The Empis (Coptophlebia) hyalea-group from Thailand, with a discussion of the worldwide distribution of this species group (Diptera: Empididae: Empidinae)

The Empis (Coptophlebia) hyalea-group is especially diversified in the Oriental region and is here partly reviewed. Twelve new species from Thailand are described and keyed, namely Empis (Coptophlebia) atratata sp. n., E. (C.) kosametensis sp. n., E. (C.) lamruensis sp. n., E. (C.) miranda sp. n., E. (C.) nahaeoensis sp. n., E. (C.) nganga sp. n., E. (C.)pakensis sp. n., E. (C.) pseudospinotibialis sp. n., E. (C.) pulchra sp. n., E. (C.) ratburiensis sp. n., E. (C.) spinotibialis and E. (C.) thapensis sp. n. The group is recorded for the first time from the Nearctic and Neotropical regions, and is presently known to be distributed in tropical and subtropical regions of Central America, Africa, Southeast Asia and Australasia, and Southwestern North America.

In this paper, the distribution of the E. (C.) hyaleagroup is increased by new records, especially for the Ori ental region for which the group is preliminarily reviewed and twelve new species from Thailand are described.

The Empis (Coptophlebia) hyalea-group
The subgenus Coptophlebia of the genus Empis is a large worldwide group of species very similar to Empis s.str. with a rather long, well sclerotized proboscis (especially the labium), a characteristic venation (R4+5 at right angle, cell dm truncate) and a well-developed pilosity on the legs of males. These sub genera are easily distinguished by the abbreviation of the first median vein (M1) in Coptophlebia. In addition, as already pointed out on several occasions, Coptophlebia and Empis s. str., like several other taxa of generic or subgeneric level within the tribe Empidini, are not monophyletic (Chvala, 1994;Daugeron 1997aDaugeron , 2000aDaugeron , b, 2001, and the name of Copto phlebia should be reserved for a small group of Palaearctic spe cies related to the type-species of the subgenus, namely E. (C.) hyalipennis Fallen, 1816. Consequently species of Coptophlebia and Empis s.str. should be studied together whenever possible.
To resolve taxonomic and phylogenetic problems monophy letic species groups within the tribe Empidini were recognized instead of the traditional genera and subgenera, for which it was not possible to assess the monophyly, and these groups were included as terminal taxa in a global phylogeny of the Empidini (Daugeron, 1997a(Daugeron, , 2000a(Daugeron, , 2000b(Daugeron, , 2001. In this way, forty monophyletic groups were recognized and tentatively included in a phylogenetic analysis (Daugeron, 2000b).
Within the subgenus Coptophlebia, eight monophyletic groups are known, including the Empis (Coptophlebia) hyaleagroup, which was recently discovered and defined on the basis of two synapomorphies (Daugeron, 2002), the presence of a desclerotized zone in the middle of the labella and an unpaired epandrium in the male hypopygium, which allow this group to be distinguished from all other species groups of Empidini.

Material and morphological terms
This study is partly based on material collected in Thailand by one of us (PG), during several successive missions (1997, 1999, 2000 and 2001) especially at the Na Haeo Field Research Station (SWU-FIRS), and deposited in the Royal Belgian Insti tute ofNatural Sciences (KBIN-IRSNB, Brussels). The undeter mined Oriental and Australasian material studied was borrowed from the Bishop Museum (Hawaii) and the Australian Museum (Sydney).
Whenever possible, the Oriental species of the genus Empis (apart from those belonging to the subgenus Planempis, which significantly differs from Empis s.str. and Coptophlebia) were studied in order to compare them with the potential new species: Types of species described by Melander (1946), De Meijere (1907, 1911 and Frey (1953) were borrowed from the National Museum ofNatural History, Smithsonian Institution (Washing ton, USNM), the Zoologisch Museum (Amsterdam, ZMAN) and the Zoological Museum (Helsinki) respectively. Nevertheless, the type material of species described by Frey, Bezzi and Yang & Yang (1997) and placed in Empis s.str. was not available for study; however these species have a complete Ml vein whereas it is abbreviated in all the new species described in this work. For the same reason, it was not possible to study the type specimens described without subgeneric place ment or in Coptophlebia by Brunetti (1913Brunetti ( , 1917Brunetti ( , 1920 and Bezzi (1904Bezzi ( , 1912Bezzi ( , 1914. In this case the original descriptions, figures and the key published by Brunetti (1920) tentatively allowed us to include these species in the E. (C.) hyalea-group and compare them with the new species described in this paper.
In addition to the five Oriental Coptophlebia species, Melander (1946) described or recognized six additional Copto phlebia from North and Central America. These species were also borrowed (Washington, USNM) and studied in detail, in order to check for the possible presence of the E. (C.) hyaleagroup in the New World.
The morpho-anatomical terms follow McAlpine (1981) except for the male genital sclerites, which are those of Sinclair et al. (1994), Cumming et al. (1995) and Daugeron (1997b). It must be noted that the epandrium is unpaired in the E. (C.) hyalea-group (Daugeron, 2002); consequently the term epandrial lamella simply refers to the lateral aspects of the epan drium. Diagnoses only refer to males.

Male
Head. Occiput black with row of short postocular bristles. Ocellar triangle prominent, black with pair of bristles. Face black. Palpus dark brown. Antenna black, first flagellomere conical, second and third flagellomeres aristiform. Proboscis black. Labrum length 1.3 times head height. Labella shorter than prementum with short bristles, desclerotized zone indistinct. Holoptic, upper ommatidia considerably enlarged.

Female
Similar to male except for the following characters: dichoptic with lower facets slightly enlarged, frons slightly wider than face. Legs entirely black with very short pilosity except for the following long pennate bris tles: fore tibia with 1 row of dorsals on apical 2/3, first fore tarsomere with 1 row of dorsals; mid femur with 1 row of dorsals, 1 row of ventrals except basally, mid tibia with 1 row of dorsals, a few ventrals basally, first mid tarsomere with 1 row of dorsals. Hind femur and tibia with 1 row of dorsals and ventrals, first hind tarsomere with 1 row of dorsals. Fore tibia with short ventral flattened bris tles on apical 3/4. Second mid tarsomere with 1-2 short dorsal pennate bristles. Abdomen with short bristles, pointed at tip with cercus longer than wide and with a few short bristles.
Etymology. From the Latin atratata meaning dressed in black.

Remark.
The species is only known from Ko Samet island, Rayong Province (Fig. 15).

Empis (Coptophlebia) kosametensis sp. n.
( Fig. 2) Dark brown species; epandrium with 1 strong spine like bristle at tip and a pair of dorsal projections bearing a brush ofbristles.
Thorax dark brown to black. Most bristles missing. Antepronotum with some short black lateral bristles. Dorsocentrals apparently uniserial ending with 2 strong bris tles in prescutellar depression. Laterotergite with fan of strong, long black bristles. Anterior and posterior spira cles brown.
Legs brown. Fore tarsomeres somewhat swollen, first fore tarsomere with 1 strong, long anteroventral bristle at base and apical circlet of strong, long bristles, other fore tarsomeres with numerous strong, long dorsal bristles especially at tip. Mid femur with 1 antero-and posteroventral rows of strong, long bristles; mid tibia with 1 dorsal row of 3 strong, long bristles (on basal 1/4, middle and apically), 1 strong, long ventral bristle on basal 1/3; first 3 mid tarsomeres with apical circlet of strong, long bristles especially first two mid tarsomeres with 1 strong, long dorsal bristle. Hind femur with short bristles dorsally, fine, slightly longer bristles ventrally; hind tibia with 1 dorsal row of 5-7 strong, long bristles, at least 1 strong, long ventral on middle; hind tarsomeres swollen; first hind tarsomere with numerous strong ventral bristles, 1 strong, long dorsal basally, 1 pair of strong, long api cally; second and third hind tarsomeres with strong dorsal bristles especially apically.
Etymology. The name of the species is derived from the typelocality.
Remark. The species is only known from Ko Samet island, Rayong Province (Fig. 15).
Thorax brown, scutum dusty, scutellum very dusty grey. All bristles black. Antepronotum with 1 fine lateral bristle. Postpronotal lobe with 1 strong, long basal bristle. Proepisternum, lateral part of prosternum without distinct bristle. Acrostichals only represented by pair of fine, short posterior bristles in front of prescutellar depression. Dorsocentrals uniserial, distinct, ending with 3 strong, long posterior bristles, 2 of which in prescutellar depres sion. Laterotergite with fan of strong, long bristles. The other strong, long bristles are as follows: 1 pre-and 1 postsutural supraalar, 3 notopleurals, 1 postalar. Anterior and posterior spiracles blackish and brown respectively.
Legs brown. Fore tibia with 1 dorsal and ventral rows of rather strong, long bristles; first fore tarsomere ven trally and dorsally covered with rather strong, long bristles; all fore tarsomeres with apical circlet of distinct bristles. Mid femur with 2 strong, long posteroventral bristles basally; mid tibia with 1 dorsal and ventral rows of 3 and 2 strong, long bristles respectively; first mid tar somere with 1 strong, long posteroventral bristle; other mid tarsomeres with distinct bristles apically. Hind tibia with 1 dorsal row of 3 strong bristles on apical 3/4; hind tarsomeres swollen; first hind tarsomere with strong, short ventral bristles especially on base; first four hind tarsomeres with 2 strong, long dorsoapical bristles.
Abdomen brown with distinct bristles on base and mar gins of first 3 tergites, sternites with pair of strong, long posterior bristles. Hypopygium (in poor condition) (Fig. 3). Cercus with fine short internal bristles (visible in dorsal view). Epan drium with pair of small bristly dorsal bumps; epandrial lamella with 1 very long ventral bristle. Hypandrium well sclerotized, very distinct in lateral view. Phallus charac teristic: rather fine, short, distinctly flared at tip (visible in dorsal and caudal views).
Female unknown.
Etymology. The name of the species is derived from the typelocality.

Empis (Coptophlebia) miranda sp. n. (Figs 4, 5)
Blackish species of large size; wing very dark, densely microtrichiate; middle of labella with a long desclerotized zone, mid tarsus of legs dark brown to brownishyellowish, mid tibia with antero-and posteroventral rows of pennate bristles apically.

Male
Head. Occiput black with row of postocular bristles. Ocellar triangle prominent, black. Face black. Palpus dark brown. Antenna black, first flagellomere long, four times pedicel length, second and third flagellomeres somewhat styliform. Labrum blackish, length twice head height. Labium dark, with some minute bristles, labella longer than prementum with long desclerotized zone on middle. Holoptic, upper ommatidia enlarged.
Abdomen dark brown. Base, second tergite, lateral part of tergites 2-5 with distinct bristles. Sternite with poste rior pair of bristles. Hypopygium (Fig. 5). Cercus somewhat S-shaped, thick in lateral view, with some distinct bristles. Epandrium with 2 distinct pointed, small anterodorsal projec tions bearing some rather long bristles; epandrial lamella not lengthened, with 1 strong, long bristle at tip and 1 row of strong, shorter bristles ventrally. Hypandrium well sclerotized, with characteristic notch at tip in caudal view. Phallus thick basally to thin apically, long, black to yel lowish apically.

Female
Similar to male except for the following characters: dichoptic with all ommatidia of equal size; irons as wide as face; first mid tarsomere whitish; legs densely covered with long pennate bristles as follows: fore and mid tibiae, all femora with ventrals and dorsals, hind tibia with dor sals, and ventrals except on basal third, first hind tarsomere with dorsals; first fore tarsomere with short dorsal pennate bristles; wing bicolor: basal third dark brown, apical 2/3 transparent; abdomen with short bristles, pointed at tip with cercus longer than wide with a few bristly hairs. Remark. The species is only known from Ko Samet island, Rayong Province (Fig. 15).

Male
Head. Occiput dusted black with row of postocular bristles. Ocellar triangle prominent, black, with pair of rather long bristles. Face dusted black. Palpus yellow. Scape and pedicel dark brown to black, first flagellomere strongly conical, lengthened, yellow at base, blackish api cally, second and third flagellomeres aristiform, black. Labrum brown, length about twice head height. Labium dark brown; prementum with 2 rows of distinct, short anterior bristles; labella black, as long as prementum, with row of distinct short bristles. Holoptic, upper ommatidia enlarged.
Abdomen brown, covered with numerous distinct bris tles on base, posterior margins of tergites, lateral margins of tergites and sternites; all sternites with posterior pair of strong, long bristles; sternites 6 and 7 feebly sclerotized; tergite 8 without bristles. Hypopygium (Fig. 6). Cercus with 1 very strong, curved spine-like bristle laterally. Epandrial lamella narrowly separated mediodorsally, higher than long, fused with cercus anterodorsally, prolonged anterodorsally as long rounded projection bearing 2 strong, short apically rounded spines, with fan of at least 5 strong, long bristles at tip, several strong, long bristles ventrally. Hypandrium membranous ventrally. Phallus long, rather thick, dark at base; thin, clear at tip. Etymology. The name of the species is derived from the type-locality.

Remarks.
All females collected at Na Haeo appear morphologically identical (especially as lateral margins of first four abdominal tergites bear pennate bristles), whereas three males belonging to another species closely related to E. (C.) nahaeoensis were also collected at Na Haeo. In addition, four males with the hypopygium identical to E. (C.) nahaeoensis but with significant differences in the pilosity of legs were collected in two localities other than Na Haeo (namely Ko Samet and Muay Don Lakon) with four females with darker wings than the females from Na Haeo and without abdominal pennate bristles. Finally a single male collected in the South of Thailand (at Ban Khlong Kua, Songkhla Province), although very close to E. (C.) nahaeoensis, is a different species. Thus E. (C.) nahaeoensis belongs to a complex of closely related species and it is best to wait for additional material or mated pairs before describing the female.
Legs dark brown. Fore tibia clear at base, with short distinct anterodorsal bristles on apical 1/2; first fore tarsomere with apical circlet of strong bristles. Mid femur with fine ventral bristles; mid tibia with 1 row of 3 strong, long dorsal bristles (on basal half, middle, apically), 2 strong, long ventral bristles (on middle, apical 1/3); first mid tarsomere with 1 ventral and anterolateral rows of bristles; first 3 mid tarsomeres with apical circlet of strong, long bristles. Hind femur dorsally and ventrally covered with fine bristles; hind tibia with numerous fine, long ventral and dorsal bristles, 1 row of about 10 very strong, long dorsals; first hind tarsomere swollen covered with numerous very strong, long dorsals and ventrals; second and third hind tarsomeres with apical circlet ofbristles.
Abdomen dark brown. Distinct bristles at base. Ter gites with distinct lateral and posterior bristles. Sternites with at least 1 pair of posterior bristles, sternite 8 with about 10 strong, long posterior bristles.
Hypopygium (Fig. 7). Cercus distinctly projected pos teriorly, pointed at tip, middle rather thick in lateral view, with numerous fine, short internal bristles. Epandrium with 2 long anterodorsal projections, bearing 1 row of minute spine-like dorsal bristles; epandrium lamella lengthened, pointed at tip with numerous strong, long bristles ventrally and at tip. Hypandrium membranous ventrally. Phallus very long, pointed at tip.
Female unknown.
Etymology. The name of the species is derived from the typelocality.
Remark. The species is known from Phang Nga Prov ince (Fig. 15).

Male
Head. Occiput black, somewhat shiny, with row of short postocular bristles. Ocellar triangle prominent, black with pair of short bristles. Face black, shiny. Palpus brown. Antenna black, first flagellomere conical, second and third flagellomeres aristiform. Labrum dark brown, length 1.5 times head height. Labium dark brown with short bristles; labella shorter than prementum. Holoptic, upper ommatidia enlarged.
Legs dark brown. Base of fore tibia somewhat deformed (especially visible in frontal view) with 1 anterodorsal row of distinct short bristles including 2 strong spine-like apicals; fore tarsomeres somewhat swollen with distinct bristles apically; first fore tarsomere with 1 strong, long anteroventral bristle basally. Mid femur with 1 dorsal and ventral rows of bristles as long as femur depth; middle of mid tibia with 1 strong, long ven tral bristle, 1 antero-and posterodorsal rows of strong, long bristles; first mid tarsomere with 1 strong, long anterolateral bristle. Hind femur with 1 dorso-and ventroposterior rows of long bristles; hind tibia with 1 ventral and dorsal rows of strong, long bristles; hind tarsomeres swollen; first hind tarsomere with long dorsal bristles; long dorsal bristles on tips of the remaining 3 hind tarsomeres.
Hypopygium (Fig. 8). Cercus thick in lateral view. Epandrium with pair of bristly bumps dorsally, tip of epandrial lamella rather rounded with row of about 10 strong, long bristles. Hypandrium well sclerotized, pointed at tip. Phallus rather long.
Female unknown.
Etymology. The name of the species is derived from the typelocality.
Remark. The species is known from Satun Province (Fig. 15).
Empis (Coptophlebia) pseudospinotibialis sp.n. (Fig. 9) Very similar to E. (C.) spinotibialis except for the fol lowing characters: occiput entirely dusty black, antenna very dusty, thorax more dusty. Fore femur and basal 1/2 of fore tibia pale brown, otherwise legs dark brown. Bris tles of legs somewhat stronger, more numerous on tarsi. Fore tibia more distinctly deformed in lateral view, S-shaped in frontal view, with slightly shorter dorsal spine-like bristle on middle. Cercus somewhat rounded at tip in dorsal view, without minute ventral projection (Fig.  9).
Female Similar to male except for the following characters: all bristles shorter; occiput blackish, not dusty; dichoptic with all ommatidia of equal size; frons wider than face, brown, subshiny; face subshiny; thorax shiny dark brown to somewhat dusty in prescutellar depression; fore tibia not deformed, only with short bristles; first fore tarsomere with strong, short ventral bristles; mid femur with very short dorsal bristles on apical 1/3; hind femur with 1 ven tral and dorsal rows of pennate bristles as long as femur depth. Hind tibia with 1 row of ventral pennate bristles as long as tibia depth at middle, shorter pennate dorsal bris tles on apical 1/2; hind tarsomeres not swollen. Abdomen pointed at tip with cercus longer than wide with a few bristly hairs.
Etymology. The name of the species is derived from that of E. (C.) spinotibialis sp. n., as these two species are very similar morphologically.
Remark. The species is known from Phang Nga Prov ince (Fig. 15).
Female unknown.
Etymology. From the Latin pulchra meaning beautiful.
Remark. The species is known from the boundary of Prachuab and Chumpon Provinces (Fig. 15).

Male
Head. Occiput dusted blackish with row of postocular bristles. Ocellar triangle prominent, black with pair of short bristles. Face black. Palpus brown. Scape and pedicel dark brown, flagellum black, first flagellomere somewhat conical, second and third flagellomeres aristiform. Labrum brown-black, length twice head height. Labium black, bare, labella as long as prementum. Holop tic, upper ommatidia enlarged.
Legs. Fore and mid legs brown, hind legs dark brown. Fore tibia with 1 antero-and posterodorsal rows of rather short, strong bristles on apical 1/2; first fore tarsomere with 1 dorsal row of strong bristles. Mid tibia with 1 strong, very long ventral bristle on middle, 1 dorsal row of 3 strong, long bristles (on basal 1/4, middle, apically); first mid tarsomere with numerous short ventral, lateral spine-like bristles. Hind femur with fine, short antero dorsal bristles on basal 1/2 and ventrals; hind tibia cov ered with numerous strong, long antero-and posterodorsal bristles, shorter ventrals; first hind tarsomere swollen, covered with numerous strong, long dorsal bristles, rather shorter spine-like ventrals; second and third hind tarsomeres with 2 strong, long dorsals apically.
Abdomen brownish. Base and posterior margin of seg ments with distinct bristles. Posterior margins of tergite and sternite 8 with 5 and 6 strong, long bristles respec tively. Hypopygium (Fig. 11). Cercus pointed at tip, middle rather thick in lateral view, with numerous short internal bristles. Epandrium with 2 strong anterodorsal projections bearing some strong dorsal spine-like bristles; epandrial lamella subtriangular, somewhat lengthened, with about 15 strong, long bristles on ventral margin and tip. Hypandrium membranous ventrally. Phallus thin, long, not pointed at tip.
Female unknown.
Etymology. The name of the species is derived from the typelocality.
Legs brown. Fore tibia somewhat deformed, with 1 strong, spine-like dorsal bristle on middle and another finer, shorter on apical 1/4 (Fig. 13); first fore tarsomere with 1 strong, long anterolateral bristle at tip, 1 strong, long ventral apically; first 3 fore tarsomeres with distinct bristles apically. Mid femur with 1 strong, long posteroventral bristle on middle; mid tibia with 1 dorsal row of 3 bristles (1 short on basal 1/4, 1 longer on middle, 1 strong, very long apically), 1 strong, long anterolateral bristle on basal 1/3; base of first mid tarsomere with 1 strong, long ventral bristle, 1 anterolateral apically. Hind femur with rather fine ventral bristles slightly longer than femur depth on basal 1/2; hind tibia covered with rather strong ventral bristles, 1 dorsal row of 4 long bristles, 1 strong, long anterodorsal bristle on apical tip; hind tarsomeres swollen; base of first hind tarsomere with pair of strong ventral bristles; all hind tarsomeres with pair of rather fine, long dorsal bristles apically.
Female unknown.
Etymology. From the Latin spina and tibia in reference to the presence of a strong spine-like bristle on the fore tibia.
Remark. The species is known from Loei Province (Fig. 15).

Male
Head. Occiput blackish, somewhat dusty, with some short postocular bristles. Ocellar triangle prominent, black, with pair of short bristles. Face black. Palpus brown. Antenna black, first flagellomere conical, second and third flagellomeres aristiform. Labrum brown to black, length twice head height. Labium brownish with short distinct bristles; labella shorter than prementum. Holoptic, upper ommatidia enlarged.
Legs dark brown. Fore tibia with anterodorsal row of about 10 rather short, strong bristles; basal tip of first fore tarsomere with 1 strong, long anterolateral bristle; first 3 fore tarsomeres with 1 strong ventroapical; last three fore tarsomeres with fine, rather long dorsals. Mid femur with 1 posteroventral row of rather long bristles; mid tibia with 1 strong, long ventral bristle on middle, 1 dorsal row of 3 strong, long bristles; apical tip of first mid tarsomere with 1 strong, long dorsal bristle. Hind femur with very short dorsals; hind tibia with 4 strong, long dorsals; hind tarsus swollen with long dorsal and spine-like short ventral bristles.
Abdomen dark brown. First 3 tergites with distinct bristles on posterior margin. Segment 8 with some distinct posterior bristles.
Female unknown.
Other material. 1 male in poor condition, same data. Etymology. The name of the species is derived from the typelocality.
Remark. The species is known from Phang Nga Prov ince (Fig. 15).

DISTRIBUTION OF THE EMPIS (COPTOPHLEBIA) HYALEA -GROUP
In the Oriental region the tribe Empidini is undoubtedly dominated by the E. (C.) hyalea-group: in addition to the 23 species not available for study and only tentatively included in this group, 31 described and 15 undescribed species are now inventoried from India, Burma, Thailand, Laos, Vietnam, South China (including Hainan Island), Malaysia, Singapore, Taiwan, Philippines, Japan (Ryukyu Islands) and Indonesia (Sumatra, Java). The group also seems well diversified in the Australasian region (with the exception of Australia and New Zealand) with 12 spe cies, 10 of which are new, recorded from Papua New Guinea and New Caledonia.
Even if the E. (C.) hyalea-group is well represented in the Oriental and Australasian regions, it seems to have a worldwide distribution with the exception of the Palaearctic region. In Africa, after reviewing all known species and studying extensive determined and undeter mined material of the tribe Empidini (Daugeron, 1997a, 2000a, 2001, Daugeron & Grootaert, 2003, four unnamed species from West Africa (Guinea, Ivory Coast and Central Africa) were found. The E. (C.) hyalea-group is also recorded from the Nearctic and Neotropical regions and includes at least the following four species: E. anthophila Melander, 1946 (New Mexico), E. asema Melander, 1902 (Texas), E. hirticrus Melander, 1927 (Arizona) and E. impar Melander, 1946 (Costa Rica). In the New World, the group therefore appears to be con fined to Southwestern North America and Central Amer ica. Finally, the distribution of the group is much more widespread than expected. In a previous paper, one of us (Daugeron, 2002) hypothesized that the E. (C.) hyalea group was at least of Miocene origin as it seemed to follow an Afro-Oriental track with an Australasian exten sion (New Caledonia). Actually, with the exception of the Palaearctic region, the group is worldwide, but appears to be mostly confined to the tropical or subtropical regions of Central America, Africa, Asia and Australasia, and Southwestern North America. Thus, the present distribu tion of the group requires a more complex explanation, and only the reconstruction of the phylogeny and a com parison with groups showing a similar distribution (e.g. the Hemiptera Fulgoromorpha of the family Lophopidae, see Soulier-Perkins, 2000) may facilitate an historical interpretation of its biogeography (Daugeron, in prep.).