by Frank E. French
The North American Dipterists' Society and the Biting Fly Workshop
will meet jointly June 1-4, 2001, at Sul Ross University, Alpine,
Texas. Collecting will be permitted in a large Nature Conservancy
tract in the nearby Davis Mts. and elsewhere in the Chihuahuan Desert
Region in Trans-Pecos Texas. For information and registration contact:
Dr. Frank E. French
Department of Biology,
Statesboro, GA 30460-8042, USA.
Fax: 912 681 0845
by Brian V. Brown
Greetings fellow Dipterists. I have been "volunteered" to
organize the NADS informal conference for the ESA meeting this year.
In order to have titles of talks placed in the ESA program, we need to
have them by the end of this month. Therefore, anyone wishing to give
a presentation please get in touch with me as soon as possible (I am
going into the field on 29 March). Another alternative available to
those unable or unwilling to come up with a title this early, but who
still want to make a presentation, is that we list our meeting in the
ESA program without titles. We could organize our session later, and
distribute the list of speakers via Fly Times in October. Note that
talks would not be listed in the ESA program. Probably we will have to
do a combination of these. For those of you who feel it is important
to be listed in the program, please contact me as soon as possible.
Others will still get a chance to speak.
Dr. Brian V. Brown
Associate Curator of Entomology
History Museum of Los Angeles County
900 Exposition Boulevard
Angeles, CA 90007 USA.
FAX (213) 746-2999
by Scott E. Brooks and Jade Savage
McGill University (Macdonald
Campus), Ste-Anne-de-Bellevue, Quebec, Canada
The 2000 NADS Informal Conference was held at Montreals Palais
des Congrès last December during the Annual Meeting of the
Entomological Society of America. Overall the conference was a
success. In contrast to past meetings, which have typically been held
during the evening, Decembers meeting was scheduled (by the ESA
organizing committee) in the afternoon. Although this was a break in a
long standing NADS tradition, the earlier time slot turned out to be a
good thing as it seemed to result a larger audience. Six talks were
given dealing with dipteran systematics and the use of Diptera in
biotic surveys. Jeff Cumming of the CNC, Ottawa presented a new
cladistic classification of the Empidoidea based on his recent work
with Brad Sinclair. Miranda Smith, a Masters student of Doug
Currie at the Royal Ontario Museum, discussed the results of her
molecular analysis of Simulium s.str. Brian Brown of the
Natural History Museum of Los Angeles County, presented data from 3
years of Malaise trapping ant- decapitating phorids at La Selva
Biological Station, Costa Rica, and provided comments on rarity and
techniques for optimizing sampling. Fiona Hunter of Brock University,
St. Catharines, discussed the results of a survey of the Niagara
Escarpment, which indicates that several families of Diptera can be
used as indicators of water quality. Steve Marshall of Guelph
University discussed and contrasted several Canadian biodiversity
projects with ongoing Neotropical surveys and explained the problems
and opportunities presented by the Costa Rican Sphaeroceridae and
Micropezidae. Brian Wiegmann discussed recent progress in the Diptera
survey of the Great Smoky Mountains National Park and urged everyone
to get involved. After the scheduled talks further discussions were
held on a wide range of topics such as updates on the next NADS field
meeting being held in the Davis Mountains of Texas, announcement of
recent Diptera products fromWashington, and a report on the bizarre
natural history of fergusoninid flies that are now in quarantine and
set for release in Florida. Following the meeting, many dipterists
gathered at Pub St. Paul in Old Montreal for refreshments and further
discussion. Thanks to all the speakers and participants for making the
2000 meeting a great one! The 2001 NADS Informal Conference, in San
Diego, California, is being organized by Brian Brown.
by Norm Woodley and Chris Thompson
Systematic Entomology Lab.,
ARS, USDA, Smithsonian Institution, Washington, D.C.
The Department of Entomology, Smithsonian Institution, passed away
peacefully in its sleep last month. It was only 38 years old. At the
same time, the Department of Systematic Biology was born and is now
under the acting leadership of Scott Miller. The Diptera program in
Washington remains the same. The regulars are still here (Ray Gagne,
Chris Thompson, Wayne Mathis, Norm Woodley, and Allen Norrbom, with
Holly Williams, Nancy Adams and Lucrecia Rodriguez downtown; Sonja
Scheffer at Beltsville; Rick Wilkerson, Dan Strickman, and Mark
Potter, with Tom Gaffigan and Jim Pecor at the Museum Support Center).
Unfortunately Kyle Apigian left for graduate school at UC Berkeley and
has not been replaced.
Downtown, we have two long-term associates this year: Amnon
Freidberg from Tel Aviv is spending his sabbatical here studying
various acalyptrates and Steve Gaimari got a 3 year NSF grant to study
the higher phylogeny of Lauxanioidea. At the Walter Reed
BioSystematics Unit, there are two post-doctoral fellows: Dina
Fonseca, who is doing molecular systematics on mosquitoes, and Anice
Mureb Sallum from Brazil, working on Anopheles systematics. Both Peter
Hibbs and Alessandra Baptista are still struggling along with their
PhD work. Isaac Winkler from Brigham Young will join the group as a
new PhD student this Fall, working with Wayne and Sonja. Also Paul
Arnaud and his wife, Madeline, are making extended visits now to
Washington that his collection is here. Lloyd Knutson spent a few
weeks in January working on Sciomyzidae. Ilam Yarom is coming in May
and will stay until the Fall studying with Amnon and Steve. We are
also expecting a stream of short term visitors, including Marc de
Meyer, Marion Kotrba, et alia. But remember we always can find space
for one more!
The NADS publication program is slow, but is progressing. MYIA 11, a
World Catalog of the Stratiomyidae by Norm Woodley, will be out by the
time this newsletter is released (publication date, 15 April). MYIA 6
which will be devoted to reviewing the life and accomplishments of
Curt Sabrosky and Bill Wirth should be out soon, too. Volume 2 of the
Diptera Data Dissemination Disk is being finalized and will be out by
The BioSystematic Database of World Diptera is up and running at the
Diptera WWW site. The Nomenclator has been online since last August.
Various support pages were added last December, and version 2 of the
Dataset for Nomenclator now includes more than 200,000 names. Coverage
includes the data from the various regional Diptera catalogs for most
families (the biggest, Tipulidae, Tachinidae, etc., are still
unfinished) and all the names from the past twenty years. A grant from
the Schlinger Foundation allowed us to purchase in digital format all
the new species reported in the Zoological Record for the last 21
years (volumes 115-136). Still there are some 40,000 species group
names to be entered. See the status page (www.diptera.org/names/bdwdss.htm)
for details. Selected specialists are now being contacted about
contributing family treatments for the next version of the database.
We hope to have another MYIA containing these family treatments out by
the end of the year. Steve and I have already finished the
Eurychoromyiidae and Marc de Meyer and I are working on the
Mormotomyidae, so expect a MAJOR contribution soon! In a related
effort, the Walter Reed BioSystematics Unit has put online a new
catalog of the mosquitoes of the World (wrbu.si.edu/www/culicidae/cataloggeneraentry.html).
A printed version is planned.
Under the leadership of Steve Gaimari, the Diptera group launched
its first major expedition. In early March, Wayne Mathis, Amnon
Freidberg, Frank Parker and Steve left for Bolivia to look for the
World's rarest fly, Eurychoromyia mallea Hendel. They will
remain until the end of the month. Allen Norrbom, Brian Brown and
Steve Marshall will replace them and collect through April. At press
time, the word from Bolivia was that collecting was good, but no Eurychoromyia
Finally, in the last six months there has been a lot of activity
concerning taxonomy and biodiversity. Last fall a new organization,
ALL Species was established in California to speed up the complete
enumeration of ALL (All Life List) organisms. See the fall issue
of the Whole Earth magazine or go to their WWW site (www.all-species.org)
for more details. Science magazine (29 September 2000) was devoted
to BioInformatics for Biodiversity, with E. O. Wilson calling for
a Global Biodiversity Map. The Convention on Biodiversity has launched
the Global Taxonomic Initiative (GTI) (www.biodiv.org/programmes/cross-cutting/taxonomy/default.asp).
Last month in Montreal, the Global Biodiversity Information Facility
was established. So, with the smell of new and more money, the bird
and butterfly people are racing to again be ahead of the pack. This
time, however, I believe the Dipterists should stand tall and take
their rightful place in front. And to do this we need only to educate
people about flies. Flies are the among the least known groups of
biodiversity, although we already know that they represent more
than 10% of the known World Biota. They are of critical importance
to Man, many being important pests of crops, others being vectors
of disease. However, many are beneficial, pollinating flowers, serving
as model systems for genetics and medicine, and as biological control
agents of pests and weeds. Compared to the birds and butterflies,
flies are more speciose, more important, and just as pretty! So
spread the message!
by Jeff Skevington
Canadian National Collection of Insects,
After spending nearly four years at the University of Queensland in
Australia I have landed back in Canada at the CNC. While in Australia
I completed a Ph.D. program under the supervision of David Yeates. My
work there had three focuses: 1) To investigate the utility of two
mitochondrial genes (12s and 16s rDNA) for developing a phylogeny of
the Syrphoidea (Skevington & Yeates, 2000). 2) To examine
phylogenetic relationships of members of the tribe Eudorylini
(Pipunculidae, Pipunculinae) using morphological characters. Two
hundred and fifty-seven species of Eudorylini from all biogeographical
regions and all known genera were examined. Sixty species were
included in an exemplar-based phylogeny for the tribe (Skevington &
Yeates, in press). 3) To revise the Australian species of Eudorylini
within this phylogenetic context. Five genera including 81 species of
Eudorylini are represented in the Australian fauna. Only 16 Australian
species were previously described in this tribe. The largest
publication resulting from this work should be out later this year
(Skevington, in press), and manuscripts on the other genera are nearly
complete. In the context of the above revisions, patterns of
hilltopping and phenology were explored for genera and species of
My work in Ottawa under the supervision of Jeff Cumming is supported
by a Natural Sciences and Engineering Research Council of Canada
Fellowship which is held through McGill University. Over the next two
years I intend to: 1) Extend my research on relationships of lower
Cyclorrhaphan flies using morphological and molecular characters. 2)
Continue my work on systematics of Australian Pipunculidae while
getting back into Nearctic pipunculid revisions. 3) Learn more about
Conopidae while working on the Costa Rican Diptera project.
If you have any pipunculids or tales about great hilltop collecting
sites, I am always interested in examining more material and finding
out about new collecting areas. Also, if you are planning on going to
Australia for the 5th International Congress of Dipterology and need
some ideas for field trips let me know and I may be able to give you
some tips. Any general questions about the Congress may also be
directed to me. The conference runs from the 30th of September to the
4th of October 2002 (more information is available from
My contact details are: Dr. Jeff Skevington, Diptera Unit, Systematic
Entomology Section, ECORC, Agriculture and Agri-Food Canada, Ottawa,
Ontario, K1A 0C6, CANADA, Phone: 613-759-1178, Fax: 613-759-1927,
web site: http://res2.agr.ca/ecorc/staff/skev-j.htm
Skevington, J. H. (in press). Revision of
Australian Clistoabdominalis (Diptera: Pipunculidae).
Skevington, J. H. & Yeates, D. K. (2000). Phylogeny of the
Syrphoidea (Diptera) inferred from mtDNA sequences and morphology with
particular reference to classification of the Pipunculidae (Diptera).
Molecular Phylogenetics and Evolution 16(2), 212-224.
Skevington, J. H. & Yeates, D. K. (in press). Phylogenetic
classification of Eudorylini (Diptera, Pipunculidae). Systematic
by Mike Irwin
Dept. Natural Res. & Environmental Sciences,
Univ.of Illinois, Urbana, Illinois, USA
Here is a partial listing of the publications and other products our
therevid team, sponsored by NSF and the Schlinger Foundation, has put
out during the past two years.
In Referred Journals:
Gaimari, S.D., and M.E. Irwin.
2000. Phylogeny, classification, and biogeography of the cycloteline
Therevinae (Diptera: Therevidae). Zool. J. Linnaean Soc. (London) 129:
Gaimari, S.D., and M.E. Irwin. 2000. Revision of the mexicana-group
of Ozodiceromyia Bigot (Diptera: Therevidae). Proc. Entomol.
Soc. Wash. 102: 561-600.
Gaimari, S.D., and M.B. Mostovski. 2000. Burmapsilocephala
cockerelli, a new genus and species of Asiloidea (Diptera) from
Burmese amber. Bull. Nat. Hist. Mus. (London), Geology Series 56:
Metz, M. A., and M. E. Irwin. 2000. A new therevid genus from
Dominican amber, the revisions of Lindneria Kröber and
Insulatitan Gen. Nov. and their phylogenetic relationships
with closely related extant Therevinae (Diptera: Therevidae). Annals
of the Entomological Society of America 93: 977-1018.
von Tschirnhaus, M. Irwin, M. Hauser, N. Evenhuis, and T. Pape.
2000. Provisional checklist of the Agromyzidae, Therevidae,
Mythicomyiidae, Sarcophagidae and Stratiomyidae (Diptera) of the
Brandberg Massif, Namibia. Cimbebasia Memoir 9: 383-384.
Webb, D. W., and M. E. Irwin. 1999. Revision of Tabuda
Walker and Tabudamima Irwin & Lyneborg, with the
description of a new genus Incoxoverpa Webb and Irwin
(Diptera: Therevidae: Therevinae). Ann. Entomol. Soc. Am. 92: 644-674.
Wiegmann, B. M., S. C. Tsaur, D. W. Webb, D. K. Yeates, and B. K.
Cassel. 2000. Monophyly and relationships of the Tabanomorpha
(Diptera: Brachycera) based on 28S ribosomal gene sequences, Annals of
the Entomological Society of America 93: 1031-1038.
Winterton, S.L. 2000. Phylogenetic revision of Acupalpa Kröber
(Diptera: Therevidae: Agapophytinae). Insect Systematics and Evolution
Winterton, S.L. & Irwin, M.E. 1999. Laxotela - a new
genus of Therevidae (Diptera) from Australia. Entomologica
Scandinavica 30: 299-310.
Winterton, S.L., Irwin, M.E. & Yeates, D.K. 1999. Systematics of
Nanexila gen. nov. (Diptera: Therevidae) from Australia.
Invertebrate Taxonomy 13: 237-308.
Winterton S.L., Irwin, M.E. & Yeates, D.K. 1999. Phylogenetics
of the Taenogera Kröber genus-group (Diptera:
Therevidae), with descriptions of two new genera. Australian Journal
of Entomology 38: 274-290.
Winterton, S.L., Merritt D., O'Toole, A., Irwin M.E. & Yeates
D.K. 1999. Morphology and Histology of the spermathecal sac, a novel
structure in the female reproductive system of Therevidae (Diptera:
Asiloidea). International Journal of Insect Morphology and Embryology,
28(4): 273- 279.
Winterton, S.L., Skevington, J.H., Irwin, M.E. & Yeates, D.K.
2000. Phylogenetic revision of Bonjeania Irwin & Lyneborg
(Diptera: Therevidae). Systematic Entomology 25: 295-324.
Yang, L., B. M. Wiegmann, D. K. Yeates, and M. E. Irwin. 2000.
Higher-level phylogeny of the Therevidae (Diptera: Insecta) based on
28S ribosomal and elongation factor 1a gene sequences. Molecular
Phylogenetics and Evolution 15: 440-451.
Yeates, D. K. and B. M. Wiegmann. 1999. Congruence and controversy:
Toward a higher-level phylogeny of the Diptera. Annual Review of
Entomology 44: 397-428.
Books, Theses, Dissertations & other one-time publications:
S. L. 1997. Systematic revision of Nanexila gen. nov. (Diptera:
Therevidae) from Australia. Masters thesis, University of Queensland.
Irwin, M. E. 1997. Therevidae. In: Solís, A. (ed.), Las
Familias de insectos de Costa Rica. INBio.
Gaimari, S. D. 1998. Phylogeny, classification, and biogeography of
the Cycloteline Therevinae (Diptera: Therevidae). Ph.D. Dissertation,
University of Illinois at Urbana-Champaign.
Power, N. 1998. Spatial and temporal distribution of Therevidae in
southeast Queensland. Honours Thesis, University of Queensland.
Holston, K. C. 1999. Answering the nomenclatural challenge with a
systematic database of Thereva names (Diptera: Therevidae). Master's
Thesis, University of Illinois at Urbana-Champaign.
Winterton, S. L. 2000. Endemic radiation of a diverse clade:
phylogenetic revision of Agapophytinae subfamily nov. (Diptera:
Therevidae) from Australasia. Ph.D. Dissertation, University of
Yang, L. 2000. Molecular phylogenetics of the Therevidae and their
position among the families of the Asiloidea (Insecta: Diptera). Ph.D
Dissertation, North Carolina State University.
Websites or other internet sites reflecting this project:
The family Therevidae has been on the WWW since April 1996 with
periodic updates and additions. The site
details our PEET research, profiles therevid research team members,
provides minutes of meetings, and recounts therevid collecting
expeditions. Therevid MANDALA became searchable via the WWW in January
2000 at http://pherocera.inhs.uiuc.edu/index.htm.
Users are guided to find information via various groupings of queries.
The site uses live databases for searching (except when site
maintenance is underway) so current data may be reflected as it is
updated or entered. The site also provides metadata information about
Mandala and invites guests to sign in at our guestbook. ASC features
therevid PEET project in gallery. In honor of the NSF's 50th
anniversary, the Association of Systematics Collections sponsored a
gallery of NSF funded projects.
The projects have one page summaries and then links to websites for
the projects. Ours includes the main home page, links to participants
in the research, and to Mandala, our database for cataloging
specimens, nomenclature, and literature.
by Rudi Mattoni, Jeremiah George and Ken Osborne
Geography, UCLA, Los Angeles, California
Rhaphiomidas terminatus abdominalis (Delhi Sand giant
flower-loving fly - DSF) is the only dipteran listed as an endangered
species under the federal Endangered Species Act. The species is
restricted to a few fragments of highly degraded habitat, but its
future viability depends on a single ca 100 ha site that is the only
site with sufficient size and overall diversity to offer any hope for
carrying the species into the future. This core site supports the sole
semblance of the Delhi dunes ecosystem that historically covered about
12,000 ha. In addition to the DSF there are a number of rare
arthropods associated with the core site, a dense population of the
restricted Los Angeles pocket mouse, and several plants and other
vertebrates representing desert adapted species. The Delhi sands
provide the sole cismontane habitat for the aggregate.
For your background information we have been running a standard ca 3
km transect sampling most of the site, starting in 1996. We scored the
following numbers since then:
||N samples (present)
Making reasonable assumptions of sighting efficiency (20%), equal
sex ratio, life span (4 days), and interpolating time between samples,
the total population size of this core habitat is >>1000. The
methodology and calculations follow the Pollard transect walk
technique. By comparison all other occupied sites, which now number
about ten, have populations of <<100. The unequivocal message
these results convey is that the only viable option to maintaining the
DSF depends on conserving the entire core habitat.
No focused effort has been made to preserve the core habitat. If
measures are not implemented prior to further development pressures,
the only significant remaining piece of the Delhi sands ecosystem will
be history. Several of the smaller sites occupied by DSF, all less
than 10 ha, deserve protection as well to assure potential higher DSF
From the long term evolutionary perspective the only other, and
nominate, subspecies (Rhaphiomidas terminatus terminatus) has
been extinct for over three decades. It was found in sandy areas along
the Los Angeles river and the El Segundo sand dunes. The sister taxon,
Rhaphiomidas trochilus, which Rogers and Mattoni regard as a
subspecies of terminatus, was recently found by Ballmer on a
small site in the San Joaquin valley. The (sub)species was thought
extinct for over fifty years prior to Ballmerís discovery.
We urge all Dipterists to join us in urging the U. S. Fish and
Wildlife Service to concentrate its efforts on preserving the umbrella
DSF and its habitat.
Rudi Mattoni, Dept. Geography, UCLA (email@example.com)
George, Lepidoptera Research Foundation (firstname.lastname@example.org)
by Doug and Ruth Craig
Department of Biological Sciences,
University of Alberta, Edmonton, Canada
Why did we go?
Reconstructed phylogenies, based both on
morphological and molecular information, are now available for the
Polynesian black fly subgenus Inseliellum, and will provide
the underpinnings for a biogeography of this important group of
aquatic insects. However, there is lack of data for many species. For
the molecular work, only 23 of 48 now known species were available. Of
particular importance was lack of material from the Marquesas and
Austral Islands that have basal species, and from Raiatea which has a
minor species radiation in higher clades. The morphological
reconstructed phylogeny, although including 32 of the then known 41
species, was based mainly on larval structures, because of lack of
data from the pupae and male adult genitalia. Larval simuliids exhibit
considerable homoplasy and obtaining male genitalia for which
homoplasy is reduced, will strengthen further phylogenetic analysis.
To remedy the inadequate taxon sampling of the molecular analyses and
lack of information on simuliid male genitalia, Ruth and I spent five
weeks (9th Oct. - 14th Nov., 2000) in Polynesia, with the main
objective to obtain larvae from the above islands and to rear pupae to
obtain male adults.
The trip was initiated by family
business in New Zealand, September, 2000, and on the outward leg from
Edmonton a few days were spent at the Bishop Museum, Honolulu. There
the opportunity was taken to examine the large holdings of Pacific
Simuliidae. One surprise was a single female specimen of
Inseliellum from Tonga - quite the range extension, and a great
excuse to eventually visit Tonga. Copies of hard-to-get maps of the
Marquesas Islands were also obtained. Hospitality by Neal Evenhuis and
Ron England was up to the usual standard. On the return trip from New
Zealand, we visited seven Polynesian islands.
A few days in Tahiti first of all to make contact with the Louis
Malard Medical Research Institute and to renew old acquaintances, and
obtain other maps. We began the collection of simuliids with a stiff
eight hour hike up the Fautaua Valley behind Papeete. This is the
'Forbidden" Valley, since permission is needed to enter the
valley which is a water catchment area for Papeete. Needless-to-say,
being as unfit as we were at that time, the trip made for an
interesting next few days. Collecting and the scenery were excellent.
From Tahiti we went directly to the Marquesas, flying into Nuku
Hiva. Surprise! This place is not what one might expect of a tropical
island. There was no greenery at all to be seen at the airport on the
north- western corner. Indeed, there was a drought throughout
Polynesia, serious enough that tourists were being denied access to
Bora Bora. Collection of the seriously anthropophilic S. buissoni
was difficult because we found it hard to find streams that had
sufficient water. That did not stop the few females from finding us!
However, they were not as bad as they might have been and application
of repellent worked well. Kept the mosquitoes off too. But, it was not
difficult to imagine what living on Nuku Hiva would be like when the
biting rate is up as high as some 18,000 per day (estimated by Y
Schan). Then, there are the ceratopogonids!
One point to be made about Nuku Hiva is that the drive from the
airport to main town of Taiohae is not to be missed. It is only some
35 km long, but takes about 2 hours, even in a serious
four-wheel-drive vehicle. Do it as a life experience, but under no
circumstance consider doing it if it has rained! Take the helicopter
and pay the $85US shot!
Hiva Oa, to the south, is in many ways similar to Nuku Hiva, but
there is not the horrendous drive from the airport to the town of
Atuona. Here, while very dry, there was more water and we hit the
cascades looking for S. adamsoni. Not one larva of any black
fly species was found on the cascades. Indeed, all the larvae were on
leaves, not rocks. All of which does not help with the problem of why
S. adamsoni shares derived characters with the larvae of rock
loving, cascade-dwelling species in Tahiti?? Hiva Oa was pleasant and
neither S. adamsoni nor S. gallinum bite humans. We
did not get to Fatua Hiva, the southern-most island, because of the
cost of hiring a boat and the lack of guarantee of getting anything
So, back to Tahiti for a night to replenish the alcohol and glacial
acetic acid and out west the next morning to Raiatea. Here we hoped to
obtain a number of poorly known species. However, all the lower
altitude streams yielded up were the common garden species of S.
malardei and S. lotii. It was not until Ruth and I did
some serious bush whacking to get to higher altitude that we managed
to get some of the oviceps-group. A major hike up the Temahani
Plateau got us up into the clouds and also some larvae of S.
castaneum and S. bogusium - both basal to their individual
clades. By this time we are beginning to feel that we have been
working too hard and did actually spend a day swimming and lying
around the place on the beach.
So, after being a bit refreshed, back via Tahiti and south to the
Austral Islands, Rurutu first. This is an unusual island in that it is
about 11 million year old and normally would have eroded to below sea
level by now, but 4 mya there was renewed vulcanicity and it was
uplifted and now has 30 metres plus high, fossil coral (= makatea)
cliffs surrounding it. This might account for the fact that absolutely
no other aquatic invertebrates, or fish, were seen in the streams
sampled. However, Simulium rurutuense larvae were there by the
million. Because of some travel problems with Air Tahiti changing
their flight schedules, we spent two more days than necessary in
Rurutu and were only granted one full day on the next eastern-most
island, Tubuai. Plenty of time to show that there were no Simuliidae
in the very small streams on that very small island (twenty minutes to
drive completely around!).
So, back to Tahiti and now quite a bit fitter, we went on a major
trip up the Vaitepiha River on the smaller part of Tahiti. A
wonderful, completely unaltered river, but being very dry, few
cascades of any use. In desperation then we collected on a seep and
have perhaps found the elusive larva of S. cheesmanae. Or, if
molecular studies don't confirm that, certainly a new species within
the "castaneum" clade. Also, some more of the new
species recently described in the "hirticranium"
group. Moral - never turn your nose up at a seep!
Well, Ruth and I discovered we were not as fit as we had thought,
but a day resting helped. Then for me it was off to deliver a seminar
at the Malard Institute on the work I had done there over the last 20
years. No, not in French - almost without exception the scientists and
technician there spoke English.
A day trip via ferry to Moorea was enough to collect from a couple
of sites there for fresh material - nothing new. Indeed, apart from
the maybe new material from the Vaitepiha River, nothing new turned up
anywhere - bottom of the barrel perhaps??
Then home here to Edmonton. Not a bad trip all in all and if I had
to rank it out of 10, I'd have no trouble assigning it a 9. Why not
the full score? Well, that would have taken about another $10,000US to
hire boats in the Marquesas to get to other islands. Maybe some other
time. But, perhaps it should have a 9.5 - already Deirdre Joy and Mike
Spironello are making good use of the chromosomes and DNA from the new
Visiting some 50 sites in Polynesia gave Ruth and I a broad over
view of the condition of running water habitats on the islands. During
the period of the trip there was a drought in Polynesia, sufficiently
serious that water restrictions were in force on Bora Bora. It was
noticeable that most streams and rivers were in their base flow mode.
Indeed, in the Marquesas Islands and elsewhere it was often difficult
to find streams with sufficient water to provide suitable habitats for
One noticeable aspect of walking up streams and rivers in Polynesia
is that, on the smaller islands, almost every stream has been altered
by human activity. This is generally in the form of a small barrage
(dam) and a pipe conducting water away. But in many instances it was
the presence of the pipe that provided walking access to the higher
reaches of the running water. In Tahiti-nui, such barrages are common
even on larger rivers, and water use and draw-down are considerable.
But, pristine rivers are still to be found on Tahiti-iti, in
particular the Vaitepiha River.
In general the streams and rivers of Polynesia appear to be in
reasonable health. On some islands running water will be a limiting
factor in any economic expansion. Rurutu is a good example, where
during the period visited, there was little running water and what
there was, was fully utilized.
One site was of particular interest. High on the Temehani Plateau,
Raiatea, a tributary of the Vaihuaru River, this locality is truly
unique. The altitude is such that vegetation is very unusual, as is
the ground- water source of the river and cascade. Unfortunately,
there is a considerable amount of human impact - mainly carving in the
soft rock, with vegetation removed and garbage left. This area would
be worthy to consider as a nature reserve, or "national park".
A formal report on this expedition, with maps, illustrations of
sites sampled, lists of habitat characteristics and species taken, is
deposited in the Bishop Museum. Tonga - here we come in 2002!
by Bill Barr
Edmonton, Alberta, Canada
Fly Times Issue 11 (October 1993) contains a note from Brian
Brown recommending hexamethyldisilizane (HMDS) for drying small flies.
A few notes of caution follow. The Materials Safety Data Sheet
(MSDS) for this chemical states that its flash point is 14 degrees
C, well below room temperature, making it quite volatile. It is
also highly flammable. "Vapor may travel considerable distance
to source of ignition and flash back" and it is advised to
"Use non-sparking tools" (wood, plastic), and "Store
in a cool dry place." HMDS is incompatible with strong oxidizing
agents and with strong acids. The MSDS does not mention: 1) best
storage is probably in an explosion-proof refrigerator, 2) work
with the chemical in a class A fume hood, 3) work with only small
volumes of HMDS, 4) keep the chemical on ice when it is out of the
refrigerator. HMDS is toxic, and the target organs are the nerves,
so I wear goggles, respirator, and gloves.
by D.L. Deonier
P.O. Box 1625 Pittsburg, KS 66762, USA
With the vast majority of the Deonier Collection of Ephydridae
deposited at the USNM several years ago, I have decided to apportion
the remnant, including several paratypes, between the following
collections: Academy of Natural Sciences of Philadelphia, California
Academy of Sciences, University of Minnesota Insect Collection, and
the Cleveland Museum of Natural History.
from Ian Walker
Department of Biology, Okanagan College,
Kelowna, B.C., Canada
The most recent edition of the Chironomus newsletter may be downloaded
from the Chironomid Home Page at: http://www.ouc.bc.ca/eesc/iwalker/intpanis/
Sturgis McKeever and Frank E. French have put together a very nice
webpage, showing beautiful pictures of these small flies, as well as
interesting evidence that they transmit trypanosomes between male
frogs. Check it out!
by Dieter Schlee
Staatliches Museum für Naturkunde,
[The following is a translation of "Schlee, D. 1978. In
Memoriam Willi Hennig 1913-1976 Eine biographische Skizze.
Entomologica Germanica 4:377-391" through the kind permission of
the publishers, Dr. E. Naegele, E.Schweizerbart'sche
Verlagsbuchhandlung, Science Publishers (http://www.schweizerbart.de).
We have not included the bibliography or the three figures (see
captions at the end) from the original paper. Eds.]
Abstract. An outline of Professor Dr. Willi Hennig's life history
and an abstract of his work on zoology and phylogenetic systematics,
etc. are compiled, including some background information originating
from his family and from the author's experience with him in his
Department of Phylogenetic Research at Ludwigsburg.
close scrutiny, the much-maligned systematics turns out to be a very
important science of its own. While its sole role in zoology was
formerly one of classification, in the course of time it was given the
great task, supported by comparative anatomy, palaeontology,
experimental studies, etc., of investigating the relationships of
animals and their development based on the theory of evolution. The
ultimate aim is a system which represents a 'natural history of
creation' in the most succinct form possible."
From a high school essay by the 18-year-old Willi Hennig.
Professor Dr.-phil. Dr. rer.nat.h.c.
Willi Hennig died during the night of November 4, 1976 in the 63rd
year of his life. A sudden cardiac arrest took him without warning
from a life that was rich in human terms and extremely productive in
scientific terms. Until the evening he had worked on the new edition
of his book "The larval forms of the Diptera" with his usual
intensity and had then reminisced with his family about their last
holiday in Crete.
He will be sorely missed by his family - his wife, his three sons
and their families; by the Department of Phylogenetic Research at
the State Museum of Natural History in Stuttgart and Ludwigsburg
which he headed since 1963; by a wide circle of international scholars
interested in phylogenetic systematics whose spokesman he was; by
many dipterologists all over the world who have lost in him the
expert with the most comprehensive knowledge; and by many researchers
and students who received valuable suggestions from his wealth of
knowledge: as co-editor1 of the "Zeitschrift für Morphologie
der Tiere", "Zoomorphologie", and "Das Tierreich",
as consultant for "Entomologica Germanica" and "Systematic
Zoology", and as author of the textbooks "Taschenbuch
der Zoologie", "Stammesgeschichte der Insekten",
the chapter "Diptera" in the "Handbuch der Zoologie",
as reviser of 14 families (among them the Muscidae and Anthomyiidae)
in the series "Die Fliegen der palaearktischen Region"
edited by E. Lindner and as author of numerous publications on phylogenetic
systematics and many specialized investigations. His published works
adds up to more than 9,000 pages and thousands of drawings he had
made himself; but even these monumental figures cannot adequately
express its scientific importance--we have to include the unusually
broad spectrum and the deep exploration of his fields of study and
to point to numerous other publications which were influenced by
his suggestions in order to give some idea of the impact of his
2. Chronology 1913-1932 and the Principles of the Phylogenetic
He was born in Dürrhennersdorf near Zittau in
Saxony on 20 April 1913 as the eldest of three sons--his father was a
railway official--and from his early years onwards his family
encouraged him to be active and to develop perseverance. In addition
to attending the village school, he was tutored in English, Latin, and
French by a retired physician. His schooling was so effective that he
was allowed to skip one grade when he transferred to grammar school in
Dresden; it can be assumed that his later interests had their roots
in his activities during these years. He not only collected beetles
and butterflies and started a herbarium (his other favourite pastime
was going for a ride in a locomotive, as he fondly recalled), but also
visited regularly the Zoological Museum in Dresden during the years he
attended grammar school (with emphasis on modern languages) in
Dresden. There he also met Dr. W. Meise and Dr. Klaus Günther.
How deeply Willi Hennig had already adopted the phylogenetic
approach and become familiar with the broad field of (comparative and
analytical) zoology at that time can be seen with surprising clarity
from an essay he wrote as assignment in the subject "German"
and entitled "First essay. 4 May 1931. The position of
systematics in zoology." Willi Hennig had just turned 18 at that
time. Fortunately, the original of this essay--written in Gothic
script in an exercise book--has survived the subsequent turmoil ,of
the war and the postwar period and is still in possession of his
family. It has been published posthumously: Hennig (1978).
In his essay he treats mammals, marsupials, birds, butterflies,
bivalves, Artemia, etc., some of which accompanied by
illustrations, provides comparisons of the systems of the entire
higher zoological categories from Aristoteles to Siebold-Leuckart, and
deals with comparative morphology, palaeontology, zoogeography,
behaviour, etc., all of which reveals his wide interests and his
occupation with zoological literature.
This essay shows his enthusiasm for systematics in terms of phylogeny,
his almost impatient need to emphasize its general importance, -to
fight against improper superficiality, to give reasons for various
misinterpretations, etc.; and it also expresses his views that relationships
and systematics are identical ("...clarification of relationships
and thus systematic affinity ...."). He gives several examples
of convergence (in a wider sense, of course, since he does not differentiate
between convergence s-str. and parallelism) being responsible for
false phylogenetic conclusions; and that this can be verified by
testing is for him an important realization. Of course, we cannot
expect to find, in this school essay, the definition of the term
"synapomorphy" which he coined much later. However, its
meaning, i.e. common possession of a unique character (which cannot
be proved to have evolved several times, etc.) as evidence of relationship,
is already foreshadowed in the following example.
"Most recently, similarities in certain
modes of behaviour have been used to clarify relationships and thus
systematic affinities: Heinroth believes that he can infer a closer
relationship between Monticola saxatilis L. and the redstarts
because of the characteristic and unique tremor of the tail which is
common to both."
The two words "characteristic" and "unique" were
not emphasized in the original, but have been underlined here to
indicate that he had searched for evidence of relationship even then
in terms of the succinct criterion which he used later (e.g. Hennig in
Hennig & Schlee, in press), namely "a particular character
state shared by all members of the group" and "not present
in any taxon outside the group."
Although he did not universally apply this kind of argumentation in
the essay, it is nevertheless interesting to note that he was aware of
it--and that he used the example as a kind of working hypothesis (in
fact, he should have cited additional characters, "conflicting
evidence", etc. both in support and in rejection of Heinroth's
interpretation). Incidentally, Willi Hennig did not mention that the
use of common primitive characters as evidence of relationship may be
deceptive; instead he emphasized the dangers of convergent evolution.
The exercise book with the 29 page essay provides further clues: it
reveals that his approach to writing, his style, his argumentation and
presentation had already been developed during his school days: Like
in his later manuscripts, the sentences must have poured forth in a
rapid stream and without requiring any subsequent editing. Some
features uncommon in a school essay [abbreviations, deletions, parts
of words squeezed into a line] seem to suggest that a sentence had to
be completed as fast as possible to make room for the next thought.
His flowing style which casually juxtaposes different points of view
or themes and his preference to disregard any formal structuring are
already evident. The use of clear ink drawings for illustration was
another practice dating back to that time. No doubt he wrote the text
without prior draft (as he would do later when key words and index
cards were often all he needed as aid to memory).
[This may explain why he left the deletions in the text, why he
placed a table with explanations as a three-page insertion within
a single sentence, separated by two commas, and why he wrote "German
essays" on the cover of his exercise book, with the title of
the essay inside, although the exercise book accommodated only that
one essay.] Incidentally, his essay was marked "very good"
and received the following comments from his teacher (Schm = Schmidt?):
"This essay is quite a creditable attempt at giving a critical
treatment of the specific field of study of the author. It is not
merely concerned with compiling facts but occasionally offers some
original ideas as well."
The first paper by the 19-year-old W. Hennig, with W. Heise as
co-author, on the snake genus Dendrophis appeared as early as
1932. A continuation of the paper, again written by these two authors,
was published in 1935.
W. Hennig's interest in bird (e.g. in the above-mentioned school
essay) and reptile taxa may very well be significant, all the more so
since he was joined by M. Heise who is well known as an ornithologist.
W. Hennig regarded birds as part of the group Reptilia and concluded
that Crocodylia and Aves are sister groups--while, for example, E.
Mayr considered this to be absolutely impossible [he called it "absurd"
although he did not question the characters shared by Crocodylia and
Aves and their interpretation as synapomorphy; see also Hennig's
(1974: 289) reply to Mayr].
1932-1939: Student Days and First Positions and Publications
After graduating from grammar school (1932) with excellent
grades (except for physical education), he started his studies in
zoology, botany, and geology at the University of Leipzig. As in his
school days, he was given special status: he did not have to meet the
general requirement stipulated by the Director of the Zoological
Institute, Mr. Meisenheimer, that the major laboratory could only be
attended after completion of the two-semester basic course. He was
permitted to attend the laboratory in his first year since he had
already acquired the necessary background knowledge during his grammar
school days. Intensive work enabled him to complete his thesis
(Contributions to the study of the copulatory apparatus of the
cyclorrhaphan Diptera) and his studies by April 1936, i.e. when he had
reached age 23.
At that time he had already published eight more papers, mainly on
Diptera, among them "Structure and relationship of the Kerguelen
fly" (1934), but also, for example, a 68-page revision of the
genus Draco (Agamidae); a total of 500 pages including his
After a brief period as a trainee at the State Museum of Zoology in
Dresden, he joined the German Entomological Institute of the
Kaiser-Wilhelm-Gesellschaft on I January 1937, initially under a grant
from the German Research Association and subsequently, as of 1939, as
a staff assistant.
By 1939 the list of his publications comprised as many as 41 titles
with a total of 1065 pages. These papers dealt mainly with taxonomy
and the copulatory apparatus of widely different families of Diptera
(among them the first seven families for E. Lindner's "Flies of
the Palaearctic region"), but were also on subjects like "On
some regularities of geographic variation in the reptile genus Draco
L.: "parallel' and .. convergent" race formation", the "Problem
of classifying higher categories", and "On the question of
the systematic position of Braula...", on beetle larvae
and the first fossils in amber (1938: Diptera: Rachiceridae; 1939:
1939-1945: Military Service and Draft of the "Principles of
a Theory of Phylogenetic Systematics'
He received a shortened
basic military training in the infantry starting in the winter of 1938
until the spring of 1939 and was conscripted when the war broke out.
He served as an infantryman in Poland, France, Denmark, and Russia
where he was wounded in 1942 and sent to several field hospitals. He
was given a six month "working leave" in Berlin and later
received instructions from the Medical Academy of the Army in Berlin
to work in the field of malaria control in Greece and Northern Italy
(specifically in Venice and environs). He continued this work not only
until the end of the war but also during the time he was in [British)
He wrote the draft of his fundamental work on the "Principles
of a theory of phylogenetic systematics" in the period between
the end of the war and his return home. It was published in 1950 (see
below) and the original manuscript is in possession of his family
(Fig. 1). He wrote it in an Italian notebook, about 21 x 31 cm in
size, with a heavy cardboard cover. Although it does not contain a
title, it begins with a two-page table of contents (which was later
edited only slightly for publication), followed by the flowing text
which for the most part fills almost whole pages. It gives the
impression that the manuscript was put on paper in an almost
continuous, uniformly calm flow (barely interrupted by occasional
splashes of his fountain pen). Chapter follows chapter almost without
any space in between--it seems additions or changes were not
contemplated. (Despite the fact that additional sheets were added, not
all of the chapters listed in the table of contents could be
accommodated in the 170-page notebook.)
This manuscript is not the only scientific work W. Hennig produced
during the war. The list of publications indicates that 25 papers
were published between 1940 and 1945, among them two on his work
in epidemic control, some revisions of the Acalyptratae for E. Lindner's
"Flies of the Palaearctic region", studies on the relationships
of "The genera combined within the 'Phytalmiidae'" and
of the Pupipara, the "Catalogue of the Diptera of Formosa",
a "Contribution to the problem of the 'relationships between
larval and imaginal systematics"', and several studies on larvae.
These publications are by no means manuscripts dating from the
prewar years, but were indeed written during the war since he used
to work continuously wherever he was and no matter where he was.
He received the necessary background material from his wife through
the army postal service. She managed to obtain the necessary material,
to look after proof-reading, etc., despite the uncertain times and
her care in bringing up their children. It seems amazing how much
the two were able to accomplish in those times.--However, some material
was lost, for example, the entire manuscript for "Bronn's Classes
and Orders of the Animal Kingdom." The original burned in a
safe in Berlin. The carbon copy was lost later during the last weeks
of the war. W. Hennig never started writing the manuscript anew.
He also encountered difficulties in getting a rather long paper on
Phlebotominae published: this manuscript could not be printed during
the war because of paper supply problems despite the fact that he
directed inquiries to several publishers, even in Austria. The
manuscript was not considered "essential to the war effort"
and printing of the manuscript was therefore not authorized. This
manuscript, too, has never been published.
The terror of war continued to have a lasting effect on him even in
his later years. He himself, seriously wounded, was one of five
survivors of an entire company and one of his brothers did not return
from Stalingrad. Even much later, particularly during the time of the
annual siren testing, he used to remark how intensely his war
experiences had made him aware of the limitations imposed on his
creative output and how much he felt obliged to use the available time
to the utmost.
1945-1950: Postwar Period, Completion of the "Principles
..." and of the "Larval Forms of the Diptera"
his family continued to live in Leipzig after they were bombed out
during the war, W. Hennig was discharged from captivity to West
Germany since there was concern that he might end up as a Soviet
prisoner of war should he return to the "Eastern Zone" of
Germany. So he stayed in Hamburg, Plön, and Göttingen--for
six weeks at a time because this was the maximum period for which
ration cards were issued in each municipality. In November 1945 he
crossed the East German border illegally to join his family in
Leipzig. Fortunately, the situation was already defused since it was
well known within the Zoological Institute that he had never supported
the Nazi regime either by being a member in any of its organizations
or by any other activities.
On 1 December 1945, at the age of 32, he was appointed Acting
Director of the Zoological Institute at the University of Leipzig. He
resigned from this position in Leipzig on 1 April 1947 in order to be
able to return to the German Entomological Institute which had been
moved to Mecklenburg.
On his return to Berlin, he expanded the manuscript of the "Principles
... of phylogenetic systematics" which he had written while in
captivity. He worked on it in a cold room and by candlelight during
the 10 months of the Berlin blockade in 1948. (The family received a
coal ration of 25 lbs. for the entire winter plus 75 lbs. extra for
the three small children; they all collected horse chestnuts and
acorns as additional fuel). Paper continued to be in short supply when
the "Principles..." were ready for printing (which explains
the absence of an index and the much too small edition).
Starting in April 1947 when he returned to Berlin, he also
continued-to work on the "Larval forms of the Diptera" which
he had begun before the war and which was published in 1948. The large
card index had been moved to the German Entomological Institute in
Mecklenburg where it was found after the war in disarray, yet still
He was now also in charge of organizing the return of the museum
holdings. The building in Berlin was intact but had not yet been
released by the [U.S.] occupying power and so the mansion of a brewery
owner in the east end of the city was used to accommodate the
collections and the library which had fortunately survived the war
without losses. W. Hennig had thus regained his old position at the
German Entomological Institute.
In addition to the "Principles" and the "Larval
forms", he published other papers on the morphology and systematics
of Diptera during the period from 1945-1950 including short essays,
i.e. "Problems of biological systematics" and "Explanation
of terms used in phylogenetic systematics" (the two essays
were published in "Forschungen und Fortschritte" (Research
and Progress] 21/23: 276-279, 25: 137-139).
During this particularly productive period he published 35 papers
in addition to the papers mentioned above. Among the 35 papers were
such universally important works like the "Critical comments
on the phylogenetic classification of the insects" (1953),
"Diptera" in the "Handbook of plant diseases"
(1953), "Wing venation and classification of the Diptera with
special reference to the fossils described from the Mesozoic"
(1954), "Systematics and phylogeny" (1957), the first
edition of 'Invertebrates I and II" in the 'Textbook of zoology"
(new editions followed later, the newest will be published shortly),
"The dipterous fauna of New Zealand as systematic and zoogeographical
problem" (1960), the important monograph of the "Muscidae"
for "Lindner", etc. The revised edition of the "Principles..."
was also completed in 1961 and sent to the United States for translation
and printing (he did not see this work again until it was published
in 1966 under the title "Phylogenetic Systematics").
Finally, the time approached for Dr. H. Sachtleben, the Director of
the German Entomological Institute, to retire. W. Hennig, who until
then was Deputy Director, was nominated as successor when the
construction of the "Berlin Wall" on 13 August 1961 abruptly
changed the situation. Until then W. Hennig had commuted daily from
his home in West Berlin to the German Entomological Institute which is
located in the Eastern Sector of Berlin in the belief that the
division of the city was only temporary-- in the same way as his
general attitude was governed by the hope that Germany would
eventually be reunified. He resigned from his position with the German
Entomological Institute after the Berlin Wall was built. He stayed on
in West Berlin for almost two years and taught at the Technical
Until his death he regarded Berlin as his true home, not only
because of his ties with the German Entomological Institute but also
because of the entire cultural atmosphere.
1963: Move to Stuttgart/Ludwigsburg
Institution in Washington and Prof. E. Hardy from the Department of
Entomology of the University of Hawaii in Honolulu offered W. Hennig a
new sphere of activity. However, he decided to stay in Germany because
of his sons' education and also because he felt a need to remain close
to the cultural heritage of the ancient Greco-Roman Europe, as he
expressed it on several occasions. Prof. A. Kaestner, Munich, and
Prof. E. Schüz, Director of the State Museum of Natural History
in Stuttgart until 1969, used their influence to secure for him a
department of phylogenetic research at the Stuttgart Museum
(Ludwigsburg Branch). Thus he moved with his family to Ludwigsburg in
Reports on the activities of his department are included in the
annual reports of the Museum and are published in: Jh. Verein vaterl.
Naturkunde Württemberg 120: 38-39, 121: 23, 122: 23-24, Jh.
Gesellschaft Naturkunde Württemberg 124: 19-21, 33-34, 125:
30-31, 126: 16-17, 127: 19-21, 128: 16-17, 129: 21-22, 130: 405-406,
131: 223, etc.
1963-1976: The State Museum of Natural History in Stuttgart
Ludwigsburg; Focus on Fossil Studies etc.; Teaching Activities
Cut off from his accustomed collections and libraries in Berlin
with their vast holdings unique in Germany, he searched for, and
found, a "new niche" for his future focus of activities:
the study of fossils in amber. Among his many previous papers, he
had published two papers on Diptera and Aphaniptera in Baltic amber
as far back as 1938 and 1939 and since that time he had wanted to
study these fossils in greater detail. He had learned that the Königsberg
fossil collection had been moved to Göttingen where Prof. A.
Seilacher assisted him in his search and arranged for the loan of
the extensive material, as did Prof. O.H. Walliser.
In the period from 1964-1972, W. Hennig thus published 17 papers on
various groups of Diptera from the Baltic amber (as well as three
papers on fossils from the Lebanon amber, see below) which in theory
and practice went far beyond everything previously published in this
His studies of the fossils in amber yielded several important results:
The book "Phylogeny of the Insects" (1969, published by
W. Kramer, Frankfurt) grew out of his extensive file cards which
he always prepared, namely those dealing with "Localities of
Palaeozoic and Mesozoic insects", "Phylogenetic evolution
of the insects", and "Aims, methods and limitations of
phylogeny." This book does not only provide a critical phylogenetic
review of the described Palaeozoic and Mesozoic fossils of all insect
groups but also an extremely important contribution to the development
of his theory of phylogenetic systematics.
In his search for amber material with fossil inclusions he came upon
amber from Lebanon (an amber which was then still considered to be
non-fossiliferous). Thanks to the collaboration with his department on
the one hand and with the members of the Geological-Palaeontological
Institute in Tübingen3 on the other, his search produced the
greatest collection of fossils trapped in this resin from the Lower
Cretaceous--the geologically oldest fossil-bearing "amber"
known to date.
In 1966 he was invited to Canada by the Entomology Research
Institute, Ottawa. Although he had suffered a mild heart attack a few
months before his scheduled departure, he felt obliged to undertake
the trip. He stayed in Canada from August to December 1967. This trip
again culminated in extensive scientific studies and publications. He
also visited scientific institutions in New York, Washington, Urbana,
and Chicago during a one month stay in the U.S.A. Shortly before the
end of his visit he suffered another heart attack. His doctors urged
him to return home immediately--yet he insisted on keeping his promise
and completing the scheduled two weeks of collecting. Fossils in amber
were again at the centre of his investigations.
Another focus of his work was the
taxonomic-systematic revision of the dipterous families Muscidae
(1955-1964) and Anthomyiidae (1966-1976) as his 13th and 14th
contributions to "The flies of the Palaearctic region"
edited by Professor E. Lindner (his friend and colleague from the
State Museum of Natural History in Stuttgart/Ludwigsburg). Professor
Hennig viewed this task, which he could have confined to taxonomy per
se, as a challenge to conduct comparative-morphological analyses to
verify not only the taxonomic, but also the phylogenetic relevance of
characters and to provide extensive documentation in the form of
drawings--always his own (more than 3,000 illustrations, often
including several figures of the same form). After examining an
enormous amount of material--which he obtained on loan on an
individual basis or inspected on location in museums during his
holidays abroad--he based his phylogenetic conclusions on broad
comparisons which went far beyond the particular group he
General Works and Supplementary Studies
He used the same
approach in his general works, for example his treatment of the
Diptera in the "Handbook of zoology." Here as well he went
far beyond mere compilation and always considered all morphological
and non-morphological criteria in terms of their suitability as
phylogenetic evidence. His didactic efforts went to the point of
redrawing all illustrations he wanted to include and providing
detailed information in the captions in an attempt to achieve a
uniform and optimally informative picture.
He also published detailed studies of specific queries whose scope
would have exceeded the framework of handbook-oriented treatments.
Here, too, he used a broad basis for comparison: His investigation of
the wing base of Nematocera also includes information on Mecoptera;
the paper on the phylogenetic implications of the hypopygium of
Lonchoptera etc. (1976, the last paper he published) also includes
information on many other groups of Diptera; he also included many
details on musculature (based on sections he had made himself) to
substantiate his comments on the views advanced by other authors.
In his last year he worked intensively on the new edition of the
"Larval forms of the Diptera" (the publication of his
working material is in preparation), on a revised edition of the
"Textbook of zoology" (the 4th completely revised edition
is in press), as well as on a general phylogenetic manuscript (its
publication is in preparation) and other papers (see list of publications).
He needed such multi-faceted work:
thus he might work on dipterous larvae in the Museum during the day
and in the evening he might, for example, get so involved in
non-insect invertebrates that the only time he had left for writing
comments on manuscripts he received from his various journals, for
opinions requested from the German Research Association etc., were the
weekends where he would write heaps of letters in an effort to
complete all these tasks quickly and thoroughly.
The lectures which he gave as Associate Professor at the Tübingen
University (since 1970) were for him a welcome change from his
primarily entomological work at the Museum. In these lectures he
discussed, for example, new theories on mollusc phylogeny or coelom
evolution. His efforts to keep up and to continuously expand his
knowledge in many fields of zoology and to discover phylogenetically
relevant evidence were far-reaching and extended also to vertebrates
(as is indicated by a manuscript for a book on which he was working).
His informed and detailed arguments and questions would occasionally
embarrass even experts in their fields.
While he undertook numerous speaking engagements and attended conferences,
etc. in his earlier years (for example, in 1963/64 he traveled to
Helsinki, London, Rome, Munich, Marburg, and Freiburg, and visited
the museums in Helsinki, Copenhagen, Paris, London, Florence, Göttingen,
Frankfurt, and Tübingen for study purposes), in his later years
he concentrated his efforts on personal discussions on a particular
issue to expand his theories and elaborate his methodology, on "filtering
out" previous suggestions on phylogenetic evaluation (e.g.
in his "Phylogeny of the insects" and "Textbook of
zoology"), and on his own work on specific extant and fossil
groups. Insofar as his papers were not intended for publication
in other journals, almost all of his papers which he wrote during
the time he worked in Stuttgart/Ludwigsburg were published in the
"Stuttgarter Beiträge zur Naturkunde", the Museum's
"Stuttgarter Beiträge zur Naturkunde" is a scientific
journal which has been published by the State Museum of Natural
History in Stuttgart since 1957 (editor: Dr. K.W. Harde); it has a
circulation of 1,000 copies and at the present time copies are still
available for purchase (through direct orders or through bookstores)
or exchange for other journals.
W. Hennig regarded a discussion of phylogenetic principles among a
small group of colleagues, particularly a written or oral discussion
with an author on the basis of his manuscript (whose level of work
appeared to him worth spending some time on) to be more important than
general speaking engagements before a large audience. He perceived
serious dangers in having to shorten his argumentations (which might
entail misunderstandings) because of the time limitations imposed by
lectures; and he regarded the elaboration of certain elements required
for lecture purposes as (tedious) repetition of issues he had already
discussed on several occasions before; and finally, he missed the
personal contact and the stimulus provided by a personal discussion.
If he felt that a person was truly interested in phylogenetic
methodology, W. Hennig would untiringly answer specific questions,
approach them from various angles, and cite numerous examples in
support of his arguments without any regard for time or his own
projects that may have been scheduled for that particular time.
He realized that many authors (unlike himself) evidently found
it difficult to transpose the phylogenetic argumentation from a
study on other animal groups used as a model to their own specific
animal group, and he therefore made every effort to plant "germ
cells" in as many animal groups as possible. This is also the
reason why he enjoyed supervising post-graduate students working
on a thesis since he expected to encounter here a combination of
mental agility, openmindedness, and a wealth of facts.
In order to ensure adequate publication and dissemination of the
findings of these theses, he used his influence as co-editor of
the "Zeitschrift für Morphologie der Tiere" / "Zoomorphologie",
together with Prof. P. Ax who shared his views, to get even voluminous
manuscripts printed and he strenuously (and successfully) opposed
efforts to restrict the size of a manuscript. He repeatedly expressed
the view that this journal should act as "a vessel for theses."
In his efforts to establish the "germ cells" he tried to
increase the precision of the phylogenetic argumentation to a level
which the author of a given manuscript could or still would reach; but
when he realized that a certain limit was reached (or could not be
reached from the very beginning), he would be satisfied with
phylogenetically less relevant argumentation and still recommend
publication (particularly in cases where the journals in question had
less sophisticated phylogenetic ambitions than the ones he himself was
involved in). He made this decision in order to preserve the
documentary value of a paper dealing, for example, with investigations
of rarely studied organs or animal groups. He would then express the
hope that the author might perhaps progress and improve upon his
phylogenetic conclusions at some later stage, or that others might
take up this particular topic and bring it to a satisfactory
He did not like to get involved in the flood of theoretical papers
on phylogeny (of every possible shade); he considered many of the
papers he had to read because he received them as manuscripts to
be intolerable on account of the frequent superficiality in their
argumentation which he could not understand, and on account of the
constant repetition of "counter-arguments" which he had
refuted before more than once on the basis of specific examples
and well-founded reasoning (which the author had not read or had
not understood for semantic or other reasons). Against this background,
he would occasionally be pleased with papers which at least to some
degree followed his approach--in hopes that the authors should not
become discouraged and should try to improve.
In addition he was convinced (as he pointed out on several
occasions) that in 10 or 20 years' time the wheat would separate from
the chaff even without his doing. He wanted to write a new book on
phylogeny after he had completed the revision of the "Larval
forms"; he felt that this would be the time to deal with
The massive attack by E. Mayr however did elicit his immediate
reaction; Mayr's remarks on monophyly in particular struck at the
heart of his phylogenetic approach. W. Hennig strenuously objected
to the dilution of this term (for example through the incorporation
of groups which are characterized only by evident non-synapomorphies)
in Mayr, as he did vis-a-vis visitors whenever he deemed it necessary.
He was also extremely amazed to see that people could be satisfied
with formulations of the type "A, B, and C have a common ancestor
and are therefore a monophyletic group" as evidence of relationship
and that even when they were given a detailed explanation (for example:
man, snail and earthworm also have a common ancestor, but this is
no proof that the three are related more closely to one another
than to other forms), they did not understand that there was a difference
between the above statement and the following formulation and meaning
of "...having an ancestor which only they share."
Sometimes he felt that the time when his ideas would be generally
understood was still in the distant future in view of the endless
discussions of elements of phylogenetic methodology which he had
explained before to the very limits of achievable clarity and which
nevertheless gave rise to spectacular misinterpretations. He overcame
his anger over such incidences or difficulties arising from other
causes which he considered small-minded and parochial (as they must
appear to him given his background, see above) by burying himself in
work, a remedy he had already adopted during the war and the postwar
Fortunately, various honours were bestowed on
him just in time--as one, might almost say with hindsight in view of
his short life--to give him optimism and confidence regarding the
impact of his work.
He was awarded the Fabricius Medal by the German Entomological
Society in 1953 (i.e. when he was only 40 years old).
In 1955 he was named Associate Member of the Finnish Entomological
Society, Helsinki, in 1959 he was named a Member of the German Academy
of Natural Scientists 'Leopoldina Halle, and in 1963 Associate Member
of the American Entomological Society, Philadelphia.
In 1968 he was awarded an honorary doctorate by the Faculty of
Mathematics and Natural Sciences of the Free University of Berlin, an
honour that had special meaning for him because of his close ties with
Berlin where, one might almost say, he had left his heart and soul and
because it was Professor Klaus Günther4, a person he held in
extremely high regard, who presented the honorary doctorate and
delivered the eulogy (Fig. 2).
E. Schuz provides an outline of the text of the eulogy and a
description of the ceremony held at the guesthouse of Dr. K.E.
Scheufelen in Stuttgart in Jh. Ges. Naturkunde Württemberg
In 1972 the Royal Swedish Academy of Sciences, Stockholm, named him
Special honours were bestowed on him in the form of the Cold Medal
of the Linnean Society, London (1974) and the Gold Medal of the
American Museum of Natural History, New York (1975). W. Hennig
received the latter medal in Stuttgart from the Director of the
American Museum, Dr. Thomas D. Nicholson.
Finally, in 1976, he was named Honorary Member of the Society of
Systematic Zoology, New York.
4. W. Hennig, The Man
In view of his enormous activities
one might be inclined to assume that he did not have any interests
outside zoology--but that was far from true. He was a music enthusiast
and loved Mozart and Handel in particular. For many years he would
listen to every Mozart opera broadcast from Salzburg, using a libretto
to follow the music. He also attended many concerts and opera
performances in Berlin. Later he traveled extensively with his wife,
in particular to southern Europe with its beautiful scenery and
cultural heritage, and in the course of these travels he became
something of an authority on ancient art history. When he learned that
someone from his department or from administration intended to go on a
trip, for example, to Rome or Florence, he would provide detailed
impromptu descriptions and useful suggestions which were greeted with
much enthusiasm; he often brought with him stacks of travel books on
art history and lent them to colleagues who expressed interest in
them. He seemed to know everything there is to know about the cultural
treasures, particularly Romanesque and Byzantine art up to 15th
century art, of all of Italy (his favourite country--he was also able
to converse in Italian), but also of Sicily, Yugoslavia, Greece,
France, and Spain, and he acquired this knowledge thanks to his
phenomenal memory and his remarkable grasp of major concepts through
He developed a particularly efficient method of getting to see all
that he deemed worthwhile on a trip. While he left the organization of
a trip entirely to his wife, he would study every available source to
learn about the cultural treasures he was interested in; he would copy
these items and note down particulars and would then show these
sketches to a tourist guide who could then guide him promptly and
without wasting any time to the places he wanted to see (Fig. 3).
He also had an opportunity to briefly visit Australia and New Guinea
as well as Thailand, the northern part of India and Nepal, and finally
Java, Bali, and Burma (where he, of course, searched for Burmese
He would not always tell the destination of his trips, mainly where
trips to distant countries were concerned. Sometimes it required many
skillful questions to wheedle his secret out of him, and Professor E.
Lindner was particularly successful in this. On one occasion Professor
Hennig returned to the office after a holiday and showed his
unsuspecting audience a photo of himself, in "tiger tops",
riding on an elephant in south Nepal. He was pleased as punch when the
ladies asked him, despite his holiday suntan, whether the photo had
been taken in the Berlin Zoo.
He liked original humour, especially humour with a Berlin flavour,
and an equally matched partner like Professor Klaus Günther could
inspire him to sparkling and witty repartees. He would recite passages
from classical literature in a spontaneous contest with Dr. Gerd von
Wahlert-- in Greek, Latin and in German, if necessary. His immense
library at home did not only contain books on zoology and the
classics, or Humboldt's accounts of his travels (a man he admired and
whose portrait on the Orinoco always hung above his desk in the
Museum), but he also had books by Karl May, for example 'Winnetou",
or the "Leatherstocking Tales" within easy reach. Who would
have expected that? -- W. Hennig also liked our dog and used to remark
to others with some pride that it had accepted him immediately and
liked to accompany us to his office in the Museum, while it would bark
at all other members of the Museum staff although it had encountered
During occasional visits to the Museum his grandchildren (he had
five) treated him with cheerful and warm affection, which was
Nobody expected that his life would end so soon and so abruptly, he
himself included judging from his remarks regarding his plans for the
future. The only indication to the contrary was an entry ("to be
transferred to Schlee, if necessary") in his personal copy of the
Diptera treatise of the "Handbook of zoology" he was using
for revision purposes.
In keeping with his wishes (he was averse to any form of personality
cult and even disliked the use of his name in connection with the
phylogenetic methodology--he felt that 'phylogenetic systematics'
should suffice), no official representatives attended his funeral. He
was buried in the mountain cemetery in Tübingen with only his
close friends and relatives present. His brother, a minister from
their home district in Saxony and, like Willi Hennig, a remarkable
person of great determination, deep commitment to his work, and
profound understanding, conducted the funeral service and found the
right words to pay true tribute to the man even without emphasizing
the role that zoology had played in his life.
Although Willi Hennig was unable to realize many of his plans, what
he had achieved is nevertheless many times more than a scientist could
dream of accomplishing in a lifetime. We can assume that the unlimited
support and absolute harmony he found in his family were essential for
his work. The role of his family, and particularly that of Mrs. Irma
Hennig, in this monumental achievement cannot be overestimated.
Mrs. Hennig's mathematics and biology studies, her continued
interest in these subjects and her active support, in addition to the
example set by the father, may well have had a guiding influence in
the career development of their sons since all three attended
university, two studied biology and chemistry and are now engaged in
research work at the Max Planck Institute, while the third son has
completed his studies as a secondary school teacher of German and
5. Postscript and Acknowledgment
Although I had an
opportunity to catch a glimpse of some of the aspects of Willi
Hennig's life in the course of the almost ten years of working with
him and talking to him on a daily basis, it was not sufficient, of
course, to gain an overall picture of his life. I am therefore
grateful to Mrs. Hennig and her sons for their kind cooperation in
answering all my questions in preparation of this paper and for review
of the manuscript. In particular, I am indebted to Mrs. Hennig for
making available W. Hennig's handwritten texts (the draft of the "Principles...",
and the high school essay), the list of publications and the
photographs. Permission to make the significant text of the high
school essay known to the general public is also greatly appreciated
(see: Ent. Germ. 4 (3/4): 193-199; Stuttgart 1978).
The complete list of Willi Hennig's
publications appeared in: Beitrage zur Entomologie [Contributions to
Entomology) 28: (00): 000-000; Berlin 1978. Only a few selected works
and relevant papers are listed below.
- Obituary by P. Ax (1977)
- List of publications in: Beiträge zur Entomologie 28 (1);
- For details see SCHLEE & DIETRICH in: Neues Jahrb. Geol.
Paleontol., Monatshefte, 1970:40-50; SCHLEE in: Stuttgarter Beitr.
Naturkunde 213: 1-72, 1970; Kosmos (Stuttgart) 1972: 460-463.
- W. Hennig also expressed this in his obituary for K. Günther,
Hennig's last publication in 1976.
- Page from W. Hennig's manuscript (1950): "Principles of
a theory of phylogenetic systematics", chapter entitled "The
- Willi Hennig with Professor Klaus Günther (left) who
presented the honorary doctorate of the Free University of Berlin at
the guesthouse of Dr. K.E. Scheufelen, Stuttgart, 1968.
- Willi Hennig and his wife in Hong Kong on January 26, 1976.