margin broadly and shallowly U-shaped; phallapodeme with posteromedial broad keel; aedeagus a simple tube, mostly parallel-sided. Female terminalia: Female ventral receptacle with operculum much smaller than extended process, trapezoidal; extended process broadly curved, widest medially. Third-instar larva: Larvae lacking well-developed prolegs on segments other than segment 12, only a short, bump-like process evident. Natural history: Larvae of Dimecoenia represent an apparent reversal in the generalized adaptive scheme of Ephydrini by inhabiting mud substrates associated with salt marshes. This has apparently resulted in the atrophy of the prominent, ventral prolegs, including the crochets, which are functionally adapted to movement within algal mats. Distribution. The composite distribution for the two species now included in Dimecoenia, D. fuscifemur Steyskal and D. spinosa (Loew), is temperate North America with specimens of D. spinosa occurring southward to Costa Rica and on some islands of the West Indies. Remarks. As characterized here, Dimecoenia now includes just two species, and these were most recently revised by Mathis and Simpson (1981). The monophyly of this clade is confirmed by the following synapomorphies: 1. The anterior margin bearing conspicuous, spine-like setae; 2. Ventral margin of antennal facial groves rounded, nearly horizontal and not steeply angled.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF88FF8E20DF65EEA920FB96.taxon	diagnosis	Diagnosis. Ephydra is distinguished from other genera of Ephydrini by the following characters: Head: Lateroclinate, fronto-orbital setae 3, well developed, subequal; development of cruciate, interfrontal setae variable, either with one well-developed pair or weak to lacking; basal flagellomere simple, lacking secondary seta inserted below arista on lateral surface; arista variable, subpectinate to macropubescent, if subpectinate, basal thickening extended about 1 / 3 of aristal length, if macropubescent, basal thickening extended over 1 / 2 of aristal length; antennal groove distinct but not deeply impressed. Thorax: 5 (1 + 4) well-developed; dorsocentral setae; presutural supra-alar seta present; intrapostalar seta present; supra-alar seta present; disc of scutellum generally concolorous with posterior portion of scutum. Costal vein ratio 0.24 – 0.40; M 1 vein ratio 0.61 – 0.81. Tarsal claws long, although length variable, and nearly straight; pulvilli greatly reduced or absent. Abdomen: Structures of male terminalia considerably modified depending on subgenus and species group (see appropriate diagnosis of subtaxon for further details). Female terminalia: Female ventral receptacle with operculum small, trapezoidal in shape; extended process relatively large, C-shaped, length 2 – 3 times width of operculum. Distribution. This speciose genus occurs primarily in temperate regions of the world as follows: New World. Widespread mostly in the Nearctic Region but extended into the northern Neotropical Region (12 ° – 65 ° N): Canada (just south of the Great Bear Lake), southward into Mexico (Oaxaca) and the West Indies (Dutch West Indies). Old World. Widespread, mostly in the Palearctic and Afrotropical (temperate) regions but extended into the northern Oriental Region: Norway to Japan, southward to the Canary Islands, South Africa, across southern Asia (Afghanistan, Iran, and Tibet) to Japan and China. Remarks. Wirth (1971, 1975) published excellent revisions of Ephydra on a worldwide basis, and his two papers should be consulted for a more detailed discussion of the natural history and for identification of extralimital species. Our treatment is essentially a synopsis of Wirth’s valuable studies, although with some modifications. Krivosheina and Ozerov (2021) recently produced an excellent review of the Russian fauna. Ephydra is the most speciose genus in the tribe Ephydrini with 34 species, and these are classified into two subgenera, Ephydra and Halephydra. The subgenus Ephydra is further divided into species groups. The subgenera and species groups are characterized and identified in the following key.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8AFF8823C46109AD7DFA5F.taxon	description	Figs 31 – 33	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8AFF8823C46109AD7DFA5F.taxon	description	Zatwarnicki 1995: 247 – 248 [world catalog].	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8AFF8823C46109AD7DFA5F.taxon	diagnosis	Diagnosis. Ephydrella is distinguished from other genera of the tribe Ephydrini by the following combination of characters: moderately large shore flies, body length 3.0 – 6.0 mm. Head: Frons with shiny, metallic mesofrons, parafrons relatively dull, appearing somewhat “ membranous ”; two well-developed, lateroclinate fronto-orbital setae; cruciate interfrontal setae lacking; paravertical setae not evident; medial and lateral vertical setae well developed. Basal flagellomere lacking a lateral seta; arista long, almost double length of basal flagellomere, basal half somewhat swollen; dorsal aristal branches not evident, at most arista appearing macropubescent. Face with larger setae at lateral and oral margins; 1 large genal seta. Thorax: Mesonotum dark colored, brown to metallic dark green or blue, similar to frons, with longitudinal light gray stripes in area of acrostichal tracks, scutellum triangular, posterior angle rounded, relatively acutely angulate, or narrowly truncate. Chaetotaxy: 1 well-developed pair of prescutellar acrostichal setae, otherwise as setulae; 4 (1 + 3) or 5 (1 + 4) well-developed dorsocentral setae, sometimes these appreciably reduced, posterior seta displaced laterally; 1 postpronotal seta; 1 presutural supra-alar seta; 1 supra-alar seta; 1 postalar seta; 2 lateral scutellar setae; prosternal setulae sparse on at least posterior portion; anepisternum with 1 long seta; katepisternum with 1 long, dorsoclinate seta. Wing generally hyaline; R stem vein bare above; costal vein ratio 0.23 – 0.25; M 1 vein ratio 0.68 – 0.76. Male hindfemur not differing markedly from fore- or midfemur, lacking stout setae; male hindtibia lacking tuft of setulae; male hindtarsi cylindrical, normal; pulvilli much reduced or absent; tarsal claws long and nearly straight. Abdomen: Male with five visible tergites, female with 6 – 7; coloration similar to that of notopleuron, often submetallic but with varying levels of microtomentum. Male terminalia symmetrical, epandrium longer than wide, bearing prominent surstyli, these often elongate, with medial, often small triangular structure between surstyli, with medial fissure; aedeagus in lateral view cylindrical, moderately slender, base with more sclerotized, distinctly curved process; phallapodeme in lateral view elongate, keel distinct, either apices rod-like, slender; gonite sheathing aedeagus, usually longer than wide, variously shaped, but usually acutely pointed apically; subepandrium narrow band than arches over base of aedeagus; hypandrium shallowly v-shaped, angle very obtuse. Female terminalia: Female ventral receptacle with small papilla-like operculum; egg guide sternites bearing two pairs of strong setae. Natural history: Bock (1987) observed that larvae of some Australian species are associated with marine algae of the genus Cladophora. Distribution. Ephydrella is thus far only found on Australia and New Zealand. Remarks. Ten species have been described thus far, all from Australia and New Zealand, and at least one species from New Zealand has not been described. The shape of the surstylus and its relative length compared to the length of the medioventral process are used extensively to identify species of Ephydrella.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8DFF8920ED62BDAC5DFD22.taxon	description	Figs 34 – 36	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8DFF8920ED62BDAC5DFD22.taxon	diagnosis	Diagnosis. Moderatelylargeshoreflies, bodylength 4.0 – 5.5 mm, generally appearing dull, densely microtomentose, grayish. Head: Frons dull, gray, brown or velvety black, uniform, some species with silvery white spots; 2 ocellar setae; 3 lateroclinate fronto-orbital setae; lacking cruciate interfrontal setae; postvertical setulae weak; lateral and medial vertical setae present. Antenna black, pedicel sometimes brownish; arista glabrous, base broad, generally black to yellowish brown. Face projected anteriorly, arched, face and gena silvery white to dark gray; face with one strong seta and several setulae; gena with 3 – 5, dorsally curved setae; facial setae aligned with parafacial suture, setae lacking along ventral martin. Clypeus prominent, clearly projected ventrally beyond ventral oral margin; oral opening large, gaping. Thorax: Mesonotum gray, brown, or velvety black, unicolorous or with stripes and spots. Five dorsocentral setae (2 + 3); two rows of acrostichal setae, prescutellar pair larger; 1 postpronotal seta; 1 presutural supra-alar seta; 2 supra-alar setae; lateral scutellar setae not arising from tubercles; shape of scutellum variable; katepisternum and anepisternum appearing pubescent, 1 – 4 katepisternal setae; 0 – 1 anepisternal seta. Wing mostly hyaline, with sharp separation of subcostal, proximity of middle cubital vein, disc shape at rear edge of wing; lacking a spur vein; halter faintly yellowish; costal vein ratio 0.21 – 0.23; M 1 vein ratio 0.72 – 0.74. Legs gray, apices of tibia and tarsi reddish brown; pulvilli present but rudimentary; tarsal claws relatively long and straight. Abdomen: Gray to greenish, microtomentose, numerous short setulae. Male terminalia: Epandrium shield-like, ovate to ellipsoidal, with ventromedial fissure; cerci comparatively elongate, length about equal to paired ventral extensions of epandrium; gonite in lateral view nearly straight, apex variously shaped, apex arrow-like, pointed or excavated subapically; aedeagus in lateral view tubular, parallel sided to tapered, curved or almost L-shaped. Female terminalia: Female ventral receptacle with operculum tube-like, much smaller than extended process, trapezoidal; extended process broadly curved, widest medially, with neck that extends into operculum. egg guide sternites bearing two pairs of weak setae. Third-instar larva: Prolegs along venter of abdominal segments. Natural history: Adults and immature stages are associated with saline habitats, usually shorelines. Sometimes the salinity is very high. Distribution. Thus far, species of Halmopota are found exclusively in the Palearctic Region and can be conveniently summarized by Turkey (H. anatolicus Canzoneri and Meneghini, H. tomentosus Canzoneri and Meneghini) and Tibet (H. chinensis Krivosheina, H. hutchinsoni Cresson, H. kozlovi Becker, H. villosa Becker) plus the following: H. mediterraneus Loew (Algeria, Egypt, Iraq, Iran, Morocco, Spain, Syria, Tibet, Turkey), H. insignis (Becker) (Russia, Ukraine), H. salinarius (Bouché) (Germany, Poland, Russia), H. septentrionalis Canzoneri and Meneghini (Bulgaria, Italy, Tadzhikistan, Turkey, Turkmenistan), and H. stackelbergi Krivosheina (Tadzhikistan). Remarks. While having sympatric species distributions is not unusual in Ephydridae, this condition for at least some species of Halmopota may suggest the need to again revise the included species. To that end, we have provided more detailed distributional data for all described congeners (see above). Although the generic name, Halmopota, ends with the letter a, it is a masculine noun, and the species’ epithets should agree in gender.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8CFF8B20F46408ADEBFE10.taxon	description	Figs 37 – 51	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8CFF8B20F46408ADEBFE10.taxon	diagnosis	Diagnosis. Neoephydra is distinguished from other genera of Ephydrini by the following characters: medium-sized to large shore flies, body length 3.00 – 5.30 mm. Head: Mesofrons with vestiture variable; lacking cruciate, interfrontal setae; lateroclinate, fronto-orbital setae either 2 or 5 – 6, not 3; basal flagellomere lacking large seta inserted on lateral surface; arista moderately short, thickened basally, with macropubescent vestiture dorsally, apical half style-like, bare; postocular setae variable; large facial setae declinate; gena moderately high to high, gena-to-eye ratio 0.30 or larger. Thorax: Chaetotaxy: Acrostichal setae, including a prescutellar pair, not well-developed; dorsocentral setae 5 (1 + 4), development variable; supra-alar seta variable; presutural supra-alar seta lacking; intrapostalar seta present, although sometimes weak. Wing generally hyaline; costal vein ratio 0.19 – 0.36; M 1 vein ratio 0.64 – 0.86. Hindtibia lacking apical seta; tarsal claws nearly straight; pulvilli essentially absent. Abdomen: Male terminalia symmetrical, epandrium longer than wide; surstyli fused medially except near apices and with 1 – 2 lateral projected processes or prongs in addition to apical prominences; aedeagus in lateral view shallowly crescent-shaped and generally quite slender, at least apically. Female terminalia: Female ventral receptacle with small papilla-like operculum. Natural History: Like many taxa of the subfamily Ephydrinae, specimens of Neoephydra inhabit diverse and what would appear to be environments inimical to life. Oliveira (1954 a) noted that Dr. Herman Lent found larvae, pupae, and adults of a Chilean species in the hot effluent of a high altitude, hot water geyser located at El Tatio (5200 m), near San Pedro de Atacama. Although the temperature of the water was not taken, Dr. Lent stated that it was sufficiently hot to cook an egg. Dr. Lent also observed a small, predatory toad, Telmatobius peruvianus Wiegmann (Anura: Leptodactylidae), whose diet consisted solely of freshly emerged, adult flies. Numerous larvae and pupae of a second species, collected in southern Brazil, were found to inhabit warm, algae-covered, and often saline water that had accumulated in depressions of large rocks near the seashore (Oliveira 1954 b, 1958). Water evaporation from these shallow depressions is rapid, accounting for the concentration of salts. Hennig (1943) and Oliveira (1954 b, 1958) described and illustrated the larvae of four species belonging to this genus. Based on these figures, larvae of Neoephydra are typical of the tribe, with eight pairs of claw-bearing prolegs on the ventral surface, the terminal pair being larger and with crochets opposable to those of the other prolegs. The posterior spiracles are borne on a long respiratory tube which bifurcates posteriorly. Distribution. Members of Neoephydra are only known from the Neotropics and some island in the south Atlantic Ocean, where they are widespread and occur in habitats similar to those of the Holarctic genera Ephydra Fallén and Setacera Cresson. Remarks. Neoephydra is the generic name for most South American species that had been placed in the genus “ Dimecoenia. ” As noted by Steyskal (1970) and Wirth (1971), the Neotropical species, which were treated as congeners of Dimecoenia (Wirth 1968), are structurally dissimilar from the Nearctic species. We came to the same conclusion as Steyskal and Wirth after studying structures of the male terminalia. To identify species, particular attention should be given to structures of the male terminalia and female ventral receptacle. Three species groups are also recognized (the araucaria, dasycephala, and neotropica groups). These groups were proposed for specimens that are dissimilar superficially from the typical Neoephydra (araucaria Group) but which have structures of the male and / or female terminalia that closely resemble those of other similar aggregates. Mathis and Marinoni (2016) chose the informal category of “ species group ” for these groups because of its flexibility, without encumbering additional, relatively fixed nomenclature, such as would be required for the subgeneric category. The monophyly of Neoephydra is established by the following characters: (1) Conformation of the male terminalia: The surstyli are fused medially except near their apices and each surstylus bears one or two additional, anterolateral prongs. This conformation is unique within the subfamily Ephydrinae. (2) Conformation of female ventral receptacle: All species groups have a small, papilla-like operculum, which is also apparently unique to females of Neoephydra. It is likely that the species groups of Neoephydra are monophyletic, although the araucaria Group lacks characterization by derived character states. The monophyly of the remaining groups is well established, as evidenced by the ap - propriate characterization heading each group’s treatment.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8EFF8520F46279AC7CFB31.taxon	description	Figs 52 – 66	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF8EFF8520F46279AC7CFB31.taxon	diagnosis	Diagnosis. Specimens of Notiocoenia are similar to those of Coenia but are distinguished from the latter and other genera of Ephydrini by the following combination of characters: Small to moderately large shore flies, body length 1.90 – 4.20 mm; mostly grayish brown to brown, generally appearing dull but subshiny to shiny over much of dorsum. Head: Wider than high from anterior view; frons mostly flat, wider than long, vestiture more or less uniform, dull, microtomentose, mostly unicolorous brown, margins of mesofrons indicated by shallowly impressed furrow, otherwise undifferentiated from parafrons, lacking large setae, cruciate or otherwise; well-developed, lateroclinate, fronto-orbital setae 2, plus 1 much smaller proclinate seta anterior of larger setae; paravertical seta small, inconspicuous, none 1 / 2 length of vertical seta; well-developed, ocellar setae 1, slightly divergent; small postocellar setae 2 – 3 setulae; medial vertical setae well developed, lateral vertical seta either well developed or lacking. Antenna normally developed, dark colored, basal flagellomere considerably longer than high, rounded apically; pedicel bearing several setae, especially on medial and ventral surfaces, bearing one large, dorsally erect seta on dorsum; arista longer than combined length of other antennal segments, macropubescent or bearing short hairs dorsally. Face protuberant, broadly transversely arched, setulose to setose; interfoveal hump evident but variously developed; vestiture densely microtomentose. Eye subspherical, slightly higher than wide. Gena short, gena-to-eye ratio 0.20 or less; well-developed, genal seta 1. Mouthparts generally retracted into oral opening, clypeus not exposed; maxillary palpus dark; prementum longer than wide; microtomentose to microtomentose, setose. Thorax: Vestiture variable, dull to shiny, generally becoming shinier posteriorly; coloration grayish brown to blackish brown. Pleural areas generally becoming paler colored ventrally; humeral callus, part of anepisternum, and katepisternum grayish tan to whitish tan, otherwise pleural areas darker brown; forecoxa gray to silvery gray, contrasting with remainder of pleural areas. Chaetotaxy as follows: Acrostichal setae when present arranged in two rows, larger setae mostly anterior of transverse suture; lacking well-developed, prescutellar seta; well-developed, dorsocentral setae 4 (1 + 3), rarely 5 (2 + 3), anterior 1 – 2 setae smaller, several smaller setae anterior of larger setae; interalar setae 2, anterior seta inserted just posterior of transverse suture, posterior seta posterior of level of posteriormost dorsocentral seta; postpronotum either bare or bearing 2 – 3 setulae, none more than 1 / 4 length of posterior, notopleural seta; presutural interalar seta 1; interalar setae 2, rarely 3, anterior seta inserted just posterior of transverse suture, posterior seta smaller, inserted just posterior of posteriormost dorsocentral seta; postsutural, supra-alar seta variable; postalar seta well developed; disc of scutellum mostly bare, bearing 2 - 3 setulae, or bare; lateral scutellar setae 2; notopleural setae 2, inserted near each ventral angle; anepisternal seta 1, inserted near middle of posterior edge, numerous smaller setae, particularly toward dorsal and posterior margins; prosternum bare; katepisternal seta 1, with 1 – 2 smaller setae around larger seta; midcoxa with 1 larger seta. Wing hyaline to palely infuscate; R stem vein bare dorsally; halter yellowish to pale brown; costal vein ratio 0.13 – 0.16; M 1 vein ratio 0.55 – 0.68. Legs variable, femora swollen to slender; coloration rufous to black; pulvilli well developed; tarsal claws short and distinctly curved; pulvilli well developed; tarsal claws short and distinctly curved; hindfemur of male not differing markedly from fore- or midfemur, lacking stout setae as above; hindtibia of male lacking tuft of setulae; hindtarsi of male cylindrical, normal. Abdomen: Dorsum either darker brown, with some bluish coloration or blackish, subshiny to shiny; tergite 5 of variable length compared to tergite 4. Male terminalia: Surstylus either fused indistinguishably with ventral margin of epandrium (paniculata Group) or fused but distinct and setose (pollinosa Group); aedeagus either a long, slender, tapered process, apex slightly coniform or produced ventrally as two symmetrical narrow processes. Female terminalia: Female ventral receptacle with tiny, flat operculum and a very large, roughly C-shaped extended process in lateral view; females sternites square to rectangular, relatively wide (Costa et al. 2024). Distribution. The composite distribution for species of this genus is Neotropical and extends from 24 ° – 53 ° S along the east and west slopes of the Andes Mountains. Remarks. Although the monophyly of Notiocoenia appears to be well established, its relationship to other taxa within Ephydrini is not as evident. Although we suggest a sister group relationship with Coenia, as indicated in the cladogram, the evidence to support this alliance is not wholly convincing. See also remarks under Austrocoenia. Following the precedent of Mathis and Marinoni (2016), we recognize two species groups, which are identified in the following key.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF83FF86227967D4AD49F981.taxon	description	Figs 67 – 74	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF83FF86227967D4AD49F981.taxon	diagnosis	Diagnosis. Paracoenia is distinguished from other genera of the tribe Ephydrini by the following combination of characters: Small to large shore flies, body length 2.10 – 4.40 mm; dark colored and often with subshiny, metallic reflections. Head: Face projected, transversely arched; paravertical setae large, subequal to vertical setae. Two well-developed lateroclinate fronto-orbital setae. Thorax: Postpronotal seta distinct, length at least 1 / 2 as long as posterior notopleural seta; dorsocentral setae 5 (1 + 4); scutellum with dorsum convex; prosternum bare. R stem vein bearing 1 – 2 setulae above, inserted beyond transverse septum; costal vein ratio 0.19 – 0.27; M 1 vein ratio 0.81 – 0.92. Hindcoxa with row of setae posteriorly along ventral margin; tarsal claws short and distinctly curved, comparatively short; pulvilli well developed. Abdomen: Male terminalia: Surstyli distinct as elongate, narrow, arm-like projections, projections oriented ventrally; a medial, triangular process between surstylar arms; gonite (sometimes called hypandrial process) well developed, sheathing aedeagus. Third-Instar larva: Body length 9.50 – 18.80 mm; body fusiform, integument translucid; 12 segmented; 3 - segmented antenna; maxillary (terminal) sensory organs (consisting of seven sensillae, separated by flattened ridges in 2 groups, 2 + 5); 3 legless thoracic segments, 1 st thoracic segment with two anterior spiracles; eight legless abdominal segments; caudal segment with anus ventrally, elongated as a long retractile, branched respiratory tube, tube length 30 – 50 % body length. Natural history: A few species of this genus occur in the effluent of hot springs, and some are uniquely found only at these sites. Moreover, some of the springs are sulfurous and otherwise inimical to most other forms of life. These species are as follows: P. calida Mathis from Wilbur Hot Springs (39 ° 2.3 ’ N, 122 ° 25.3 ’ W) in Colusa County, California; P. quatei (Wirth) from Sulphur Mountain Spring (34 ° 25.7 ’ N, 119 ° 05.6 ’ W) in Ventura County, California; P. beckeri (Kuntze) from Acque Albule (41 ° 54.2 ’ N, 12 ° 29.8 ’ E), a sulfurous spring near Tivoli, Roma, Italy. Adults and immature stages of these three species were collected only at these respective and specific sites despite efforts to sample them from nearby habitats and elsewhere (Wirth 1954 for P. quatei; Giordani Soika 1956 a for P. beckeri; and Mathis 1975 for P. calida). Wirth (1954) described the 3 rd- instar larva of P. quatei, which has small pseudopods along the ventral surface of segments, and these lack the long crochet-like setae, found more typically in larvae of Ephydra (Aldrich 1912). A fourth possible species that may occur in hot springs is P. ampla Mathis, which is only known from Los Angeles, Los Angeles County, California. M. C. VanDuzee collected the holotype male in April of 1915. We wonder if the specific site were a hot spring, such as Alvarado Hot Springs, and if more specimens could be found there or at another hot springs in the area. These species were perhaps preadapted to live in association with hot springs, as other congeners, such as P. wirthi Mathis (Tecopa Hot Springs) and P. turbida (Curran) (Yellowstone hot springs) have been collected in and on the effluent of some hot springs in addition to occurring in cold-water wetlands. The “ preadaption ” may involve the algal food source that lives in these hot springs and other aquatic sources. Remarks. The lineage comprising Paracoenia and related genera in our data and analysis the sister group to the remaining taxa of Ephydrini. This lineage plus the remaining taxa of Ephydrini, as here delimited, is characterized by the following character states (some have become modified secondarily): 1. Number of dorsocentral setae: Although other genera of the subfamily Ephydrinae sometimes have five pairs of dorsocentral setae (e. g. Notiocoenia Mathis and Austrocoenia Wirth), the anterior pair (or pairs) is weakly developed. There are five well-developed pairs only in members of Ephydrini (the anterior pair is presutural; specimens of Cirrula gigantea have the anterior four pairs of dorsocentral setae are weakly developed, a condition we interpret to be secondary). 2. Development of intrapostalar seta: In most species of the family, the intrapostalar seta is either lacking or is very much reduced, less than one-half the length of the postalar seta. In members of this lineage, the intrapostalar seta is frequently as long. 3. Setal vestiture of proepisternum: Throughout most of the family the proepisternum is bare of setae (although frequently it is thinly to densely microtomentose). In members of this lineage, there are numerous setulae that are generally conspicuously evident.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF84FF81220D6649AC18FD9D.taxon	description	Figs 75 – 81	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF84FF81220D6649AC18FD9D.taxon	diagnosis	Diagnosis. Paraephydra is distinguished from other genera of Ephydrini by the following combination of characters: Moderately small to medium-sized shore flies, body length 2.40 – 3.80 mm; setation normally developed, not generally appearing pilose. Head: Mesofrons shiny, with metallic luster, differentiated from microtomentose parafrons; cruciate interfrontal setae l; two well-developed lateroclinate setae antennal groove distinct but not deeply impressed; basal flagellomere lacking large seta inserted on lateral surface; arista as long or slightly longer than combined length of first 3 antennal segments, gradually tapered from base to apex, with subpectinate, dorsally branching rays on basal 2 / 3; postocular setae normally developed, not conspicuous; larger facial setae extended from interfoveal hump with 1 – 2, distinctly porrect to anaclinate. Thorax: Females with one prescutellar, acrostichal setae; well-developed dorsocentral setae 4 (1 + 3); supra-alar seta present; presutural supra-alar seta lacking; intrapostalar seta either weakly developed or lacking; disc of scutellum concolorous with posterior portion of scutum; females lacking dense patch of setae between posterior 2 dorsocentral setae. Costal vein ratio 0.23 – 0.43; M 1 vein ratio 0.72 – 0.79. Hindtibia with apical, anteroventral seta, length equal to or larger than width of tibia at widest point. Abdomen: Male terminalia: symmetrical; epandrium longer than wide, narrowed ventrally, fused almost imperceptibly with base of united surstyli; surstyli fused medially except at near apex; posterior surstylar process only slightly longer than lateral process; both processes apical; gonite, hypandrium, and apparently aedeagus fused to form one compact structure, curved anteriorly, wide basally, tapered to rounded apex. Female terminalia: Female ventral receptacle with operculum flat, disc-like. Natural History: Like other ephydrines, Paraephydra occurs in wetlands. In southern Chile (Osorno Province), we collected specimens of P. stauros in a sedge meadow near the margins of small but apparently permanent ponds. Nothing is known about the immature stages or the microhabitat of the genus. Distribution. Neotropical; widespread but uncommon, from Puerto Rico south through Brazil to Chile. None of the congeners is known to be sympatric. Remarks. Paraephydra was proposed to accommodate two closely related species, P. freitasi (Oliveira) and P. stauros Mathis. Sexual dimorphism is evident in the chaetotaxy of Paraephydra. Females, unlike males, have a prescutellar acrostichal seta that is larger than other acrostichal setulae. Based on this character, Oliveira (1954 c) described P. freitasi in the genus Ephydrella, as that genus, unlike species of Neoephydra, lacks these setae.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF84FF8320E2657FAA4FF92D.taxon	description	Figs 82 – 86	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF84FF8320E2657FAA4FF92D.taxon	diagnosis	Diagnosis. Setacera is distinguished from other genera of Ephydrini by the following characters: Moderately small to large shore flies, body length 2.46 – 5.85 mm. Head: Mesofrons shiny, with metallic luster; cruciate, interfrontal setae lacking or weakly developed; lateroclinate, fronto-orbital setae 2 pairs; fronto-orbits shiny with metallic luster concolorous with mesofrons; basal flagellomere with prominent seta on lateral surface below aristal insertion; arista with subpectinate branching along dorsal surface from between onehalf to 2 / 3 of aristal length; dorsum of interfoveal facial ridge sloping very gradually; ridge projecting markedly forward in many species attaining broad apex from which arched face extends ventrally at nearly a right angle, face receding to oral margin in other species; antennal groove distinct but not deeply impressed; postocular setae normally developed, not conspicuous; larger facial setae declinate. Thorax: Dorsocentral setae 5 (1 + 4); presutural supra-alar seta 1, generally subequal to posterior notopleural setae in species from Western Hemisphere. Costal vein ratio 0.28 – 0.30; M 1 vein ratio 0.84 – 0.90. Tarsal claws shallowly curved, nearly straight, comparatively long; pulvilli lacking or greatly reduced. Abdomen: Structures of male terminalia symmetrical but unusually complicated by addition of several secondary processes and prongs; epandrium elongate; well-developed surstyli generally fused medially, projecting from ventral margin of epandrium. Female terminalia: Female ventral receptacle with operculum as high as wide, broadly rounded dorsally; extended process more or less J-shaped. Natural History: The immature stages of Setacera and Ephydra closely resemble each other, and Johannsen (1935) considered them to be the most highly specialized of the family. Like larvae of Ephydra, those of Setacera are characterized by long, terminal respiratory tubes and by eight pairs of short, conical, abdominal prolegs, of which the last pair is the largest, with claws opposable to those of the other prolegs. Johannsen (1935) published a figure of the cephalopharyngeal skeleton of S. needhami, a species described inadvertently from the immature stages (Cresson 1935), and Foote (1982) described and illustrated the immature stages of S. atrovirens. Unlike Ephydra, specimens of Setacera occur primarily in freshwater habitats, although Johannsen (1935) reared a specimen of S. atrovirens from a puparium collected in a brine pool near Ithaca, New York. Where members of Setacera do occur, even within what appears to be their preferred habitat, specimens are not collected frequently, and collection of a good series usually requires diligent persistence. Most species seem to prefer lentic aquatic systems, especially where a layer of floating algae has accumulated on the water’s surface. This is the typical habitat of most species of Ephydrini, and their crochetbearing prolegs are apparently an adaptation to this habitat, allowing movement through and attachment to the algae. Distribution. Among the genera of Ephydrini, Setacera is by far the most widely occurring, genus with species being found in all major faunal realms. The Neotropics, however, have a depauperate fauna. Remarks. Some members of Setacera exhibit sexually dimorphic features that are probably secondary. Males of these species bear prominent hair-tufts of varying lengths at tibial apices and often on the coxae. The extent and length of tufts, or their absence, are excellent species-level characters. Some species now included in Setacera were previously placed in the genus Ephydra Fallén, and some recent authors still prefer the precedent of Setacera as an included subgenus of Ephydra (Giordani Soika 1956 b, Dahl 1959). Setacera is indeed closely related to Ephydra, as evidenced by the similarity of adults and immatures of both genera. Setacera, however, can be consistently distinguished in both sexes from all other genera of Ephydrini and its monophyly corroborated by the following synapomorphies: (1) Basal flagellomere seta: Aside from the arista, there are usually no other large structures emanating from the basal flagellomere. Specimens of Setacera, however, have a large seta inserted just below the aristal insertion on the lateral surface. (2) Vertico-orbits: Within the tribe Ephydrini, the vertico-orbits are generally either shiny or densely microtomentose and grayish, appearing dull. This area, in specimens of Setacera, is uniquely invested with a dense patch of microtomentum that appears velvety. Velvety areas occur elsewhere in a few species of the tribe (parafrons in Cirrula gigantea Cresson; frons and orbits in Ephydra auripes Aldrich) but not in the specific area as described for Setacera. (3) Genal seta: This seta is usually very prominent, arising below the eye. Although this seta is still larger than surrounding ones in specimens of Setacera, its comparative size is smaller, and for convenience, we have compared it with the length of the arista. (4) Cruciate interfrontal setae: Although some species of the tribe Ephydrini do not have these setae, most genera have at least a few species in which they occur. Consequently, our interpretation of the general groundplan of the tribe is for their presence, and their absence in Setacera is apparently unique, i. e. a synapomorphy. (5) Prescutellar acrostichal setae: As with the preceding characters, these setae are generally present in Ephydrini. We know of no specimens of Setacera, however, where they are present, and we interpret this apparent loss to be synapomorphic. Mathis (1982 b) recognized five monophyletic species groups that are relatively finely divided (for example, the aurata and trina groups could be combined) and are characterized as follows. The aldrichi Group is the sister group of the pacifica group, and it is characterized and its monophyly established by: (1) Configuration of aedeagus: As before, the aedeagus is typically broadly rounded apically and almost as wide as long. Males of the aldrichi Group have a somewhat pointed aedeagus that we interpret to be apomorphic. (2) Configuration of epandrium: Males of the aldrichi Group have an anteroventral, digitiform process, apparently a unique condition, and one that we interpret to be apomorphic.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
0D42878FFF84FF8320E2657FAA4FF92D.taxon	description	The aurata Group (S. aurata (Stenhammar )) is the sister group of the trina Group, as evidenced by the distance between the cerci and setae on sternite 9. This distance is generally not greater than the height of the cerci. In females of this species group, the setae on sternite 9 re inserted farther ventrad, making the distance between them and the cerci conspicuously loner than the cercal height. Length of female sternite 8. The length is three to four times its width, not the more generalized five or more times. Length of female tergite 8. Tergite 8 in females is very long and partially accounts for the ventral position of sternite 9. This longer dimension of tergite 8 is a synapomorphy for this lineage. The aurata Group is distinguished from the trina Group by the shape of the epandrium. Although it is not uncommon for the epandrium to have appendages of various shapes, this is the only lineage to have a bluntly rounded, parallel-sided, posterolateral process arising from each side. The breviventris Group (S. breviventris (Loew), S. multicolor (Soika), S. viridis Miyagi) occurs in the Old World and is characterized and its monophyly established by the following synapomorphy: Development of the presutural supra-alar seta weak. In most Ephydrini, this seta is well developed, usually as long as the presutural seta. In species of this group, however, this seta is weak and is considerably smaller than the presutural seta. The micans Group (S. atrovirens (Loew), S. micans (Haliday )). This species group has a Holarctic distribution. The group is characterized and it monophyly established by the following synapomorphies: (1) Configuration of aedeagus. The aedeagus among most ephydrines is nearly as wide as long, and its apex is broadly rounded. In males of the micans Group the aedeagus is three to four times longer than wide, and its apex is acutely pointed. (2) Shape of female ventral receptacle. Usually, the operculum is nearly as high as the extended process, sometimes more so, and its width is subequal to its height. In females of the micans Group, the operculum is relatively small, both its width and height, especially compared to the size of the extended process. The monophyly of the pacifica Group (S. durani Cresson, S. jamesi Mathis, S. needhami Johannsen, S. pacifica (Cresson), S. pilicornis (Coquillett), S. trichoselis Mathis) is established by: (1) Configuration of vertico-orbits: In Setacera this band is more or less broad and usually has a subanterior swelling. But in members of the pacifica Group this band is very narrow and is sometimes difficult to detect. The narrowed aspect of this character is interpreted to be a synapomorphy. (2) Configuration of female ventral receptacle: For most ephydrines, the operculum is typically wider than high. For females of the pacifica Group, however, the height is subequal to its width, an apomorphic character. The trina Group (S. freidbergi Mathis, S. meneghinii Canzoneri, S. trina Collin). See our comments under the aurata Group for character evidence that these two group a closely related. The monophyly of the trina Group is established by the shape of the male gonite, which is unique for this group. A second character is the shape of the epandrium, which is similar in the species of this group, having the ventrolateral angles explanate and slightly recurved.	en	Mathis, Wayne N., Sepúlveda, Luciane Marinoni and Tatiana A. (2025): Phylogeny and taxonomy of the shore-fly tribe Ephydrini with comments on related tribes in Ephydrinae (Diptera: Ephydridae). Zoologia (e 24044) 42: 1-42, DOI: 10.1590/S1984-4689.v42.e24044, URL: https://doi.org/10.1590/s1984-4689.v42.e24044
