The Simuliidae form a family of small, sturdy-bodied midges with a notorious
reputation in some parts of the world as bloodsucking pests. About 1,491
species are currently recognized; however, a large number of additional
unnamed species, distinguished chromosomally, have been identified by cytologists.
Most morphologically recognized species seem to be complexes of "sibling
species" or "cytospecies," which are biologically and chromosomally
distinct but anatomically very similar to each other. Modern studies of
Simuliidae must take special account of this problem.
Members of the family are usually called black flies, but in New Zealand
they are widely known as sand flies. Bloodsucking is not a universal habit,
but females of most species feed on warm-blooded vertebrates, often preferring
either birds or mammals. In Australia and New Zealand a few species of Austrosimulium are serious mammal biters; A. pestilens is a livestock pest that undergoes
major periodic outbreaks in Queensland, and A. ungulatum can be a
vicious man-biting pest in parts of New Zealand. Except in the Marquesas,
where they can be a severe local scourge, simuliids are rarely more than
a minor biting nuisance in the Pacific Islands. There is no evidence that
black flies are involved in any human disease in Australasia or Oceania,
but they are believed to play a part in the transmission of Onchocerca
gibsoni, a filarial parasite of Australian cattle. They have been incriminated
as vectors of Leucocytozoon (a blood protozoan of birds) in penguins in
New Zealand and are among the vectors of myxomatosis virus of rabbits in
Australia.
Early stages are aquatic and virtually confined to running waters. Larvae
and pupae are attached to submerged substrates, most often trailing vegetation,
stones, or the fixed bedrock of cascades and waterfalls. Those of some Polynesian
species have been found in an unusual microhabitat, lodged between closely
packed fallen leaves trapped in water current. In tropical Africa juveniles
of some species live on mayfly nymphs or crabs in an obligate relationship
(phoresy), but no phoretic forms have been found in Australasia or Oceania.
Surprisingly, simuliids have failed to colonize Tonga, Samoa, and Hawaii,
island groups with flowing streams potentially able to support their development.
The simuliid fauna of the Australasian/Oceanian Regions, at least at the
morphospecies level, is not very rich (120 spp.), but there is a clear relationship
of its components to particular geographical areas. Nearly all species from
these areas belong to Simulium s. l. or to the locally endemic genus Austrosimulium. The Australian fauna, in addition to these 2 genera,
contains a few primitive forms assignable to the Prosimuliini. The generic
affinities of these primitive taxa are obscure, but they do not belong to
the Holarctic genus Cnephia Enderlein, in which they were formerly
placed. They are treated here as generically unassignable, pending their
comprehensive study in relation to the prosimuliines of other regions.
Only Austrosimulium is represented in the small New Zealand fauna and only Simulium s. l. in the much larger fauna of New Guinea. The same subgenera
of Simulium are found in New Guinea as in Australia, but only 1 species
(S. ornatipes) is known to be common to the 2 areas, despite their
Pleistocene connection, their proximity, and the tropical character of northern
Queensland. The fauna of Melanesia and Polynesia shows evidence of derivation
from only the west and contains 2 endemic Simulium subgenera, Hebridosimulium and Inseliellum; species from Micronesia, known only from adults,
are here provisionally assigned to the latter subgenus.
Distribution in the Pacific Islands is patchy, not only because of absence
from some apparently suitable islands, but because black fly colonization
is precluded on the innumerable small coral islands and atolls by absence
of running water. Craig (1983) mapped the disjunct distribution and recognized
several chromosomally and morphologically differentiated taxa in Polynesia,
which are described in Craig & Craig (1987) and Craig (1987). Simulium
ornatipes and S. neornatipes in Australia and New Caledonia, respectively,
represent cytological species complexes (Bedo 1977, 1984).
Refs.: Smart & Clifford (1965, keys New Guinean spp.), Crosskey
(1967a, classif. W Pacific Simulium; 1985, nomencl.; 1988 world checklist),
Dumbleton (1973, keys, rev. Austrosimulium), Colbo (1976, key Australian Simulium), Crosby (1981, key NZ immat.), Craig (1983, phylog. Polynesian
spp.; 1987, keys, rev. Society Is ssp.),
aurantiacum Tonnoir, 1925a: 234 (Simulium). Australia (NSW);
Australia (Qld, Tas, Vic).
fergusoni Tonnoir, 1925a: 238 (Simulium). Australia (NSW);
Australia (SA).
fuscoflava Mackerras & Mackerras, 1948: 236 (Cnephia tonnoiri ssp.). Australia (Qld).
orientalis Mackerras & Mackerras, 1950: 170 (Cnephia tonnoiri ssp.). Australia (Qld); Australia (NSW, Tas, Vic).
pilfreyi Davies & Györkös, 1988: 107 (Cnephia).
Australia (ACT).
strenua Mackerras & Mackerras, 1950: 170 (Cnephia). Australia
(Qld).
terebrans Tonnoir, 1925a: 237 (Simulium). Australia (Vic);
Australia (NSW).
tonnoiri Drummond, 1931: 6 (Simulium). Australia (WA).
umbratorum Tonnoir, 1925a: 237 (Simulium). Australia (Vic).
AUSTROSIMULIUM Tonnoir, 1925a: 230. Type species: Simulia australensis Schiner, 1868, orig. des.
AUSTROSIMULIUM Tonnoir, 1925a: 230 (as gen.). Type species: Simulia
australensis Schiner, 1868, orig. des.
albovelatum Dumbleton, 1973: 524. NZ (NZ).
alveolatum Dumbleton, 1973: 522 (laticorne ssp.). NZ (NZ).
australense Schiner, 1868: 15 (Simulia). NZ (NZ).
caecutiens Walker, 1848: 113 (Simulium). Nomen nudum.
caecutiens Hutton, 1874: 165 (Simulium). Nomen nudum.
tillyardi Tonnoir, 1923d: 11 (Simulium). NZ (NZ).
bicorne Dumbleton, 1973: 538. NZ (NZ).
campbellense Dumbleton, 1973: 535. NZ (Campbell I).
cornutum Tonnoir, 1925a: 243. Australia (Vic); Australia (NSW, Tas).
crassipes Tonnoir, 1925a: 242. Australia (Vic); Australia (NSW, Qld)
dugdalei Craig, Craig & Crosby, 2012: 120. NZ (NZ).
dumbletoni Crosby, 1976: 17. NZ (NZ).
extendorum Craig, Craig & Crosby, 2012: 123. NZ (Stewart I).
fiordense Dumbleton, 1973: 518 (multicorne ssp.). NZ (NZ).
fulvicorne Mackerras & Mackerras, 1950: 184. Australia (Qld).
laticorne Tonnoir, 1925a: 254. NZ (NZ).
longicorne Tonnoir, 1925a: 254. NZ (NZ).
mirabile Mackerras & Mackerras, 1948: 266. Australia (Qld).
montanum Mackerras & Mackerras, 1952: 107. Australia (NSW); Australia
(Vic).
multicorne Tonnoir, 1925a: 254. NZ (NZ).
stewartense Dumbleton, 1973: 522. NZ (Stewart I).
tillyardianum Dumbleton, 1973: 513 (n. n. for tillyardi Tonnoir).
NZ (NZ).
tillyardi Tonnoir, 1925a: 253. NZ (NZ). [Preocc. Tonnoir, 1923.]
tonnoiri Craig, Craig & Crosby, 2012: 162. NZ (NZ).
ungulatum Tonnoir, 1925a: 250. NZ (NZ).
unicorne Dumbleton, 1973: 543. NZ (NZ).
vailavoense Craig, Craig & Crosby, 2012: 153. NZ (NZ); N (Stewart I).
vexans Mik, 1881: 201 (Simulium). NZ (Auckland Is); NZ (Campbell
I).
NOVAUSTROSIMULIUM Dumbleton, 1973: 484. Type species: Simulium
bancrofti Taylor, 1918, orig. des.
bancrofti Taylor, 1918b: 168 (Simulium). Australia (Qld);
Australia (NSW, Tas, Vic, WA).
furiosum Skuse, 1889b: 1365 (Simulium). Australia (NSW); Australia
(Qld, Tas, Vic, WA).
weindorferi Tonnoir, 1925a: 248. Australia (Tas).
simile Tonnoir, 1925a: 249. Australia (Tas).
austrosimile Smart, 1944: 133 (unnec. n. n. for simile Tonnoir).
magnum Mackerras & Mackerras, 1955: 109. Australia (Qld).
pestilens Mackerras & Mackerras, 1948: 260. Australia (Qld);
Australia (NSW, NT).
torrentium Tonnoir, 1925a: 247. Australia (Tas); Australia (ACT,
NSW, Vic).
ssp. torrentium Tonnoir, 1925a: 247. Australia (Tas); Australia (NSW, Vic).
ssp. hilli Mackerras & Mackerras, 1949: 401 (torrentium spp.).
Australia (ACT).
victoriae Roubaud, 1906c: 521 (Simulium). Australia (Vic);
Australia (NSW, Qld, Tas).
tasmaniense Tonnoir, 1925a: 245. Australia (Tas).
SIMULIUM Latreille, 1802: 426. Type species: Rhagio colombaschensis Fabricius, 1796, mon.
EUSIMULIUM Roubaud, 1906b: 521. Type species: Simulia aurea Fries, 1824, mon.
bonninense Shiraki, 1935: 21 (Eusimulium). Bonin Is.
trukense Stone, 1964: 631. Micronesia; Guam.
GOMPHOSTILBIA Enderlein, 1921a: 199 (as gen.). Type species: Gomphostilbia
ceylonica Enderlein, 1921, orig.des.
ambonense Takaoka, 2003: 183. Indonesia (Maluku).
anggiense Takaoka, 2003: 371. Indonesia (Irian Jaya).
arfakense Takaoka, 2003: 324. Indonesia (Irian Jaya).
asperum Takaoka, 2003: 162. Indonesia (Maluku).
baliemense Takaoka, 2003: 403. Indonesia (Irian Jaya).
barretti Smart & Clifford, 1965: 603. PNG (PNG).
biakense Takaoka, 2003: 253. Indonesia (Irian Jaya).
bifurcatum Takaoka, 2003: 149. Indonesia (Irian Jaya).
catleyi Smart & Clifford, 1965: 608. PNG (Bismarck Arch); PNG
(PNG).
centrale Smart & Clifford, 1965: 611. PNG (PNG); PNG (Bismarck
Arch).
curvum Takaoka, 2003: 154. Indonesia (Maluku).
demutaense Takaoka, 2003: 407. Indonesia (Irian Jaya).
hagai Takaoka, 2003: 240. Indonesia (Maluku).
halmahereanse Takaoka, 2003: 247. Indonesia (Maluku).
heldsbachense Smart & Clifford, 1965: 594. PNG (PNG).
hemicyclium Smart & Clifford, 1965: 604. PNG (Bismarck Arch);
PNG (PNG).
hiroshii Takaoka, 1994: 1093. Solomon Is; PNG (Bougainville I).
inernum Takaoka, 2003: 142. Indonesia (Irian Jaya).
irianense Takaoka, 2003: 258. Indonesia (Irian Jaya).
irayense Takaoka, 2003: 344. Indonesia (Irian Jaya).
jayapuraense Takaoka, 2003: 300. Indonesia (Irian Jaya).
kalimerahense Takaoka, 2003: 135. Indonesia (Irian Jaya).
kamimurai Takaoka, 2003: 191. Indonesia (Maluku).
keravatense Smart & Clifford, 1965: 597. PNG (Bismarck Arch);
PNG (PNG).
kuingingiense Smart & Clifford, 1965: 606. PNG (PNG).
malukuense Takaoka, 2003: 263. Indonesia (Maluku).
manokwariense Takaoka, 2003: 270. Indonesia (Irian Jaya).
manuselaense Takaoka, 2003: 295. Indonesia (Maluku).
motoyukii Takaoka, 2003: 351. Indonesia (Irian Jaya).
munumense Smart & Clifford, 1965: 599. PNG (PNG); PNG (Bismarck
Arch).
nami Smart & Clifford, 1965: 612. PNG (PNG).
norforense Takaoka, 2003: 378. Indonesia (Irian Jaya).
nudipes Takaoka, 2003: 337. Indonesia (Irian Jaya).
palauense Stone, 1964: 633. Belau.
pamahaense Takaoka, 2003: 319. Indonesia (Irian Jaya).
peregrinum Mackerras & Mackerras, 1950: 175. Australia (Qld).
prafiense Takaoka, 2003: 363. Indonesia (Irian Jaya).
ransikiense Takaoka, 2003: 287. Indonesia (Irian Jaya).
rhopaloides Craig, Englund & Takaoka, 2006: 12. Solomon Is.
seramense Takaoka, 2003: 437. Indonesia (Maluku).
sorongense Takaoka, 2003: 275. Indonesia (Irian Jaya).
sutubense Takaoka, 2003: 384. Indonesia (Irian Jaya).
tafulaense Takaoka, 2003: 259. Indonesia (Maluku).
takae Takaoka, 2003: 391. Indonesia (Irian Jaya).
wakrisense Takaoka, 2003: 399. Indonesia (Irian Jaya).
watetoense Takaoka, 2003: 281. Indonesia (Maluku).
HEBRIDOSIMULIUM Grenier & Rageau, 1961: 96 (as gen.). Type
species: Simulium jolyi Roubaud, 1906, orig. des.
adsonense Craig in Craig et al., 2006: 76. Vanuatu.
anatolicum Craig, 1987: 382. French Polynesia (Society Is).
banksi Craig in Craig et al., 2006: 25. Vanuatu.
calllipygium Craig in Craig et al., 2006: 18. Vanuatu.
johnfrumi Craig in Craig et al., 2006: 30. Vanuatu.
jolyi Roubaud, 1906a: 142. Vanuatu.
laciniatum Edwards, 1924f: 572. Fiji.
lucyae Craig in Craig et al., 2006: 61. Vanuatu.
pankumuense Craig in Craig et al., 2006: 39. Vanuatu.
paradisium Craig in Craig et al., 2006: 63. Vanuatu.
seatopygium Craig in Craig et al., 2006: 65. Vanuatu.
spiroi Craig, Currie & Hunter in Craig et al., 2006: 79. Vanuatu.
subparadisium Craig in Craig et al., 2006: 69. Vanuatu.
supercilium Craig in Craig et al., 2006: 72. Vanuatu.
tuberculum Craig in Craig et al., 2006: 45. Vanuatu.
undulatum Craig in Craig et al., 2006: 74. Vanuatu.
INSELIELLUM Rubtsov, 1974: 275 (as gen.). Type species: Simulium
oviceps Edwards, 1933, orig. des.
adamsoni Edwards, 1932b: 109. French Polynesia (Marquesas).
adelaideae Craig, 2004: 3. French Polynesia (Society Is).
admixum Craig, 1987: 397. French Polynesia (Society Is).
arlecchinum Craig, 1987: 398. French Polynesia (Society Is).
bogusium Craig, 1997: 889. French Polynesia (Society Is).
buissoni Roubaud, 1906c: 521. French Polynesia (Marquesas).
caesariatum Craig & Joy, 2000: 1256. French Polynesia (Society Is).
castaneum Craig, 1987: 399. French Polynesia (Society Is).
cataractarum Craig, 1987: 400. French Polynesia (Society Is).
cheesmanae Edwards, 1927b: 242. French Polynesia (Society Is).
clibanarium Craig, 1997: 889. French Polynesia (Society Is).
concludium Craig, 1997: 886. French Polynesia (Society Is).
connae Craig, 1997: 868. French Polynesia (Society Is).
dojeycorium Craig, 1997: 871. French Polynesia (Society Is).
dussertorum Craig, 1997: 885. French Polynesia (Society Is).
englundi Craig, 2004: 10. French Polynesia (Marquesas).
evenhuisi Craig, 1997: 869. French Polynesia (Society Is).
exasperans Craig, 1987: 392. French Polynesia (Society Is).
fararae Craig & Joy, 2000: 1253. French Polynesia (Society Is).
fossatiae Craig, 1997: 872. French Polynesia (Society Is).
gallinum Edwards, 1932b: 108. French Polynesia (Marquesas).
guamense Stone, 1964: 633. Guam.
hirticranioum Craig & Joy, 2000: 1255. French Polynesia (Society Is).
hispidum Craig & Joy, 2000: 1256. French Polynesia (Society Is).
hukaense Séchan in Klein et. al., 1983: 24.
French Polynesia (Marquesas).
uahukae Lavondes & Pichon, 1972: 158. Nomen nudum.
jnabsium Craig, 1997: 879. French Polynesia (Society Is).
joyae Craig in Craig & Joy, 2000: 1254. French Polynesia (Society Is).
lonckei Craig, 1997: 880. French Polynesia (Society Is).
lotii Craig, 1987: 388. French Polynesia (Society Is).
opunohuense Craig, 1987: 391. French Polynesia (Society Is).
malardei Craig, 1987: 387. French Polynesia (Society Is).
maraaense Craig, 1997: 870. French Polynesia (Society Is).
mataverense Craig & Craig, 1987: 364. Cook Is. [Cytotype "damn"
Craig, 1983: 535.]
mesodontium Craig, 1987: 403. French Polynesia (Society Is).
middlemissae Craig, 1997: 891. French Polynesia (Society Is).
mumfordi Edwards, 1932b: 109. French Polynesia (Marquesas).
neoviceps Craig, 1987: 403. French Polynesia (Society Is).
oviceps Edwards, 1933a: 37. French Polynesia (Society Is).
pallidicranium Craig & Joy, 2000: 1255. French Polynesia (Society Is).
pichoni Craig, Fossati & Séchan, 1995: 792. French Polynesia
(Marquesas Is).
proctorae Craig, 1997: 887. French Polynesia (Society Is).
pseudocorium Craig & Joy, 2000: 1250. French Polynesia (Society Is).
pufauense Craig, 1997: 877. French Polynesia (Society Is).
rivieri Craig, Fossati & Séchan, 1995: 794. French Polynesia
(Marquesas Is).
rurutuense Craig, 1997: 864. French Polynesia (Austral Is).
sechani Craig & Fossati, 1995: 795. French Polynesia (Marquesas
Is).
shannonae Craig, 1997: 881. French Polynesia (Society Is).
sublonckei Craig, 2004: 6. French Polynesia (Society Is).
tahitiense Edwards, 1927b: 242. French Polynesia (Society Is).
teruamanga Craig & Craig, 1987: 358. Cook Is. [Cytotype "teruamanga"
Craig, 1983: 535.]
uaense Séchan in Klein et al., 1983: 24. French
Polynesia (Marquesas).
MOROPS Enderlein, 1930e: 93 (as gen.). Type species: Wilhelmia
pygmaea Enderlein, 1922 [preocc., = Simulium wilhelmlandae Smart,
1944], orig. des.
PSELAPHOCHIR Enderlein, 1936b: 120 (as gen.). Type species: Pselaphochir
oculata Enderlein, 1936, orig. des.
aropaense Takaoka, 1995: 240. PNG (Bougaiunville I).
aureonigrum Mackerras & Mackerras, 1950: 178. Australia (Qld).
avilae Smart & Clifford, 1965: 566. PNG (PNG).
botulus Smart & Clifford, 1965: 535. PNG (PNG).
brandti Smart & Clifford, 1965: 589. PNG (PNG).
cervus Smart & Clifford, 1965: 540. PNG (PNG).
clathrinum Mackerras & Mackerras, 1948: 248. Australia (Qld);
Australia (NSW).
clavum Smart & Clifford, 1965: 541. PNG (PNG).
cristatum Smart & Clifford, 1965: 553. PNG (PNG).
dycei Colbo, 1976: 256. Australia (Qld); Australia (NT).
evelynae Smart & Clifford, 1965: 545. PNG (PNG).
faheyi Taylor, 1927: 71. Australia (Qld); Australia (NT).
farciminis Smart & Clifford, 1965: 532. PNG (PNG).
fimbriatum Smart & Clifford, 1965: 544. PNG (PNG).
gagiduense Smart & Clifford, 1965: 573. PNG (PNG).
gorokae Smart & Clifford, 1965: 558. PNG (PNG).
inornatum Mackerras & Mackerras, 1950: 181. Australia (Qld);
Australia (NSW, Vic).
inuitkiense Smart & Clifford, 1965: 547. PNG (Bismarck Arch).
josephi Smart & Clifford, 1965: 580. PNG (PNG).
kainantuense Smart & Clifford, 1965: 564. PNG (PNG).
kaiti Smart & Clifford, 1965: 582. PNG (Bismarck Arch).
kerei Takaoka & Suzuki, 1994: 207. Solomon Is.
kawagishii Takaoka & Suzuki, 1995: 263. Solomon Is.
kietaense Takaoka, 1995: 242. PNG (Bougaiunville I).
kokodae Smart & Clifford, 1965: 580. PNG (PNG).
lalokiense Smart & Clifford, 1965: 576. PNG (PNG).
lawnhillense Colbo, 1976: 264. Australia (Qld); Australia (NT).
longirostre Smart, 1972: 613 (n. n. for rostrata Smart &
Clifford). PNG (PNG).
rostrata Smart & Clifford, 1965: 587. PNG (PNG). [Preocc. Lundström,
1911.]
mackerrasorum Colbo, 1976: 261. Australia (Qld); Australia (NT, WA).
mafuluense Smart & Clifford, 1965: 586. PNG (PNG).
matokoense Smart & Clifford, 1965: 585. PNG (PNG).
melatum Wharton, 1949: 406. Australia (NSW); Australia (Qld, Vic).
mendiense Smart & Clifford, 1965: 562. PNG (PNG).
minji Smart & Clifford, 1965: 555. PNG (PNG).
mussauense Delfinado, 1971: 137. PNG (Bismarck Arch).
nicholsoni Mackerras & Mackerras, 1948: 251. Australia (Qld);
Australia (NSW, Vic).
noroense Takaoka & Suzuki, 1995: 269. Solomon Is; PNG (Bougainville I).
oculatum Enderlein, 1936b: 121 (Pselaphochir). PNG (PNG).
pangunaense Takaoka, 1995: 250. PNG (Bougaiunville I).
papuense Wharton, 1948: 359. PNG (PNG); Solomon Is.
peggyae Takaoka, 1995: 249. PNG (Bougaiunville I).
pohaense Takaoka & Suzuki, 1995: 257. Solomon Is.
purosae Smart & Clifford, 1965: 560. PNG (PNG).
raunsimnae Smart & Clifford, 1965: 569. PNG (Bismarck Arch);
PNG (Bougainville I, PNG).
rounae Smart & Clifford, 1965: 539. PNG (PNG).
saihoense Smart & Clifford, 1965: 549. PNG (PNG).
selewynense Takaoka & Suzuki, 1995: 262. Solomon Is.
sherwoodi Stone & Maffi, 1971: 299. Solomon Is.
solomonense Takaoka & Suzuki, 1995: 254. Solomon Is.
standfasti Colbo, 1976: 259. Australia (Qld).
tafae Smart & Clifford, 1965: 578. PNG (PNG).
torresianum Mackerras & Mackerras, 1955: 106. Australia (Qld).
wamenae Smart & Clifford, 1965: 584. PNG (PNG).
wantoatense Smart & Clifford, 1965: 537. PNG (PNG).
wilhelmlandae Smart, 1944: 132 (n. n. for pygmaea Enderlein).
PNG (PNG).
pygmaea Enderlein, 1922a: 70 (Wilhelmia). PNG (PNG). [Preocc.
Zetterstedt, 1838.]
yulae Takaoka, 1995: 244. PNG (Bougaiunville I).
NEVERMANNIA Enderlein, 1921a: 199 (as gen.). Type species: Simulium
annulipes Becker, 1908 [= Simulium ruficorne Macquart, 1838],
orig. des.
CHELOCNETHA Enderlein, 1936b: 117 (as gen.). Type species: Chelocnetha
biroi Enderlein, 1936 [= Simulium ornatipes Skuse, 1890], orig.
des.
neornatipes Dumbleton, 1969b: 213. New Caledonia (New Caledonia).
norfolkense Dumbleton, 1969b: 220 (ornatipes ssp.). Norfolk
I.
ornatipes Skuse, 1890d: 632. Australia (NSW); Australia (Qld, SA,
Vic, WA), PNG (PNG).
biroi Enderlein, 1936b: 117 (Chelocnetha). Australia (NSW).
Subgenus WALLACELLUM Takaoka
WALLACELLUM Takaoka, 1983: 20. Type species: Simulium carinatum Delfinado, 1969, orig. des.
alfurense Takoka, 2003: 14. Indonesia (Maluku); Indonesia (Irian Jaya).