Table Of ContentCraig, D. A.; Craig, R. E. G.; Crosby, T. K. 2012.
Simuliidae (Insecta: Diptera). Fauna of New
Zealand 68: 336 pp.
The Copyright notice printed on page 4 applies to the use
of this PDF.
This PDF is not to be posted on websites.
Links should be made to:
FNZ.LandcareResearch.co.nz
EDITORIAL BOARD
Dr R. M. Emberson, c/- Department of Ecology, P.O. Box 84, Lincoln University,
New Zealand
Dr M. J. Fletcher, Director of the Collections, NSW Agricultural Scientific
Collections Unit, Forest Road, Orange, NSW 2800, Australia
Dr R. J. B. Hoare, Landcare Research, Private Bag 92170, Auckland, New
Zealand
Dr M.-C. Larivière, Landcare Research, Private Bag 92170, Auckland, New
Zealand
Mr R. L. Palma, Natural Environment Department, Museum of New Zealand
Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand
SERIES EDITOR
Dr T. K. Crosby, Landcare Research, Private Bag 92170, Auckland, New
Zealand
Fauna of New Zealand
Ko te Aitanga Pepeke o Aotearoa
Number / Nama 68
Simuliidae
(Insecta: Diptera)
Douglas A. Craig1, Ruth E. G. Craig1, Trevor K. Crosby2
1 Department of Biological Sciences, University of Alberta, Edmonton,
Alberta T6G 2E9, Canada
2 Landcare Research, Private Bag 92170, Auckland 1142, New Zealand
1 [email protected], 2 [email protected]
M anaaki
W henua
P R E S S
Lincoln, Canterbury, New Zealand
2012
4 Craig, Craig & Crosby (2012): Simuliidae (Insecta: Diptera)
Copyright © Landcare Research New Zealand Ltd 2012
No part of this work covered by copyright may be reproduced or copied in any form or by any means (graphic,
electronic, or mechanical, including photocopying, recording, taping information retrieval systems, or otherwise)
without the written permission of the publisher.
Cataloguing in publication
Craig, Douglas A.
Simuliidae (Insecta: Diptera) / Douglas A. Craig, Ruth E. G. Craig, and Trevor K. Crosby. –
Lincoln, N.Z. : Manaaki Whenua Press, Landcare Research, 2012.
(Fauna of New Zealand, ISSN 0111-5383 (print), ISSN 1179-7193 (online) ; no. 68).
ISBN 978-0-478-34734-0 (print)
ISBN 978-0-478-34735-7 (online)
I. Craig, Ruth E. G. II. Crosby, Trevor K., 1946– III. Title. IV. Series.
UDC 595.771(931)
Suggested citation:
Craig, D. A.; Craig, R. E. G.; Crosby, T. K. 2012. Simuliidae (Insecta: Diptera). Fauna of New Zealand
68: 336 pp.
Molecular section: Craig, D. A.; Cywinska, A. 2012: Molecular analysis of New Zealand
Austrosimulium (Diptera: Simuliidae) species. Pp. 60–65, 205–208, 298–301 in Craig, D. A.; Craig,
R. E. G.; Crosby, T. K. 2012: Simuliidae (Insecta: Diptera). Fauna of New Zealand 68, 336 pp.
Dr R. J. B. Hoare of the Editorial Board oversaw the reviewing of the manuscript and accepted the revised
version. Prepared for publication by the series editor and the authors using computer-based text processing,
layout, and printing at Landcare Research, Private Bag 92170, Auckland, New Zealand.
To access on-line extracts from this series visit:
http://fnz.landcareresearch.co.nz/
Māori text by H. Jacob, Ōtaki.
Published by Manaaki Whenua Press, Landcare Research, P.O. Box 40, Lincoln, Canterbury, N.Z.
Website: http://www.mwpress.co.nz/
Printed by PrintLink Ltd, Wellington
Date of publication 29 June 2012
Front cover: Head of female Austrosimulium dumbletoni Crosby from Jackson Bay, WD (Photographer: D. A. Craig).
Publication of the Fauna of New Zealand series is the result of a research investment by the Ministry
for Science and Innovation.
Fauna of New Zealand 68 5
POPULAR SUMMARY HE WHAKARĀPOPOTOTANGA
Class Insecta
Order Diptera
Suborder Nematocera
Family Simuliidae
Genus Austrosimulium
Subgenus Austrosimulium
Black flies, sandflies, te namu
Known in New Zealand as “sandflies” or “Te Namu” and
elsewhere in the world mainly as “black flies”, Simulii-
dae are iconic New Zealand insects. Virtually every New
Zealander has been bitten by female simuliids at some Illustration / Whakaahua: Austrosimulium ungulatum
time or other, as have the many overseas tourists who have Tonnoir. Brass sculpture, Milford Sound Visitor Centre,
Fiordland. Artist: Elizabeth Thomson, 1991 (Photographer:
visited Fiordland.
Andrew Baird, Invercargill).
Simuliids of New Zealand belong to the genus Austro-
simulium known only from New Zealand, Tasmania, and
mainland Australia. The nearest relative is the genus Te Namu
Paraustrosimulium in South America. This relationship
has been confirmed by both morphological and molecular Koia tēnei ko te namu, ko tōna ingoa Pākehā i Aotearoa
examination. Therefore these simuliids would appear nei, ko te “ngaro onepū”, i tāwāhi, ko te “ngaro pango”. He
to be of Gondwanan provenance, when South America, ngārara ingoa araara te namu i Aotearoa — ko wai o tātou
Australia, New Zealand, Tasmania, Africa, India, and kāore anō i ngaua e te namu uwha? Tae atu anō ki te hunga
Antarctica were co-joined in the super continent Gond- haramai i tāwāhi ki Te Whakataka-kārehu-a-Tamatea — ko
wana that broke up between 120–80 million years ago. rātou anō e rongo kino ana i te ngau a te namu.
In Australia there is a small segregate of species in the Nō te puninga Austrosimulium ngā simuliid o Aotearoa,
subgenus Novaustrosimulium, with all their remaining waihoki ko Aotearoa, ko Tahimania me te tuawhenua o
Austrosimulium species in the subgenus Austrosimulium. Ahitereiria anake ōna kāinga e mōhiotia ana. Ko ōna uri
Questions still remain, however, as to exactly when New tino tata, ko te puninga Paraustrosimulium, i Amerika ki
Zealand simuliids arrived here, but this study indicates te Tonga. Kua whakaūngia tōna hononga ā-whakapapa
they are not much older than 5 million years. ki tērā i runga i te hanga o ngā tinana me ngā mātaitanga
Simuliid larvae require running water and in New rāpoi ngota. Nō reira, ko Te Uri Māroa pea te kāinga tu-
Zealand are more or less ubiquitous, occurring in almost auri o ēnei puninga e rua, i te wā e whenua kotahi ana a
all running water habitats. Although there are 19 species Amerika ki te Tonga, a Ahitereiria, a Aotearoa, a Tahima-
of Austrosimulium in the country, the females of only 3 nia, a Āwherika, a Īnia, me Te Kōpakatanga ki te Tonga.
species are serious biters of humans. Males never bite. The Nō te 120–80 miriona tau ki muri i wehewehe ai a Te Uri
females bite to obtain nutrients to produce eggs: females Māroa. Tērā tētahi wehenga iti i Ahitereiria o ētahi momo
that do not take a blood meal can still lay a smaller number nō te puninga whāiti Novaustrosimulium, engari ko ērā atu
of eggs. Markedly few males have ever been collected in o ōna momo o te puninga Austrosimulium, nō te puninga
the field despite the equal abundance of males and females whāiti Austrosimulium. Kāore anō i kitea nōnahea rawa i
in their immature stages. tau mai ai ngā namu o Aotearoa ki konei. Engari ko tā tēnei
In the North Island the main biter is the New Zealand rangahautanga nei e tohu ana, nō te takiwā o te 5 miriona
black fly, Austrosimulium australense, found the length tau ki muri, paku neke atu rānei.
(continued overleaf) (haere tonu)
6 Craig, Craig & Crosby (2012): Simuliidae (Insecta: Diptera)
and breadth of that island, but less so in the South Island Me wai rere e ora ai ngā punua a te namu. Me kī, kei
and Stewart Island. Another species in the North Island ngā wāhi katoa o Aotearoa, me he wai e rere ana i reira.
that bites to some extent is A. tillyardianum, but it is not Tekau mā iwa ngā momo Austrosimulium kei Aotearoa,
found in large numbers, and then only south of Auckland. engari ko ngā uwha o ētahi momo e toru noa iho e kaha
In the South Island the two main biters are A. tillyardianum, mōhiotia ana mō tā rātou ngau i te tangata. Kāore rawa ngā
mainly in the north and then south through Canterbury, toa e ngau. He ngau tā te uwha, kia kaha ai tana whānau
while in Westland and Fiordland the simuliid of notoriety hua. Arā ētahi uwha kāore e kai toto, engari ka whakaputa
is the West Coast black fly, A. ungulatum. The females hua tonu; heoi anō, he iti ake ngā hua ka whānau mai i a
of A. ungulatum will fly long distances to obtain a blood rātou. He ruarua noa ake ngā toa kua kohia, tēnā i ngā
meal and their ferocity has generated many horror stories uwha, ahakoa e āhua rite ana te maha o ngā punua toa me
from early settlers in New Zealand, and more recently, ngā punua uwha.
the annual million or so overseas tourists. Few of the lat- I Te Ika-a-Māui, ko te Austrosimulium australense te
ter leave New Zealand unscathed if they visit Fiordland, namu kaha te ngau, kitea ai i te whāroatanga atu o tēnei
and many tend to react badly to bites since they have not motu. Kei Te Waka-a-Māui, kei Rakiura anō, engari kāore
previously been exposed to the biting of this species. It has e pērā te kaha o te kitea. Ko tētahi atu namu ngau kei Te
been noted that, similar to the quote regarding weather, at- Ika-a-Māui, ko te A. tillyardianum, heoi anō, kāore e tini
tributed to Mark Twain (Samuel L. Clemens), that in New ngerongero, ka mutu kei te tonga anahe o Tāmakimakaurau.
Zealand “Everybody talks about sandflies, but nobody does E rua ngā momo namu matua o Te Waipounamu e rongonui
anything about them”. ana mō te ngau. Ko tētahi ko A. tillyardianum, kei ngā
New Zealand Austrosimulium can be placed into two whenua mai i Te Tauihu o te Waka ahu atu ki Waitaha. Ko
main species-groups, based on the presence (ungulatum tērā atu, a A. ungulatum, e rongonui ana i Te Tai Poutini
species-group) or absence (australense species-group) of a me Te Whakataka-kārehu-a-Tamatea. Ko ngā uwha o A.
basal tooth on the tarsal claws of the female. Elsewhere in ungulatum, ka rere ki tawhiti ki te kimi toto hei kai mā
the world those possessing the tooth are normally known as rātou, ka mutu he ngārara niwha tonu. Nō konā tana uru ki
bird feeders, whereas those lacking the tooth are normally ngā kōrero paki a ngā Pākehā tuatahi i whakanohonoho i
more mammophilic. New Zealand simuliids are oppor- Aotearoa, tae atu ki ā te hunga tāpoi o ngā tau tata ake — ko
tunistic and females will generally take blood meals from tōna kotahi miriona nei e taetae mai ana i tāwāhi i ia tau. O
whatever is available. It is well established, however, that ngā wae tāpoi ka haere ki Te Whakataka-kārehu-a-Tamatea,
whereas Fiordland crested penguins are heavily attacked by he tokoiti ka hoki ki ō rātou kāinga i rāwāhi me te kore
A. dumbletoni, a toothed-clawed member of the ungulatum e mamae o te tinana e te ngau a namu. Ka mutu, ka āhua
species-group, its females studiously avoid biting humans. kino te ngau ki a rātou, he kore nō ō rātou tinana i waia ki
The simuliids that attack penguins are known to be a vector te ngau a tērā momo namu. Kua puta te kōrero āhua pērā
for a bird “malaria”, Leucocytozoon, but are not known to i tā Mark Twain (Samuel L. Clemens), tāna kōrero mō te
be a vector for any disease to people. huarere, e mea ana, i Aotearoa nei “ko te namu kei ngā
Molecular analysis indicates that most New Zealand ngutu o te katoa, engari kāore tētahi tangata kotahi nei i te
simuliid species are of relatively recent origin, probably kori ake ki te rongoā i te raruraru”.
evolving during the 5 glaciations over the last 500 000 Ka noho mai ngā Austrosimulium o Aotearoa ki ētahi
years, and this is in agreement with some other such inves- huinga momo e rua — ko ērā e whai niho ana i te pūtake
tigations of the New Zealand fauna. There are, however, o ngā matihao o te uwha (te huinga ungulatum) me ērā
lineages in the ungulatum species-group which are high karekau he niho pērā (te huinga australense). I ētahi atu
altitude, cold water species; and this specialised habitat in wāhi o te ao, ko te nuinga o ngā mea he niho pēnei ō rātou,
the Southern Alps is a little more than 5 million years old. he kai i te toto o te manu, ko ngā mea kāore he niho pēnei,
That age is in general agreement with molecular evidence he kai i te toto o te whāngote. Engari ko te nuinga o ngā
from this study. simulid o Aotearoa, he whai i te ara māmā e ngata ai te
A common question is “What did New Zealand sim- hiahia, nō reira ka whāia e ngā uwha te toto o te mea e tata
uliids feed on before humans arrived?” Of little doubt it ana. Heoi anō, kua āta kitea i ētahi atu rangahau, ahakoa e
would be on the vast number of birds that now are greatly muia ana ngā tawaki e te A. dumbletoni, he huānga matihao
depleted — gone are the days of the “dawn chorus” — and whai niho nō te rōpū ungulatum, kāore ngā uwha e ngau
probably also the large numbers of seals present then. i te tangata. Ko ngā namu e urutomo ana i ngā tawaki, ko
Indeed, searching along beaches for a blood meal from rātou ngā kaikawe i tētahi momo malaria kei te iwi manu,
either birds or seals still appears inherent behaviour of ko te Leucocytozoon te ingoa. Heoi, e ai ki ngā mātauranga
New Zealand adult female simuliids — and probably the
reason for their name “sandflies”. (haere tonu)
Fauna of New Zealand 68 7
Worldwide, simuliids are notorious for their disease o nāianei, kāore e whakawhitia mai ana tētahi mate e tērā
transmission, in particular onchocerciasis, or river blind- momo namu ki te tangata.
ness, in Africa and South America, but more widely for E mea ana ngā tātaritanga rāpoi ngota, ehara i te mea nō
the nuisance value of their bites, especially in the northern tuauriuri whāioio te nuinga o ngā namu o Aotearoa. Nō roto
North Hemisphere. In Canada, simuliids are celebrated in kē mai i ngā kapinga e 5 o te ao i te tio i te 500,000 tau kua
the “The Black Fly Song” written by Wade Hemworth in mahue ake nei. Ā, e tautoko ana tēnei i ngā rangahautanga
1949. New Zealand has two sculptures featuring simuliids i ētahi atu o ngā mea oreore a Tāne. Heoi anō, arā ētahi
— probably unique in the world — that are remarkably kāwai o te huinga ungulatum he noho ki ngā wāhi teitei,
accurate and most appropriate for the region. One is a ki te wai makariri. Ko te wāhi o Kā Puke Māeroero koirā
set of brass adults in the visitor centre at Milford Sound, tō rātou kāinga whāiti, kua paku neke atu i te 5 miriona
Fiordland, crafted by Elizabeth Thomson in 1991; the tau e tū ana. E tautoko ana tēnei i ngā kitenga rāpoi ngota
other is a giant 2 m long fly suspended on the outside of o tēnei mahi rangahau.
the Bushman’s Cafe, Pukekura, Westland. Ko tētahi pātai ka kaha te uia ake, “He aha te kai a
ngā namu o Aotearoa i mua i te taenga mai o te tangata?”
Kāore e kore ko te huhuatanga o te manu i te mata o te
Contributors Doug and Ruth (nee Heath) Craig were born whenua i ērā wā. Kua tino kore tērā āhua iāianā, kua
in Nelson, New Zealand, and attended their local Colleges. ngaro te pūorooro o te ngahere i ngā manu korihi o te ata.
Doug went to the then Canterbury College, University of Ā, he kekeno anō pea te kai i ngā rā o nehe, inā te tini o
New Zealand, Christchurch in 1959, where, in his first year, te kekeno i ērā wā. Heoi anō, te āhua nei kei te mau tonu
he failed both his major subjects, Botany and Zoology, but te whanonga a ngā uwha kātua o ngāi simuliid mā, kei te
passed well in Physics! Ruth started university a year later kimi tonu pea i ngā toto o te manu tai, o te kekeno i tātahi.
and they met in Botany I when Doug repeated the subject. Ā, koia anō pea i tapaina ai ki te reo Pākehā i Aotearoa nei
Both were doing biology degrees with a view to becoming ko te “ngaro onepū”.
secondary schoolteachers, as the demographic bulge of Puta noa i te ao, e mōhiotia ana te namu mō tana kawe
children from the post-World War 2 ‘baby boom’ was fast i ngā tini mate. Ko tētahi ko te onchocerciasis, i Āwherika
approaching and teachers were in short supply. The New me Amerika ki te Tonga, ko te ‘kāpō awa’ tōna ingoa
Z.ealand Department of Education provided very good kārangaranga. E whakakinongia ana anō hoki te namu mō
post-primary studentships to attend university in return for tana ngau, ā, ko te taha raki o te Tuakoi Raki tētahi tino
teaching an equal number of years. Ruth continued along wāhi i pērā ai. I titoa anō he waiata mōna i Kānata i te tau
that path and taught at Christchurch Girls’ High School. 1949, e Wade Hemworth, ko “Te Waiata Ngaro Pango”
Doug, however, became beguiled by the aquatic insects, te ingoa. E rua ngā whakapakoko o te namu i Aotearoa,
the Blephariceridae, and after finishing his B.Sc. (Hons) he mahi ahurei tonu pea huri i te ao. Me kōrero te tika o
degree in the then Zoology Department, became Vida te hanga o ngā mea nei, me te hāngai ki te rohe kei reira
Stout’s first Ph.D. student and worked on the biology of nei rātou. Ko tētahi, he huinga kātua, he mea hanga ki te
those insects. When Vida took a sabbatical leave, and then kuratea. Kei te whare manuhiri i Piopiotahi tērā, he mea
a leave of absence in Sweden, Bob Pilgrim took over the waihanga e Elizabeth Thomson i te tau 1991. Ko te tuarua,
supervisory details. As far as the secondary school teach- he namu kaitā, e rua mita rawa te rahi, e iri mai ana i waho
ing went, that was paid off by some ‘temporary assistant o te Wharekai Bushman i Pukekura, i Te Tai Poutini.
lectureships’ and buying the remainder out. Doug and Ruth
were married in 1962. Immediately Doug finished his Ph.D. Translation by H. Jacob
in October 1966 (with the now autonomous University Ōtaki
of Canterbury), both left for Edmonton, in Canada; for
Doug this was a sabbatical replacement position in the
Department of Entomology, University of Alberta, and an
opportunity for overseas experience. It was meant to last
for just a single year, but with another year available, and
then a position vacant, Doug was hired as departmental
morphologist. So then with a family of two children
(Jacqueline and Michael), two cats, and a mortgage, the
stay became permanent. Good working conditions and
colleagues helped too! Initially Doug worked with WHO
funding on the embryology of simuliids in relation to
8 Craig, Craig & Crosby (2012): Simuliidae (Insecta: Diptera)
Doug and Ruth Craig at 7th International Congress of Dipterology, Costa Rica, 2010.
their control in Africa where the female of these insects once, Vanuatu twice, and Fiji a number of times. It seemed
transmits river blindness. Ruth became a technician in only natural to eventually do a taxonomic revision of New
Plant Science until daughter Jackie was born and then, Zealand Simuliidae. Having daughter Jacqueline, now an
after a child-rearing stint, became a sessional instructor in anthropologist living in Auckland, and a grand-daughter
the Department of Zoology for a number of years. With there too, made the decision relatively easy.
parents and relatives still in New Zealand, return visits Contributor Trevor Crosby was born in Cambridge, New
were a necessity, albeit expensive with a family, so trips Zealand, and grew up in the central North Island towns of
were restricted to various sabbatical leaves. Stopovers Raetihi and Piriaka (attending Taumarunui High School),
were, however, made in Polynesia, Rarotonga, and Fiji to with a year at Whangaehu (Wanganui Boys’ College). His
collect simuliids, and that resulted in discovery of many family then moved to Karamea on the West Coast, and
new species, which kindled an interest in biogeography of Trevor went to the University of Canterbury, Christchurch
Pacific Simuliidae. Inbetween times a childhood interest (in receipt of a post-primary studentship similar to his
of mucking about in running water was refined, and Doug fellow contributors above) and gained his B.Sc. (Hons)
spent a decade on proximate hydrodynamics of simuliid in Zoology. The opportunity to do a Ph.D. on the simuliid
larvae and other aquatic insects. An earlier, but important Austrosimulium tillyardianum put secondary teaching on
aspect to that was a detailed understanding of the shape and hold (subsequent employment with DSIR became approved
structure of the larvae, garnered from having one of the first service). He spent a year in Wellington as a DSIR science
Scanning Electron Microscopes (SEM) in Western Canada. editor (bulletins and occasional publications), before
Originally full of glowing vacuum tubes — now just cold moving to Auckland to join Entomology Division, DSIR
transistors — that laboratory, established in 1971, is still as curator of NZAC, a position he held until 2009. He
going strong, albeit with shiny new instruments. Doug has been leader of the invertebrate biosystematics group
retired in 1997 after 31 years teaching, having had quite (1980–1983, 1997–2007), and has been involved with the
enough of the changes that are taking place in universities Fauna of New Zealand since its inception, being its editor
nowadays. As an Emeritus Professor and still active with since 1998. Collaboration with colleague Graeme Ramsay
research grants, Doug, with Ruth (now a retired Registered saw the publication in 1992 of the Bibliography of New
Nurse), in the last decade have travelled widely across the Zealand terrestrial invertebrates 1775–1985, and later that
Pacific collecting simuliids for taxonomic revisions. They year the database “BUGS on-disc” was published on CD as
have been as far west as Palau in Micronesia, completely part of New Zealand’s first science CD, which now forms
around all the Polynesian island chains, some more than part of the backbone of the online BUGZ literature website.
Fauna of New Zealand 68 9
Trevor has had expert witness responsibilities in forensic
entomology: his co-authored paper on using entomological
evidence to prove importation of cannabis into New
Zealand was awarded the 1987 Philip Allen Memorial
Award by the Forensic Society of Great Britain; and the
calculation of possible time of death using calliphorid
maggots has assisted a number of New Zealand Police
investigations. Since 1994 he has provided training in
entomology and identification for phytosanitary purposes
in the South Pacific (particularly Vanuatu and Samoa)
and Asia (India, Cambodia, Lao PDR, Myanmar, Viet
Nam, and most recently Brunei). His interest in education
has continued through being a school Board of Trustee
Trevor preparing Fauna of New Zealand 68, 5 June 2012
member: Gladstone Primary (1998–2004), and Western
(Photograph: Birgit Rhode)
Springs College (2001–present), and with his wife Bev he
has run a Dyslexia Parent Support Group since 2000. Bev
and children (Cameron, Karl, and Erin) have supported
his various endeavours, but sometimes remind him of the
family events he has missed while away on consultancies.
Dedicated to the memory of Lionel Jack Dumbleton
(29 December 1905 – 25 September 1976). Despite
being incapacitated by a stroke shortly before he was to
retire in 1970, Jack, with great determination completed
his major work on New Zealand Simuliidae (1973),
which provided a sound taxonomic framework for our
research (see Lowe & Ramsay (1977) for his obituary).
“…no sooner had the sun risen, and we issued from our tent to wash by the river side, than those peculiarly
vexatious pests, the sand-flies (namu), commenced their attacks on our bare hands and feet. The sand-fly is a
small black insect, and swarms in such myriads, that one is never free from their vengeance, if remaining for
a single instant in the same position: whilst sketching, my hands are frequently covered with blood, and their
numbers being inexhaustible, one at last gets weary of killing them.” (p. 21), “The horrid sand-flies attacked us
to-day more unmercifully than ever, and in such clouds that I should imagine them to be a species very nearly
allied to those that constituted the fourth plague of Egypt. (p. 23)”
Account about biting of Austrosimulium australense near the Waikato River on the AK/WO area boundary on
29 September 1844. Angas, G. F. 1847: Chapter I: pp. 21 & 23, Journey into the interior of New Zealand — The
Waikato. In Savage Life and Scenes in Australia and New Zealand Vol. II. London, Smith, Elder & Co.
“The blood-sucking ones are all females: the male is a perfect gentleman, for it is rarely seen and never bites.”
Anon (1961) New Zealand Science Review 19: 75.
10 Craig, Craig & Crosby (2012): Simuliidae (Insecta: Diptera)
Frontispiece. Female adult of Austrosimulium tillyardianum feeding on the foot of DAC
at Roding River, Nelson in 2007. Note the diuretic fluid extruded from the fly’s anus — a
mechanism for rapidly concentrating the blood meal. (Photographer: D. A. Craig).
