Professional Documents
Culture Documents
www.mapress.com/zootaxa/
Copyright © 2006 Magnolia Press
ZOOTAXA 1206
ISSN 1175-5334 (online edition)
Abstract
The Chinese species of the genus Ochthera Latreille are reviewed. Two species, Ochthera japonica
Clausen and O. macrothrix Clausen, are recorded from China for the first time. Two species, O.
guangdongensis sp. nov. and O. hainanensis sp. nov., are described as new. A key to the species
from China is given. The distribution of the genus in China is discussed.
Introduction
The genus Ochthera Latreille is one of the most distinctive genera of shore flies, being
easily recognized by the greatly swollen raptorial forelegs, the pair of medial facial tuber-
cles, and by two pairs of scutellar setae. Larvae and adults are predaceous, larvae have
been found on muddy and sandy shores, preying on midge larvae (Simpson, 1975). The
genus occurs worldwide with 37 known species (Mathis & Zatwarnicki, 1995). There are
seven species known from the Palearctic Region and eight from the Oriental. Only three
species have been recorded from China (Mathis & Zatwarnicki, 1995): Ochthera circu-
laris Cresson, 1926; Ochthera pilimana Becker, 1903; and Ochthera sauteri Cresson,
1932.
In this paper, we review the species of the genus Ochthera from China. Two species
are recorded from China for the first time, and two species are described as new. A key to
the seven known species from China is presented and their distribution in China is dis-
cussed.
Voucher specimens are deposited in the Entomological Museum of the China Agricul-
tural University, Beijing. The following abbreviations are used for spines, setae, and bris-
Taxonomy
1. Male with sensory structure at apex of fore basitarsomere, fore tibial spur about as long
as fore tibia ................................................................................................................... 2
Male without sensory structure at apex of fore basitarsomere, fore tibial spur about 1/3
as long as fore tibia ....................................................................... O. japonica Clausen
2. Face with 2 medial facial tubercules acutely pointed at apex ....................................... 3
Face with 2 medial facial tubercules rather rounded at apex ....................................... 4
3. Male with sensory structure consisting of 3 black setae at apex of fore basitarsomere,
mid femur with 3 strong setae at middle ................................... O. macrothrix Clausen
Male with sensory structure consisting of 4 or 5 black setae at apex of fore basitarsom-
ere, mid femur with weak setae at middle ....................................................................5
4. Male with sensory structure consisting of 3 setae at apex of fore basitarsomere; clypeus
high than wide, acutely pointed at apex ....................................... O. circularis Cresson
Male with sensory structure consisting of 4 setae at apex of fore basitarsomere;
clypeus about as high as wide, slightly pointed at apex ..... O. guangdongensis sp. nov.
5. Male with sensory structure consisting of 5 black setae at apex of fore basitarsomere;
clypeus small; gonites slightly short and thick ..........................O. hainanensis sp. nov.
Male with sensory structure consisting of 4 black setae at apex of fore basitarsomere;
clypeus large; gonites distinctly tapering toward .......................................................... 6
6. Clypeus conical with distinctly constricted apex; surstylus wide at apex .......................
..........................................................................................................O. pilimana Becker
Clypeus triangular without distinctly constricted apex; surstylus narrow and smooth at
apex .................................................................................................. O. sauteri Cresson
Ochthera circularis Cresson, 1926: 254. Type locality: Taiwan (Takao). Clausen, 1977: 480–482
[revision].
Diagnosis. Clypeus higher than wide, acutely pointed at apex; face with 2 medial facial
tubercules, rather rounded at apex; fore tibial spur about as long as fore tibae; male with
sensory structure at apex of fore basitarsomere consisting of 3 black setae with thickened
base and arising from a pit. Surstylus with distinct medial process in posterior view; cercus
short and thick; gonites distinctly tapering toward apex.
1206
FIGURES 1–4. Ochthera circularis Cresson, Male. 1. Head, anterior view; 2. fore basitarsomere,
lateral view; 3. genitalia, posterior view; 4. genitalia, lateral view.
Description. Male body length 4.7–4.9 mm, wing length 4.2–4.3 mm.
Head (Fig. 1). Subshiny black, with thin pale gray microtomentum; frons nearly pol-
ished with dense brownish yellow microtomentum at lateral margin. Setulae on head pale,
setae black; 1 lateroclinate vt, 1 proclinate fr longer than vt, 12 subequal and proclinate orb
thin and very short. Face with 2 medial facial tubercules rather rounded and subshiny
FIGURES 5–9. Ochthera circularis Cresson, Male. 5. gonites and gonal arch, internal view; 6.
gonite, lateral view; 7. aedeagus, lateral view. Female. 8. Ventral receptacle, lateral view; 9. abdo-
men, ventral view.
shiny black with pale microtomentum; tarsi brownish yellow except tarsomere 5 and hind 1206
basitarsomere black. Setulae and setae on leg black. Fore femur much swollen with row of
very short dense v (except 2 basal v slightly longer) and 4 long pv, and with 1 long av
basally and 2 long av apically; mid and hind femora each with one row of av. Fore tibia
with 1 long apical spur subshiny black, about as long as fore tibae; mid tibia apically with
2 setae; hind tibia with 1 seta apically. Fore basitarsomere (Fig. 2) thickened with dense
yellow ventral setae, with sensory structure consisting of 3 black setae at apex; hind basi-
tarsomere distinctly dilated with yellow ventral setae. Wing hyaline; veins brown except
veins on basal portion of wing brownish yellow. Alula pale yellow with pale setulae; halter
yellowish.
Abdomen subshiny black with pale microtomentum; each tergite with dense pale
microtomentum on posterior margin. Setulae on abdomen pale. Male genitalia (Figs. 3–7):
Surstylus with distinct medial process in posterior view; cercus short and thick; gonites
distinctly tapering toward apex. Female genitalia. Ventral receptacle (Fig. 8); abdomen
(Fig. 9).
Specimens examined. 2 %%, Guangdong: Ruyuan county, Nanling National Nature
Reserve, 23. VII. 2003, Xingyue Liu; 3 %%, Guangxi: Maoershan National Nature
Reserve, 21. VII. 2003, Shuwen An; 2, Guizhou: Dashahe Nature Reserve, 24. VIII. 2004,
Yajun Zhu; 1 %, Henan: Neixiang county, Baotianman National Nature Reserve, 13. VII.
1998, Xueyou Hu; 2 &&, Henan: Neixiang county, Baotianman National Nature Reserve,
14. VII. 1998, Xueyou Hu; 1 &, Hunan: Changsha, 15. VIII. 1985, Naizhong Chen.
Distribution. China (Guangdong, Guangxi, Guizhou, Henan, Hunan, Taiwan); Viet-
nam, India, Nepal, Sri Lanka, Indonesia, Malaysia, Philippines.
Diagnosis. Face with 2 medial facial tubercules obtuse and subshiny black at apex; face
with distinct carina; clypeus about as high as wide, slightly pointed at apex. Male with sen-
sory structure consisting of 4 small black setae at apex of fore basitarsomere. Surstylus
narrow apically with distinct medial process in posterior view; cercus rather short and
thick; gonites slightly tapering toward apex.
Description. Male body length 3.8 mm, wing length 3.3 mm.
Head (Fig. 10). Subshiny black, with golden microtomentum; frons nearly polished,
with dense yellow microtomentum at lateral margin. Setulae on head pale, setae black; 1
lateral vt, 1 long proclinate fr, 10–14 subequal and proclinate orb thin and very short. Face
with silvery microtomentum, with 2 medial facial tubercules obtuse and subshiny black at
apex; face with conspicuous carina; clypeus with silvery microtomentum about as high as
wide, slightly pointed at apex. Antenna black; arista black with 3 dorsal branches. Probos-
cis shiny blackish with pale setulae; palpus yellow with pale setulae.
1206
FIGURES 10–13. Ochthera guangdongensis sp. nov., Male. 10. Head, anterior view; 11. fore basi-
tarsomere, lateral view; 12. genitalia, posterior view; 13. genitalia, lateral view.
Thorax subshiny black, with yellow microtomentum except antero-lateral margin with
grayish white microtomentose. Pleura black with silvery microtomentum. Setulae and
setae on thorax black. Notopleural area and anepisternum with pale setulae. 1 posterior
npl; 1 sa; 1 psa longer than sa; scutellum with 2 pairs of sc. Anepisternum with 1 long thin
seta; katepisternum with 1 strong seta on upper portion. Legs subshiny black with pale
microtomentum; tarsi brownish yellow except tarsomere 5 and hind basitarsomere black.
Fore femur much swollen with row of very short dense v, one row of 4 long pv and 2 long
cal spur subshiny black about as long as fore tibia; mid tibia apically with 1 seta. Fore 1206
basitarsomere (Fig. 11) with sensory structure consisting of 4 small black setae; fore tar-
someres 1–2 with dense yellow ventral setae; hind basitarsomere distinctly thickened with
yellowish ventral setae. Wing hyaline; veins brown except veins on basal portion of wing
brownish yellow. Alula pale yellow with pale setulae; halter pale yellowish.
Abdomen subshiny black with grayish microtomentum, each tergite with dense pale
microtomentum at lateral margin. Setulae on abdomen pale. Male genitalia (Figs. 12–16):
Surstylus narrow apically with distinct medial process in posterior view; cercus rather
short and thick; gonites slightly tapering toward apex. Female genitalia. Ventral receptacle
(Fig. 17); abdomen (Fig. 18).
FIGURES 14–18. Ochthera guangdongensis sp. nov., Male. 14. gonites and gonal arch, internal
view; 15. gonite, lateral view; 16. aedeagus, lateral view. Female. 17. Ventral receptacle, lateral
view; 18. abdomen, ventral view.
Diagnosis. Face with 2 medial facial tubercules acutely pointed at apex; face with distinct
carina; clypeus small, triangular without distinctly constricted apex; tarsi brownish yellow;
male with sensory structure consisting of 5 black setae at apex of fore basitarsomere. Sur-
stylus narrow and smooth apically; cercus rather short and thick; aedeagus rather large
with thick apex; gonites slightly short and thick.
Description. Male body length 3.8 mm, wing length 3.0 mm.
Head (Fig. 19). Subshiny black, with yellowish microtomentum; frons nearly pol-
ished, with dense yellow microtomentum at lateral margin. Setulae on head pale, setae
black; 1 lateral vt, 1 proclinate long fr, 10–15 subequal and proclinate orb thin and very
short. Face with bicuspie medial facial tubercules, subshiny black and acutely pointed at
apex, face with conspicuous carina; clypeus small triangular without distinctly constricted
apex. Antenna black; arista black with 3 dorsal branches. Proboscis shiny blackish with
pale setulae; palpus yellow with pale setulae.
Thorax subshiny black, with yellow microtomentum. Setulae and setae on thorax
black, setae somewhat setulae-like; notopleural area and anepisternum with pale setulae. 1
posterior npl; 1 sa; 1 psa longer than sa; scutellum with 2 pairs of sc. Anepisternum with 1
long thin seta; katepisternum with 1 strong seta on upper portion. Legs subshiny black
with pale microtomentum; tarsi brownish yellow except tarsomere 5 and hind basitarsom-
ere black. Fore femur much swollen with row of short dense v, one row of 4 long pv and 2
long strong av. Mid and hind femora each with one row of strong av. Fore tibia with 1 long
apical spur subshiny black; mid tibia apically with 1 seta, hind tibia apically with 1 seta.
Fore basitarsomere (Fig. 20) thickened with sensory structure consisting of 5 black setae at
apex, and with dense yellow ventral setae; hind basitarsomere distinctly swollen. Wing
hyaline; veins brown except veins on basal portion of wing brownish yellow. Alula pale
yellow with pale setulae. Halter pale yellow.
1206
FIGURES 19–25. Ochthera hainanensis sp. nov., Male. 19. Head, anterior view; 20. fore basitar-
somere, lateral view; 21. genitalia, posterior view; 22. genitalia, lateral view; 23. gonites and gonal
arch, internal view; 24. gonite, lateral view; 25. aedeagus, lateral view.
Diagnosis. Clypeus obtuse and rounded; face with 2 medial facial tubercules distinctly
separated, obtuse at apex; fore tibial spur about 1/3 as long as fore tibia; male without sen-
sory structure at apex of fore basitarsomere. Surstylus rather wide without distinct medial
process; cercus slender; aedeagus narrowing toward apex and with constricted base bear-
ing 2 denticles; gonite rather narrow apically.
Description. Male body length 4.8–5.0 mm, wing length 4.1–4.3mm.
Head (Fig. 26). Subshiny black, with golden microtomentum; frons with thin yellow-
ish brown microtomentum except lateral margin with dense brown microtomentum. Setu-
lae on head pale and setae black; 1 lateral vt, 1 long and several short proclinate fr, vt
longer than fr, 10 subequal proclinate orb thin and short. Face with 2 medial facial tuber-
cules distinctly separated, obtuse and subshiny black at apex; clypeus obtuse and rounded.
Antenna black; arista black with 3 dorsal setulae. Proboscis shiny blackish with brown set-
ulae; palpus black brownish with pale setulae.
Thorax subshiny black, with golden microtomentum; mesonotum golden microtomen-
tose except antero-lateral margin with grayish white microtomentose; anepisternum with
silvery microtomentum. Setulae and setae on thorax black; notopleural area with yellowish
brown setulae, anepisternum with pale setulae; 1 long posterior npl; 1 sa; 1 psa longer than
sa; 1 presc as long as sa; scutellum with 2 pairs of setae. Anepisternum with 2 long thin
setae; katepisternum with 1 strong seta on upper portion. Legs subshiny black with pale
microtomentum; tarsi brownish black. Setulae on legs yellow, setae black. Fore femur
1206
FIGURES 26–29. Ochthera japonica Clausen, Male. 26. Head, anterior view; 27. fore basitarsom-
ere, lateral view; 28. genitalia, posterior view; 29. genitalia, lateral view.
FIGURES 30–34. Ochthera japonica Clausen, Male. 30. gonites and gonal arch, internal view; 31.
gonite, lateral view; 32. aedeagus, lateral view. Female. 33. Ventral receptacle, lateral view; 34.
abdomen, ventral view.
pale microtomentum at lateral margin. Setulae on abdomen pale. Male genitalia (Figs. 28– 1206
32): Surstylus rather wide without distinct medial process; cercus slender; aedeagus nar-
rowing toward apex and with constricted base bearing 2 denticles; gonite rather narrow
apically. Female genitalia. Ventral receptacle (Fig. 33); abdomen (Fig. 34).
Specimens examined. 1 %, Beijing: China Agricultural University, 25. VII. 1975,
Fasheng Li; 1 %, Heilongjiang: Sujinlitong, 20. VIII. 1970; 1 %, Shenyang: Labour Park,
24. VII. 2003, Dong Zhang; 4 %%, Shenyang: Labour Park, 26. VII. 2003, Dong Zhang; 1
%, Shenyang: Labour Park, 16. VII. 2003, Dong Zhang; 1 %, Xinjiang: Wushen, 18. VII.
1959, Changfa Li; 1 &, Beijing China Agricultural University, 10. x. 1950, Jikun Yang.
Distribution. China (Beijing, Heilongjiang, Shenyang, Xinjiang); Japan, Russia.
Diagnosis. Face with 2 medial facial tubercules acutely pointed at apex; face with distinct
carina; clypeus as high as wide, with rounded to slightly pointed apex; male with sensory
structure consisting of 3 black setae at apex of fore basitarsomere; fore tibial spur about as
long as fore tibia; mid femur with 3 large ad at middle. Surstylus distinctly curved inward
apically in lateral view; cercus short and thick; aedeagus rather large, nearly quadrate with
apical portion narrowing and slightly curved; gonite finger-like apically.
Description. Male body length 4.3 mm, wing length 3.9 mm.
Head (Fig. 35). Subshiny black, with thin yellow microtomentum; frons with dense
yellowish microtomentum except lateral margin with brownish yellow microtomentum.
Setulae on head yellow, setae black; 1 lateral vt and 1 short medial vt, 1 proclinate strong
fr, fr longer than vt; 15–20 subequal and proclinate orb thin and very short, 3 slightly
longer than others. Face black with dense golden microtomentum, and with 2 medial facial
tubercules, acutely pointed at apex; face with distinctly carina; clypeus as high as wide,
rounded to slightly pointed apex. Antenna black; arista black with 3 dorsal branches. Pro-
boscis shiny blackish with yellow setulae; palpus brownish with yellow setulae.
Thorax subshiny black; mesonotum golden microtomentose, with brown stripes
between acr and dc and laterally along dc rows, and with grayish spot intrascutal suture.
Setulae and setae on thorax black; notopleural area with golden microtomentum, anepis-
ternum with pale microtomentum; 1 long posterior npl; 1 sa; 1 psa longer than sa; 1 long
presc as long as sa; scutellum with 2 pairs of sc. Anepisternum with 1 long black and 2
short yellow setae; katepisternum with 1 strong seta on upper portion. Legs subshiny black
with pale to golden microtomentum; tarsi golden except tarsomere 5 and hind basitarsom-
ere black. Fore femur much swollen, with one row of very short v, one row of 3 long pv,
basally with 1 short av and 2 long av; mid femur with 3 strong ad at middle. Fore tibia with
FIGURES 35–38. Ochthera macrothrix Clausen, Male. 35. Head, anterior view; 36. fore basitar-
somere, lateral view; 37. genitalia, posterior view; 38. genitalia, lateral view.
1206
FIGURES 39–43. Ochthera macrothrix Clausen, Male. 39. gonites and gonal arch, internal view;
40. gonite, lateral view; 41. aedeagus, lateral view. Female. 42. Ventral receptacle, lateral view; 43.
abdomen, ventral view.
Abdomen subshiny black with golden microtomentum, each tergite with pale microto-
mentum at posterior-lateral margin. Setulae on abdomen pale. Male genitalia (Figs. 37–
41): Surstylus distinctly curved inward apically in lateral view; cercus short and thick;
aedeagus rather large, nearly quadrate with apical portion narrowing and slightly curved;
gonite finger-like apically. Female genitalia. Ventral receptacle (Fig. 42); abdomen (Fig.
43).
Specimens examined. 1 %, Yunnan: Menghai, 06. III. 1999, Ding Yang; 1 &, Yunnan:
Mengla, 09. III. 1999, Ding Yang.
Distribution. China (Yunnan); India, Australia.
Diagnosis. Face with 2 medial facial tubercules, acutely pointed at apex; face with distinct
carina; clypeus conical with distinctly constricted apex. Tarsi yellow; male with sensory
structure consisting of 4 black setae at apex of fore basitarsomere. Surstylus without dis-
tinct medial process in posterior view; cercus short and thick; aedeagus subtriangular in
lateral view; gonites distinctly tapering toward apex.
FIGURES 44–47. Ochthera pilimana Becker, Male. 44. Head, anterior view; 45. fore basitarsom-
ere, lateral view; 46. genitalia, posterior view; 47. genitalia, lateral view.
Head (Fig. 44). Subshiny black, with golden microtomentum; frons nearly polished. 1206
With dense brown microtomentum at lateral margin. Setulae on head pale, setae black; 1
lateroclinate strong vt, 1 proclinate long fr, fr longer than vt, 10–20 subequal and procli-
nate orb thin and very short. Face with bicuspid medial facial tubercules acutely pointed
and subshiny black at apex, face with conspicuous carina; clypeus conical with distinctly
constricted apex. Antenna black; arista black with 3 dorsal hairlike branches. Proboscis
shiny blackish with pale setulae; palpus yellow with pale setulae.
FIGURES 48–52. Ochthera pilimana Becker, Male. 48. gonites and gonal arch, internal view; 49.
gonite, lateral view; 50. aedeagus, lateral view. Female. 51. Ventral receptacle, lateral view; 52.
abdomen, ventral view.
Diagnosis. Face with 2 medial facial tubercules, acutely pointed at apex; face with distinct
carina; clypeus higher than wide, without constricted apex. Tarsi brownish yellow; male
with sensory structure consisting of 4 black setae at apex of fore basitarsomere. Surstylus
without distinct medial process in posterior view; cercus slender; aedeagus subtriangular
in lateral view; gonites distinctly tapering toward apex.
Description. Male body length 3.5 mm, Wing length 3.7 mm.
Head (Fig. 53). Subshiny black, with brownish yellow microtomentum; frons nearly
polished, with dark brown microtomentum at lateral margin. Setae on head black; 1 lateral
vt, 1 proclinate long fr, fr longer than vt, 10–20 subequal and proclinate orb thin and very
short. Face with golden microtomentum, and with bicuspid medial facial tubercules
than wide, without constricted apex. Antenna black; arista black with 3 dorsal branches. 1206
Proboscis shiny blackish with pale setulae; palpus yellow with pale setulae.
FIGURES 53–56. Ochthera sauteri Cresson, Male. 53. Head, anterior view; 54. fore basitarsom-
ere, lateral view; 55. genitalia, posterior view; 56. genitalia, lateral view.
FIGURES 57–61. Ochthera sauteri Cresson, Male. 57. gonites and gonal arch, internal view; 58.
gonite, lateral view; 59. aedeagus, lateral view. Female. 60. Ventral receptacle, lateral view; 61.
abdomen, ventral view.
Abdomen subshiny black with golden microtomentum, tergites 1–3 with dense brown
microtomentum on lateral margin. Setulae on abdomen yellow. Male genitalia (Figs. 55–
subtriangular in lateral view; gonites distinctly tapering toward apex. Female genitalia. 1206
Ventral receptacle (Fig. 60); abdomen (Fig. 61).
Specimens examined. 1 %, India: Kar. St. Mudigere, 6. Apr. 1980. W. N. Mathis & A.
Freidberg; 2 &&, India: Kar. St. Mudigere, 6. Apr. 1980. W. N. Mathis & A. Freidberg.
Distribution. China (Taiwan); India.
Remarks. The specimens of O. sauteri from Taiwan were unavailable for our
research. We studied the identified specimens of this species from India that were kindly
provided by Dr. W. N. Mathis.
Discussion on zoogeography
China belongs faunistically to two zoogeographical realms – the Palearctic Realm and the
Oriental Realm, consisting of the following zoogeographical regions (Zhang, 1998): South
China Region, Central China Region, South West China Region, Qinghai-Xizang Region,
Neimeng-Xinjiang Region, North China Region and North East China Region. The first 3
belong to the Oriental Realm, the latter 4 Regions belong to the Palearctic Realm. The
genus Ochthera is mainly found in the Oriental Realm, except for one species, O. japon-
ica, that occurs in the Palearctic Realm (Fig. 62). Six species, O. circularis, O. guangdon-
gensis sp. nov., O. hainanensis sp. nov., O. macrothrix, O. pilimana and O. sauteri, occur
in the South China Region, which consists of Fujian, Guangdong, Guangxi, Hainan, Tai-
wan, Yunnan. This pattern of distribution may indicates that these six species prefer tropi-
cal climates. Ochthera circularis Cresson can also be found in the Central China Region,
including West Henan and Guizhou. Obviously, the South China Region is the diversity
center of the Chinese species, having the greatest number of species occurring there. The
Acknowledgements
We are very grateful to Prof. Fasheng Li and Mr. Xueyou Hu (Beijing), Dr. W. N. Mathis
(Washington, D. C.), and Dr. A. Freidberg (Tel Aviv) for collecting the specimens, and to
Mrs. Lili Zhang and Mrs. Mengqing Wang (Beijing) for their kind help in many ways. Our
sincere thanks are due to W. N. Mathis (Washington, D. C.) for kindly providing the iden-
tified specimens of Ochthera sauteri, collected from India, for our research. This research
is supported by the National Natural Science Foundation of China (No. 30225009).
References
Becker, Th. (1903) Gyptische Dipteren gesammelt und beschrieben. Mitteilungen aus dem Zoolo-
gischen Museum in Berlin, 2 (3), 67–195.
Clausen, P.J. (1977) A revision of the Nearctic, Neotropical, and Palearctic species of the genus
Ochthera, including one Ethiopian species, and one new species from India. Transactions of
the American Entomological Society, 103, 451–530.
Cresson, E.T. Jr. (1926) Descriptions of new genera and species of Diptera (Ephydridae and Micro-
pezidae). Transactions of the American Entomological Society, 52, 249–274.
Cresson, E.T. Jr. (1932) Studies in the Dipterous family Ephydridae. Paper IV. Transactions of the
American Entomological Society, 58, 1–34.
Hennig , W. (1943) bersicht ber die bisher bekannten Metamorphosestadien der Ephydriden, mit
Neubeschreibungen nach dem Material der Deutschen Limnologischen Sundaexpedition.
(Diptera: Ephydridae). Arbeiten ber morphologische und taxonomische Entomologie aus Ber-
lin-Verlag, 10 (2–3), 105–138.
Mathis, W.N. & T. Zatwarnicki. (1995) World catalog of shore flies (Diptera: Ephydridae). Mem-
oirs on Entomology, International, 4, 1–423.
Simpson, K.W. (1975) Biology and immature stages of three species of Nearctic Ochthera (Diptera:
Ephydridae). Proceedings of the Entomological Society of Washington, 77(1), 129–155.
Zhang, Y. Z. (1998) The second revision of zoogeographical Regions of China. Acta Zootaxono-
mica Sinica, 23 (supplement), 207–222.