& r W w t A . ^ [Link],. .

Systematic Entomology (1990) 15, 153-217

A revision of the Holarctic species of Serromyia Meigen

(Diptera: Ceratopogonidae)

A. BORKENT and B. BISSETT Biosystematics Research Centre,

Agriculture Canada, Research Branch, Central Experimental Farm, Ottawa, Ontario, Canada

Abstract. This revision recognizes sixteen extant species of Serromyia

in the Holarctic Region including seven described as new: [Link]
Borkent, [Link] Borkent, [Link] Borkent, [Link] Remm,
[Link] Borkent, [Link] Borkent and [Link] Borkent. In
addition, eleven new synonyms are indicated. Keys and descriptions of
the male and female adults, and their known distributions, are given for
all species. The egg and first instar larva of Serromyia nudicolis Borkent
[Link]., the pupa of [Link] (Meigen) and the fourth instar larvae of an
unidentified Serromyia are also described.
All available fossil specimens were examined, some redescribed, and
two named as new: [Link] Borkent and [Link] Borkent.
New combinations are [Link] (Heyden), Atrichopogon trichopus
(Thomson) and Monohelea scirpi (Kieffer).
Limited phylogenetic interpretation indicates that all extent Holarctic
species, with the exception of [Link] Delecolle and Braverman,
form a monophyletic group. Six Baltic amber species are not members
of this clade. Metacanthohelea is recognized as the sister group of the
genus Serromyia.

Introduction Meillon & Downes, 1986; Giles & Wirth, 1982),

the Holarctic species remain very poorly under-
The genus Serromyia Meigen is a rather distinc- stood. A number of new species from the
tive taxon within the Ceratopogonidae. Within
Nearctic Region were known to us and the
the Holarctic Region its members can be easily
identification of Palaearctic species was a jumble
recognized with the naked eye, or perhaps with
of confused synonymy and conflicting, generally
a hand lens, by their swollen hind femora which
uninterpretable, descriptions.
bears strong ventral spines.
The second motive for initiating a revision
We began a revision of this genus with two
was to examine the phylogenetic relationships
purposes in mind. The first was to provide the
between the species of Serromyia. Because we
basis for accurate identification of the known
were initially confident that the genus was
species in the Holarctic Region. Although the
monophyletic and exhibited a reasonable degree
species outside the Holarctic Region have
of interspecific structural variation, we con-
been recently revised (Meillon & Wirth, 1983;
sidered the genus a good candidate for phylo-
Correspondence: Dr A. Borkent, 2330—70th genetic analysis. On the whole, our expectations
St. SE, Salmon Arm, British Columbia, VIE 4M3, were only partially met. One major difficulty
Canada. was the current status of our understanding

153
154 A. Borkent and B. Bissett

of the phylogenetic relationships between the AMNH: Department of Entomology Collec-

genera of Ceratopogonidae. Enough uncer- tion, American Museum of Natural History,
tainties exist that virtually all other Cerato- Central Park West at 79th St., New York,
pogonidae must be used in outgroup New York, 10024, U.S.A.
comparisons. BMNH: Department of Entomology, British
The genus is relatively well represented Museum (Natural History), London SW7
by Eocene and Oligocene fossils and we 5BD, U.K.
rrm- incorporated these into our revision to BPBM: Department of Entomology Collection,
better understand the diversification of the ex- Bernice P. Bishop Museum, P.O. Box
tant species in the Holarctic Region. 19000A, Honolulu, Hawaii 96819, U.S.A.
CASC: Department of Entomology, California
Academy of Sciences, Golden Gate Park,
San Francisco, California 94118, U.S.A.
tutorials and Methods
CNCI: Canadian National Collection of Insects,
Specimens examined. This study was based Biosystematics Research Centre, Agriculture
on the examination of 657 males, 1154 females, Canada, Ottawa, Ontario K1A 0C6, Canada.
five pupae and nine larvae of Serromyia. These CUIC: Cornell University Insect Collection,
included representatives of all named species in Department of Entomology, Cornell Univer-
t able 1 noted with an asterisk. We were un- sity, Ithaca, New York 14850, U.S.A.
fortunately unable to examine some African HNHM: Zoological Department, Hungarian
species housed in the South African Institute Natural History Museum, Baross utca 13,
for Medical Research, Johannesburg. H-1088, Budapest, Hungary.
Generally, the genus was poorly represented INHS: Illinois Natural History Survey
in most of the collections from which we re- Insect Collection, 607 E. Peabody Drive,
quested loans. We were able to study the imma- Champaign, Illinois 61820, U.S.A.
ture^ of only two Serromyia identified to species. ISNB: Institute Royal des Sciences Natu-
The eggs and first instar larvae of [Link] relles de Belgique, Rue Vautier 29, B-1040,
sp.n. were secured from field-collected female Bruxelles, Belgium.
adults which were decapitated to induce egg IZBE: Institute of Zoology and Botany,
laying. A pupa of [Link] (Meigen) and three Vanemuise str. 21, Tartu 202400, U.S.S.R.
larvae and four pupae of unidentified Serromyia KUPC: Katedra Systematicke Zoologie,
were also available from the collections of Prirodovedecke Fakulty, University Karlovy,
Thieneniann and Strenzke (ZSMC). Vinicna 7, 128 44 Praha 2, Czechoslovakia.
Filially, to obtain better outgroup compari- LACM: Los Angeles County Museum of
sons for phylogenetic analysis and to estimate Natural History, 900 Exposition Blvd., Los
potential for parallelism of character states, we Angeles, California 90007, U.S.A.
examined adult material of virtually every MCZC: Entomology Department, Museum of
described genus of Ceratopogonidae. Comparative Zoology, Harvard University,
We were able to examine type material of all 26 Oxford St., Cambridge, Massachusetts
Serromyia species described in this analysis un- 02138, U.S.A.
less otherwise noted. Details about the location MNHN: National Collection of Insects,
of type material as well as sex and details of Museum National d'Histoire Naturelle, 45
labelling are recorded under each species. rue Buffon, Paris 75005, France.
We were able to examine most of the material MZW: Museum of the Earth, Al. Na Skarpie
housed in the HMHN and interpreted by Remm 20/26, 00-488 Warsaw, Poland.
(IU73L). MZLU: Department of Zoology, Zoological
I'he abbreviations used to represent various Museum, Helgonavagen 3, S-223 62 Lund,
naisei.'iiis from which material was received and Sweden.
when: specimens are deposited are those pro- MZSF: Museum Zoologique, Universite de
vided \jy Amett & Samuelson (1986). Private Strasbourg, 29, Blvd. de la Victoire, F-67
colKni vns are abbreviated according to the Strasbourg, France.
proa-:!'ue used in that work. Those used in this NHMW: Naturhistorisches Museum Wien,
study are as follows: Postfach 417, Burgring 7, 1040 Wien, Austria.
 Revision of Holarctic species of Serromyia Meigen 155

Table 1. Checklist of Serromyia species of the world. For each species the sex examined in this study is noted
with an asterisk. The type locality is given for each species.

M F

aethiopiae Clastrier & Wirth 1961: 219. Gambia.

agathae Meillon & Wirth 1983: 388. Republic of South Africa.
alpheus Heyden 1870: 251 (Ceratopogon). Rott, Federal Republic of Germany. Miocene. New
combination.
anomalicornis (Loew) 1850: 30 (Ceratopogon). Baltic amber. Eocene.
atra (Meigen) 1818: 84 (Ceratopogon). Latvia.
albitarsis Kieffer 1919: 71. Hungary.
micronyx Kieffer 1919: 70. Hungary. New synonym.
nitens Goetghebuer 1920: 73. Belgium. New synonym.
spinosipes Kieffer 1919: 72. Hungary. New synonym.
barberi Wirth 1952: 205. California.
bicolor Borkent [Link]. Federal Republic of Germany.
borealis Borkent [Link]. Alberta.
colorata Statz 1944: 150. Rott, Federal Republic of Germany. Miocene.
austera Statz 1944: 150. Rott, Federal Republic of Germany. Miocene. New synonym.
spinofemorata Statz 1944: 150. Rott, Federal Republic of Germany. Miocene. New synonym.
crassifemorata Malloch 1914: 218. Illinois.
esakii Tokunaga 1940: 218. Caroline Islands.
femorata (Meigen) 1804: 28 (Ceratopogon). Europe.
armata (Meigen) 1818: 83 (Ceratopogon). Germany.
flavicrus (Kieffer) 1906: 63 (Palpomyia). New name for flavipes Gimmerthal. New synonym.
flavipes Gimmerthal 1847: 144 (Ceratopogon), not Meigen 1804: 28. England. New synonym.
foersteri (Meigen) 1838: 21 (Ceratopogon). England.
inermipes Kieffer 1919: 73. England.
festiva Kieffer 1911: 346. Seychelles.
heveli Giles and Wirth 1982 : 442. Sri Lanka.
ledicola Kieffer 1925a: 156. Estonia.
europaea Clastrier 1963: 61. Austria.
macronyx Goetghebuer 1933: 355. Belgium.
maculipennis Giles & Wirth 1982: 444. Sri Lanka.
mangrovi Delecolle & Braverman 1987: 57. Egypt.
meiswinkeli Meillon & Wirth 1983: 390. Republic of South Africa.
morio (Fabricius) 1775: 800 (Culex). England.
nudipennis Kieffer 1913: 10. England. New synonym.
neethlingi Meillon & Wirth 1983: 392. Republic of South Africa.
nocticolor Kieffer 1914: 268. Republic of South Africa.
nudicolis Borkent [Link]. Maine.
pacifica Remm [Link]. Sakhalin Island, East Siberia, U.S.S.R.
pendleburyi Macfie 1934: 280. Malaya.
polonica Szadziewski 1988: 135. Baltic amber. Eocene.
punctata Giles & Wirth 1982: 446. Sri Lanka.
reyei Debenham 1970: 161. North Territory, Australia.
rossi Meillon & Wirth 1983: 396. Republic of South Africa.
rufitarsis (Meigen) 1818: 83 (Ceratopogon). Europe.
gelida Kieffer 1925: 429. Latvia. New synonym.
bispinosa Goetghebuer 1936a:321. Belgium. New synonym.
dipetala Remm 1965: 182. Estonia. New synonym.
ryszardi Borkent [Link]. Baltic amber. Eocene.
sierrensis Borkent [Link]. California.
silvatica Meillon & Downes 1986: 175. Republic of South Africa.
sinuosa Borkent [Link]. Baltic amber. Eocene.
spinigera (Loew) 1850: 30 (Ceratopogon). Baltic amber. Eocene.
elongatus Meunier 1904: 231 (Ceratopogon). Baltic amber. Eocene.
156 A. Borkent and B. Bissett

Table 1 (continued)

M F

stuckenbergi Meillon & Wirth 1983: 398. Republic of South Africa.

subinermis Kieffer 1919: 73 (as var. of spinosipes). Hungary.
succinea Szadziewski 1988: 135. Baltic amber. Eocene.
tecta Borkent [Link]. Federal Republic of Germany.
vockerothi Borkent [Link]. Manitoba. «
zuluensis Meillon and Wirth 1981: 589. Republic of South Africa.

NHRS: Naturhistoriska Riksmuseet, Sektionen In general, Serromyia adults were collected

for Entomologi, 104 05 Stockholm, Sweden. in habitats where either pools, seeps or small
OX AM: Hope Entomological Collections, creeks were found. Although we found adults
University Museum, Parks Road, Oxford at any time of the day, during full daylight
OX1 3PW, U.K. adults of some species occurred very close to
PEHC: P. Havelka, Landesanstalt fur the ground. In these instances the net edge was
Umweltschutz Baden-Wurttemberg, Postfach held on the ground while sweeping vegetation.
21 13 10, 7500 Karlsruhe 21, F.R.G. At dusk, individuals were more common in
RYSC: R. Szadziewski, Department of In- vegetation well above ground level.
vertebrate Zoology, University of Gdansk, The invariable reaction of Serromyia adults
Czolgistow 46, 81-378 Gdynia, Poland. when swept up was to feign death and fall to the
UCDC: Department of Entomology, University bottom of the net, remaining there sometimes
of California, Davis, California 95616, for several minutes. They would then slowly
U.S.A. crawl from amongst other insects and debris. It
USNM: United States National Entomological was therefore important to observe the net catch
Collection, Department of Entomology, U.S. for several minutes to determine whether adults
National Museum of Natural History, were present. Individuals found resting on veg-
Washington, DC 20560, U.S.A. etation had a similar reaction of collapsing and
UZMH: Division of Entomology, Zoological falling to the ground when disturbed (see bio-
Museum, SF-00100, Helsinki 10, Finland. nomic section under [Link]). This may ex-
ZMAN: Instituut voor Taxonomische Zoologie, plain why adults were often collected near
Universiteit van Amsterdam, Plantage ground level during daylight hours.
Middenlaan 64, Amsterdam, The Nether- Further specific bionomic information which
lands. influenced collecting success is given with the
ZMAS: Zoological Museum, Academy of account of the pertinent species.
Science, Universitetskaya, Naberzhnayal, B- We found that light trapping with a UV light
164, Leningrad, U.S.S.R. failed to attract Serromyia. In one location at
ZMUC: Zoologisk Museum, Universitet- Mary's Peak, 21 km W Corvallis, Oregon, such
sparken 15, DK 2100, Kobenhavn, Denmark. a trap was placed within 5 m of resting females
ZMUH: Zoologisches Institut und Zoologisches of [Link] Wirth and still did not attract indi-
Museum, Universitat Hamburg, Papendamm viduals. However, several European species
3, 2 Hamburg 13, F.R.G. have been collected using light trapping
ZSMC: Zoologische Sammlung des Bayerischen methods.
Staates, Munchhausenstrasse 21, D-8000 We attempted to collect immatures of
Munchen 60, Bayern, F.R.G. Serromyia, but efforts to extract larvae or pupae
from moss and bottom samples by various flo-
tation techniques failed.
Collecting, preservation and examination
Adults were preserved in 70% ethanol. They
techniques
were then examined in one of several ways.
We had the greatest success in obtaining both Most specimens were mounted on microscope
individuals and species by sweeping vegetation. slides using a method developed by Leo Forster
 Revision of Holarctic species of Serromyia Meigen 157

of our centre. Adults had their wings removed used by ceratopogonid workers, our approach
and placed in 15% acetic acid. The head and was to search for any character that might be
abdomen were dissected from the thorax and all useful for recognizing taxa. Several characters
were placed in 10% KOH which was then heated not previously used are therefore incorporated
in a hot water bath. When fully cleared these into this analysis. Some characters which have
were placed with the wings in the acetic acid. been traditionally viewed as 'good taxonomic'
All parts were then taken through successive characters were found to be of no value and are
baths of 100% 2-propanol, 2-propanol layered either relegated to the generic description or
over clove oil, pure clove oil (where the are ignored.
antennae and left legs were further separated This examination of specimens for significant
from the head and thorax respectively) and characters provided the rational basis for the
finally into Canada Balsam on the slide. The measurements and ratios given in Tables 2 - 1 3 .
antennae and legs were sometimes removed Antennal and wing measurements were taken
while the specimen was in the Canada Balsam. as described by Grogan & Wirth (1979).
Many adults were critical point dried and Aedeagus length/width was the maximum
glued to pins. Such dried material allowed median, longitudinal distance (median base to
examination of pruinosity and some other tip of aedeagus) divided by the distance between
character states. We left no material preserved the tips of lateral apodemes of the aedeagus.
in alcohol because such material becomes Length of female hind claws was the minimum
bleached, distorted, and unfit for study in only a distance between the base of the claw and its
few years. tip, regardless of the degree of curvature of the
We also examined specimens of most species claw.
cleared, dissected and placed in glycerine. This On many of the labels of material housed
allowed for careful side by side comparison of in the CNCI, there are numbers referring to
many structures of two or more species. These further collecting data or bionomic information.
specimens were subsequently slide mounted in These are numbers beginning with either 'JAD'
Canada Balsam at the end of the study. or 'CD', referring to specimens collected or
At the beginning of this study we searched dealt with by either Mr J. Antony Downes or
for character states which could be used to the senior author, respectively. Such further
recognize species of Serromyia using pinned, information is recorded on data sheets housed
glycerine immersed and slide mounted material. in the Diptera Unit of our Centre.
Although conscious of characters previously Most photomicrographs were taken with

Table 2. Descriptive statistics for the antennal ratio (ftagellomeres 1 0 - 1 3 / 1 - 9 ) of male Serromyia.

Species n Mean Min. Max. SD

barberi 21 1.04 0.91 1.12 0.054

borealis 3 0.89 0.82 0.93
crassifemorata 9 0.95 0.90 1.00 0.029
nudicolis 14 1.08 0.98 1.20 0.061
sierrensis 1 1.16 1.16 1.16
vockerothi 2 0.90 0.89 0.91
ledicola 25 1.03 0.86 1.15 0.071
atra 14 1.03 0.93 1.11 0.066
bicolor 2 1.14 1.03 1.25
femorata 15 1.22 1.04 1.56 0.121
mangrovi 2 1.02 1.00 1.05 —

morio 13 1.11 1.04 1.21 0.043

pacifica 1 1.17 1.17 1.17
rufitarsis 8 1.01 0.92 1.10 0.056
subinermis 8 1.15 1.06 1.21 0.047
tecta 2 1.02 1.00 1.03 -
158 A. Borkent and B. Bissett

Table 3. Descriptive statistics for ratio of antennal flagellomeres 10/11 of male Serromyia.

Species n Mean Min. Max. SD

barberi 21 0.76 0.68 0.83 0.038

borealis 3 0.92 0.83 0.98 —

crassifemorata 9 0.97 0.81 1.16 0.090

nudicolis 15 0.84 0.77 0.90 0.045
sierrensis 1 0.80 0.80 0.80 —

vockerothi 2 0.93 0.90 0.95 —

ledicola 26 0.71 0.58 0.84 0.060

atra 15 0.74 0.63 0.81 0.046
bicolor 3 0.85 0.82 0.88 —

femorata 16 0.94 0.83 1.04 0.055

mangrovi 2 0.54 0.53 0.55 -

morio 13 0.89 0.78 0.99 0.064

pacifica 1 1.03 - —

rufitarsis 8 0.78 0.72 0.87 0.054

subinermis 13 1.22 1.12 1.37 0.086
tecta 2 0.91 0.87 0.96

Table 4. Descriptive statistics for the wing length of male Serromyia (in mm).

Speck's n Mean Min. Max. SD

barberi 18 2.15 1.50 2.32 0.193

borealis 3 1.62 1.50 1.80 —

crassifemorata 9 1.65 1.38 1.84 0.145

nudicolis 14 1.95 1.80 2.36 0.147
sierrensis 1 1.86 1.86 1.86 —

vockerothi 2 1.64 1.62 1.66 —

ledicola 25 2.35 1.70 2.88 0.265

atra 17 1.66 1.36 1.84 0.115
bicolor 3 2.21 2.02 2.38 —

femorata 22 2.49 2.12 2.84 0.182

mangrovi 2 1.21 1.20 1.22 -

morio 13 2.31 2.02 2.46 0.127

pacifica 1 2.24 2.24 2.24 -

rufitarsis 8 1.56 1.48 1.72 0.078

subinermis 11 2.25 1.82 2.52 0.190
tecta 2 1.69 1.62 1.76 —

either a Reichert Zetopan or a Nikon Optiphot Terms and abbreviations for structures
microscope equipped with interference contrast
optics. The larval structures were photographed In general we use the terms provided by Downes
using a scanning electron microscope. Most & Wirth (1981) for adults. Although not in
structures were measured using a micrometer in keeping with some of the terms traditionally
a Nikon compound microscope. Wing charac- used by ceratopogonid workers, their work pro-
ters were measured using a micrometer in a vides names for structures which are consistent
Wild M8 steroscope. Subsequent analysis of with those utilized in the rest of the Diptera.
data was through computer programs. General terms for larval structures follow
 Revision of Holarctic species of Serromyia Meigen 159

Table 5. Descriptive statistics for the number of elongate strong bristles on the hind coxa of male Serromyia

Species n Mean Min. Max. SD

barberi 22 0 0 0 0
borealis 3 0 0 0 —

crassifemorata 9 0 0 0 0
nudicolis 16 0 0 0 0
sierrensis 1 0 0 0 —

vockerothi 3 0 0 0 —

ledicola 27 1.41 0 4 1.01

atra 15 1.13 0 3 0.99
bicolor 3 0 0 0 —

femorata 22 0 0 0 0
mangrovi 2 0 0 0 —

morio 14 0.29 0 2 0.61

pacifica 1 1 1 1 —

rufitarsis 10 0 0 0 0
subinermis 11 0 0 0 0
tecta 2 0 0 0 -

Pruinosity

Strong bristles

Strong bristles

Figs 1A—D. A, scutum of male Serromyia femorata in lateral view; B, thorax of female [Link] in lateral view;
C, foreleg of male [Link]; D, hindleg of male [Link].
160 A. Borkent and B. Bissett

Teskey (1981) and detailed characters (i.e. some groups have bristles that are more
chaetotaxy) as well as pupal characters follow markedly developed than in any species of
Saunders (1924) and Lawson (1951). Serromyia. The strong bristles as used here to
We use some terms in the keys and descrip- describe character states refer to those bristles
tions that require further comment. Some adult which stand out from the remaining setae as
Serromyia lack pruinosity on the scutum and in thicker and often appear to be more darkly
such specimens the scutum is brilliantly shiny pigmented. Figs 1C, D indicate examples of
and highly reflective in pinned material (the typical strong bristles on the fore and hind leg,
humeral pits are not distinct from the surround- respectively. The lateral strong bristles on the
ing shiny cuticle) and utterly lacking fine spicules hind coxa are shown in Fig. IB. Sometimes
when viewed laterally in slide-mounted speci- strong bristles are broken off at the base and
mens. Specimens which are reported to exhibit then the enlarged socket needs to be searched
pruinosity on the scutum are somewhat dull for, for accurate interpretation of keys and
in reflected light (so that the humeral pits ap- descriptions.
pear as discrete, shiny patches). When slide Leg coloration is illustrated somewhat dia-
mounted, such specimens have a short coat of grammatically. Leg pigmentation intensity dif-
fine spicules visible in lateral view (Fig. 1A). fered between specimens and varied according
We looked for pruinosity amongst the dorso- to preparation technique. Illustrations therefore
central setae on the laterally mounted thorax are meant to show extent of pigmentation for a
using interference contrast optics. given species with the intensity of pigmentation
When one examines the brilliantly shining not necessarily comparable between species.
scutum of a pinned specimen with a dissecting Parameres are described as either rounded or
microscope, one can be confident of the lack of tapered apically. Rounded parameres are some-
pruinosity; but when the scutum is dull, care times shrivelled and may appear pointed. How-
must be taken. A somewhat dirty specimen may ever, in most such instances the two parameres
appear dull when in fact it lacks pruinosity. In look different from one another. Parameres
most of these instances the humeral pits would which are tapered apically are always of clearly
also be dull. We generally preferred to examine defined and characteristic form, as illustrated in
slide-mounted material to ensure correct in- the drawings of the male genitalia.
terpretation of the state of pruinosity. Ratios and some structures discussed in this
We use the term 'strong bristles' in describing study are abbreviated as follows:
armature of the legs. Among ceratopogonids, L: length

Table 6. Descriptive statistics for hind femur length/width of male Serromyia.

Species n Mean Min. Max. SD

barberi 10 6.51 5.62 7.06 0.575

borealis 2 5.67 5.57 5.77 -

crassifemorata 9 6.10 4.16 7.06 0.937

nudicolis 16 5.96 5.38 6.64 0.388
sierrensis 1 6.19 6.19 6.19 -

vockerothi 3 5.90 5.74 6.05 —

ledicola 15 6.51 5.87 7.36 0.445

atra 13 6.55 5.00 7.00 0.539
bicolor 3 7.64 7.52 7.76 —

femorata 19 6.47 5.23 7.52 0.517

mangrovi 2 4.12 4.08 4.15 —

morio 13 7.15 6.32 7.50 0.612

pacifica 1 6.00 6.00 6.00 -

rufitarsis 10 6.68 5.90 7.06 0.367

subinermis 10 6.82 6.46 7.26 0.287
tecta 2 6.28 6.25 6.32 -
 Revision of Holarctic species of Serromyia Meigen 161

W: width this study limited species determination to the

AR: antennal ratio; length of flagellomeres interpretation of morphological discontinuities.
10—13 divided by length of flagellomeres 1 —9 Although this study has little phylogenetic
for males and flagellomeres 9—13 divided by interpretation, readers should know that our
length of flagellomeres 1—8 for females approach is cladistic, as more fully described by
CR: costal ratio; costa length/wing length as Borkent (1984). We emphasize the importance
measured from MA of outgroup comparisons for the determination
HC/Ta 5 : length of hind leg claw divided by of character state polarity and recognize that
length of hind leg tarsomere 5 cladistic analyses are often only as good as are
MA: arculus those comparisons. The general lack of cladistic
Ta: tarsomere information in this study is due to a paucity of
interpretable character states.
Traditionally, the genus Serromyia has gen-
Rearing
erally been considered a member of the tribe
We were able to obtain eggs and first instar Stilobezziini, but most recently Wirth & Grogan
larvae of Serromyia nudicolis by decapitating (1988) have dealt with the historic difficulty of
females collected in the field. We found that distinguishing the Stilobezziini from the
females kept in the laboratory for 2—3 days Ceratopogonini by combining the two tribes.
before decapitation would be more likely to lay Serromyia, therefore, is now considered to be a
eggs. Resultant first instars did not survive on a member of the Ceratopogonini. Nevertheless,
nematode culture offered to them. The tech- the relationships between the genera in these
nique followed that successfully implemented two tribes remains problematic and the combin-
for other species of Ceratopogonidae by Linley ing of the two groups can only be regarded as a
(1985) using Panagrellus redivivus. interim measure until phylogenetic relationships
become better understood.
A consequence of this lack of understanding
Criteria for species recognition and
of phylogenetic relationships between the gen-
phylogenetic analysis
era is that outgroup comparisons must include
Reproductive isolation is the criterion for ob- virtually all other genera of Ceratopogonidae.
jective species recognition (Mayr, 1969: 26), As such, virtually every character state that we
but lack of bionomic and field observations in examined within Serromyia was also present

Table 7. Descriptive statistics for ratio of aedeagus length/width of male Serromyia.

Species n Mean Min. Max. SD

barberi 21 0.63 0.54 0.75 0.057

borealis 3 0.68 0.65 0.71 —

crassifemorata 9 0.70 0.65 0.81 0.054

nudicolis 13 0.65 0.56 0.73 0.047
sierrensis — — — — —

vockerothi 1 0.61 0.61 0.61 —

ledicola 22 0.46 0.39 0.54 0.041

atra 17 0.57 0.49 0.62 0.038
bicolor 3 0.52 0.48 0.59 —

femorata 20 0.54 0.41 0.62 0.053

mangrovi 2 0.61 0.56 0.65 —

morio 14 0.62 0.56 0.68 0.039

pacifica 1 0.70 0.70 0.70 —

rufitarsis 11 0.61 0.50 0.64 0.040

subinermis 11 0.59 0.49 0.72 0.063
tecta 2 0.55 0.55 0.56 —
162 A. Borkent and B. Bissett

Table 8. Descriptive statistics for antennal ratio of female Serromyia.

Species n Mean Min. Max. SD

barberi 50 1.23 1.04 1.38 0.076

borealis 2 0.96 0.93 0.99 —

crassifemorata 12 0.96 0.88 1.04 0.051

nudicolis 42 1.12 1.00 1.21 0.044
sierrensis 1 1.24 1.24 1.24 —

vockerothi 1 1.20 1.20 1.20 —

ledicola 33 1.27 1.06 1.40 0.074

atra 5 1.17 1.08 1.21 —

bicolor 2 1.10 1.04 1.16 —

femorata 15 1.21 1.10 1.28 0.051

mangrovi 2 1.14 1.10 1.19 -

morio 17 1.23 1.14 1.38 0.078

pacifica 1 1.13 1.13 1.13 —

rufitarsis 3 1.22 1.20 1.23

subinermis 10 1.25 1.17 1.39 0.1)72
tecta 2 1.06 0.96 1.15 —

Table 9. Descriptive statistics for the ratio of antennal flagellomeres 8/9 of female Serromyia.

Species n Mean Min. Max. SD

barberi 53 0.62 0.55 0.68 0.031

borealis 3 0.78 0.77 0.80 —

crassifemorata 13 0.77 0.69 0.95 0.074

nudicolis 43 0.68 0.62 0.82 0.041
sierrensis 1 0.61 0.61 0.61 —

vockerothi 2 0.60 0.59 0.60 —

ledicola 34 0.60 0.54 0.68 0.042

atra 5 0.69 0.65 0.76 —

bicolor 2 0.74 0.74 0.74 —

femorata 15 0.63 0.58 0.70 0.038

mangrovi 2 0.66 0.65 0.67 -

morio 17 0.63 0.59 0.68 0.026

pacifica 1 0.66 0.66 0.66 -

rufitarsis 4 0.68 0.62 0.72 -

subinermis 12 0.61 0.50 0.67 0.049

tecta 2 0.71 0.68 0.74 —

at least somewhere in the outgroup and the Classification and descriptive format
polarity (plesiomorphic/apomorphic) was
therefore uninterpretable. Because the Serromyia species we examined
Fresh clues to interpret the phylogenetic had so few interpretable apomorphies, we have
relationships between the genera in the arranged species as follows: the Nearctic species
Stilobezziini and Ceratopogonini indicate that in alphabetical order, the Holarctic species S.
current concepts need to be modified (Borkent, ledicola Kieffer, followed by the Palaearctic
in prep.) and we have therefore not used the species in alphabetical order.
phylogenetic interpretations proposed by All new species are to be attributed to the
Remm (1975) or Szadziewski (1988). senior author.
 Revision of Holarctic species of Serromyia Meigen 163

The characters described are those that are The diagnosis of the genus allows recognition
either useful for species discrimination or once the specimen is determined to belong to
for phylogenetic interpretation. Because we the family and the species diagnoses allow sep-
examined as many species of Serromyia as poss- aration from other Serromyia species.
ible on a world basis, the descriptions reflect When given localities on specimen labels or
variation of character states throughout the in past literature are now named differently, the
genus. current locality is given in square brackets.
Extent of leg pigmentation is shown diagram- We have listed the previous descriptions of
matically in the illustrations. Significant vari- species of Serromyia under the appropriate
ation is represented by two drawings for a name but only have done so when certain of
particular leg. that author's concept. For European workers,

Table 10. Descriptive statistics for the wing length of female Serromyia (in mm).

Species n Mean Min. Max. SD

barberi 50 2.11 1.68 2.54 0.181

borealis 3 1.65 1.60 1.72 —

crassifemorata 13 1.63 1.42 1.98 0.156

nudicolis 44 1.84 1.62 2.48 0.151
sierrensis 2 2.11 1.94 2.28 —

vockerothi 2 1.75 1.72 1.78 —

ledicola 36 2.25 1.66 2.74 0.234

atra 6 1.66 1.58 1.82 —

bicolor 2 2.08 2.06 2.10 —

femorata 19 2.35 1.96 2.68 0.209

mangrovi 2 1.42 1.39 1.45 -

morio 19 2.07 1.56 2.28 0.171

pacifica 1 2.16 2.16 2.16 -

rufitarsis 4 1.62 1.56 1.68 -

subinermis 10 1.97 1.72 2.16 0.134

tecta 2 1.56 1.55 1.57 —

Table 11. Descriptive statistics for the number of elongate strong bristles on the hind coxa of female Serromyia.

Species n Mean Min. Max. SD

barberi 54 0 0 0 0.00
borealis 3 0 0 0 —

crassifemorata 13 0 0 0 0.00
nudicolis 43 0.05 0 1 0.21
sierrensis 2 0 0 0 —

vockerothi 3 0 0 0 —

ledicola 32 2.53 0 4 1.11

atra 7 1.57 1 2 —

bicolor 2 0 0 0 —

femorata 20 0.05 0 1 0.22

mangrovi 2 0 0 0 -

morio 17 0.53 0 2 0.72

pacifica 1 4 4 4 —

rufitarsis 4 0 0 0 —

subinermis 12 0 0 0 0.00
tecta 2 0 0 0 -
164 A. Borkent and B. Bissett

Table 12. Descriptive statistics for the ratio of hind Ta]/Ta 2 of female Serromyia.

Species n Mean Min. Max. SD

barberi 54 2.14 1.91 2.79 0.150

borealis 3 1.92 1.83 1.97 —

crassifemorata 13 1.91 1.66 2.03 0.092

nudicolis 45 2.25 2.00 2.42 0.124
sierrensis 2 2.18 2.10 2.25 —

vockerothi 2 2.42 2.23 2.63 —

ledicola 39 2.40 2.11 3.12 0.191

atra 7 1.84 1.71 1.98 —

bicolor 2 2.50 2.43 2.56 —

femorata 19 2.34 2.19 2.78 0.121

mangrovi 2 1.84 1.84 1.84 -

morio 18 2.27 1.93 2.48 0.186

pacifica 1 2.32 2.32 2.32 -

rufitarsis 3 2.01 1.91 2.10 -

subinermis 12 2.45 2.27 2.69 0.109

tecta 2 2.07 2.03 2.11 —

Table 13. Descriptive statistics for the ratio of length of hind claw/length of hind tarsomere 5 (HC/Ta 5 ) of female
Serromyia.

Species n Mean Min. Max. SD

barberi 47 1.34 1.19 1.65 0.090

borealis 3 0.38 0.36 0.42 —

crassifemorata 13 0.35 0.30 0.44 0.040

nudicolis 40 0.84 0.62 1.06 0.103
sierrensis 2 1.33 1.27 1.39 —

vockerothi 2 1.03 0.98 1.08 —

ledicola 35 1.28 1.07 1.49 0.109

atra 4 0.33 0.31 0.38 —

bicolor 2 1.20 1.16 1.24 —

femorata 18 1.16 1.04 1.29 0.068

mangrovi 2 1.30 1.25 1.35 -

morio 19 1.14 0.97 1.35 0.107

pacifica 1 1.17 1.17 1.17 -

rufitarsis 3 0.76 0.67 0.89 -

subinermis 11 1.13 0.88 1.39 0.177

tecta 2 0.40 0.38 0.43 —

most descriptions could only be reinterpreted literature which may extend the range of a
when a given author's original material was given species are discussed under 'distribution
available for examination. and bionomics' or 'taxonomic discussion' of
Type specimens in the Canadian National that species.
Collection are given numbers in a reference text
and these numbers are reported here with the
Serromyia Meigen
description of the type locality of each named
species as 'CNC No.'. Serromyia Meigen 1818: 83. Type-species
Distribution maps are based entirely on Ceratopogon femoratus Meigen (by
material actually examined during the course of monotypy). Generic name cited in specific
this study. Further possible records from the synonymy.
 Revision of Holarctic species of Serromyia Meigen 165

Prionomyia Stephens 1829: 237. Type-species Ceratolophus Kieffer). Type-species Cerato-

Ceratopogon femoratus Meigen (subsequent pogon femoratus Meigen (automatic).
designation by Westwood 1840: 126). Diagnosis. Male adults: with hind femur
Ceratolophus Kieffer 1899: 69, not Barboza de markedly swollen, bearing at least two rows of
Bocage 1873. Type-species Ceratopogon strong, stout bristles ventrally, on more than
femoratus Meigen (by monotypy). apical 0.58 of hind femur. Female adults: with
Johannseniella Williston 1907: 1 (new name for hind femur markedly swollen, bearing at least
Ceratolophus Kieffer). Type-species Cerato- two rows of strong, stout bristles ventrally, with
pogon femoratus Meigen (automatic). base of claws straight or rounded and distal
Ceratolophana Strand 1928: 48 (new name for portion curved. Pupae: with only very short,

Foreleg Midleg

Figs 2A-F. Pigmentation of male fore and mid legs; A - B , Serromyia barberi-, C, S. borealis] D, [Link];
E, [Link]; F, [Link].
166 A. Borkent and B. Bissett

rounded abdominal tubercles and with body broadly separated ([Link] Delecolle &
setae very small. Fourth instar larvae: head Braverman, [Link] Debenham) or narrowly
very small in relation to body and all head separated to contiguous, bare; male antenna
capsule setae simple (not bifurcating) including with flagellomeres separate or with all or some
setae s, o. of flagellomeres 1 — 10 fused, with well-
developed plume, with flagellomeres 10—13 or
11 — 13 elongate; female antenna with each of
Description
flagellomeres 9—13 more elongate than each of
Adults. Moderately sized (wing length of flagellomeres 1—8, though in some species 9 is
males = 1.0—2.9 mm, of females = 1.2—2.7 only slightly longer than 8, sensilla coeloconica
mm); general body coloration patterned or dark absent; 5 palpal segments; third palpal segment
brown, either bare or with light pruinosity; eyes with shallow pit or lacking sensory pit but

Foreleg Midleg

Figs 3A-F. Pigmentation of male fore and mid legs; A, Serromyia vockerothi; B - C , [Link]; D, [Link];
E, [Link]'. F, [Link].
 Revision of Holarctic species of Serromyia Meigen 167

bearing on surface a patch of sensilla which are lacking short inner tooth, or with only a single
bulbous apically (as in Rowley & Cornford, elongate claw present, with spur near base
1972: Fig. 2); female mandible lacking or with present or absent (lacking in [Link],
7—14 teeth; thorax dull or shiny (with or without [Link]); hind femur markedly swollen,
light pruinosity), scutum lacking anteromedial with ventral patch of strong bristles arranged as
tubercle; humeral pits shallow, evident in a single row of 1—3 bristles basally, forming two
pruinose species as shiny bare patches; legs rows at mid length, scattering distally into an
either without strong bristles or variously indistinct pattern up to 4 bristles wide; hind
armed; female with claws of fore and mid legs tibia basally arcuate to fold against swollen hind
equal and short, with base of claws straight or femur, with dorsal row or rows of bristles; hind
rounded and distal portion curved (Figs first tarsomere with single ventral row of
16D—G); hind leg claws either equal, short, palisade setae, with additional stout subbasal

Foreleg Midleg

Figs 4A-F. Pigmentation of fore and mid legs; A, male Serromyia femorata; B, male [Link]; C - D , male
[Link]; E, female [Link]; F, female [Link].
168 A. Borkent and B. Bissett

setae present or absent; some species with fourth base; abdominal tergite 2 with anteromedial
tarsomeres bearing pair of sinuate setae triangular apodeme; male genitalia rotated
([Link], [Link] Meillon & Wirth, in some individuals of all species, lacking
[Link] Meillon & Wirth, [Link] apicolateral processes on tergite 9; with well-
Kieffer); wing with well-developed microtrichia, developed gonocoxite, gonostylus; most species
female wing lacking or with scattered macro- with anteromedial apodeme ventral to gonocoxal
trichia at apex; C R = 0 . 4 1 - 0 . 7 3 for males, apodeme; parameres separate or fused, fused
0.52—0.85 for females; cells r! and r2+3 dis- or articulated basally with gonocoxal apodeme;
tinct, cell rj/cell r 2+ 3 = 0.39-1.42 for males, aedeagus articulated laterally with base of go-
0.33—1.14 for females; stem of media very short nocoxite, some species with apex of aedeagus
to distinct, M2 indistinct to well developed at directed ventrally; male cercus large, situated

Foreleg Midleg
 Revision of Holarctic species of Serromyia Meigen 169

on lateral margin of segment 10, directed poster- Pupa. Length 3.1 (for [Link]) or 3.6—4.2
iorly; female genitalia with posterior margin of mm; general coloration light brown; dorsum of
sternite 8 somewhat bilobed to completely cleft thorax, dorsum and ventrum of abdomen
([Link] Clastrier & Wirth), with each lobe covered with short spicules; all body setae very
bearing a sometimes indistinct patch of setae; short, simple, tapering; operculum with apical
sternite 9 divided medially, either truncate or medial knob (similar to that described by Kettle
with anteromedial margin projecting medially; & Lawson, 1952: Fig. 90); anteromedial seta
segment 10 bearing 2 - 7 strong bristles ventrally, present, with sensory pit located just ventrally,
cercus well developed; 1—2 spermathecae, most anterodorsal seta single with associated sensory
species also with an additional blind sperma- pit, 2 dorsolaterals present, ventrolateral and
thecal duct, pigmented apically; spermathecae ventromedian setae absent, thoracic dorsals
ovoid to ellipsoidal, with or without surface with i, ii on abutting low tubercles, iv postero-
pores; a few members of some species with lateral to v; respiratory organ (as shown in
three fully developed spermathecae. Kettle & Lawson, 1952: Fig. 75, but somewhat

Figs 6A-F. Male genitalia C, F; aedeagus A, D; paramcrc B, E; A - C , Serromyia barberi; D - F , [Link].

170 A. Borkent and B. Bissett

wider at apex) dark brown except light brown at rest of head, ventral side with small medial
very base, length 0.85-1.26 mm, L/W = expansion and ventral suture, posterior margin
2.2—2.9, with 6 (for [Link]) or 8—11 spiracular of frontal sclerite slightly acuminate in middle,
openings, each on a slight protuberance; frontal suture terminating at pronotal sensory
metathorax indentate medially but not com- pit; sensory pits medium-sized, sensory pit
pletely divided (Kettle & Lawson, 1952: Fig. j around collar large, anterior one somewhat
102); setae on abdomen situated on low, smaller, all setae simple, of average length and
rounded tubercles, those on segment four dis- thickness, setae p short with small base, setae o
tributed as follows (Fig. 16B): 2 d.a.s.m., 3 with approximated bases abutting subgenal ring,
d.p.m., 4 l.p.m. (3 on one tubercle), 3 vent.; setae y of medium length, u and v with abutting
caudal spine with dark apex, directed laterally bases, seta w situated more anteriorly and a
at about 60° from longitudinal axis. short distance from subgenal ring, setae q, s, t,
Larva (partially after Glukhova, 1979). Head x present; antenna prominent with short, wide
short-oval, medium brown, collar weakly segments, segment 2 bearing 2-segmented short
developed, very narrow, barely darker than the rod and 4 small sensilla; eyes of medium size,

Figs 7A-F. Male genitalia C, F; aedeagus A, D; paramere B, E; A - C , Serromyia crassifemorata; D-F,

S. nudicolis.
 Revision of Holarctic species of Serromyia Meigen 171

somewhat spherical; labrum short, wide, Distribution and bionomics

sensillae as reported for [Link] below,
mandible short, brown, external surface con- The genus Serromyia is virtually cosmopolitan
cave, one denticle basally, maxillary palpus but has not been recorded from the Neotropical
short, hypostoma spherical, moderately convex, Region or from New Zealand. Several records
lacking denticles; labium narrow, anteriorly in the literature extend distributions of des-
acuminate plate; epipharynx with 2 sets of cribed species in Europe but we have unfortun-
plates; body very thick in relation to head, ately have been unable to confirm these.
lacking pigmentation, anal segment (segment Without reidentification, such records cannot
10) with very short setae, 4 dorsal setae not be used: Havelka (1978) reported [Link]
paired. from North Africa, and Remm (1973a) noted
Eggs. Markedly elongate, rounded anteriorly, [Link] from Mongolia. Remm (1967) reported
somewhat tapered posteriorly (Fig. 16A); in [Link] and S. morio (Fabricius) from the
S. nudicolis with longitudinal rows of spicules Caucasus.
(see under that species); egg shell known only We studied a series of eight females from
for [Link] (see under that species). Japan, labelled 'Japan, Hokkaido, Daisetzusan,

Figs 8A—F. Male genitalia C, F; aedeagus A, D; paramere B, E; A - C , Serromyia vockerothi; D - F , [Link].

172 A. Borkent and B. Bissett

Aizankei, 17-VII-1986, D.M. Wood CD812' above, members of the genus are certainly
(CNCI). These were very similar to the female expected to be present in China.
of [Link] Kieffer, but, considering the Although Fox (1946) considered
similarity between females of most Serromyia Johannseniella fluviatilis Lutz from the Neo-
species, could well represent another, unnamed tropical Region to be a species of Serromyia,
species. Further collecting will hopefully obtain Ortiz (1958) correctly assigned the species to
an associated male which will allow interpret- the genus Culicoides Latrielle.
ation of this species. Because so little bionomic information is
Yu (1978) included the genus Serromyia in available, it is uncertain what general statements
his key to Chinese genera, though no species can be made about the genus. All the Holarctic
has yet been identified from that country. We species seem to be associated with bogs, fens,
have examined an undescribed species from wet meadows, streams or small rivers. Further-
Taiwan (one female) and considering the more, south of treeline, all species are restricted
material of uncertain status from Japan noted to wooded regions (i.e. absent from prairie).
 Revision of Holarctic species of Serromyia Meigen 173

We have discussed further, more detailed Szadziewski & Krzywinski (1988) reported un-
habitat observations under each of the pertinent identified Serromyia from flowers of Anthriscus
species. silvestris in Poland.
Female adults of nearly all species of Information on immatures is very limited.
Serromyia ([Link] Malloch excepted) We were able to observe the eggs and first
have serrate mandibles, and limited obser- instar larvae of [Link] as discussed below
vations indicate that they capture and feed on under that species. Strenzke (1950) and
other insects (Edwards, 1920, 1923, 1926; Gad, Thienemann (1950) found larvae of [Link]
1951; Downes, 1955, 1978; see below under and [Link] in mosses at lake margins. Kettle
barberi, femorata). Edwards (1920) also noted & Lawson (1952) found Serromyia femorata
that the females of [Link] prey on the larvae in mud associated with marshlands. Be-
males during mating (see below under that cause of uncertainty of identification, it should
species). Females probably also take nectar be noted that the larvae collected by these
from flowers (see below under [Link]). authors may actually belong to another species
174 A. Borkent and B. Bissett

of Serromyia. One pupa in the Strenzke (1950) species as an unplaced species of Sphaeromiini.
collection was a member of [Link] (see below However, we have examined the female type,
under that species). housed in the NHRS, and found it belongs in
Atrichopogon Kieffer. This change results in a
new combination for the species.
Taxonomic discussion
Kieffer (1901) provided a brief description
This study indicates that the genus Serromyia (as part of a key) of a new species Serromyia
contains thirty-two extant and eight fossil scirpi based on a female from Germany and
species, reflecting some changes in taxonomic subsequently (1913, 1919) gave more complete
status and the description of several new species. descriptions (also as part of keys). Remm (1988)
A list of included species is given in Table 1. most recently also included the name in his
Wirth & Grogan (1988) included Ceratopogon Palaearctic catalogue as a species of Serromyia.
trichopus Thomson 1869 from China, in Several characters, however, suggest that
Serromyia while Wirth (1973) considered the [Link] is not actually a member of Serromyia.

A &

Figs 11A-F. Male genitalia C, F; aedeagus A, D; paramere B, E; A - C , Serromyia rufitarsis; D-F,

S. subinermis.
 Revision of Holarctic species of Serromyia Meigen 175

Kieffer (1913, 1919) noted that the fork of the communis Fabricius 1805 and [Link]
cubitus (as posticale) was markedly distal to the Meigen 1804 as members of Serromyia but there
position of the crossvein r-m (as transversale) is, in our opinion, no evidence for this. Remm
and that the wings were spotted. Combined (1988) placed C. communis in Ceratopogon
with the presence of short, equal claws on the (considering that Fabricius' description of
fore and mid legs and an elongate hind claw C. communis was not a new name but fol-
(equal to the length of the fifth tarsomere), lowed Meigen's earlier 1804 description) and
this description cannot apply to any known [Link] in Dasyhelea Kieffer. The date of
Serromyia, but does fit a general description of submission for the catalogue in Remm (1988)
several species of Palaearctic Monohelea. We was the end of 1982 and Szadziewski (1986),
therefore transfer the name to that genus as a after examining the type specimens, has shown
new combination: Monohelea scirpi (Kieffer). that the Ceratopogon palustris is actually a
Serromyia fuligipennis Clastrier has recently species of Forcipomyia. Havelka (1978) also
been placed in a new genus Congohelea Wirth interpreted Ceratopogon candidatus Winnertz
& Grogan and we agree that it was incorrectly 1852 as a member of Serromyia but considered
placed in Serromyia. the name unavailable because of lack of use
Havelka (1978) considered Ceratopogon (50-year rule). Remm (1988) correctly placed
176 A. Borkent and B. Bissett

this species as a synonym of Ceratopogon The species outside the Holarctic Region are
niveipennis Meigen. well described and keyed. Debenham (1970)
Stephens (1829) named Prionomyia pusilla described those in the Australian Region, Giles
as a new species among other members & Wirth (1982) dealt with those in the Oriental
of Serromyia (all of which he placed in and Australasian Regions and Meillon & Wirth
Prionomyia) but, without any description, the (1983) described and keyed those from the
name is clearly a nomen nudum. Afrotropical Region. The recently described
Some confusion has surrounded the author- [Link] Meillon & Downes from South Africa
ship and validity of the generic name Serromyia. (Meillon & Downes, 1986) has not been incor-
Although first published in synonymy with porated into a key but is the only Serromyia in
Ceratopogon, it is available under I.C.Z.N. the Afrotropical Region in which the female
Code Article 11(e). has equal hind claws.

Figs 13A-F. Photomicrographs of Serromyia colorata; A, hind leg (from holotype of [Link])', B, hind
claw (from holotype of [Link]); C, hind femur (from holotype of [Link]); D, fore, mid, and hind legs
(from holotype of [Link]); E, abdomen (from holotype of [Link]); F, antenna and hind claw (from
holotype of [Link] femorata).
 Revision of Holarctic species of Serromyia Meigen 25

There have been several characters which Kieffer, 1919; Goetghebuer, 1934; Zihali-
have been discussed and used extensively Sebess, 1940). However, it may be that freshly
by previous authors to recognize species of collected material from Europe exhibits differ-
Serromyia in the Palaearctic. Goetghebuer ences in colour that were not detected in this
(1922b, 1934) noted the difference in number study. The legs of specimens in the material we
of acrostichal and dorsocentral setae between examined were various shades of yellow, brown
[Link] and [Link]. Although the differ- and black.
ences are probably statistically significant, there Extent of leg coloration has also been used by
is overlap in the number of setae. We did not a number of authors to recognize species of
include this character in our study because we Serromyia. Winnertz (1852) considered such
could not count it consistently. pigmentation to be intraspecific variation. The
Several authors have noted differences in leg results of this study indicate that extent of
coloration (red, yellow, brown) which we have pigmentation can be used to recognize some
been unable to confirm (e.g. Meigen, 1818; species.

Figs 14A-F. SEM photomicrographs of Serromyia nudicolis; A - D , egg; E, first instar head capsule in
anterodorsal view; F, first instar head capsule in anterolateral view.
178 A. Borkent and B. Bissett

Zihali-Sebess (1940) used the ratio of wing [Link] had a range from 4.09 to 7.73 (n= 37).
vein m/r-m to distinguish some Serromyia Szadziewski (1988) recorded some values for
species but we found no statistically significant the fossil material, however, that exceeded these
differences between the species. values.
Szadziewski (1988), in his study of Baltic Meillon & Wirth (1983) and Wirth & Grogan
amber ceratopogonids, used the ratio of hind (1988) noted in their generic diagnoses that
femur length/width to distinguish some fossil male genitalia are 'concealed beneath the tip of
Serromyia species. Because of marked variation the pregenital terga' [= tergite 8]. Although
within extant species, we found the character to true for some specimens of some species, we
be of very limited value for males (Table 6) and have been unable to confirm this for living or
of no value for characterizing females, where freshly killed specimens. It is possible that such
total range values for all Holarctic species was a position is an artefact of preservation.
3.97—7.73. Intraspecific variation for female: Meillon & Wirth (1983) mistakenly drew the
of some species was also large; for example female genitalia of Serromyia aethiopiae as

Figs 15A- F. SEM photomicrographs of first instar larva of Serromyia nudicolis; A, anterior portion of head
capsule in anteroventral view; B, mouthparts in ventral view; C, mouthparts in ventral view; D, antenna and
labrum in anterolateral view; E, right maxillary palp; F, terminal abdominal segments in dorsal view.
 Revision of Holarctic species of Serromyia Meigen 179

having a very broad sternite 8. The lateral may also be due to drawing the specimen at an
margin of sternite 9 was included as part of angle.
sternite 8, which is, in fact, more narrow Mayer (1957) noted that members of the
posteriorly. Heteromyiini, Sphaeromiini and Palpomyiini
It is likely that spermathecal shape differs all have 3 anterior and 4 posterior dorsal setae
between at least some species of Serromyia but on abdominal segment four. Each of these two
we were unable to exami ! in a consistent groups of setae actually included a sensory
enough manner to categorize those differences. pit. In the somewhat dirty pupal material of
Apparent shape changes dramatically with dif- Serromyia examined we could not be certain
ferent orientations of the spermatheca within whether sensory pits were present or not. The
the body cavity. different sensilla counts, therefore, may not be
The only previous description of the pupal significant.
stage has been by Kettle & Lawson (1952), who Descriptions of larvae have been previously
characterized the pupa of [Link] (perhaps provided by Kettle & Lawson (1952) and
misidentified to species; not seen), and by Mayer Glukhova (1977, 1979). In addition, the first
(1957), who tabulated some setal distributions. instar larva of [Link] is described below.
Our description of the pupa above is based on a Our diagnosis above should allow recognition
few specimens (n=5) from the Strenzke and of the genus. The simple anal segment with very
Thienemann collections (ZSMC). One of these short setae is unusual but not unique, being
contained a pharate adult of [Link] but the characteristic in some Paradasyhelea Macfie,
others could not be confidently identified. Stilobezzia Kieffer and Sphaeromias Curtis.
Kettle & Lawson (1952) drew the respiratory We examined three fourth instar larvae of an
organ of the pupa of [Link] with the apex unidentified Serromyia from the Strenzke and
more narrow than that of any of the specimens Thienemann collections (ZSMC). These
we studied. This may indicate real variation but matched in nearly every detail the excellent
description by Glukhova (1979:160). Glukhova,
however, in drawing seta w in her ventral view
of the head capsule, failed to draw it in her
lateral view. The seta is just dorsal of the
subgenal ring. We failed to confirm the obser-
vation that the posterior margin of the eye was
slightly triangularly elongated. The specimens
we examined had eyes which were generally
spherical with a somewhat ragged edge. In
addition, we could not locate sensory pits r and
k.
We were unable to locate the specimens used
by Kettle & Lawson (1952) in their description
of [Link] larvae. Considering the uncer-
tainty in species identification at that time, these
may have been the larvae of one or more other
Serromyia species.
The description of the egg above is based
mostly on the undeveloped ova within preserved
female abdomens. Otherwise, only the egg of
[Link] is known (described below). A
recent investigation of the surface sculpturing
Fig. 16. A, egg shell of Serromyia nudicolis; B, of eggs of Culicoides (Kariya et al., 1989) show
setal distribution on fourth abdominal segment of significant differences between species, indi-
pupa of unidentified Serromyia; C - G , medial view of cating that this character may provide useful
posterior claw of female right fore leg: C, Metacan- information in distinguishing the eggs of differ-
thohelea cogani; D, [Link]; E, [Link]; F, 5. ent Serromyia species.
maculipennis; G, [Link]. The way the egg shell chorion is split is similar
180 A. Borkent and B. Bissett

to that reported for Culicoides circumscriptus intimated the possibility of splitting the group
Kieffer by Becker (1961) and Bezzia varicolor into two or more genera but could find no
(Coquillett) by Thomsen (1937), suggesting that evidence to support clear distinctions. We too
this is a widespread characteristic at least within could find no evidence to support the recog-
the Ceratopogoninae. Otherwise, so few egg nition of more than one genus.
shells have been described that no further Wirth & Grogan (1988) recently described
comparisons are possible. the new monotypic genus Metacanthohelea from
Meillon & Wirth (1983) noted the variation Africa. Further examination of two males and
among species of southern hemisphere Old two females of [Link] indicates that most
World Serromyia as compared to the relative character states which were indicated as dis-
homogeneity of species in the Holarctic. They tinguishing this genus also occur in at least some

Figs 17A-D. Distributions of: A, Serromyia barberi; B, [Link] (*), [Link] ( • ) ; C, [Link];
D, [Link] (*), S. vockerothi (•).
 Revision of Holarctic species of Serromyia Meigen 181

species of Serromyia (see discussion b e l o w portion straight (Fig. 16C); female hind claws
u n d e r ' P h y l o g e n y ' ) . T h e k e y p r o v i d e d by W i r t h short and equal, with well-developed inner tooth;
& G r o g a n (1988) is t h e r e f o r e revised as follows: male with less than 0.43 of apex of hind femur
bearing strong bristles
22. Female claws with rounded or straight base, distal Metacanthohelea Wirth & Grogan
portion curved (Fig. 16D—G); female hind claws
either single and elongate or, if short and equal, Boorman & Rowland (1988) recently
lacking inner tooth; male with more than 0.58 p r o v i d e d a g e n e r i c k e y to t h e c e r a t o p o g o n i d
of apex of hind femur bearing strong a d u l t s of G r e a t B r i t a i n . H o w e v e r , m a n y f e m a l e
bristles Serromyia Meigen
s p e c i m e n s of Serromyia f o u n d t h e r e will n o t
22. Female claws with marked bend at base, distal k e y o u t p r o p e r l y b e c a u s e of t h e p r e s e n c e of

Figs 18A-C. Distributions of: A, Serromyia ledicola; B, [Link] (•), [Link] (*); C, [Link].
182 A. Borkent and B. Bissett

macrotrichia on the wing tip. The key will basis using Wirth & Grogan (1988) with the
work if all specimens, regardless of sex, are modifications suggested above allowing for
treated as if they were male. better separation of Metacanthohelea and
Serromyia.
Some terms in the keys presented below
require special attention or interpretation and
the reader is directed to the above section
Keys
'Terms and abbreviations for structures'.
Members of the genus Serromyia may be Males are generally easier to identify than
recognized as such in the Holarctic Region females, especially those of species from the
using Downes & Wirth (1981) or on a worldwide Palaearctic.

11
Figs 19A-D. Distributions of: A, Serromyia femorata;
[Link] (*).
B, [Link]: C, [Link]; D, [Link] (•),
 Revision of Holarctic species of Serromyia Meigen 183

Key to adult males of Nearctic Serromyia 3. Cells ri and r 2 + 3 pale to dark brown; hind
species claw longer than tarsomere 5 (HC/Ta 5 = 1.07—
1.65) . 4
1. Mid femur yellow with apical portion more darkly
pigmented 2 Cells r] and r 2 + 3 dark brown; hind claw equal to
or shorter than tarsomere 5 (HC/Ta s = 0.62-
Mid femur uniformly dark brown 4 1.08) . " 6
2. Hind coxa with 0 - 4 elongate lateral setae (Fig. 4. Cells T] and r 2 + 3 dark brown; fore and mid tibiae
IB); fore femur with or without strong ventral brown (Sierra Nevada) sierrensis
bristles; scutum pruinose (as in Fig. 1A); adults
present 13 July to 6 November Cells r! and r 2 + 3 pale to light brown; fore and mid
(broadly Holarctic) ledicola tibiae pale or with just apex of mid tibia darkly
pigmented 5
Hind coxa lacking elongate lateral seta; fore femur
with or without strong bristles, if present situated 5. Coxa of hind leg with one or more strong lateral
at least anteriorly; scutum pruinose or lacking setae (as in Fig. IB); scutum pruinose (broadly
pruinosity; adults present 2 June to 3 July 3 Holarctic) ledicola

3. Fore femur with strong bristles anteriorly and Coxa of hind leg lacking strong lateral setae;
posteriorly; scutum pruinose (Canada, northern scutum lacking pruinosity (western
United States) vockerothi Nearctic) barberi

Fore femur with one strong bristle anteroapically; 6. Ratio of antennal flagellomeres 8/9 = 0.62-0.82
scutum lacking pruinosity (Sierra Nevada) (eastern Nearctic) nudicolis
sierrensis Ratio of antennal flagellomeres 8/9 = 0.59-0.60
4. Paramere with a distinctly pointed apex (Fig. 6B); (Canada, northern States) vockerothi
west of continental divide (western
Nearctic) barberi
Key to adult males of Palaearctic Serromyia
Paramere with a rounded (sometimes difficult to
see) apex (Figs 6E, 7B, 7E); east of continental species
divide 5 1. Body pale yellow (Egypt) mangrovi
5. Fore femur with stout bristles at least anteriorly Body dark brown to black 2
and posteriorly (eastern Nearctic) nudicolis
2. Parameres tapered apically to a definite point
Fore femur lacking strong bristles 6
(Figs 9B, 11B, E) 3
6. Scutum pruinose; aedeagus with lateral prongs
Parameres rounded apically, shrivelled in some,
(Fig. 6D) (Alberta, Manitoba) borealis
but apex not pointed (Figs 8E, 9E, 10B, E, 12B) .5
Scutum lacking pruinosity; aedeagus lacking 3. Scutum lacking pruinosity; each paramere
lateral prongs (Fig. 7A) (eastern Nearctic) bifurcate (Fig. 11B) apically (Europe). . . rufitarsis
crassifemorata
Scutum pruinose; each paramere ending in a single
point (Figs 9B, 11E) 4
Key to adult females of Nearctic Serromyia 4. Apex of paramere bent laterally at about a 45-90°
species angle (Fig. 11E) (Europe) subinermis
1. Claws of hind leg equal, markedly shorter than Apex of paramere straight, sword-like (Fig. 9B)
tarsomere 5 2 (Federal Republic of Germany, Caucasus) bicolor
Hind leg claw single and elongate, with one small 5. Thorax shiny; cells r t and r 2 + 3 dark brown;
s ; de tooth; claw equal to or longer than tarsomere aedeagus with side prongs directed anterolaterally
,r. 3 (Fig. 10A) 6
2. Mandible serrate; fore and mid tibiae darkly Thorax pruinose; cells ^ and r 2 + 3 light to pale
pigmented (Fig. 4F); scutum pruinose (Alberta, brown; aedeagus with side prongs directed
Manitoba) borealis dorsolaterally, laterally or posterolaterally (Figs
8D,9D, 10D, 12A) 7
Mandible reduced, lacking teeth; fore and mid
tibiae pale or yellow; scutum lacking pruinosity 6. Fore femur with three rows of strong bristles; wing
(eastern Nearctic) crassifemorata length = 2.0—2.5 mm (Europe) morio
184 A. Borkent and B. Bissett

Fore femur with one ventral row of strong bristles Fig. 4F) 7
and 0—2 anterior or posterior strong bristles; wing
length = 1.3-1.9 mm (Europe) atra Fore tibia only partially pigmented (Figs 4E, 5B,
( . H. I ) 8
7. Fore femur with three rows of strong bristles; hind
femur with anterior strong bristles (as in Fig. ID) 7. Hind coxa lacking strong lateral bristles
(Europe) femorata (Europe) subinermis (in part)
Hind coxa with about 4 strong lateral bristles
Fore femur lacking strong bristles or with one row
(eastern Palearctic) pacifica
of strong bristles in some composed of only a few
spines, some with 1 - 2 additional strong posterior 8. Coxa of hindlcg with 0 - 4 strong lateral bristles;
bristles; hind femur lacking anterior strong collected 20 July to 6 November (broadly
bristles 8 Holarctic) ledicola
8. Each of fore and mid femora and tibiae at most Coxa of hindlcg lacking strong lateral bristles;
only partly pigmented (Fig. 3B, C); parameres collected before 14 August 9
markedly swollen in apical half (Fig. 8E);
9. Wing slightly infuscatcd, with cells and r 2+ 3
gonostylus narrow and pointed apically (Fig. 8F)
dark brown (Europe) subinermis (in part)
(broadly Holarctic) ledicola
Wing pale, with cells ri and r 2+ 3 pale or slightly
Fore and mid femora and tibiae entirely brown (as
pigmented (Europe) femorata
in Fig. 3D); parameres sausage shaped, apical half
about equal in diameter for entire length (Figs 10. Mid femur entirely dark brown; HC/Ta s =
10E, 12B); gonostylus with rounded apex (Figs 0.67-0.89 (Europe) rufitarsis
10F, 12C) 9
Mid femur pale basally; HC/Ta 5 = 0.97-1.32
9. AR = 1.17; wing length 2.2 mm; apex of gonostylus (Europe) morio
markedly swollen (Fig. 10F) (eastern
Palearctic) pacifica
Serromyia barberi Wirth
AR = 1.00-1.03; wing length 1.6-1.8 mm; apex
Serromyia barberi Wirth 1952: 205. Holotype,
of gonostylus at most only slightly swollen (Fig.
12C) (Federal Republic of Germany) tecta 8 adult, pinned, labelled 'Eureka, Cal. 22.5',
'HS Barber Collector', 'Holotype Serromyia
barberi W . W . Wirth' (USNM); allotype 9
labelled as for holotype but collected [Link]
Key to adult females of Palaearctic
(USNM); paratypes: \16, 29 from type
Serromyia species
locality collected either 22.v or [Link] (USNM);
1. Body pale yellow (Egypt) mangrovi U.S.A.: 1 9 , Cal., Humboldt Co., Hely Creek,
Body dark brown to black 2 [Link].1948 (USNM); 1 $ , Cal., San Luis
Obispo Co. biacl. Lake Canyon (not found).
2. Hind claws equal, similar to fore and mid leg Diagnosis. Male: only Nearctic species with a
claws 3
paramere with a distinctly pointed apex.
Hind claws unequal (a single claw present), longer Female: only Nearctic species with the fore and
than fore and mid leg claws 4 mid tibae pale or with only apex of mid tibia
3. Scutum pruinose; hind coxa lacking strong lateral pigmented, with the scutum lacking pruinosity,
bristles (Federal Republic of Germany) . . . . tecta and with a single elongate hind claw.
Description. Male adult. Descriptive statistics:
Scutum lacking pruinosity; hind coxa with
1 - 2 strong lateral bristles (as in Fig. IB) see Tables 2—7.
(Europe) atra Head: dark brown; antennal flagellomere 10
with plume arranged in more than one whorl;
4. Palpus mostly pale, with only segments 4 and 5
palpus dark brown.
darkly pigmented (Federal Republic of Germany,
Thorax: dark brown; scutum bare of
Caucasus) bicolor
pruinosity.
Palpus uniformly dark brown 5 Legs: coloration pattern as indicated in Fig.
5. Scutum pruinose 6 2A, B; strong bristles on femora, tibiae dis-
tributed as follows: 1 present anteriorly, to 21
Scutum lacking pruinosity 10
present anteriorly, ventrally, posteriorly on fore
6. Fore tibia nearly entirely darkly pigmented (as in femur, present or absent posteriorly on fore
 Revision of Holarctic species of Serromyia Meigen 185

tibia, present or absent anteriorly, ventrally, British Columbia, with no flowing water nearby.
posteriorly on mid femur, present or absent Although rarely abundant, a large collection
posteriorly on mid tibia, present ventrally on of six males and 100 females was made at Mary's
hind femur, present or absent anteriorly on Peak (Parker's Creek), 21 km W Corvallis,
hind femur, dorsally on hind tibia; ventral Oregon on [Link].1985. In most other areas,
bristles on hind femur arising from slightly however, it often took 1—2 h to locate one
developed tubercles; hind tai straight; ta 4 setae specimen.
straight or slightly curved. During broad daylight S. barberi adults were
Wing: pale with veins of cells r t , r 2 + 3 darkly most commonly collected low in the vegetation
pigmented. and the edge of the net was held on the ground,
Abdomen: dark brown. while sweeping, to obtain these individuals.
Genitalia (Fig. 6C): gonostylus with outer However, some specimens were collected by
margin evenly curved, tapering gradually to sweeping at arm level, especially along the
slightly swollen, rounded or somewhat pointed margins of streams. Adults were restricted
apex; paramere (Fig. 6B) slender, with pointed to a limited area within a habitat so that, for
apex; aedeagus (Fig. 6A) lacking lateral prongs, example, the margin of a stream could be swept
distal portion a simple, slender projection, for several hundred metres without finding a
extreme apex directed ventrally. specimen and then suddenly two or three would
Female adult. Descriptive statistics: see be collected in the space of 5 m. We are unable
Tables 8—13. Similar to male except for usual to report any generalization which would predict
sex differences and as follows: where these specimens might be located.
Head: mandible serrate. In general, females were collected more
Legs: coloration pattern as indicated in Fig. frequently than males and this is reflected in
4E; strong bristles absent on femora, tibiae their relative numbers in the collections made
except ventrally on hind femur; claw of hind leg and in the museum collections examined.
single, elongate, with basal tooth. Observations of female feeding were made at
Wing: pale or pale with veins of cells r1? r 2 + 3 two localities. At 6.5 km SE of Hebo, Oregon,
darkly pigmented; macrotrichia restricted to female S. barberi flew out of stream margin veg-
apical margin. etation throughout the afternoon to dart into
Genitalia (as in Fig. 12D): sternite 9 trunc- small chironomid swarms which formed in dark
ate medially to somewhat truncate but with enclaves in the stream bank. At least some of
anteromedial margin developed, pointed; two these were of males of Nilotanypus Kieffer.
spermathecae, additional spermathecal duct Female S. barberi assumed a ragged up and
terminating in pigmented apex. down flight, somewhat mimicking the chiro-
Distribution and bionomics. S. barberi nomid flight pattern, and then left the swarm to
is known from the southern half of British return to the vegetation. We succeeded in
Columbia south to California (Fig. 17A). The collecting one female with a chironomid male
only other species west of the continental divide but could make no detailed observations as to
are S. sierrensis from the Sierra Nevada of how the prey was captured or held. At Parker's
California and S. ledicola which has been col- Creek on Mary's Peak, 21 km W Corvallis,
lected just west of the continental divtde in Oregon, females were quiescent until after dusk
Montana and Colorado. Adults of [Link] when they could be collected along the narrow
have been collected from 17 May to 11 August. gravel road which bisected the stream where
The most southerly locality of S. barberi was a many had been resting earlier. They were es-
specimen labelled 'S. Fork Santa Ana R., Calif., pecially common between 09.00 and 09.30 p.m.
18 June, 1945'. This disjunct indicates further Females flew in directed flight at the height of
profitable collecting in southern California. about 2—2.5 m until they met with chironomid
We found S. barberi most common in areas of swarms which formed along this road. They
seeps and along the margins of small streams, then joined the swarm and assumed a similar
nearly always associated with flowing water. A ragged up and down flight. We were unable to
few specimens were found along the margins of make further observations on prey capture or
larger rivers. However, we found one female at feeding. We placed white sheets under such
the margin of a fen, 3 km E Salmon Arm, swarms but no S. barberi females landed on
186 A. Borkent and B. Bissett

these, as may happen with prey capture in other except ventrally on hind femur, arising from
groups (Downes, 1978). slightly developed tubercles; hind Ta! straight
W. W. Wirth collected a male at Revelstoke, to with slight basal curvature; Ta 4 setae straight
British Columbia, on an Umbelliferae, where or with slight curve.
the specimen was presumably feeding on flower Wing: pale with veins of cells r l 5 r 2 + 3 slightly
nectar. pigmented.
We examined a gynandromorph female from Abdomen: dark brown.
Cowichan Lake, B.C. (CNCI) in which flagello- Genitalia (Fig. 6F): gonostylus with outer
meres 1—8 of the right antenna are male-like margin evenly curved, tapering gradually for
and plumose. basal half, with rounded, slightly to moderately
Taxonomic discussion. One character, gener- swollen apex; paramere (Fig. 6E) with apical
ally of good diagnostic value within the genus, half somewhat swollen, somewhat rounded, but
exhibited a puzzling degree of variation in male with apex difficult to discern; aedeagus (Fig.
S. barberi. The number of strong bristles on the 6D) with lateral prongs, directed laterally, distal
fore femur varied from 1 to 21, representing portion a simple, slender projection markedly
individuals with a near total lack of strong longer than lateral prongs, extreme apex
bristles on all the legs to very bristly specimens. directed ventrally.
In some areas, such as southern Vancouver Female adult. Descriptive statistics: see
Island, variation ranged virtually continuously Tables 8—13. Similar to male except for usual
from 2 to 21 bristles, while on Mary's Peak, sex differences and as follows:
21 km W Corvallis, Oregon the senior author Head: mandible serrate.
collected 6 males on [Link].1985 with 1, 1, 1, 10, Legs: coloration pattern as indicated in
13 and 17 bristles on the fore femur. We were Fig. 4F; claws of hind leg equal, small.
unable to find any correlation with size, geo- Wing: pale with veins of cells r 1 ; r 2 + 3 darkly
graphical or temporal distribution, although it pigmented; macrotrichia absent.
should be noted that only sixty-six males were Genitalia (Fig. 12D): sternite 9 truncate
available for study. medially; two spermathecae, additional sper-
The sex of one of the paratypes from Eureka mathecal duct terminating in pigmented apex.
was incorrectly given as male by Wirth (1952). Distribution and bionomics. [Link] is
We were unable to locate the paratype from known from Alberta to Manitoba (Fig. 17B).
Black Lake Canyon. Adults have been collected from 15 June to 4
Material examined. Seventy-two males and July.
268 females. We collected this species at two localities. At
Derivation of specific epithet. The name bar- 10 km E Spruce Grove, Alberta, two males
beri refers to the collector of the type series, H. were swept from the upper branches of short
S. Barber. (3—4 m) spruce trees in a well-developed
sphagnum bog. At the type locality, both males
and females were swept from grassy tussocks,
Serromyia borealis Borkent sp.n.
very near to the ground, in a black spruce bog.
Diagnosis. Male: only Nearctic species with Adults were restricted to a small patch of ground
mid femur completely pigmented brown and about 15 m in diameter, which was a little drier
aedeagus with lateral prongs. Female: only than the surrounding area.
Nearctic species with claws of hind leg equal, Taxonomic discussion. The association of
short (male-like) and mandible well developed, males and females was based on the number of
with teeth. each collected at the type locality. It should be
Description. Male adult. Descriptive statistics: noted, however, that one female of S. vockerothi
see Tables 2—7. was also collected at this site.
Head: dark brown; antennal flagellomere 10 Types. Holotype, 8 adult on microscope
with plume arranged in more than one whorl; slide, labelled 'Holotype Serromyia borealis
palpus dark brown. Borkent, CNC No. 20128, 8 , 3.2 km. N.
Thorax: dark brown; scutum pruinose. Nordegg, Alta., 28-VI-1985, A. Borkent
Legs: coloration pattern as indicated in Fig. CD379', allotype on microscope slide labelled
2C; strong bristles absent on femora, tibiae as for holotype (CNCI); paratypes: 98, 99
 Revision of Holarctic species of Serromyia Meigen 187

labelled as for holotype (CNCI, USNM); simple, slender projection, extreme apex
CANADA: I S , Alberta, Nordegg, [Link]. 1921 directed ventrally.
(CNCI); 28, Alberta, 10 km E Spruce Grove Female adult. Descriptive statistics: see
(Wagner Bog), [Link].1985 (CNCI); 19, Tables 8—13. Similar to male except for usual
Manitoba, Churchill, [Link].l947 (CNCI). sex differences and as follows:
Derivation of specific epithet. The name Head: mandible vestigial, no teeth evident.
borealis (northern) refers to the perceived Legs: coloration pattern as indicated in Fig.
boreal habitat of this species. 5B; strong bristles absent on femora, tibiae
except ventrally on hind femur; claw of hind leg
single, elongate, with basal tooth.
Serromyia crassifemorata Malloch
Wing: macrotrichia absent or restricted to
Serromyia crassifemorata Malloch 1914: 218. apical margin.
Lectotype designated by Frison (1927), 9 Genitalia (as in Fig. 12D): sternite 9 truncate
adult, pinned, labelled '1789', 'Serromyia medially to somewhat truncate but with
crassifemorata Malloch (Type)', ' 9 ' , 'Lecto- anteromedial margin developed, pointed; two
type 9 Serromyia crassifemorata Malloch', spermathecae, additional spermathecal duct
'Property of the Illinois Natural History Sur- terminating in pigmented apex.
vey' (1NHS); paralectotype 9 on pin from Distribution and bionomics. [Link]
same locality (as indicated by Malloch, 1914), is known from southern Ontario and Quebec,
with additional labels stating: '#1789 IL, Mt. south to Tennessee and Georgia (Fig. 17B).
Carmel 28 May 1884 from leaves of Quercus' Adults have been collected from 2 May to 22
'Serromyia crassifemorata Det. A. Borkent' June. Johannsen (1928, 1943) reported this
(INHS). species from Lewiston, New York, and, though
the species almost certainly does occur in New
Diagnosis. Male: only Nearctic species York, we failed to examine any material from
lacking strong bristles on legs, with mid femur that state.
uniformly dark brown, with aedeagus lacking The nonfunctional mandibles of the female,
side prongs and paramere with rounded apex. unique in the genus, indicates that it does not
Female: only Nearctic species with short, equal feed on other insects.
hind claws and reduced, nonserrate mandibles. The localities from which this species have
Description. Male adult. Descriptive statistics: been collected suggest that S. crassifemorata
see Tables 2—7. occurs in both lotic and lentic habitats. W. W.
Head: dark brown; antennal flagellomere 10 Wirth collected a series of females in Virginia in
with plume arranged in more than one whorl; Osmunda bogs (a genus of fern, characteristic
palpus light to dark brown. of wet woodland and some open swamps) and a
Thorax: medium to dark brown; scutum bare few from a stream margin at Alexandria,
of pruinosity. Virginia. We collected one specimen from
Legs: coloration pattern as indicated in Fig. Rondeau Provincial Park in southern Ontario
2D; strong bristles absent on femora, tibiae in a malaise trap located in a wet swampy area.
except ventrally on hind femur, present or Taxonomic discussion. The recognition of the
absent dorsally on hind tibia; ventral bristles on previously unknown male is based on the as-
hind femur arising from slightly developed sociation of twenty males with two females of
tubercles; hind Ta! straight; Ta 4 setae straight [Link] from Cobden, Illinois, col-
or with slight curve. lected 7—9.v. 1918. One additional male was
Wing: slightly infuscated, with veins of cells from Roaring Spring, Jonesboro, Illinois.
r x , r 2 +3 darkly pigmented. Wirth & Grogan (1981) provided a descrip-
Abdomen: dark brown. tion of the female of [Link].
Genitalia (Fig. 7C): gonostylus with outer Material examined. Twenty-one males and
margin evenly curved, tapering gradually to seventy-six females.
slightly swollen, rounded to somewhat pointed Derivation of specific epithet. The name
apex; paramere (Fig. 7B) with apical half crassifemorata means thickened femora and
sausage shaped, rounded apex; aedeagus (Fig. presumably refers to the thick hind femora of
7A) lacking lateral prongs, distal portion a this and all other species in the genus.
188 A. Borkent and B. Bissett

Serromyia nudicolis Borkent sp.n. basal tooth.

Wing: a few macrotrichia restricted to apical
Diagnosis. Male: only Nearctic species with margin.
mid femur completely darkly pigmented, with Genitalia (as in Fig. 12D): sternite 9 some-
numerous strong bristles on legs (at least fore what truncate but with anteromedial margin
femur with anterior and posterior strong developed, pointed; two spermathecae, ad-
bristles) and paramere with rounded apex. ditional spermathecal duct terminating in
Female: only Nearctic species with a ratio of pigmented apex.
antennal flagellomeres 8/9 = 0.62-0.82, and an First instar larva. Total length = 0.74-0.91
elongate, single hind claw, with HC/Ta 5 = mm (n=4).
0.62-1.06. Head: length = 78.4-84.0 |xm (n = 5), length/
Description. Male adult. Descriptive statistics: width = 1.40—1.53 (n = 3); capsule medium
see Tables 2—7. brown, with well developed, darkly pigmented
Head: dark brown; antennal flagellomere 10 egg burster (Fig. 14F); eye spot pigmented
with plume arranged in more than one whorl; black, shaped as simple, elongate oval, oriented
palpus dark brown. dorsoventrally; antenna large, flat disc, with 7
Thorax: dark brown; scutum pruinose. short sensilla at anterolateral margin of base
Legs: coloration pattern as indicated in Fig. (Figs 14E, 15D), setae x on medial margin of
2E; strong bristles of femora, tibiae distributed antenna! base (Fig. 15D); labrum with antero-
as follows: anteriorly, posteriorly on fore femur, lateral group of 4 short, thick sensilla with very
present or absent dorsally, ventrally on fore fine spicules at bases, a more medial, short seta,
femur, posteriorly on fore tibia, present or an elongate ventrolateral seta, an apical setiform
absent anteriorly, dorsally on fore tibia, at least sensillum (Figs 15B, C, D); mandible sickle
some present but present or absent anteriorly, shaped, elongate, extending in adducted state
dorsally, ventrally, posteriorly on mid femur, to level of setae o, apex longitudinally corru-
posteriorly on mid tibia, present or absent gated but mandible not seen well enough to
anteriorly, dorsally on mid tibia, anteriorly, describe further; maxilla with 1 elongate seta at
dorsally, ventrally on hind femur, dorsally on base of anteriorly projecting lobe, palpus
hind tibia; ventral bristles on hind femur arising reduced with 8 sensilla distributed as follows: 1
from slightly developed tubercles; hind Ta! small aboral, 2 large, stout aboral sensilla, 2 on
straight or with slight basal curvature; Ta 4 setae anterolateral lobe, 3 on anteromedial lobe (Figs
straight or with slight curve. 15C, E); labium with single, anteriorly pro-
Wing: slightly infuscated with veins of cells jecting tooth (Fig. 15C); otherwise following
r t , r2+3 darkly pigmented. setae present (Figs 14E, F, 15A, D): t, s, q, p,
Abdomen: dark brown. w, u, v, o, y (j could not be seen), distribution
Genitalia (Fig. 7F): gonostylus with outer similar to that given by Glukhova (1979) for
margin evenly curved, tapering gradually for fourth instar but seta u directly dorsal to v;
basal half, with pointed apex or with slightly sensory pits z, k, r, m, n absent; epipharynx
swollen, somewhat rounded apex; paramere with lateral arm well developed, with barely
(Fig. 7E) with apical half sausage shaped (some- discernible, posteriorly directed comb.
times somewhat shrivelled), apex rounded; Thorax, abdomen: lacking pigmentation;
aedeagus (Fig. 7D) lacking lateral prongs, distal thorax with well-developed collar; proleg
portion a simple, slender projection, extreme absent; perianal setae short, poorly developed,
apex directed ventrally. restricted to lateral, dorsal surface of segment 9
Female adult. Descriptive statistics: see (Fig. 15F); no evidence of anus, anal papillae.
Tables 8—13. Similar to male except for usual Egg. Dark brown, elongate, slightly curved
sex differences and as follows: (Fig. 14A), length = 592-632 \im, width =
Head: mandible serrate. 7 9 - 8 7 nm, length/width = 6 . 8 - 8 . 0 («=10);
Legs: coloration pattern as indicated in Figs with short tubercles arranged in somewhat
4E, 5A; strong bristles of femora, tibiae distrib- scattered, longitudinal rows (Figs 14A, B)
uted as follows: 1 - 2 apically on mid tibia (easily seen at x 100 under compound micro-
(absent on one specimen), ventrally on hind scope); tubercles longer around micropyle,
femur; claw of hind leg single, elongate, with arranged in circular pattern, amongst which are
 Revision of Holarctic species of Serromyia Meigen 189

very small, blunt tubercles (Figs 14C, D); egg those which failed to develop) were pale or pale
shell with circular cap at very anterior end and brown. Larvae hatched about 12-15 days after
dorsal, longitudinal fissure restricted to anterior eggs were laid on agar plates and held at about
1/3 (Fig. 16A). 20°C. Emerging first instars were very active,
Distribution and bionomics. [Link] is moving with a slow snake-like motion charac-
known from Ontario to Newfoundland, south teristic of many other ceratopogonid larvae. We
to Tennessee and Georgia (Fig. 17C). The introduced nematodes (as described in Materials
southern localities are restricted to the and Methods) but, in spite of observing one
Appalachian Mountains and probably reflect larva feeding, failed to rear these to even the
the cool, wet habitat of this species. Adults second instar.
have been collected from 23 May to 30 July, but Taxonomic discussion. Wirth (1952: 206,
seem most common, at least in southern Ontario 1965) identified members of S. nudicolis as
and Quebec and the northeastern states, from [Link].
mid June to mid July. Types. Holotype, 8 adult on microscope
We collected adults of this species in moist slide, labelled 'Holotype Serromyia nudicolis
deciduous or mixed deciduous—coniferous Borkent, CNC No. 20126, 8 , Maine, 33.5 km S
woods, always in the presence of small running Ashland CD587, 2-VII-1986, A. Borkent'
seeps or streams. In some but not all instances (CNCI); allotype on microscope slide labelled
these waters led to wet meadows or fens. as for holotype (CNCI); paratypes: 118, 179,
We made some observations on male adult 2 first instar larvae, 1 egg, 1 egg shell labelled as
behaviour at the type locality, 33.5 km S for holotype (CNCI, USNM); CANADA: 7 9 ,
Ashland, Maine. At this place the vegetation Ontario, Iroquois Falls, various dates ranging
had been recently cut and trees and shrubs were [Link]—[Link].l987 (CNCI); 19, Ontario, Black
only about 2—3 m high. We were able to collect Lake, 15 km SW Lanark, [Link].1975 (CNCI);
males from the very tops of small Abies 19, Ontario, Algonquin Park, [Link].l960
balsamea. Sampling at regular intervals led to (USNM); 2 9 , 2 first instar larvae, 13 egg shells,
repeated collections, indicating that the males Ontario, 10 km SW Richmond, [Link]. 1987
were actively seeking out these sites. In general, (CNCI); 8 9 , Ontario, Ottawa, [Link].1946,
we were able to collect mostly females of [Link]. 1946, [Link].l951, [Link].1964, [Link].1980
S. nudicolis and these observations of tree (CNCI); 2 9 , Ontario, Waubamik, [Link]. 1915
topping may indicate that the males were scarce (USNM); 19, Ontario, Finland, [Link].1960
in collections because they swarm, or at least (CNCI); 3 9 , Ontario, Maynooth, [Link].1953
congregate, at the apices of tree tops where we, (CNCI); 19, Ontario, Midland, [Link].1956
and most other collectors, have been unable to (CNCI); 19, Quebec, Mistassini, [Link].1956
collect at most sites. (CNCI); 2 9 , Quebec, Beechgrove, [Link].l955,
We were able to observe on two separate [Link].1962 (CNCI); 2 9 , Quebec, Gatineau
occasions female avoidance behaviour. We in- Park, Black Lake, [Link].1985, [Link].l985
itially found each of two females hanging on the (CNCI); 4 9 , Quebec, Gatineau Park,
underside of leaves about 1 m above the ground. Bourgeois Lake, [Link].1987 (CNCI); 9 9 , 1 first
Upon being slowly approached to a distance of instar larva, 6 egg shells, Quebec, 5 km W Old
about 50 cm, each of these females dropped to Chelsea, [Link].1987 (CNCI); 5 9 , Quebec, Old
the ground. Net collected specimens of this and Chelsea, [Link].1959, [Link].1961, [Link].l964
of every other Nearctic species we collected, (CNCI); 19, Quebec, Kam. Co., Parke Re-
exhibited a similar behaviour of dropping to the serve, [32 km S Riviere du Loup], [Link].l957
net bottom when disturbed. (CNCI); 2 9 , Quebec, Meach Lake, [Link].1916,
The eggs and first instar larvae of S. nudicolis [Link].l916 (CUIC); 18, 6 9 , New Brunswick,
were obtained by decapitating field caught fe- Kouchibouguac National Park, vii.1977,
males and placing these in petri dishes on a [Link].1977, [Link].1978 (CNCI); 3 9 ,
shallow layer of agar. Eggs from such females Nova Scotia, Cape Breton Highlands National
were laid at the rate of 12—16 per minute with Park, North Mountain, [Link].1983 (CNCI);
some laid in a group and abutting laterally, 19 , Newfoundland, mountains east of Codroy,
while others were scattered on the agar. Fertile [Link].1905 (MCZC); 19, Terra Nova
eggs were dark brown, while non-fertilized (i.e. National Park, Newfoundland, [Link]. 1961
190 A. Borkent and B. Bissett

(USNM); U.S.A.: 6 9 , Wisconsin, Washburn mid femur yellow with apical portion more darkly
Co., [Link]. 1953 (USNM); 2 9 , Michigan, pigmented and scutum lacking pruinosity.
Gladwin Co., [Link].1958, [Link].1943 (USNM); Female: only Nearctic species with fore and mid
1 9 , Michigan, Kalkaska Co., [Link].1951 tibiae dark brown and with an elongated, single
(USNM); 1 9 , Michigan, Livingston Co., E.S. hind claw with the claw longer than Ta 5 .
George Reserve, [Link].l950 (USNM); 1 9 , Description. Male adult. Descriptive statistics:
Michigan, Saginaw Co., [Link].l952 (USNM); see Tables 2—6.
19 , Michigan, Alpena Co., [Link]. 1941 (USNM); Head: dark brown; antennal flagellomere
2 9 , Michigan, Wexford Co., 14. vi. 1952 10 with plume arranged in more than one whorl;
(USNM); 109, Michigan, Crawford Co., palpus dark brown.
[Link].1951, [Link].1953 (USNM); 5 9 , Michigan, Thorax: dark brown; scutum bare of
Cheboygan Co., [Link].l953 (USNM); 3 9 , pruinosity.
Michigan, Midland Co., [Link]. 1952, [Link].1952 Legs: coloration pattern as indicated in Fig.
(USNM); 1 9 , New York, Caroline-Harford, 2F; strong bristles of femora, tibiae distributed
[Link]. 1904 (CUIC); 129 , New York, Tompkins as follows: one anteroapically on fore femur,
Co., Ringwood Reservoir, 16—[Link]. 1963 one anteriorly, one posteriorly on mid femur,
(USNM); 39 , New York, Orleans Co., Albion, ventrally on hind femur, dorsally on hind tibia;
11 .vi. 1963 (USNM); 5 9 , New York, St ventral bristles on hind femur arising from
Lawrence Co., Cranberry L., 25. vi. 1963 slightly developed tubercles; hind Ta t straight;
(USNM); 4 9 , New York, St Lawrence Co., Ta 4 setae straight or with slight curve.
Benson Mines, [Link].1963 (USNM); 18, 149, Wing: slightly infuscated with veins of cells
New York, Lewis Co., Whetstone Gulf, 20— xx, r 2 + 3 darkly pigmented.
[Link].1963 (USNM); 19, New York, North Abdomen: dark brown.
Beach, L.I., 30.V.1930 (MCZC); 19, New York, Genitalia: broken from specimen, lost.
Slaterville [nr Candor], [Link].1904 (INHS); 28, Female adult. Descriptive statistics: see
129, 1 first instar larva, 14 egg shells, Vermont, Tables 8—13. Similar to male except for usual
5 km E Danby, [Link]. 1986 (CNCI, INHS, sex differences and as follows:
USNM, RYSC, NHMW); 19 , New Hampshire, Head: mandible serrate.
Third Connecticut Lake [10 km N Idlewilde], Legs: coloration pattern as indicated in Fig.
[Link]. 1952 (USNM); 18, New Hampshire, 4F; strong bristles of femora, tibiae distributed
Franconia (AMNH); 7 9 , Maine, 14 km NW as follows: single bristle at apex of mid tibia,
Camden, [Link].1986 (CNCI, RYSC); 1 9 , ventrally on hind femur; claw of hind leg single,
Maine, Katahdin, 4.V.1959 (USNM); 1 9 , elongate, with basal tooth.
Connecticut, Redding, 27.v. 1933 (USNM); 1 9 , Wing: slightly infuscated with veins of cells
Pennsylvania, Centre Co., Pine Grove Mills, r t , r23 darkly pigmented; a few macrotrichia
[Link]. 1975 (CASC); 19, Pennsylvania, Centre restricted to apical margin.
Co., [Link]. 1972 (CASC); 19, Massachusetts, Genitalia (as in Fig. 12D): sternite 9 truncate
Burgess, [Link].1876 (USNM); 48, 4 9 , Massa- medially to somewhat truncate but with an-
chusetts, Holliston 26.v, [Link].l956 (MCZC, teromedial margin developed, pointed; two
USNM); 19 , West Virginia, Cranberry Glades spermathecae, additional spermathecal duct
[= 23 km E Richwood], [Link].l955 (USNM); terminating in pigmented apex.
19, Tennessee, Gatlinburg, [Link].1947 (USNM); Distribution and bionomics. [Link] is
18, 6 9 , North Carolina, Highlands, 23.v. 1957, known only from the Sierra Nevada of
29.v. 1957 (CNCI); 19 Georgia, Unicoi State California (Fig. 17D). Adults have been col-
Park (nr Helen), 17.v. 1979 (CNCI). lected from 2 June to 3 July.
Taxonomic discussion. This is the only species
Derivation of specific epithet. The name described as new in which the holotype is a
nudicolis (nude, penis) refers to the aedeagus female. The single male was missing its genitalia,
which lacks side prongs. and although we are confident that it is cor-
rectly associated with the two females, we are
reluctant to designate such a damaged specimen
Serromyia sierrensis Borkent sp.n.
as the holotype.
Diagnosis. Male: only Nearctic species with The male of [Link] has an unusually high
 Revision of Holarctic species of Serromyia Meigen 191

antennal ratio and this feature, once more ma- basal half, with rounded or somewhat pointed
terial becomes available, may prove to be dis- apex; paramere (Fig. 8B) with apical half
tinctive, at least within the western Nearctic. sausage shaped (sometimes somewhat
CASC provides type numbers for all its holo- shrivelled), apex rounded; aedeagus (Fig. 8A)
types and has assigned [Link] no. 16472. with lateral prongs directed posterolaterally,
Types. Holotype, 9 adult on microscope somewhat reduced in some, distal portion a
slide, labelled 'Holotype Serromyia sierrensis simple, slender projection, markedly longer
Borkent, 9 , CAL [California] Placer Co., E. than lateral prongs, extreme apex directed ven-
end Bear Val., 2-VI-1964, P.H. Arnaud, Jr.', trally.
'Serromyia barberi Det. W. Wirth, '76', Female adult. Descriptive statistics: see
(CASC); allotype 8 on microscope slide. U.S.A: Tables 8—13. Similar to male except for usual
Cal., Nevada Co., Sagehen Creek, [Link]. 1974 sex differences and as follows:
(CASC); paratype: 1 9 , Cal., Sierra Co., Head: mandible serrate.
Webber Lake, [Link].l964 (CASC). Legs: coloration pattern as indicated in Fig.
Derivation of specific epithet. The name 5B; strong bristles of femora, tibiae distributed
sierrensis refers to the type locality in the Sierra as follows: absent or 1 present on fore femur, 1
Nevada of California. present apically on mid tibia, ventrally on hind
femur; claw of hind leg single, elongate, with
basal tooth.
Serromyia vockerothi Borkent sp.n.
Wing: a few macrotrichia restricted to apical
Diagnosis. Male: only Nearctic species with margin.
mid femur yellow and pigmented only apically, Genitalia (as in Fig. 12D): sternite 9 some-
with strong bristles on fore femur (at least pre- what truncate medially but with anteromedial
sent anteriorly and posteriorly) and with scutum margin developed, pointed; two spermathecae,
pruinose. Female: only Nearctic species with a additional spermathecal duct terminating in
ratio of antennal flagellomeres 8/9 = 0.59—0.60 pigmented apex.
and an elongate, single hind claw about the Distribution and bionomics. [Link] is
same length as Ta 5 . known from Alberta to southern Quebec and
Description. Male adult. Descriptive statistics: from one locality in Minnesota (Fig. 17D).
see Tables 2—7. Adults have been collected from 4 to 26 June.
Head: dark brown; antennal flagellomere 10 The holotype and allotype were collected near
with plume arranged in more than one whorl; the margin of a shallow, small (i.e. 7—8 m by
palpus dark brown. 2 m, woodland pool with thick moss and rich
Thorax: dark brown; scutum pruinose. vegetation growing around its margin. The
Legs: coloration pattern as indicated in Fig. single female from 3.2 km N Nordegg was taken
3A; strong bristles of femora, tibiae distributed in a black spruce bog (habitat described more
as follows: anteriorly, posteriorly on fore femur, fully under [Link]).
present or absent ventrally on fore femur, pos- Taxonomic discussion. Although we are
teriorly on fore tibia, present or absent an- reasonably confident of the conspecificity of the
teriorly, dorsally on fore tibia, anteriorly, three males and the allotype, we are less sure
ventrally, posteriorly on mid femur, posteriorly that the two female specimens from Alberta are
on mid tibia, present or absent on mid tibia, accurately interpreted. It may be that they are
anteriorly, ventrally on hind femur, present or actually members of [Link] or of another
absent posteriorly on hind femur, dorsally on undescribed species. Further collecting, espec-
hind tibia; ventral bristles on hind femur arising ially of males in Alberta, is required.
from slightly developed tubercles; hind Ta! The allotype is somewhat damaged and is
straight or with slight basal curvature; Ta 4 setae missing all tarsomeres and flagellomeres 6 - 1 3 .
straight or with slight curve. When first collected, this female was believed
Wing: slightly infuscated with veins of cells to be a specimen of [Link] and was kept
ri, r 2 + 3 darkly pigmented. alive in the hopes of obtaining some eggs. Un-
Abdomen: dark brown. fortunately, she died and was damaged before
Genitalia (Fig. 8C): gonostylus with outer being properly preserved.
margin evenly curved, tapering gradually for A male from Abbotsford, Quebec ([Link].
192 A. Borkent and B. Bissett

1937, CNCI) may be a member of this species dated [Link].33 (ISNB).

but lacked the lateral prongs on the aedeagus. Serromyia europaea Clastrier 1963: 61.
One of the paratypes of S. vockerothi had some- Holotype, 8 adult on microscope slide, la-
what reduced lateral prongs, indicating that belled 'Autriche [Austria] Heiligenblut
lack of lateral prongs may be within the variation 10—VIII —1960 a la lumiere Serromyia
of the species. Further material is required to europaea Holotype 8 2356, Serromyia ledi-
determine whether the Abbotsford specimen is cola Det. A. Borkent' (MNHN).
S. vockerothi or represents an undescribed Ceratopogon femoratus: authors, not Meigen.
species. Staeger 1839: 598 (in part).
Types. Holotype, 8 adult on microscope Zetterstedt 1838: 822 (in part), 1850: 3665 (in
slide, labelled 'Holotype Serromyia vockerothi part). Coquillett 1900: 396.
Borkent CNC No. 20127, 6 , 10 km S Victoria Serromyia femorata: of authors not Meigen.
Beach, Manitoba, [Link]. 1985, A. Borkent Wirth 1965: 136. Havelka 1976: 236 (in part).
CD368', allotype 9 on microscope slide la- Havelka & Caspers 1981: 30 (in part).
belled as for holotype (CNCI); paratypes: Diagnosis. Male: only Holarctic species with
CANADA: \ 8 , Quebec, Mt. St. Hilaire, 5 0 0 - fore femur, at most, with a few ventral strong
700 ft, [Link].l963 (CNCI); 19, Alberta, bristles, mid femur pigmented only at apex,
Edmonton, 1932 (CASC); 1 9 , Alberta, 3.2 km scutum pruinose, and parameres with apical
N Nordegg (Wagner Bog), [Link].1985 (CNCI); half markedly swollen and rounded (in Nearctic,
U.S.A.: I d , Minnesota, Itasca Park [30 km N only species with short aedeagus, L/W = 0.39—
Park Rapids], [Link]. 1938 (USNM). 0.54). Female: only Holarctic species with
Derivation of specific epithet. The name palpus uniformly dark brown, fore and mid
vockerothi is proposed in appreciation of our tibiae pigmented only apically, hind coxa with
friend and colleague Dr J. Richard Vockeroth. 0—4 lateral strong bristles, scutum pruinose,
His outstanding collecting efforts in all the and an elongate, unequal hind claw with HC/
Diptera are reflected in this study. He collected Ta 5 = 1.07-1.49.
specimens of nearly every Nearctic Serromyia Description. Male adult. Descriptive statistics:
species (including this one) as well as important see Tables 2—7.
series of [Link], [Link] and [Link] Head: dark brown; antennal flagellomere 10
from Europe. In addition, his unstinting willing- with plume arranged in more than one whorl;
ness to share his vast knowledge of Diptera, his palpus dark brown.
knowledge of the literature and location of often Thorax: dark brown; scutum pruinose.
difficult to find museum specimens added Legs: coloration pattern as indicated in Figs
immeasurably to this study. 3B, C; strong bristles of femora, tibiae dis-
tributed as follows: present or absent ventrally
on fore femur, present or absent anteriorly,
ventrally, posteriorly on fore tibia, present or
Serromyia ledicola Kieffer
absent ventrally on mid femur, present or absent
Serromyia ledicola Kieffer 1925a: 156. Neotype, anteriorly, ventrally, posteriorly on mid tibia,
here designated, 8 adult, on microscope slide, ventrally on hind femur, dorsally on hind tibia;
labelled 'Estonia Hiiumaa, H. Remm, 8—8— ventral bristles on hind femur arising from
53, Serromyia ledicola K det. H. Remm, slightly developed tubercles; hind Tai straight;
Neotype CNC No. 20129'. (CNCI). Remm Ta 4 setae straight or with slight curve.
1969: 214 (in part). Wing: pale or pale with veins of cells r 1? r 2 + 3
Serromyia macronyx Goetghebuer 1933: 355. darkly pigmented.
Lectotype, here designated, 8 adult, pinned Abdomen: dark brown.
but subsequently mounted on microscope Genitalia (Fig. 8F): gonostylus with outer
slide, labelled 'La Panne 7.9.33 M. Goet- margin evenly curved, tapering gently for basal
ghebuer', '[Link].N.B. 18.073Collet det., M. half, with pointed apex; paramere (Fig. 8E)
Goetghebuer', 'macronyx', 'Type Lectotype with apical half markedly swollen, apex
8 M. Goetghebuer. Serromyia ledicola Det. rounded; aedeagus (Fig. 8D) with lateral
A. Borkent' (ISNB); paralectotype 9 adult prongs, directed posterolaterally, distal portion
on microscope slide from type locality but a simple, slender projection, about twice as
 Revision of Holarctic species of Serromyia Meigen 193

long as lateral prongs, extreme apex directed localities with climatic extremes: Adak and
ventrally. Popof Islands in the Aleutians of Alaska and
Female adult. Descriptive statistics: see Lake McDonald, Montana (35 km NE
Tables 8—13. Similar to male except for usual Columbia Falls). If these are excluded, the
sex differences and as follows: earliest collection we examined in North
Head: mandible serrate. America was from 1 August. The only other
Legs: coloration pattern as indicated in Figs Nearctic species which overlaps with [Link]
4E, 5C; strong bristles absent on femora, tibiae is S. barberi of which a few specimens have been
except ventrally on hind femur; claw of hind leg collected as late as 11 August. In Europe a
single, elongate, with basal tooth. similar pattern is apparent. With the exception
Wing: pale or pale with veins of cells r1? r 2 + 3 of a sample from Obergurgl, Tirol, Austria,
light brown; a few to many macrotrichia taken near the foot of a glacier from 20 July to 3
restricted to apical margin. September, the earliest sample was from
Genitalia (as in Fig. 12D): sternite 9 truncate 1 August. Only the sympatric Palaearctic species
medially to somewhat truncate but with antero- [Link] and S. morio overlap temporally with
medial margin developed, pointed; two late records of 14 and 16 August respectively.
spermathecae, additional spermathecal duct [Link], from the Sinai, has been recorded
terminating in pigmented apex. as late as 18—19 November.
Distribution and bionomics. [Link] is Taxonomic discussion. We attempted to
known in Europe from Sweden and Estonia locate the Kieffer holotype female of [Link],
south to England and northern Italy and in collected by A. Dampf, but failed. Considering
North America from Alaska to Newfoundland that so many of Kieffer's types are known to
south to Colorado and Virginia (Fig. 18A). have been lost or destroyed, it is likely that the
Adults have been collected from 13 July to 6 type of [Link] is also gone. We have there-
November. fore designated a neotype from the same area
Remm (1973a) noted the presence of from which the holotype was originally collected
[Link] in Mongolia, collected at the end of (Mavli bog on Dago [= Hiiumaa Island],
July. We have not examined the specimens but Estonia [U.S.S.R.]).
the records are consistent with the conclusion North American workers have uniformly
that S. ledicola is a Holarctic species. considered Nearctic material of this species
The two disjunct southern Nearctic records (sometimes also including material of other
are from Gould, Colorado and Reddish Knob, species) as [Link]. However, it is clear
Augusta Co., Virginia. from examination of the type of [Link] that
The collections from Alaska, particularly this was a misinterpretation.
along the Aleutian Island chain, support the Remm (1981) provided the synonymy listed
conclusion that [Link] is a Holarctic species. here and we have been able to partially confirm
Although not yet recorded from the far eastern this through examination of the types of
Palaearctic, it is certainly to be expected there. [Link] and [Link]. He correctly states
The record of [Link] (as Ceratopogon that the late flight period may be taken as
femoratus) from Popof Island, Alaska by evidence of the synonymy, even though as far
Coquillett (1900) is actually of a specimen of as [Link] is concerned, some [Link]
S. ledicola. and [Link] also fly into August. However, we
The few available records of habitat indicate have been unable to confirm that the holotype
that this species is associated with boggy areas, of [Link] was collected late in the season
sometimes in the vicinity of seeps or small and correspondence with Dr Remm questioning
streams but also in areas with only standing him about this reached him after his untimely
waters. Goetghebuer (1936b) suggested that death. Regardless, the rather general de-
this species (as [Link]) was restricted to scription by Kieffer (1925a) does fit the concept
eutrophic lentic habitats in southern Belgium. of S. ledicola as presented here (female with
S. ledicola is virtually unique in the Holarctic long hind claw and yellow legs lacking spines)
fauna in being a late season emerger. Although and we accept Remm's suggestion of ledicola
the overall emergence period is from 13 July to being the oldest available name.
6 November, the earlier records are from Goetghebuer (1933) noted that there were
194 A. Borkent and B. Bissett

three males and fifteen females in the type atra Det. A. Borkent' (HNHM).
series of S. macronyx but we were only able to Serromyia spinosipes Kieffer 1919:72. Lecto-
examine the single male and female noted type, here designated, 8 adult on microscope
above. slide, labelled 'spinosipes typus [in red] Kieff.
Remm (1969) provided a key to the Serromyia det. Kieffer, 8 , Budapest Kertesz, Purchd.
species in the European portion of the U.S.S.R. from Budapest Mus. B.M. 1922-72. Lecto-
Based on specimens he sent to us, his concept type, Serromyia atra Det. A. Borkent, Return
of female ledicola included specimens of to Brit. Mus. (N.H.)' (BMNH); paralectotype
S. morio. 9 adult labelled as for lectotype but identified
We have examined Zetterstedt's collections as Serromyia morio (BMNH). New synonym.
and found a male and female of this species in Serromyia nitens Goetghebuer 1920: 73. Lecto-
material identified as [Link]. In addition, type, here designated, 8 adult on microscope
Zetterstedt (1850) mentioned that he had slide, labelled 'Wemmel [just north of
collected material in September, which can only Brussels, Belgium], 4 - 6 - 1 8 [June 4, 1918],
refer to [Link]. [Link].N.B. 18.073 Coll. et det., M.
One of the specimens identified by Staeger Goetghebuer, nitens Goetghe., Lectotype,
(in ZMUC) as Ceratopogon femoratus was in 8 , Serromyia atra Det. A. Borkent' (ISNB);
fact [Link]. paralectotype: 9 labelled as for lectotype but
One of the paratypes of S. europaea (from date probably incorrectly given as 4 June
Norway) (MNHN) is actually a male of 1917 (ISNB). New synonym.
S. femorata. Diagnosis. Male: only Palaearctic species with
Material examined. 85 males and 109 females. fore femur with only one longitudinal, ventral
Derivation of specific epithet. The name row of strong bristles plus 0—2 strong bristles
ledicola (Ledum (labrador tea), dweller) prob- anteriorly, and with the prongs on the aedeagus
ably refers to the boggy habitat of [Link], directed anterolaterally. Female: only Palae-
where Ledum grows and from which Kieffer arctic species with equal hind claws and a
(1925a) recorded this species. scutum bare of pruinosity.
Description. Male adult. Descriptive statistics:
see Tables 2—7.
Serromyia atra (Meigen) Head: dark brown; antennal flagellomere 10
with plume arranged in more than one whorl;
Ceratopogon ater Meigen 1818: 84. Neotype, palpus dark brown.
here designated, 8 adult on microscope slide, Thorax: dark brown; scutum bare of
labelled 'Ceratopogon ater Meigen, 8 , Ser- pruinosity.
romyia atra Mg. H. Remm, Neotype, Latvia, Legs: coloration pattern as indicated in Fig.
Jumurda Lake [10 km E. Ergli], E. Remm, 3D; strong bristles of femora, tibiae distributed
2 1 - 6 - 6 7 CNC No. 20130 Serromyia atra as follows: ventrally on fore femur, one bristle
Det. A. Borkent' (CNCI). present or absent anteriorly on fore femur,
Prionomyia atra: Stephens 1829: 238. ventrally on mid femur, present or absent
Ceratolophus ater: Kertesz 1902: 157. anteriorly, posteriorly on mid femur, 0—3
Serromyia ater: Kieffer 1906: 65. bristles anteriorly on hind femur, ventrally on
Serromyia atra: Kieffer 1919: 74. hind femur, dorsally on hind tibia; ventral
Serromyia micronyx Kieffer 1919: 70. Neotype, bristles on hind femur arising from slightly
here designated, 8 adult on microscope slide, developed tubercles; hind Ta t with slight basal
labelled 'Serromyia micronyx Kieffer, curvature; Ta 4 setae straight or with slight curve.
Neotype, 6 , 1967. V. 16, leg. Mihalyi, Wing: slightly infuscated with veins of cells
Kiskomarom [15 km. W. Marcali], Hungary, r l 5 r 2 + 3 darkly pigmented.
erdo, Serromyia atra Det. A. Borkent' Abdomen: dark brown.
(HNHM). New synonym. Genitalia (as in Fig. 10C): gonostylus with
Serromyia albitarsis Kieffer 1919: 71. Neotype, outer margin evenly curved, tapering gradually
here designated, 8 adult, labelled 'Serromyia for basal half, with somewhat pointed to
albitarsis Kieffer, Neotype, 1965.V. 19, leg. swollen, rounded apex; paramere (as in Fig.
Mihalyi, Ocsa, Hungary, lapret, Serromyia 10B) with apical half markedly swollen, apex
 Revision of Holarctic species of Serromyia Meigen 195

rounded (but shrivelled in some); aedeagus (as ater was based on material from Megerle.
in Fig. 10A) with lateral prongs, directed antero- Megerle's material was destroyed in the fire of
laterally, distal portion a simple, gradually 1848 at the Vienna Museum (Pont, 1986). We
tapering projection, about twice as long to examined three specimens from the Winthem
markedly longer than lateral prongs, very apex collection (in NHMW) which were labelled as
directed ventrally. 'ater', but unfortunately not in Meigen's hand-
Female adult. Descriptive statistics: see writing. These could therefore not be considered
Tables 8—13. Similar to male except for usual as type material and the type material of [Link]
sex differences and as follows: is considered to be lost.
Head: mandible serrate. The description of both S. micronyx and
Legs: coloration pattern as indicated in Fig. S. albitarsis by Kieffer (1919) as having females
5D; strong bristles of femora, tibiae distributed with a brilliantly shiny thorax and short, equal
as follows: present or absent ventrally on fore, hind claws can only refer to [Link] as described
mid femora, ventrally on hind femur; claws of here. The neotypes are proposed to ensure
hind leg equal, small. stability of the names applied to Palaearctic
Wing: macrotrichia restricted to apical species.
margin. It is clear from Kieffer's (1919) list of nine
Genitalia (as in Fig. 12D): sternite 9 truncate localities for [Link] that he based his de-
medially to somewhat truncate but with antero- scription on a number of specimens. The only
medial margin developed, pointed; two available material of Kieffer's type series
spermathecae, additional spermathecal duct was the lectotype and a paralectotype housed
terminating in pigmented apex. in the BMNH. The single paralectotype of
Pupa. Described under generic description. [Link], however, is a female adult of
Distribution and bionomics. [Link] is known S. morio.
from England to Latvia, south to Belgium and Kieffer's (1919) description of [Link]
Hungary (Fig. 18B). Adults have been collected does not quite fit the description of [Link] here
from 13 May to 6 July. in that the femora and tibiae of [Link] are nearly
Little can be said about the bionomics of this completely darkly pigmented. Kieffer (1919)
species. Goetghebuer (1936b) noted that adult reported [Link] with the fore and mid
[Link] were associated with rivers in northern femora and tibiae as mostly yellow. However,
Belgium but we were unable to confirm his the lectotype here designated fits the description
identifications. Two specimens from Britain had of [Link] in this study.
labels which noted that the specimens came One female pupa (ZSMC, material from
from fens (Chippenham, Wood Walton Strenzke, 1950) we examined contained a
(Hunts)). Strenzke (1950) collected a pupa from pharate adult and was somewhat dirty. What
the shore of Schohsee, F.R.G. few differences between this specimen and other
Taxonomic discussion. The association of the available material of Serromyia pupae we could
males and females was based on specimens detect (size and number of spiracular openings)
collected at the same place and time in Latvia are noted in the generic description. The label
(by Remm), Belgium (by Goetghebuer: types on the slide noted 'femorata (?) morio?' and
of [Link]) and Amsterdam (by Meijere, 1946: '246' (therefore collected at Schohsees accord-
reported as [Link]). ing to Strenzke's (1950) records).
Neotypes have been designated for Cera- The male genitalia of [Link] are indistinguish-
topogon ater, Serromyia micronyx and able from those of S. morio. However, the
Serromyia albitarsis because type specimens characters given in the keys and significant dif-
could not be located and have probably been ferences in the ratio of antennal flagellomeres
destroyed. 10/11 indicates that these are separate species.
The description of Ceratopogon ater by Material examined. 59 males, 11 females and
Meigen (1818) and the accompanying plate 1 female pupa.
drawn by Meigen (in Morge, 1975) showing the Derivation of specific epithet. The name ater
extent of leg pigmentation, corresponds to the (black) presumably refers to the overall dark
concept of [Link] as proposed where. Meigen appearance of adults of this species noted by
(1818) noted that his description of Ceratopogon Meigen.
196 A. Borkent and B. Bissett

Serromyia bicolor Borkent sp.n. the uniquely (amongst Holarctic Serromyia)

bicoloured palpus.
Diagnosis. Male and female: only Palaearctic The male of S. bicolor is unusual in a number
species with palpal segments 1 - 3 pale, segment of characters and is the most distinctive of all
4 light brown and segment 5 dark brown. the Holarctic species (other than mangrovi).
Description. Male adult. Descriptive statistics: Colour pattern of both male and female palpus,
see Tables 2 - 7 . the peculiar apex of the aedeagus, the sword-
Head: dark brown; antennal flagellomere 10 like parameres and a markedly slender (for a
with plume arranged in more than one whorl; species of Serromyia in the Holarctic) hind
palpus with segments 1—3 pale, segment 4 light femur of the male are all unique to this species.
brown, segment 5 dark brown. Havelka & Caspers (1981) partially described
Thorax: dark brown; scutum pruinose. this species under the name S. subinermis Kieffer
Legs: coloration pattern as indicated in Fig. (males are those of [Link] as recognized here,
3E; strong bristles of femora, tibiae distributed females are actually those of S. bicolor and
as follows: ventrally on mid femur, ventrally [Link]). However, the name subinermis
on hind femur, dorsally on hind tibia; ventral cannot apply to the present species as Kieffer
bristles on hind femur arising from slightly (1919) noted that S. subinermis had spines
developed tubercles; hind T a j straight; Ta 4 setae (strong bristles) on the fore and mid femora in
with slight curve or some slightly sinuate near the female and none in the male. Male [Link]
their apex. have strong bristles on the mid femur only
Wing: slightly infuscated with veins of cells and in the female these were either present or
r 1 ; r 2 + 3 darkly pigmented. absent. As far as we can discern, there are no
Abdomen: dark brown. Palaearctic names which could apply to this
Genitalia (Fig. 9C): gonostylus sinuous at species and have accordingly described it as
midlength, tapering gradually for basal half, new.
with swollen apex; paramere (Fig. 9B) blade- In their drawing of the male genitalia of
like, with pointed apex, directed posteriorly; [Link] (as [Link]), Havelka & Caspers
aedeagus (Fig. 9A) lacking lateral prongs, distal (1981) misinterpreted the apex of the aedeagus
portion truncate, with jagged margin, with as elongate and slender.
posterolateral lobes, extreme apex directed Types. Holotype, 8 adult on microscope
posteriorly. slide, labelled 'Holotype Serromyia bicolor
Female adult. Descriptive statistics: see Borkent, Bonn, Federal Republic of Germany,
Tables 8 - 1 3 . Similar to male except for usual June 7, 1976, P. Havelka, A551' (PEHC);
sex differences and as follows: allotype, 9 adult on microscope slide, labelled
Head: mandible serrate. as for holotype but collected 24 May 1976
Legs: coloration pattern as indicated in (PEHC); paratypes: 28 from holotype locality,
Fig. 5B; strong bristles of femora, tibiae dis- dated 10.v. 1976 (CNCI), 11.v. 1976, (PEHC);
tributed as follows: present or absent ventrally 19 U.S.S.R., Caucasus, Dilizan, [Link].l969
on mid femur, ventrally on hind femur; claw of (CNCI).
hind leg single, elongate, with basal tooth. Derivation of specific epithet. The name
Wing: macrotrichia restricted to anteroapical bicolor (two-coloured) refers to the distinctive
margin. bicoloured palpus of males and females of this
Genitalia (as in Fig. 12D): sternite 9 truncate species.
medially, with anteromedial margin developed,
pointed; two spermathecae, additional sperma-
thecal duct terminating in pigmented apex.
Serromyia femorata (Meigen)
Distribution and bionomics. [Link] is
known from Bonn, West Germany, and from Ceratopogon femoratus Meigen 1804: 28.
Dilizan in the Caucasus Mountains of the Lectotype, here designated, 8 adult on
Armanskaja S.S.R. (Fig. 18B). Adults have microscope slide, labelled 'Lectotype Cera-
been collected from 10 May to 7 June. topogon femoratus 8 Meigen, ex coll. Meigen
Taxonomic discussion. We have associated 291/40 (158a), prep. J. Clastrier, 1985 baume
the single female with the males on the basis of phenole, Museum Paris, Serromyia femorata
 Revision of Holarctic species of Serromyia Meigen 197

Det. A. Borkent' (MNHN); paralectotypes: Clastrier 1963: 61 (in part).

2 9 labelled as for holotype (MNHN). Staeger Ceratopogon morio: authors, not Fabricius.
1839: 598 (in part). Zetterstedt 1838: 822 (in Staeger 1839: 598 (in part). Zetterstedt 1850:
part), 1850: 3665 (in part), Schiner 1864: 584 3666 (in part).
(in part).
Chironomus femoratus: Fabricius 1805: 45. Diagnosis. Male: only Palaearctic species with
Prionomyia femorata: Stephens 1829: 237. three rows of strong bristles on the fore femur,
Ceratolophus femoratus: Kieffer 1899: 69. scutum pruinose and a pale wing. Female: only
Johannseniella femoratus: Williston 1907: 1. Palaearctic species with palpus darkly pig-
Serromyia femorata: Kieffer 1901: 157. Kieffer mented, the scutum pruinose, nearly the entire
1906: 65. Goetghebuer 1934: 61 (in part). fore and mid tibiae yellow with only very apicies
Edwards 1926: 410 (in part). pigmented in some, strong lateral bristles absent
Ceratopogon armatus Meigen 1818: 83. Lecto- from hind coxa and a pale wing.
type, here designated, 6 adult on microscope Description. Male adult. Descriptive statistics:
slide, labelled 'Lectotype Ceratopogon see Tables 2—7.
armatus 6 Meigen, ex coll. Meigen 291/40 Head: dark brown; antennal flagellomere 10
(161), prep. J. Clastrier, 1985 baume phenole, with plume arranged in more than one whorl;
Serromyia femorata Det. A. Borkent, palpus dark brown.
Museum Paris' (MNHN). Thorax: dark brown; scutum pruinose.
Prionomyia armata: Stephens 1829: 238. Legs: coloration pattern as indicated in Figs
Ceratolophus armatus: Kertesz 1902: 157. 3F, 4A; strong bristles of femora, tibiae distrib-
Serromyia armata: Kieffer 1906: 65. uted as follows: anteriorly, ventrally, posteriorly
Ceratopogon foersteri Meigen 1838: 21. Neo- on fore femur, dorsally, posteriorly on fore
type, here designated, 8 adult on microscope tibia, present or absent anteriorly on fore tibia,
slide, labelled 'Neotype Serromyia foersteri anteriorly, ventrally, posteriorly on mid femur,
Meigen, 8 , Newcastle-u-Lyme, Staff., posteriorly on mid tibia, present or absent an-
England 3-VI-1960, J. R. Vockeroth CNC teriorly or dorsally on mid tibia, anteriorly,
No. 20249, Serromyia femorata Det. A. ventrally on hind femur, dorsally on hind tibia;
Borkent' (CNCI). ventral bristles on hind femur arising from
Ceratolophus foersteri: Kertesz 1902: 157. slightly developed tubercles; hind Taj straight
Serromyia foersteri: Kieffer 1906: 65. or with slight basal curvature; Ta 4 setae straight
Ceratopogon flavipes Gimmerthal 1847: 144, or with slight curve.
not Ceratopogon flavipes Meigen 1804: 28. Wing: pale or slightly infuscated with veins of
Neotype, here designated, 8 adult on micro- cells r1? r 2 + 3 very lightly pigmented.
scope slide, labelled 'Neotype Ceratopogon Abdomen: dark brown.
flavipes Gimmerthal, 8 , Oxford, England, 6- Genitalia (Fig. 9F): gonostylus with outer
V-1953, J. R. Vockeroth, Neotype CNC No. margin evenly curved, tapering gradually
20272, Serromyia femorata, Det. A. Borkent' for basal half to tapering gradually to rounded
(CNCI). New synonym. or somewhat swollen apex; paramere (Fig.
Palpomyia flavicrus Kieffer 1906: 63. New name 9E) with apical half markedly swollen, apex
for Ceratopogon flavipes Gimmerthal. New rounded; aedeagus (Fig. 9D) with lateral
Synonym. prongs, directed posterolaterally, to dorso-
Serromyia flavicrus: Remm 1988: 35. lateral^, distal portion a simple, slender pro-
Serromyia inermipes Kieffer 1919: 73. Neo- jection, about twice to markedly longer than
type, here designated, adult on microscope lateral prongs, extreme apex directed ventrally.
slide, labelled 'Neotype Serromyia inermipes Female adult. Descriptive statistics: see
Kieffer, Brockenhurst, Hants [= Hampshire], Tables 8—13. Similar to male except for usual
England 3-V-1951, J. R. Vockeroth, CNC sex differences and as follows:
No. 20273, Serromyia femorata Det. A. Head: mandible serrate.
Borkent' (CNCI). Legs: coloration pattern as indicated in Figs
Ceratopogon rufitarsis: authors, not Meigen. 5B, C; strong bristles of femora, tibiae distrib-
Meigen 1818: 83 (in part). uted as follows: present or absent ventrally on
Serromyia europaea: authors, not Clastrier. fore femur, present or absent ventrally on mid
198 A. Borkent and B. Bissett

femur, present or absent at extreme apex of The specimens which Edwards (1920) noted
mid tibia, ventrally on hind femur; claw of hind as preying on Cricotopus pulchripes Verr. and
leg single, elongate, with basal tooth. Bezzia ornata (Meigen) were examined and are
Wing: macrotrichia absent or a few restricted indeed members of [Link].
to very apical margin. The following are literature records of prey
Genitalia (as in Fig. 12D): sternite 9 trunc- or feeding for which we were unable to examine
ate medially to somewhat truncate but with the original material and confirm the identifi-
anteromedial margin developed, pointed; two cations. Edwards (1923) noted Trichocladius
spermathecae, additional spermathecal duct sp. (Chironomidae) as prey. Gad (1951) re-
terminating in pigmented apex (one specimen ported [Link] attacking lsohelea sociabilis
with three fully developed spermathecae (Goetghebuer) and Culicoides impunctatus
present). Goetghebuer. Goetghebuer (1949) reported
Distribution and bionomics. [Link] is that S. femorata males fed on flowers and females
known from northern Fennoscandia south to preyed on Metriocnemus (Chironomidae).
northern Spain, northern Yugoslavia and Edwards (1926) noted that male [Link]
Moscow (Fig. 19A). The locality in the Kola swarm at mid-day and Downes (1955) reported
Peninsula shown on the map is based on a female swarms composed of 3—20 individuals
specimen with no further specific distributional of [Link] under the tips of alder branches
data. Adults have been collected from 6 May to at Loch Lomond, Scotland. Although these
14 August. records are consistent with the distribution
Havelka (1978, 1979) and Gil Collado (1957, of this species, we were unable to examine
1985) noted records of [Link] (Meigen) any of this material and thereby confirm the
from Spain. We have examined one female identifications.
from Santander housed in the MZLU but have Taxonomic discussion. The type of Cera-
been unable to confirm these previous records. topogon foersteri Meigen is not present in Paris
Remm (1967) reported two specimens of (Clastrier, pers. comm.) or in Halle (Dorn,
[Link] from the Causcasus and these pers. comm.) and we have accordingly desig-
identifications, too, could not be confirmed. nated a neotype for the species. Meigen's (1838)
Strenzke (1950) noted the presence of larvae description is so general that this species might
of S. femorata in lake-side mosses in Germany actually be one of any number of European
but we have been unable to confirm these species. His drawing of the female (in Morge,
identifications (material was examined, see 1975) shows a unique leg coloration with apically
under generic description). darkened fore and mid femora and completely
We examined five pairs of [Link] col- pigmented fore and mid tibiae. However, some
lected in copula from Norway, Sweden and pinned specimens of [Link] may have
Britain which allowed for confident association somewhat darkened tibiae (as in Fig. 5C).
of the sexes. Previous workers (Goetghebuer, 1920, 1934;
One of the mated pairs (from Sweden) had Remm, 1988) have considered this species to be
the male abdomen broken between the sixth a synonym of S. femorata and the designation of
and seventh abdominal segments with the the neotype here will establish this.
male's genitalia firmly attached to that of the The type of Serromyia inermipes Kieffer could
female. This indicates a mating behaviour not be located. Remm (1988) considered the
similar to that recorded for various species of name a synonym of [Link] although the
Heteromyiini, Sphaeromiini and Palpomyiini general description given by Kieffer (1919)
(Downes, 1978). Edwards (1920) noted that he could apply to several European species. We
collected mating pairs of [Link] in which have designated a neotype to confirm Remm's
the females were feeding on males. We were synonymy.
unable to locate the specimens upon which these Remm (1988) recently placed Ceratopogon
observations were based and therefore cannot flavipes Gimmerthal as a species of Serromyia.
confirm the identification. The presence of Based on the brief description given by
the broken male abdomen and attached male Gimmerthal (1847), in which he noted that the
genitalia noted above would correspond to the underside of the hind femora of the two female
presence of such a feeding behaviour. syntypes were spinose, this is probably correct.
 Revision of Holarctic species of Serromyia Meigen 199

The rest of the description, however, is so gen- bristles on hind femur arising directly from flat
eral that it does not allow for a confident place- cuticle or from very slightly developed tubercles;
ment of the name. The mention that the fore hind Tai with marked basal bend; Ta 4 setae
and mid 'Schenkel' [probably = femur] and straight or with slight curve.
'Gelenke' [probably = base of tibia] were brown Wing: pale.
is approximately a condition occurring in some Female adult. Descriptive statistics: see
female [Link]. The designation of the Tables 8 - 1 3 .
neotype here places Ceratopogon flavipes Legs: strong bristles of femora, tibiae dis-
Gimmerthal as a synonym of [Link]. tributed as follows: apically on mid tibia,
Goetghebuer (1922a) noted that the syntype mventrally on hind femur.
series of Ceratopogon femoratus was composed Genitalia: sternite 9 truncate medially.
of one male and four females. We examined Distribution and bionomics. S. mangrovi is
only the male and two females. known only from the Sinai Peninsula, Egypt.
The male paratype of [Link] from Nor- The specimens we examined were collected
way and the male paralectotype of S. rufitarsis from 4/5 June to 18/19 November at a light
were actually specimens of [Link]. trap.
Edwards (1926) suggested that female [Link]- Taxonomic discussion. Although Delecolle
morata lack macrotrichiae on the wing mem- & Braverman (1987) designated only nine
brane but this is true for only some specimens. males and nine females as paratypes, they also
Material examined. 199 males and 246 examined an additional thirteen males and 122
females. females from the Sinai.
Derivation of specific epithet. The name Material examined. Two males and two
femoratus (pertaining to the femora) pre- females collected at E-Shira el Gharkana
sumably refers to the swollen hind femur of and Ras Muhammad, Sinai Peninsula, Egypt
members of this and several other genera of (deposited in CNCI).
Ceratopogonidae. Derivation of specific epithet. The name
mangrovi refers to the mangrove swamp where
Serromyia mangrovi Delecolle & the specimens were collected.
Braverman
Serromyia mangrovi Delecolle & Braverman
1987: 57. Holotype, 9 adult, not seen, from Serromyia morio (Fabricius)
EGYPT: Sinai, Ras Muhammad (MZSF); Culex morio Fabricius 1775: 800. Neotype, here
allotype, 8 adult, not seen, labelled as for designated, 8 adult on microscope slide,
holotype (MZSF); paratypes (not seen): labelled 'Neotype Serromyia morio Fabricius,
EGYPT: 9<J, 9 9 , Sinai (MZSF, MNHN). Oxford, England 13-V-1953, J. R. Vockeroth
Diagnosis. Male and female: only Palaearctic CNC No. 20131' (CNCI).
species with body coloration uniformly pale Ceratopogon morio: Meigen 1818: 84. Staeger
yellow. 1839: 598 (in part). Zetterstedt 1850: 3666 (in
Description. As given by Delecolle & part).
Braverman (1987). Additional character states Prionomyia morio: Stephens 1829: 238.
as follows: Ceratolophus morio: Kertesz 1902: 158.
Male adult. Descriptive statistics: see Tables Serromyia morio: Kieffer 1906: 65. Edwards
2-7. 1926: 410. Goetghebuer 1934; 62.
Head: antennal flagellomere 10 with plume Serromyia nudipennis Kieffer 1913: 10. Neo-
arranged in a single whorl. type, here designated, 8 adult on microscope
Thorax: scutum pruinose. slide, labelled 'Neotype Serromyia nudi-
Legs: strong bristles of femora, tibiae dis- pennis Kieffer, Oxford, England 13-V-1953,
tributed as follows: 1 anteriorly, 1 posteriorly J. R. Vockeroth CNC No. 20248, Serromyia
on apex of fore femur, posteriorly on fore tibia, morio Det. A. Borkent' (CNCI). New
1 anteriorly, 1 posteriorly on apex of mid femur, synonym.
2 ventrally on mid tibia, 1 anteriorly, ventrally Ceratopogon femoratus: authors, not Meigen.
on hind femur, dorsally on hind tibia; ventral Schiner 1864: 584 (in part).
200 A. Borkent and B. Bissett

Serromyia femorata: authors, not Meigen. Tables 8 - 1 3 . Similar to male except for usual
Edwards 1926: 410 (in part). Goetghebuer sex differences and as follows:
1934: 61 (in part). Head: mandible serrate.
Serromyia atra: authors, not Meigen. Legs: coloration pattern as indicated in Figs
Goetghebuer 1934: 61. 4E, 5E; strong bristles of femora, tibiae dis-
Serromyia spinosipes: authors, not Kieffer. tributed as follows: absent or present ventrally
Kieffer 1919: 72. on fore femur, absent or present on mid femur,
Serromyia ledicola: authors, not Kieffer. Remm ventrally on hind femur; claw of hind leg single,
1969: 214 (in part). elongate, with basal tooth.
Ceratopogon flavicornis: authors, not Staeger. Wing: macrotrichia with very few to many,
Zetterstedt 1850: 3667. restricted to apical margin.
Genitalia (as in Fig. 12D): sternite 9 trunc-
Diagnosis. Male: only Palaearctic species with ate medially to somewhat truncate but with
fore femur with three longitudinal rows of strong anteromedial margin developed, pointed; 2
bristles and with the prongs on the aedeagus spermathecae, additional spermathecal duct
directed anterolaterally. Female: only Palae- terminating in pigmented apex (one specimen
arctic species with the scutum lacking pruinosity, with 3 fully developed spermathecae).
the mid femur distinctly paler basally, and with Distribution and bionomics. [Link] is known
an elongate, single hind claw. from England to Estonia south to France,
Description. Male adult. Descriptive statistics: Austria and Greece (Fig. 19B). Adults have
see Tables 2—7. been collected from 7 May to 16 August. The
Head: dark brown; antennal flagellomere 10 extension into August was due to two speci-
with plume arranged in more than one whorl; mens, one female from Hiiumaa Is., Estonia, 8
palpus dark brown. August and a male from Hamburg, Federal
Thorax: dark brown; scutum bare of Republic of Germany, 16 August. Otherwise
pruinosity. the latest date of collection is 18 July. It is
Legs: coloration pattern as indicated in Fig. possible that the two August specimens were
4A; strong bristles of femora, tibiae distributed mislabelled.
as follows: anteriorly, ventrally, posteriorly on The most southerly locality of S. morio is
fore femur, dorsally, posteriorly on fore tibia, Lake Ochrid, Macedonia, Yugoslavia. Edwards
present or absent anteriorly on fore tibia, (1928) recorded this species from Calvi, Corsica,
anteriorly, ventrally, posteriorly on mid femur, but the single female was actually a specimen of
posteriorly on mid tibia, present or absent S. subinermis.
anteriorly, dorsally on mid tibia, anteriorly, Remm (1967) reported [Link] from the
ventrally on hind femur, dorsally on hind tibia; Caucasus, but we have been unable to confirm
ventral bristles on hind femur arising from this identification.
slightly developed tubercles; hind Ta! straight In Zetterstedt's collection at Lund there were
or with slight basal curvature; Ta 4 setae straight three instances of a male and female placed on
or with slight curve. the same pin. On the label of one of these,
Wing: slightly infuscated with veins of cells Zetterstedt specifically noted that they were
ri, r 2+ 3 darkly pigmented. caught in copula. We were therefore able to
Abdomen: dark brown. confidently associate the sexes.
Genitalia (Fig. 10C): gonostylus with outer Little can be said about the bionomics of this
margin evenly curved, tapering gradually species. Zetterstedt (1850) stated that males
for basal half, with swollen, rounded apex; swarm near marshes in Lund, but unfortunately
paramere (Fig. 10B) with apical half markedly his identifications of S. morio included both this
swollen, apex rounded (but shrivelled in some); species and [Link].
aedeagus (Fig. 10A) with lateral prongs, di- Goetghebuer (1936b) suggested that S. morio
rected anterolaterally, distal portion a simple, was restricted to eutrophic lentic habitats in
gradually tapering projection, about twice as southern Belgium but we were unable to confirm
long to markedly longer than lateral prongs, this identification from his material.
extreme apex directed ventrally. Labels on some specimens we examined noted
Female adult. Descriptive statistics: see the following: 'from saline meadow', 'coastal
 Revision of Holarctic species of Serromyia Meigen 201

flat' (from Bar, Yugoslavia), 'at stream', 'at (fool) may refer to the unusual appearance of
light'. Two female specimens deserve special this species as compared to the other species
mention; they have the intriguing statement Fabricius placed in Culex.
'reared from nest of mole' (from near Beacons-
field, Bucks., U.K.).
Serromyia pacifica Remm sp.n.
Strenzke (1950) and Thienemann (1950) re-
corded larvae of S. morio from lake-side mosses Diagnosis. Male: only Palaearctic species
in Germany but we have been unable to confirm with an AR = 1.17, the scutum pruinose, the
these identifications. One pupa from Strenzke's fore femur entirely dark brown, with one row of
collection labelled 'Strenzke No. 246 Serromyia ventral strong bristles and 1—2 posteriorly, and
femorata (?) morio?' is actually a specimen of the parameres rounded apically. Female: only
[Link] (see under that species and under generic Palaearctic species with palpus dark brown,
discussion). Strenzke (1950: 349) also recorded nearly entire fore tibia darkly pigmented,
a specimen as [Link] (?) from Ploner Sees, scutum pruinose, mid coxa with strong, lateral
but we have not been able to examine this bristles, and with a single elongate hind claw,
specimen and confirm the identification. longer than Ta 5 .
Taxonomic discussion. The type of [Link] Description. Male adult. Descriptive statistics:
has been destroyed and only the label is left see Tables 2—7.
(Zimsen, 1964). The type locality was given by Head: dark brown; antennal flagellomere 10
Fabricius (1775) as 'Anglia' [= England] and with plume arranged in more than one whorl;
was collected on 13 May. We have accordingly palpus dark brown.
designated a specimen from Oxford, U.K., Thorax: dark brown; scutum pruinose.
collected at the same time of year, as neotype. Legs: coloration pattern as indicated in Fig.
The type of S. nudipennis could not be located 3D; strong bristles of femora, tibiae distributed
and is probably also destroyed. Kieffer's (1913) as follows: ventrally, posteriorly on fore femur,
rather general description of the female of this posteriorly on fore tibia, anteriorly, ventrally,
species does not allow for a definite application posteriorly on mid femur, posteriorly on mid
of the name to any known European species. tibia, ventrally on hind femur, dorsally on hind
However, most of the characters he does record tibia; ventral bristles on hind femur arising from
correspond to the concept of S. morio as recog- slightly developed tubercles; hind Ta! with slight
nized here (i.e pale wings, elongate hind claw). basal curvature; Ta 4 setae straight or with slight
Kieffer (1913) recorded this species from Bitche curve.
(France). We examined two females from Wing: slightly infuscated with veins of cells
France but these were somewhat damaged. We r t , r 2 + 3 darkly pigmented.
therefore are designating a neotype (male) from Abdomen: dark brown.
the same place and date as the neotype of Genitalia (Fig. 10F); gonostylus with outer
[Link] (from England). margin every curved, tapering gradually for
We examined a male of morio identified by basal half, with swollen, rounded apex;
Zetterstedt as Ceratopogon flavicornis which paramere (Fig. 10E) with apical half somewhat
was a misinterpretation of Staeger's description. swollen, apex rounded (somewhat shrivelled);
Ceratopogon flavicornis is actually a member of aedeagus (Fig. 10D) with lateral prongs, di-
the genus Bezzia (syntypes examined). Simi- rected dorsolaterally, distal portion a tapering,
larly, some specimens identified by Meijere, simple projection, markedly longer than lateral
Schiner, Edwards and Goetghebuer as prongs, extreme apex directed ventrally.
[Link] were in fact [Link]. Goetghebuer Female adult. Descriptive statistics: see
also misidentified a specimen of S. morio as Tables 8—13. Similar to male except for usual
[Link]. Remm had misidentified a female as sex differences and as follows:
S. ledicola. Head: mandible serrate.
The single paralectotype of [Link] from Legs: coloration pattern as indicated in Fig.
Budapest (BMNH) is a female adult of [Link]. 4E; strong bristles of femora, tibiae distributed
Material examined. I l l males and 129 as follows: present or absent ventrally on mid
females. femur, ventrally on hind femur; claw of hind leg
Derivation of specific epithet. The name morio single, elongate, with basal tooth.
202 A. Borkent and B. Bissett

Wing: a few macrotrichia restricted to apical Serromyia rufitarsis: Kieffer 1906: 65.
margin. Serromyia gelida Kieffer 1925b: 429. Neotype,
Genitalia (as in Fig. 12D): sternite 9 truncate here designated, 8 adult on microscope slide,
medially, with anteromedial margin developed, labelled 'Serromyia gelida Kieffer, Neotype,
pointed; two spermathecae, additional sperma- Latvia, Sivera Lake, E. Remm, 17—6—67
thecal duct terminating in pigmented apex. CNC No. 20250, Serromyia dipetala R. det.
Distribution and bionomics. [Link] is H. Remm, Serromyia rufitarsis Det. A.
known from eastern Siberia (Fig. 18C). Adults Borkent' (CNCI). New synonym.
examined were collected from 25 May to 27 Serromyia bispinosa Goetghebuer 1936a: 321.
June. Lectotype, here designated, 8 adult on micro-
The specimens from the type locality were scope slide, labelled 'La Panne [ = D e Panne,
collected in mire. The material from Yakut Belgium], 8 , 1 6 - 6 - 3 6 , M. Goetghebuer',
A.S.S.R. was taken from a taiga bog. 'Serromyia bispinosa', '[Link] N.B. 18.073
Taxonomic discussion. The description pro- Coll. et det., M. Goetghebuer', ' 8 Type
vided here is based on the single male and two Lectotype Goetghebuer, Serromyia rufitarsus
females housed in the CNCI and a manuscript Det. A. Borkent' (ISNB); paralectotype
description provided by Mrs E. Remm. labelled as for lectotype except with ' 9 '
[Link] will also be described in a separate (ISNB). New synonym.
paper, currently in press (Academy of Sciences, Serromyia dipetala Remm 1965: 182. Holotype,
Tartu) and written by H. Remm before his 8 adult, not seen, from Estonian S.S.R.,
untimely death in 1986. Mrs E. Remm (pers. Valga District, shores of Lake Yakhiyavr, 11
comm.) indicated that the species description July 1952 (IZBE); paratypes: ESTONIA
could be included in the present revision, to S.S.R.: 111c?, 699 (not seen) from Valga,
ensure that this work is inclusive, and conse- Vyru, Pyarnu, Tartu and Kharyu districts, 13
quently whichever paper appears first will pro- June to 28 July, 1951-61; LATVIA S.S.R.:
vide the date and citation for [Link]. 13<3\ 6 9 , Daugavpils and Kraslava districts,
Types. Holotype, 8 adult, from Sakhalin 2 1 - 2 4 June 1961 (IZBE). New synonym.
Island, vicinity of Juzhno-Sakhalinsk, Ceratopogon morio: authors, not Fabricius.
[Link]. 1970, mire (IZBE); paratypes: 6cJ, Staeger 1839: 598 (in part).
79 labelled as for holotype (IZBE, CNCI);
U.S.S.R.: 3 9 , Kunashir island, Juzhno-Kurilsk Diagnosis. Male: only Palaearctic species in
vicinity, [Link]. 1970 (IZBE); 1 9 , Primorye which the parameres bifurcate apically. Female:
Territory, NP 'Kedrovaya Pad', [Link].1970 only Palaearctic species with a shiny scutum, a
(IZBE); 19 , from Amur Territory, Klimoutzy, completely darkly pigmented mid femur and a
25.v. 1957 (ZMAS); 18, 49 , Yakut A.S.S.R., single elongate hind claw.
Yakutsk vicinity, [Link].1968 (IZBE). Description. Male adult. Descriptive statistics:
Derivation of specific epithet. The name see Tables 2—7.
pacifica refers to the proximity to the Pacific Head: dark brown; antennal flagellomere 10
Ocean, where this species was collected. with plume arranged in more than one whorl;
palpus dark brown.
Thorax: dark brown; scutum bare of
Serromyia rufitarsis (Meigen) new status
pruinosity.
Ceratopogon rufitarsis Meigen 1818: 83. Lecto- Legs: coloration pattern as indicated in Fig.
type, here designated, 8 adult on microscope 4B; strong bristles of femora, tibiae distributed
slide, labelled 'Lectotype Ceratopogon as follows: present or absent posteriorly on fore
rufitarsis 8 Meigen, ex coll. Meigen 293/ femura (one bristle), present or absent pos-
40 (160b), prep. J. Clastrier, 1985 baume teriorly on mid femur, ventrally on hind femur,
phenole, Museum Paris' (MNHN); paralec- dorsally on hind tibia; ventral bristles on hind
totype 8 adult labelled as for holotype but femur arising from slightly developed tubercles;
with '160a' instead of '160b' and identified as hind Tat straight or with slight basal curvature;
Serromyia femorata (MNHN). Ta 4 setae straight or with slight curve.
Prionomyia rufitarsis: Stephens 1829: 238. Wing: slightly infuscated with veins of cells
Ceratolophus rufitarsis: Kertesz 1902: 158. r t , r 2 + 3 darkly pigmented.
 Revision of Holarctic species of Serromyia Meigen 203

Abdomen: dark brown. description by Kieffer (1925b) of a male with

Genitalia (Fig. 11C): gonostylus with outer brilliantly shiny thorax and a single spine on the
margin evenly curved, tapering gradually for fore femur could apply to either [Link] or
basal half, with rounded to somewhat pointed S. rufitarsis. On the basis of the general descrip-
apex; paramere (Fig. 11B) with apical portion tion of leg pigmentation, we have placed the
bifurcate, the ventral arm slender, the dorsal, species here. However, the type locality of
posteriorly directed arm tapering sharply to [Link] was given as northern Norway (just
point; aedeagus (Fig. 11 A) with lateral prongs, north of Tromsoe [=Tromso]) which is
directed laterally, dorsolateral^ or slightly markedly farther north than any locality of
anterolaterally, distal portion a tapering projec- [Link] (Fig. 19C) or [Link] (Fig. 18B). The
tion about twice as long to markedly longer only species with far northern distributions are
than lateral prongs, extreme apex directed [Link] (Fig. 19A) and possibly [Link]
ventrally. (Fig. 19B) and [Link] (Fig. 18A). These
Female adult. Descriptive statistics: see species, however, have either a pruinose scutum
Tables 8—13. Similar to male except for usual or have many spines on the legs. It may be that
sex differences and as follows: S. rufitarsis does occur in northern Norway but
Head: mandible serrate. only future collecting will confirm this. How-
Legs: coloration pattern as indicated in Fig. ever, the name [Link] now must be considered
5F; strong bristles of femora, tibiae distributed a synonym of [Link].
as follows: present or absent ventrally on mid Remm (1965) noted that his [Link] dif-
femur, ventrally on hind femur; claw of hind leg fered from [Link] in lacking strong bristles
single, elongate, with basal tooth. (his spines) on the fore and mid femora but we
Wing: very few macrotrichia restricted to found this character to be variable, with a few
apical margin. strong bristles present in some individuals (in-
Genitalia (as in Fig. 12D): sternite 9 trunc- cluding some of those identified by Remm as
ate medially to somewhat truncate but with [Link]).
anteromedial margin developed, pointed; two Goetghebuer (1922a) noted that the syntype
spermathecae, additional spermathecal duct series of Ceratopogon rufitarsis consisted of two
terminating in pigmented apex. males. The male paralectotype of [Link] is
Distribution and bionomics. [Link] is actually a specimen of [Link].
known from England to Estonia south to Although Goetghebuer (1936a) based his de-
Belgium and Hungary (Fig. 19C). Adults have scription on only one male and one female of
been collected from 26 May to 23 July. [Link], it was uncertain which represented
Remm (1965) recorded specimens from a few the holotype. Accordingly we have designated
additional localities from Estonia and Latvia (as the male as lectotype.
[Link]), which are also probably members Material examined. Thirteen males and ten
of this species. females.
The only available bionomic information is Derivation of specific epithet. The name
from Remm (1965) who noted that this species rufitarsis (red tarsi) refers to the red-yellow tarsi
(as [Link]) was most plentiful from 1 to 10 reported for this species by Meigen (1818) (see
July and could be found not only in shrubs and generic taxonomic discussion for comments on
thickets near water but also in damp meadows reports of red coloration by previous authors).
some distance from any larger body of water.
Taxonomic discussion. Although we were un-
Serromyia subinermis Kieffer
able to examine any type material of [Link],
Dr H. Remm kindly sent other material of this Serromyia subinermis Kieffer 1919: 73 (as
species. Both the specimens and the detailed variety of [Link]). Neotype, here
description by Remm (1965) leave little doubt designated, 8 adult, labelled 'Serromyia
that this name is a junior synonym of S. rufitarsis. subinermis Kieffer, 8 , Neotype, Ocsa,
The material sent by Remm also allowed correct Hungary, lapret, 1965.V. 19, leg. Mihalyi'
association of males with females. (HNHM).
The type of Serromyia gelida Kieffer could Serromyia femorata: authors, not Meigen.
not be located and is presumed to be lost. The Zetterstedt 1838: 822 (in part), 1850: 3665
204 A. Borkent and B. Bissett

(in part). Edwards 1926: 410 (in part). 4F, 5E; strong bristles of femora, tibiae dis-
Goetghebuer 1934: 61 (in part). tributed as follows: 0—4 ventrally on fore femur,
Serromyia morio: Edwards 1928: 173, not 0 - 2 ventrally on mid femur, 0—2 apically on
Fabricius. mid tibia, ventrally on hind femur; claw of hind
leg single, elongate, with basal tooth.
Diagnosis. Male: only Palaearctic species with Wing: a few to many macrotrichia restricted
the parameres curved subapically and with the to apical margin.
pointed apex directed laterally. Female: only Genitalia (as in Fig. 12D): sternite 9 trunc-
Palaearctic species with the palpus dark brown, ate medially to somewhat truncate but with
scutum pruinose, wing slightly infuscated with anteromedial margin developed, pointed; two
veins of cells r l5 r 2 + 3 darkly pigmented, lacking spermathecae, additional spermathecal duct
strong lateral bristles on the hind coxa, and with terminating in pigmented apex.
a single elongate hind claw. Distribution and bionomics. [Link]
Description. Male adult. Descriptive statistics: is known from England to Estonia south to
see Tables 2 - 7 . Corsica, Yugoslavia and Hungary (Fig. 19D).
Head: dark brown; antennal flagellomere 10 Adults have been collected from 10 April to 5
with plume arranged in more than one whorl; July.
palpus dark brown. Taxonomic discussion. We have applied the
Thorax: dark brown; scutum pruinose. name S. subinermis to this species by designating
Legs: coloration pattern as indicated in Figs a neotype. The type material of this species is
4C, D; strong bristles of femora, tibiae dis- almost certainly lost.
tributed as follows: anteriorly, posteriorly on Kieffer's (1919) brief description of this
fore femur, present or absent ventrally on fore species noted that the fore and mid femora of
femur, posteriorly on fore tibia, present or the female had one or two spines while the male
absent dorsally, anteriorly, posteriorly on mid lacked spines altogether. Our description of the
femur, present or absent dorsally, ventrally on female fits Kieffer's concept but that of the
mid femur, posteriorly on mid tibia, present or male does not. It may well be that Kieffer
absent anteriorly, dorsally, posteriorly on mid incorrectly associated the sexes as he did for
tibiae, anteriorly, dorsally, ventrally on hind [Link] (see under [Link] and [Link]).
femur, dorsally on hind tibia, ventral bristles on We prefer to retain a given European name
hind femur arising from slightly developed where possible, rather than propose a new one.
tubercles; hind Tai straight or with slight basal We have examined specimens identified by
curvature; Ta 4 setae straight or with slight curve. Edwards and Goetghebuer as [Link], which
Wing: slightly infuscated with veins of cells in fact are members of [Link]. Edwards
r1? r2+3 darkly pigmented. (1928) recorded S. morio from Corsica but it is
Abdomen: dark brown. actually a specimen of [Link].
Genitalia (Fig. 11F): gonostylus with outer The association of the sexes of [Link]
margin evenly curved, tapering gradually to is somewhat tentative. None of the males and
pointed apex; paramere (Fig. 11E) with sub- females we examined were collected at the
apical portion somewhat expanded dorso- same place and time. Males and females col-
ventrally (expansion best seen in lateral or lected in Hejobaba (8 km W Leninvaros),
posterior view), with pointed apex directed Hungary, in May and males and females ident-
laterally or anterolaterally; aedeagus (Fig- 11D) ified by H. Remm as S. spinosipes from Estonia
with lateral prongs, directed posterolaterally to were considered correctly associated members
laterally, distal portion a simple, somewhat of this species.
broadened projection, about twice as long to In addition, we are not certain that all the
markedly longer than lateral prongs, extreme females placed here are conspecific. Although
apex directed ventrally. some appear to be correctly associated with the
Female adult. Descriptive statistics: see males, some may belong to another, unnamed
Tables 8 - 1 3 . Similar to male except for usual species. Further collecting and study are
sex differences and as follows: required to clarify this problem.
Head: mandible serrate. Material examined. Twenty-five males and
Legs: coloration pattern as indicated in Figs forty-one females.
 Revision of Holarctic species of Serromyia Meigen 205

Derivation of specific epithet. The name additional spermathecal duct terminating in

subinermis (somewhat unarmed) probably pigmented apex.
refers to the lack of spines on the fore and mid Distribution and bionomics. [Link] is known
femora which Kieffer (1919) attributed to males from only from the type locality in the Federal
of this species. Republic of Germany (Fig. 19D). The date of
collection on the labels is given as 6.5.1935 and
Dr R. Contreras-Lichtenberg (NHMW) (pers.
Serromyia tecta Borkent sp.n.
comm.) has informed us that this means 6 May
Diagnosis. Male: only Palaearctic species 1935. This clarification was added to the labels
lacking spines on fore and mid femora and on the slide material.
tibiae, each of which is entirely dark brown, Taxonomic discussion. Males and females of
A R = 1.00—1.03, pruinose thorax and with S. tecta are very similar to those of Nearctic
parameres rounded apically and swollen in [Link] and they may be conspecific. How-
apical half. Female: only Palaearctic species ever, males have different antennal ratios.
with small, equal hind claws and a pruinose Larger series of specimens from a variety of
thorax. localities are needed to determine whether this
Description. Male adult. Descriptive statistics: is due to geographical variation or is indeed
see Tables 2—7. indicative of genetic discontinuity.
Head: dark brown; antennal flagellomere 10 Types. Holotype, 8 adult on microscope
with plume arranged in more than one whorl; slide, labelled 'Holotype Serromyia tecta
palpus dark brown. Borkent, 8 , Traun [Austria], Czerny, 6.5.1935,
Thorax: dark brown; scutum pruinose. May 6, 1935' (NHMW); allotype on microscope
Legs: coloration pattern as indicated in Fig. slide labelled as for holotype (NHMW); para-
3D; strong bristles absent on femora, tibiae types: 38,29 labelled as for holotype (NHMW,
except ventrally on hind femur, present or CNCI).
absent dorsally on hind tibia; ventral bristles on Derivation of specific epithet. The name tecta
hind femur arising from slightly developed (disguised) refers to the long hidden identity of
tubercles; hind Ta x straight or with slight basal this European species.
curvature; Ta 4 setae straight or with slight curve.
Wing: slightly infuscated with veins of cells
Fossil species
r 1? r 2 + 3 darkly pigmented.
Abdomen: dark brown. Several species of fossil Serromyia have been
Genitalia (Fig. 12C): gonostylus with outer described, all from Holarctic localities, and
margin evenly curved, tapering gradually these shed some light on the history of the
for basal half, with rounded, somewhat genus. We were fortunate in being able
swollen apex; paramere (Fig. 12B) with apical to examine all of these, except one male of
half somewhat swollen, with rounded apex [Link] (Loew). Szadziewski (1988) has
(shrivelled in some); aedeagus (Fig. 12A) with recently described the Baltic amber fauna
lateral prongs, directed laterally to postero- of Ceratopogonidae and included six taxa of
laterally, distal portion a simple, slender pro- Serromyia. These are: [Link] (Loew),
jection, markedly longer than lateral prongs, [Link] (Loew), [Link] Szadziewski,
extreme apex directed ventrally. [Link] Szadziewski and two unnamed
Female adult. Descriptive statistics: see species (sp. A and B). His excellent and com-
Tables 8—13. Similar to male except for usual prehensive study provides descriptions of these
sex differences and as follows: species and we add here only a few further
Head: mandible serrate. details. Because of the good preservation and
Legs: coloration pattern as indicated in Fig. presence of some important character states,
5D; claws of hind leg equal, small. we have given names to Szadziewski's (1988)
Wing: a few macrotrichia restricted to very species A and B.
apical margin.
Genitalia (as in Fig. 12D): sternite 9 some-
Serromyia anomaiicornis (Loew)
what truncate medially but with anteromedial
margin developed, pointed; two spermathecae, Ceratopogon anomaiicornis Loew 1850: 30.
206 A. Borkent and B. Bissett

Types as indicated by Szadziewski (1988). Distribution and bionomics. [Link] is

Serromyia anomalicornis: Szadziewski 1988: known from compression fossils from the type
142. locality at Rott, Siebengebirge, Rheinland,
Federal Republic of Germany. Although earlier
Diagnosis and description. As indicated by thought to be Oligocene in age the Aquitanian
Szadziewski (1988). Additional character states epoch is now dated as Miocene (Szadziewski,
as follows: male antennal flagellomere 10 setae pers. comm.).
in a single whorl; male, female with Ta 4 with Taxonomic discussion. The above description
straight setae. is based on the examination of the holotypes of
Distribution. From Baltic amber. the species listed above and the single female
Material examined. One male and one female. described by Statz (1944: 149) as Serromyia sp.
Derivation of specific epithet. We were un- We were unable to find any significant differ-
certain why the name anomalicornis (unusual ences between these specimens, based on the
horn) proposed by Loew (1850) was applied to character states used in recognizing extant
this species. species of Serromyia. We accordingly consider
all to be conspecific.
Statz (1941) provided the names Serromyia
Serromyia colorata Statz
spinofemorata and Serromyia austera under
Serromyia colorata Statz 1944: 150. Holotype photographs of these, but failed to give any
9 , both halves of split rock present, mounted diagnostic features. The names are therefore
on wooden block, labelled 'Serromyia nomina nuda. However, the descriptions by
colorata Stz. Original!', '3527', 'Serromyia Statz (1944) do validate these names. Sphon
colorata Sz. nov. sp.' (LACM). (1973) incorrectly considered the earlier naming
Serromyia austera Statz 1944: 150. Holotype 9 , by Statz (1941) to invalidate his later use of the
both halves of split rock present, mounted on names (see ICZN Art. 23:m).
wooden block, labelled 'Serromyia austera Statz (1944) stated that the antennae of
Stz. Original!', '3526', 'Serromyia austera S. colorata, [Link] and [Link] femorata each
Sz.', 'Serromyia colorata Det. A. Borkent' had 14 segments but each actually has 13
(LACM). New synonym. flagellomeres. The macrotrichia visible at the
Serromyia spinofemorata Statz 1944: 151. wing apex reported for S. colorata were actually
Holotype 9 , only one half of split rock pres- those at the very margin.
ent, mounted on wooden block, labelled We consider some of the slight colour differ-
'Serromyia spinofemorata Statz Original!', ences noted by Statz (1944) to be artefacts of
'3528', 'Serromyia spinofemorata Sz.', preservation.
'Serromyia colorata Det. A. Borkent' Material examined. Four females.
(LACM). New synonym. Derivation of specific epithet. The name
colorata (colour) presumably refers to the dif-
Diagnosis. Female: apical portion of fore ferences in color Statz (1944) attributed to this
femur and tibia pigmented, hind claw elongate species.
with HC/Ta 5 = 1.09-1.38.
Description. Male adult. Not known.
Serromyia polonica Szadziewski
Female adult. Legs: coloration pattern with
apical portion of fore and mid femora, basal Serromyia polonica Szadziewski 1988: 135.
portion of fore and mid tibiae darkly pigmented Types as indicated by Szadziewski (1988).
(Fig. 13D); strong bristles absent on femora,
tibiae except present ventrally on hind femur Diagnosis and description. As indicated by
(Figs 13A, C); hind femur length = 0.86-1.07 Szadziewski (1988); additional character state
mm; claw of hind led single, elongate (Figs 13B, as follows: Ta 4 with straight setae.
F), with basal tooth, HC/Ta 5 - 1.09-1.38. Distribution. From Baltic amber (Eocene).
Wing: wing venation not discernible. Taxonomic discussion. We could not confirm
Genitalia (as in Fig. 13D): sternite 9 truncate the presence of the prescutal pits as reported by
medially, two spermathecae present (additional Szadziewski (1988). If well developed, they
spermathecal duct not discernible). would be unique in the genus.
 Revision of Holarctic species of Serromyia Meigen 207

Material examined. Two females (all the type Type. Holotype, 8 adult in amber, in paper
material). envelope labelled 'Diptera Nematocera Cera-
Derivation of specific epithet. The name topogonidae Serromyia sp. A, Szadziewski 1988
polonica presumably refers to the type locality 8 . MZ 14972, Serromyia sinuosa Borkent'
in Poland. (MZW).
Derivation of specific epithet. The name
sinuosa (windings) refers to the sinuate setae
Serromyia ryszardi Borkent sp.n.
found on the fourth tarsomeres of this species.
Serromyia sp. B Szadziewski 1988: 139.
Serromyia spinigera (Loew)
Diagnosis and description. Described by
Szadziewski (1988: 139) as Serromyia sp. B. Ceratopogon spiniger Loew 1850: 30. Types as
Additional character states as follows: antennal indicated by Szadziewski (1988).
flagellomere 10 with plume arranged in a single Serromyia spinigera: Kieffer 1906: 2.
whorl; strong bristles of femora, tibiae distri- Szadziewski 1988: 140.
buted as follows: anteriorly, ventrally, posteriorly Ceratopogon elongatus Meunier 1904: 242.
on fore femur, anteriorly, dorsally, posteriorly Types as indicated by Szadziewski (1988).
on fore tibia, anteriorly, ventrally, posteriorly Serromyia elongatus: Szadziewski 1988: 140.
on mid femur, dorsally, posteriorly on mid
tibia, anteriorly, dorsally, ventrally on hind Diagnosis and description. As indicated by
femur, 2—3 rows dorsally on hind tibia; aed- Szadziewski (1988). Additional character state
eagus with trifid apex; Ta 4 with straight setae. as follows: Ta 4 with straight setae.
Distribution. From Baltic amber (Eocene). Distribution. From Baltic amber (Eocene).
Taxonomic discussion. The trifid aeadeagus Material examined. Two females.
of [Link] is somewhat similar to that of Derivation of specific epithet. The name
several extant species in the Holarctic (see Figs spinigera (spine bearer) presumably refers to
8A, D, 9D), with the lateral prongs and the the noted spines on the hind femur of this
apex directed in a posterior direction. species.
Type. Holotype, 8 adult in amber, in paper
envelope labelled 'Diptera Nematocera Cera-
Serromyia succinea Szadziewski
topogonidae Serromyia sp. B, Szadziewski 8 .
MZ 16110, Serromyia ryszardi Borkent' (MZW). Serromyia succinea Szadziewski 1988: 136.
Derivation of specific epithet. The name Types as indicated by Szadziewski (1988).
ryszardi is named in honour of the many contri-
butions that Ryszard Szadziewski had made to Diagnosis and description. As indicated by
our understanding of ceratopogonid systematics Szadziewski (1988).
and, in particular, his comprehensive work on Distribution. From Baltic amber (Eocene).
Baltic amber material. Taxonomic discussion. Flagellomere 10 of
the male could not be seen clearly. Both male
and females had only straight setae on ta 4 .
Serromyia sinuosa Borkent sp.n.
The male was correctly reported to have
Serromyia sp. A Szadziewski 1988: 138. a few macrotrichia at the wing tip (discussed
below under 'Phylogeny').
Diagnosis and description. Described by Material examined. One male and two females
Szadziewski (1988: 138) as Serromyia sp. A. (all type material).
Additional character states as follows: antennal Derivation of specific epithet. The name
flagellomere 10 with plume arranged in a single succinea (amber) clearly refers to the state of
whorl; Ta 4 with sinuate setae. preservation of this species.
Distribution. From Baltic amber (Eocene).
Taxonomic discussion. The sinuate setae on
Discussion of fossil material
the fourth tarsomere are unique amongst the
Baltic amber and extant Holarctic species of Our examination of many specimens of extant
Serromyia. species as well as of available fossils of extinct
208 A. Borkent and B. Bissett

species allows for some improvements in the name available until such time as further
key given by Szadziewski (1988). In particular, specimens become available for study and a
our studies show that the ratio of the second neotype can be designated.
radial cell to the first is highly variable and a
poor tool for recognizing extant species. This
suggests that the same would be true for the Phylogeny
fossil species. The suggested changes replace A fundamental question to the interpretation
the use of that character in the key and are as of phylogenetic relationships within Serromyia,
follows: concerns the monophyly of the genus itself.
Throughout the course of this study the
presence of a swollen hind femur bearing a
Couplet 6
ventral patch of strong bristles arranged as a
Fourth tarsomerc with both straight and one pair of single row of 1—3 bristles basally, forming 2
sinuate setae [Link] rows at mid length and scattering distally into
Fourth tarsomerc with only straight setae 7 an indistinct pattern up to 4 bristles wide, was
interpreted as a derived character state. This
character state appears to be unique within the
Couplet 8
Ceratopogonidae and Culicomorpha.
Fore and mid femur with strong bristles anteriorly, Wirth & Grogan (1988), however, recently
ventrally and posteriorly [Link] described the new monotypic genus Metacan-
Fore and mid femur with only a few strong bristles thohelea which exhibits this same character state.
ventrally [Link] On the basis of this modification, we suggest
that these two together form a monophyletic
Because S. colorata is known as a compression group (Fig. 20).
fossil, it was not included in the key by Wirth & Grogan (1988) suggested several
Szadziewski (1988). If keyed out, it would run characters which may serve to distinguish
to [Link]. Although [Link] has darker Metacanthohela from Serromyia. Our study
leg pigmentation restricted to the apex of the indicates that many of these also occur in at
fore femur and tibia and [Link] females least some species of Serromyia. The proposed
have completely dark fore legs, it is unlikely distinguishing character of Metacanthohelea,
that this can be used to distinguish them followed by the character states within
Virtually all Baltic amber ceratopogonids lack Serromyia are given below:
contrasting pigmentation and this may be an
artefact of preservation (Szadziewski, 1988). — slightly arcuate hind tibia: indistinguishable from
We can suggest no other distinguishing feature. that of [Link], [Link], [Link], [Link],
Another fossil, described by Heyden (1870) [Link], [Link]. The degree of bend of
the base of the hind tibia seems strongly correlated
as Ceratopogon alpheus, is probably a member
with the thickness of the hind femur. Thicker hind
of Serromyia. The male specimen was collected
femora are associated with a stronger bend at the
from the brown coal deposits at Rott, base of the tibia.
Siebengebirge, Federal Republic of Germany.
The figure clearly shows plumose antennae and — reduced number of hind femur spines: within
markedly swollen hind femora and we know of Serromyia, the number of spines (here called strong
bristles) is generally correlated with the size of the
no other Nematocera with such a combination
hind femur. However, the extent to which the hind
of characters. Some Monohelea and Schizohelea
femur bears strong bristles does vary. Female [Link],
also have swollen hind femora but not to such a [Link] and [Link] have the same or a smaller
degree. We therefore recognize this species as a percentage of the hind femur covered with strong
new combination: Serromyia alpheus (Heyden). bristles when compared with [Link].
Unfortunately, the type was reported as lost The male of [Link] has the distal 0.39-0.43 of
(Szadziewski, 1988), and we too have tried the hind femur bearing spines. Within Serromyia the
without success to locate it at the BMNH and least amount of the hind femur bearing strong bristles
OXUM. It is possible that this male was is exhibited by [Link] with a minimum of 0.58.
conspecific with the females described as Otherwise, all Serromyia have an even more exten-
[Link]. However, we prefer to leave the sively spinose femur.
 Revision of Holarctic species of Serromyia Meigen 209

— female with equal hind claws: also present in The first suggests that the plesiomorphic state
[Link], [Link], S. crassifemorata, S. borealis, is the total lack of strong bristles (as in most
[Link]. However, in each of these Serromyia species, Ceratopogonidae), developing to a state where
each claw does not bear an inner tooth, which is strong bristles are restricted to the distal portion
present in Metacanthohelea coqani. of the hind femur (as in Metacanthohelea), and
— female with relatively short hind fourth ultimately to the extensive strong bristles of
tarsomere: we failed to find any significant differences Serromyia. This hypothesis would suggest
between the hind fourth tarsomeres of Metacan- that Metacanthohelea is the sister group of
thohelea and species of Serromyia. Serromyia.
— eyes broadly separated: also present in [Link], The second hypothesis suggests that the
[Link], [Link] and [Link]. plesiomorphic condition is a single, but exten-
— male antenna with flagellomeres 1 - 8 fused: sive row of strong bristles (as in Echinohelea
flagellomeres fused in [Link] and [Link] or Macfie, some Austrohelea Wirth & Grogan,
at least partially fused basal flagellomeres in all some Fanthamia de Meillon and some
Holarctic species other than [Link]. Stilobezzia), which develop into the more
complex pattern typical of most species of
— parameres fused: also fused in [Link], [Link]
and [Link].
Serromyia. This suggests that Metacanthohelea
cogani is merely an autapomorphic member of
— sensory pit on palpal segment 3: also present in Serromyia, with a somewhat reduced distri-
male and female S. agathae and 5. nocticolor and in at bution of strong bristles on the hind femur.
least the male of [Link].
In our opinion, the first hypothesis seems
— radically different genitalia: although the male most likely and we therefore recognize Meta-
genitalia of Metacanthohelea are somewhat different canthohelea as a valid genus (Fig. 20). We are
from those of most Serromyia species, they are similar unable to suggest an apomorphy for Meta-
to those of [Link]. In addition, some Serromyia also
canthohelea, but, considering the genus is
have rather different genitalia when compared
monotypic, it must be monophyletic.
to those in the rest of the genus (e.g. [Link],
S. mangrovi). In spite of a diligent search for character
states which might be useful in interpreting the
From the above comparisons, it appears that phylogenetic relationships between species of
the only possible distinguishing character of Serromyia, only a few clues were discovered.
Metacanthohelea is the reduced extent to which In all extant Holarctic species of Serromyia,
the male hind femur bears strong bristles. In other than [Link], the tenth flagellomere
addition, re-examination of the original material exhibits a series of whorls of elongate bristles.
(Wirth, pers. comm., and ourselves) indicates In all other Ceratopogonidae, including all
that the female claws are distinctive. They are Serromyia examined from the Oriental,
bent sharply at their base and with the distal Australian and Afrotropical Regions, the tenth
portion straight (Fig. 16C), in comparison to flagellomere has a single whorl of elongate
species of Serromyia, where the base of the bristles, similar to that of the preceding fla-
claws are either straight or evenly curved and gellomeres. The multi-whorled condition is
the distal portion of the claw is always curved therefore logically interpreted as a synapo-
(Figs 16D—G). In addition, the claws of the morphy (Fig. 20). This character state is also
hind leg of [Link], although equal and shorter reflected in the ratio of antennal flagellomere
than the fifth tarsomere, bear a well-developed 10/11 (Table 3), where [Link] has a single
inner tooth. In those Serromyia species which whorl and a low ratio and all other species have
have equal hind claws, the inner tooth is lacking. a higher ratio and have more than one whorl.
Of the above character states we can inter- Considering the monophyly of the Holarctic
pret only the degree to which the male hind species (except [Link]), another character
femur bear strong bristles for phylogenetic an- state may give further resolution. All Palae-
alysis. Outgroup comparisons indicate that most arctic species (except [Link]) and several
genera of Ceratopogonidae lack thick strong Nearctic species exhibit lateral prongs on the
bristles on the hind femur but that some bear a aeadeagus. This tripartite condition is also
single row of thick spines. Two alternate hy- present in various configurations, in some
potheses are therefore possible. African species: [Link], [Link],
210 A. Borkent and B. Bissett

•2
2.
£
o ^§
t
s
3 Qj
-S? Co .y o
o s
o Vo-" oc:
OJ CO
•S CD < c
35 o s
o <T> I
0j
2 I
c
o CO
c + CO Co Co

aedeagus lacking side prongs

flagellomere 10 with several

whorls of long setae

male with > 0 . 5 8 of hind femur

with strong bristles

2 - 4 rows of strong bristles

on hind femur

Fig. 20. Cladogram showing relationships within Serromyia and its sister lineage. Black rectangles and
accompanying descriptions represent synapomorphic character states.

[Link], [Link] and [Link]. presence of long claws in the genus and similar
This includes all those Serromyia south of the genera such as Monohelea and Stilobezzia.
Holarctic in which the body coloration is entirely Although the presence of short claws is probably
dark brown and for which the male is known. apomorphic within Serromyia, incompatibility
This may allow for interpretation of the two with other character states suggests that this
character states found within the Holarctic character state is susceptible to homoplasy and
lineage where the aedeagus either has lateral argues against such a grouping. [Link], an
prongs or is lacking these. Using the African African species known only from the female,
species as the outgroup, those species lacking has short hind claws. It is unlikely that the male
lateral prongs in the Nearctic would form a will have the synapomorphy of possessing more
monophyletic group: S. barberi, S. crassi- than one whorl of setae on flagellomere 10, a
femorata and S. nudicolis (Fig. 20). synapomorphy grouping all Holarctic species
Although based on phenetic similarity, it (except [Link]), including four species
seems likely to us that S. morio and [Link] are which have short hind claws: [Link], [Link],
sister species. The male genitalia of these two [Link] and [Link]. Considering the
species were inseparable. overall morphological similarity between
Szadziewski (1988: 142) has suggested that [Link] and [Link], these two are probably
those species where the females have short, sister species. However, as argued above,
equal hind claws may form a monophyletic S. crassifemorata is more closely related to two
group, which he called the crassifemorata group. long-clawed species than to other short-clawed
This hypothesis seems unlikely to us. It does species. In addition, also as suggested above,
seem clear that the character state of having the short-clawed [Link] is probably most closely
short claws is derived from a plesiomorphic related to [Link], a species with an elongate
longer, single claw, based on the widespread hind claw. Finally, two of the fossil species
 Revision of Holarctic species of Serromyia Meigen 211

[Link] and [Link] have short hind Serromyia males. However, this character state
claws and assuming these are correctly associ- cannot be interpreted as synapomorphic, as
ated with the males, definitely do not belong to male wing macrotrichia also occur in a number
the clade including the extant Holarctic species of other ceratopogonid genera.
(three species of which have short claws). We Previous workers have used some characters
conclude, therefore, that short hind claws have states to recognize new taxa (particularly genera)
evolved independently at least three times which show homoplasy within Serromyia. If
amongst the extant Holarctic lineage and that it Serromyia is indeed monophyletic, this may
is a poor indicator of relationship. The question indicate that those character states are sus-
may be further tested by discovery of the male ceptible to homoplasy in other groups of cera-
of [Link] and further resolution of phylo- topogonids as well and that they should be
genetic relationships amongst the Holarctic interpreted with caution. The character states
lineage. are as follows: separate or fused male flagello-
As noted above, [Link] is not part of meres; terminal three or four male flagello-
clade formed by all other Holarctic species. meres elongate; presence or absence of sensory
This, its distinctive morphology (as compared pit on third palpal segment; female mandible
to Holarctic species), and its presence in the vestigal or well developed; distance between
Sinai, may indicate an Afrotropical or Oriental eyes; presence or absence of pruinosity on scutum;
phylogenetic connection for this species. base of M2 indistinct or well developed; presence
Our examination of fossil material provided or absence of macrotrichiae on male or female
some clues about the history of the genus. All wings; presence or absence of strong bristles on
males of the fossil species had a single whorl of legs; presence or absence of sinuate setae on
setae on flagellomere 10, indicating that fourth tarsomeres; short and equal or elongate
they were not members of that clade which and single hind female claws; straight or basally
includes all extant Holarctic species (except curved hind first tarsomere; presence or absence
S. mangrovi). Either the Holarctic species of stout spine near base of hind first tarsomere;
diversified since the Eocene (Baltic amber age) female with sternite eight bilobed or completely
or they evolved elsewhere. cleft; separate or fused parameres; 1—3 func-
S. ryszardi was unique in the fossil material in tional spermathecae.
exhibiting 2—3 rows of strong dorsal bristles on Debenham (1987) recently described the new,
the hind tibia. The only extant Serromyia species monotypic genus Chimaerohela and suggested
which has a similar condition is [Link] that it was closely related to Serromyia. This
from South Africa. This character state is prob- conclusion was based on shared similarities
ably a valid synapomorphy. Only some members between Chimaerohelea and some species of
of Echinohelea have a similar condition. Serromyia. These stated similarities require
Two other clues indicate that the Baltic am- comment, based on our study of Serromyia.
ber Serromyia may not be immediately related The following gives the character state which
to the assemblage of extant Holarctic species. Debenham (1987) suggested was shared by
The sinuate setae on the fourth tarsomere of Chimaerohelea and various Serromyia, followed
[Link] (Baltic amber) is shared only with by our observations:
some more southerly members of the genus:
— fusion of the veins between cells ^ and r 2 + 3
[Link] (Australasian), [Link] (Afro-
present in [Link], male [Link], [Link] and
tropical) and [Link] (Afrotropical). It is possibly [Link]: our examination showed that
uncertain, however, which character state is no Serromyia species showed such fusion cxcept as
derived. Both conditions of sinuate and straight intraspecific variation in a few species ([Link],
setae on the fourth tarsomere are widespread [Link], [Link]).
throughout the Ceratopogonini.
— a single spermatheca present in [Link]: this
A second character state is shared by is the only species of Serromyia with a single
[Link] (Baltic amber) and [Link] (Afro- spermatheca. However, the character state appears
tropical). Males of both have a few macrotrichia to be homoplastic in numbers of other genera of
at the wing apex. Although the presence of Ceratopogonini (Alluaudomyia Kieffer (1-2),
wing macrotrichia are widespread on the wings Brachypogon Kieffer (1-2), Ceratoculicoides Wirth
of females, they are not present on any other & Ratanaworabhan (1-2), Kolenohelea de Meillon
212 A. Borkent and B. Bissett

& Wirth (1—2), Macrurohelea Ingram & Macfie (1 — distributions to be confidently interpreted. The
2), Stilobezzia Kieffer (1-3)) and is probably a poor only Holarctic species, [Link], is widespread
indicator of relationship. in the Nearctic but appears to be absent from
— wing fold extending basally from point where M2 most of the area west of the continental divide
curves toward M^ we confirm that [Link] is the in more southerly latitudes.
only Serromyia with a barely discernable, slightly Within the Palaearctic, all the European
pigmented line extending basally. species which were well represented in collec-
— male flagellomeres 4 - 1 1 fused in [Link] tions, seem to be broadly distributed in central
(named [Link] here), possibly [Link], and Europe. In southern Europe, however, species
some other genera of Ceratopogonini: we found fused appear to be restricted to mountainous regions.
flagellomeres present in [Link], [Link] and Although [Link] is the only species defi-
at least partial fusion in all Holarctic species other nitely known from northern Fennoscandia, it
than [Link]. seems likely that at least [Link] (based on its
— aedeagus similar to [Link], [Link], presence in Scotland) and [Link] (based on
[Link] (named [Link] here): we failed to see its northern distribution in North America) will
distinctive similarities between Chimaerohelea and also be found there.
any species of Serromyia. Szadziewski (1988) has suggested a European
origin for the genus Serromyia. We can see no
Of the characters discussed by Debenham logical basis for such a conclusion. We recognize
(1987), the only character state which may that zoogeographic interpretation must be based
possibly be interpreted as a synapomorphy is on an understanding of the phylogenetic re-
the dark line extending basally from M 2 . We lationships within the group under consider-
could find no other Ceratopogonini with such a ation. Until further interpretation of the
character state. However, this would indicate relationships between the known species is
that the monopoly of Serromyia plus Metacan- available, it will be impossible to identify the
thohelea would be in doubt. We consider this to early lineages of Serromyia and thereby suggest
be unlikely and that the above similarities are where the genus may have originated.
all due to homoplasy. Szadziewski (1988) also interpreted his
We are unable, however, to present an 'crassifemorata group' as exhibiting a recent
alternative hypothesis of the phylogenetic Euro-North American distribution. As indi-
position of Chimaerohelea. Further research is cated above, however, evidence suggests that
require to interpret the position of this and this group is polyphyletic (grouped on the basis
many other genera of Ceratopogonini. of parallelism). As such, no zoogeographic
interpretation could be validated.
The fossil data indicate that the extant
Zoogeography
Holarctic lineage was probably not present in
The above phylogenetic interpretation is so Europe during the Eocene. We are inclined to
sketchy that it is presently impossible to inter- think that the extant Holarctic species, as a
pret the historical zoogeography of the extant monophyletic group, are most closely related to
Holarctic species. However, some distributions those species from Africa which are darkly
allow for some general zoogeographic state- coloured. However, there is no logical evidence
ments to be made. to support this at present, other than some of
The distributions of all species of Serromyia the unpolarized shared similarities between
in the Nearctic are restricted to temperate Holarctic species (other than [Link]) and
habitats. The southern extensions are otherwise some of the African species noted above.
present only in the mountainous regions. No
Serromyia are known from the Neotrophical
Acknowledgments
Region.
Within the Nearctic, [Link] and [Link]- This study was the result of 4 years of investi-
femorata are restricted to eastern North gation. A few years ago, when the senior author
America. S. barberi is present only west of was new to the study of Ceratopogonidae, Dr
the continental divide while the remaining Willis W. Wirth gave generously of his time and
species are too poorly represented for their expertise to help in initiating my studies. He
 Revision of Holarctic species of Serromyia Meigen 213

already had a draft key of the Nearctic species thorough reviews of this manuscript: Dr William
of Serromyia (including [Link]) as well as L. Grogan, Dr Evert E. Lindquist and Dr
some notes on the species. He unstintingly gave Michael Sharkey, Dr Ryzard Szadziewski, Dr
these to us. We express our thanks to him for Richard J. Vockeroth and Dr Willis W. Wirth.
his magnanimous approach and support. Their suggestions and criticisms greatly im-
Leo Forster mounted on microscope slides proved the text.
nearly all the specimens used in this study. His Finally, the senior author thanks his wife
remarkably clear and carefully prepared mounts Annette and his three children for sharing sev-
added immeasurably to the information gath- eral field trips and assisting in many of the tasks
ered during this work. He also helped with demanded by these excursions. His son Chris
some field collecting. Without his assistance this also helped with an attempt to interpret strong
study would not have compiled the information bristle variation in [Link] males.
it has. His retirement at the end of 1988 was
sorely felt!
We also appreciate the assistance of the staff References
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Arnett, R.H. & Samuelson, G.A. (1986) The Insect
Biology Research Centre, Agriculture Canada.
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Barry Flahey drew and inked all the genitalia Brill/Flora and Fauna Pub., Gainesville, Florida.
drawings. His skills in illustrating these small Becker, P. (1961) Observations on the life cycle and
creatures are much appreciated. immature stages of Culicoides circumscriptus Kieff.
Ms Mary Dillon kindly identified some (Diptera, Ceratopogonidae). Proceedings of the
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We also extend our thanks to Mrs Evi Remm, Plates 1,2.
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Our thanks also extends to all those curators
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who lent specimens of Serromyia: Dr R.
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pogonidae (Diptera, Nematocera). Part XIV: The
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genus Serromyia Meigen. Proceedings of the
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Systematic Entomology (1990) 15, 218

Erratum
Vossbrinck, C.R. & Friedman, S. (1989) A 28s ribo-
somal RNA phylogeny of certain cyclorrhaphous
Diptera based upon a hypervariable region. Sys-
tematic Entomology, 14, 417—431.

The tsetse species listed as Glossina simulans should

be Glossina morsitans.
160 A. Borkent and B. Bissett

Teskey (1981) and detailed characters (i.e. some groups have bristles that are more
chaetotaxy) as well as pupal characters follow markedly developed than in any species of
Saunders (1924) and Lawson (1951). Serromyia. The strong bristles as used here to
We use some terms in the keys and descrip- describe character states refer to those bristles
tions that require further comment. Some adult which stand out from the remaining setae as
Serromyia lack pruinosity on the scutum and in thicker and often appear to be more darkly
such specimens the scutum is brilliantly shiny pigmented. Figs 1C, D indicate examples of
and highly reflective in pinned material (the typical strong bristles on the fore and hind leg,
humeral pits are not distinct from the surround- respectively. The lateral strong bristles on the
ing shiny cuticle) and utterly lacking fine spicules hind coxa are shown in Fig. IB. Sometimes
when viewed laterally in slide-mounted speci- strong bristles are broken off at the base and
mens. Specimens which are reported to exhibit then the enlarged socket needs to be searched
pruinosity on the scutum are somewhat dull for, for accurate interpretation of keys and
in reflected light (so that the humeral pits ap- descriptions.
pear as discrete, shiny patches). When slide Leg coloration is illustrated somewhat dia-
mounted, such specimens have a short coat of grammatically. Leg pigmentation intensity dif-
fine spicules visible in lateral view (Fig. 1A). fered between specimens and varied according
We looked for pruinosity amongst the dorso- to preparation technique. Illustrations therefore
central setae on the laterally mounted thorax are meant to show extent of pigmentation for a
using interference contrast optics. given species with the intensity of pigmentation
When one examines the brilliantly shining not necessarily comparable between species.
scutum of a pinned specimen with a dissecting Parameres are described as either rounded or
microscope, one can be confident of the lack of tapered apically. Rounded parameres are some-
pruinosity; but when the scutum is dull, care times shrivelled and may appear pointed. How-
must be taken. A somewhat dirty specimen may ever, in most such instances the two parameres
appear dull when in fact it lacks pruinosity. In look different from one another. Parameres
most of these instances the humeral pits would which are tapered apically are always of clearly
also be dull. We generally preferred to examine defined and characteristic form, as illustrated in
slide-mounted material to ensure correct in- the drawings of the male genitalia.
terpretation of the state of pruinosity. Ratios and some structures discussed in this
We use the term 'strong bristles' in describing study are abbreviated as follows:
armature of the legs. Among ceratopogonids, L: length

Table 6. Descriptive statistics for hind femur length/width of male Serromyia.

Species n Mean Min. Max. SD

barberi 10 6.51 5.62 7.06 0.575

borealis 2 5.67 5.57 5.77 —

crassifemorata 9 6.10 4.16 7.06 0.937

nudicolis 16 5.96 5.38 6.64 0.388
sierrensis 1 6.19 6.19 6.19 -

vockerothi 3 5.90 5.74 6.05 —

ledicola 15 6.51 5.87 7.36 0.445

atra 13 6.55 5.00 7.00 0.539
bicolor 3 7.64 7.52 7.76 —

femorata 19 6.47 5.23 7.52 0.517

mangrovi 2 4.12 4.08 4.15 —

morio 13 7.15 6.32 7.50 0.612

pacifica 1 6.00 6.00 6.00 —

rufitarsis 10 6.68 5.90 7.06 0.367

subinermis 10 6.82 6.46 7.26 0.287
tecta 2 6.28 6.25 6.32 —
 Revision of Holarctic species of Serromyia Meigen 175

Kieffer (1913, 1919) noted that the fork of the communis Fabricius 1805 and [Link]
cubitus (as posticale) was markedly distal to the Meigen 1804 as members of Serromyia but there
position of the crossvein r-m (as transversale) is, in our opinion, no evidence for this. Remm
and that the wings were spotted. Combined (1988) placed C. communis in Ceratopogon
with the presence of short, equal claws on the (considering that Fabricius' description of
fore and mid legs and an elongate hind claw C. communis was not a new name but fol-
(equal to the length of the fifth tarsomere), lowed Meigen's earlier 1804 description) and
this description cannot apply to any known [Link] in Dasyhelea Kieffer. The date of
Serromyia, but does fit a general description of submission for the catalogue in Remm (1988)
several species of Palaearctic Monohelea. We was the end of 1982 and Szadziewski (1986),
therefore transfer the name to that genus as a after examining the type specimens, has shown
new combination: Monohelea scirpi (Kieffer). that the Ceratopogon palustris is actually a
Serromyia fuligipennis Clastrier has recently species of Forcipomyia. Havelka (1978) also
been placed in a new genus Congohelea Wirth interpreted Ceratopogon candidatus Winnertz
& Grogan and we agree that it was incorrectly 1852 as a member of Serromyia but considered
placed in Serromyia. the name unavailable because of lack of use
Havelka (1978) considered Ceratopogon (50-year rule). Remm (1988) correctly placed

Figs 12A-D. A aedeagus, B paramere, C male genitalia of Serromyia tecta; D, female genitalia of [Link].