A REVISION OF THE GENUS SYMPHOROMYIA FRAUENFELD

A REVISION OF THE GENUS SYMPHOROMYIA FRAUENFELD
(DIPTERA: RHAGIONIDAE)'
I.
INTRODUCTION. SUBGENERA AND SPECIES-GROUPS.
REVIEW OF BIOLOGY
WILLIAMJ. TURNER
Department of Entomology, Washington State University, Pullman
Abstract
Can. Ent. 106: 851-868 (1974)
The Holarctic genus Symphoromyia, the biting snipe flies, is discussed and a taxonomic history of the group is reviewed. Five subgenera including Symphoromyia s. str., Paraphoromyia Becker, and three new subgenera, Axinicera,
Pogonaria and Ochleromyia, are established and a key for their separation and
recognition of included species-groups is presented. Each subgenus and its
associated species-groups are discussed and compared with respect to distribution,
morphology, and evolutionary trends. Known aspects of biology for the genus
are presented to include: ( 1 ) a generalized life cycle; (2) immatures; ( 3 ) natural
enemies; (4) female biting activity; ( 5 ) male swarm behavior.
The species of Symphoromyia are known as the biting snipe flies because of
the blood sucking habits of the females. The genus is easily separated from other
rhagionids by the vertically deep third antennal segment and usually elongate,
swollen first segment, the 0-2-1 tibia1 spur formula and the well developed biting
mouthparts in the females. Species are known from Europe and Asia but ?he
preponderance of forms occur in western North America. The genus consists of
three recognized European species (Szilady 1934), 28 known Nearctic ones
(James 1965; Turner and Chillcott 1973), and some six described fossil forms
(Hennig 1967). The actual number of extant species, in North America at least,
is much greater, for specific characters are often cryptic and, in many cases,
nominal species may actually consist of two or more closely related (and as yet
undescribed) ones.
Frauenfeld (1867) originally described Symphoromyia with the European species
Atherix melaena Meigen as type, and included the species crassicornis Panzer and
immaculata Meigen. Symphoromyia was recognized as distinct from other closely
related genera, especially Atherix and Ptiolina, by the enlarged, kidney shaped third
antennal segment and the open anal cell. In an attempt to divide the European
species into natural groups, Becker (1921) erected the genus Paraphoromyia for
crassicornis Panzer with no other species referred to it. He distinguished it from
Syrnphoromyia by the swollen first antennal segment, the very broad female
front, close proximity of the antennal bases, and the bare parafacial areas.
Since then, Paraphoromyia has been considered a genus by some authors (Becker
1922; SCguy 1926; Szilady 1932; Steyskal 1953; Stackelberg 1954). More often
the distinctions separating it from Symphoromyia have not been recognized as
important and it has been either reduced to subgeneric rank (Szilady 1934;
RozkoSnf and Spitzer 1965) or synonymized under Symphoromyia (Lindner
1922, 1925; Nagatomi and Kanmiya 1969). No further attempt has been made
to separate the species of Symphoromyia into subgenera or species-groups, even
for those in the Nearctic region.
The European species of Symphoromyia have been reviewed completely by
Lindner ( 1925) . RozkoSnf and Spitzer ( 1965) treated those species occurring
'Scientific paper 4200, College of Agriculture Research Center, Washington State University. Research was
conducted under Project 0209.
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in Czechoslovakia while Nagatomi and Kanmiya ( 1969) examined crmsicornis in
Japan. These last two papers provide useful figures of the male genitalia of crmsicornis and melaena. Aldrich (191 5 ) provided the last and only major revision
of the Nearctic forms. Even when Leonard (1930) monographed all rhagionids
occurring here, the portion dealing with Symphoromyia remained essentially that of
Aldrich. Until now there has been no comprehensive treatment of the Holarctic
elements of the genus, although the late J. G . Chillcott had begun a major generic
revision of world species shortly before his death in 1967.
This paper is the first of a series treating the species of this widely distributed,
somewhat diverse but poorly known genus of flies. It was felt that initial establishment of subgenera and recognition of species-groups must necessarily precede
any subsequent discussion of included species. Becker's trend for separating the
European species is revived and a proposal is made to segregate the remaining
Syrnphoromyia species into several well defined subgenera, three of which are new.
Male genitalic characters arc used to supplement the generally used external
features.
While the species of Symphoromyia may at first appear to constitute a homogeneous assemblage, they can be segregated into definite clusters of species. Based
on shared morphological features, these associations of species can then be combined into subgenera separated by definite morphological gaps or discontinuities.
While each subgenus evidently represents a distinct phyletic line within the genus,
it will often include one or more independent evolutionary trends which are associated with the respective species-groups. Failure to recognize these natural
groups or their related trends has resulted in poor understanding of relationships
and subsequent taxonomic confusion by previous workers.
The currently recognized species of Symphoromyia (James 1965; Szilady
1934) have been arranged into five subgenera (Table I ) . Only onc (Paraphoromyia) has close relatives in both the Palaearctic and boreal Nearctic regions. The
subgenus Syrnphoromyia s. str. includes two European species which form a distinct
unit and are apparently not closely related to any Nearctic group. All remaining
subgenera are limited to the Nearctic and especially to western North America.
Two (Axinicera and Pogonaria) have their greatest numbers of species in California and the last (Ochleromyia) is widely distributed throughout the Cordilleran
region. With the exception of Symphorornyia and Paraphoromyia, all subgenera
contain a number of undescribed species and close siblings, many with narrow
geographic ranges.
In this paper the species have been segregated into subgenera and speciesgroups essentially on limited qualitative morphological features. These externally
visible characters appear modified at the subgeneric level but remain relatively
constant within each group with certain exceptions (e.g., species-specific modifications of male genitalia). Important features concern the head and its appendages,
especially the relative size and shape of the antenna1 segments, distance between
the eyes usually relative to head width, and the presence or absence of hair. The
exposed male terminalia, particularly the ninth tergum and dististylus of the hypopygium, show striking differences between subgenera in general plan and yet remain
relatively stable within a particular subgenus. Additional characters of minor
importance occur on the abdomen and legs as processes, groups of bristle-like
hairs, and pile or pollinosity. Female terminalia, especially the shape of the disticerus, show differences, but these have proved significant only at the specific level
and will not be considered here.
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Table I.
853
Subgenera, species-groups, and species of Symphoromyia
Symphoromyia Frauenfeld
immaculata (Meigen), 1804
melaena (Meigen), 1820
pilosa (Meigen), 1820
Paraphoromyia Becker
Crassicomis Group
algens Leonard, 1931
crassicornis (Panzer), 1809
grisea (Meigen), 1820
griseola (Fallin), 1814
montana Aldrich, 1915
Fulvipes Group
cinerea ~ohnson,1903
currani Leonard, 1931
fulvipes Bigot, 1887
hirta Johnson, 1897
flavipalpis Adarns, 1904
pleuralis Curran, 1930
trivittata Bigot, 1887
fera Coquillett, 1894
Plagens Group
limata Coquillett, 1894
plagens Williston, 1886
latipalpis Bigot, 1887
picticornis Bigot, 1887
Varicornis Group
varicornis (Loew), 1872
modesta Coquillett, 1894
trucis Coquillett, 1894
Axinicera n. subgen.
Johnsoni Group
barbata Aldrich, 1915
inurbana Aldrich, 1915
johnsoni Coquillett, 1894
pilosa Aldrich, 1915
securifera Coquillett, 1904
Sackeni Group
sackeni Aldrich, 1915
Ochleromyia n. subgen.
Kinkaidi Group
kinkaidi Aldrich, 1915
Atripes Group
atripes Bigot, 1887
pullata Coquillett, 1894
Pogonaria n. subgen.
Pachyceras Group
cervivora Turner & Chillcott, 1973
pachyceras Williston, 1886
comata Bigot, 1887
Inconspicua Group
inconspicua Turner & Chiillcon, 1973
nana Turner & Chillcott, 1973
truncata Turner & Chillcott, 1973
Plumbea Group
inquisitor Aldrich, 1915
plumbea Aldrich, 1915
Cruenta Group
cruenta Coquillett, 1894
Association of the sexes in Symphoromyia was difficult for earlier workers
because of dimorphism in many secondary sex characteristics and the limited series
of specimens, particularly males. This resulted in numerous synonyms creating
further taxonomic problems. An attempt was made in the following key to
present basic features common to the subgenera and to use characters not totaIly
sex limited. Supplemental characters distinctive for each sex or the exceptional
species or species-group are also given.
Genus SYMPHOROMYlA Frauenfeld
Symphoromyia Frauenfeld, 1867: 496; Coquillett, 1894: 53; Aldrich, 1915: 114;
Lindner, 1925: 39; SCguy, 1926: 115; Leonard, 1930: 77; Szilady, 1932:
48; Steyskal, 1953: 239; James, 1965: 342; Stuckenberg, 1973: 651. Typespecies: Atherix melaena Meigen, 1820, Syst. Beschr. 2: 109 (original
designation).
Sympheromyia, Schiner, 1 868 : 9 10 (lapsus).
Paraphoromyia Becker, Lindner, 1922: 135; 1925: 39; Nagatomi and Kanmiya,
1969: 189.
Symphoromyia (as subgenus), Szilady, 1934: 254; RozkoSn)i and Spitzer, 1965:
363.
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FIGS. 1-5. Heads of Symphoromyia spp. in three-quarter view (A, male; B, female).
1, S. (Symphoromyia) immaculara (Meigen); 2, S. (Paraphoromyia) fulvipes Bigot; 3, S. (Axinicera) johnsoni Coquillett; 4, S. (Pogonaria) pachyceras Williston; 5, S. (Ochleromyia)
kinkaidi Aldrich.
Small to medium sized, varying from 4.5 to 10.5 mrn. Integumental color black or
dark reddish brown, occasionally brown, yellowish brown, or yellowish orange. Surface
typically dusted with gray, bluish gray, brownish gray, yellow, or brown pollen, infrequently
polished. Hair or bristle-like hairs present over most of body, appearing white, black, or
golden, frequently mixed.
Male. Head (Figs. IA-5A). Semiglobular and broadly oval, wider than thorax, ocellar
triangle prominent. Eyes large, prominent, nearly obscuring frontal area, contiguous or
narrowly separated; facets unequal or rarely similar. Face broadened beneath antennae,
usually bare, separated by deep clypeofrontal sutures from wide parafacial areas, latter
smoothly continuous with genae beneath eyes; parafacials and genae densely haired, although
former sometimes bare. Occipital surface with short and sparse bristles; ocellar triangle and
upper postocular area usually with long and prominent hair. Antenna 3-segmented, inserted
below middle, bases proximal, rarely distant; first segment either subglobular or elongate and
thickened, typically densely haired; second segment subglobose with irregular or incomplete
ring of short bristles; third segment simple and vertically deep, attached above middle, hatchetor kidney-shaped in profile, invested with fine micropubescence, especially about aristal base;
arista dorsal and subapical. Proboscis poorly developed, with only fleshy labellum. Palpus
2-segmented, second segment longer than first, elongate-conical and pointed apically or thinly
cylindrical, usually densely bristled.
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August 1974,
Thorax. Subquadrate; integument dark brown or black with uniform black or gray
pollinosity; notum bi- or trivittate or unmarked. Pleural areas variously haired, hypopleura
bare. Notum and scutellum densely haired, latter particularly behind.
Wing. Venation complete and strong, outline normal for family. Membrane gray
hyaline or infuscate brown, either wholly or on apical half, stigma slightly darker and not
strongly differentiated. R above bristled beyond humeral crossvein; costa with dense tuft of
hairs ventrally at base. Other veins bare. Anal cell closed or narrowly open. Halter
varying from black or dark brown to pale yellow; knob elongate-ovoid, shorter than stem.
Squamae small and weakly haired.
Legs. Dark brown or black or yellow, latter sometimes restricted to knees, lightly
pollinose or not; structure and bristling typically unspecialized; coxa and femora with long
hair, in some species bristle-like hairs grouped into single thorn-like structures on anterior
surface of front and mid coxae; tibiae and tarsi with short, close-set setulae. Two tibia1 spurs
present on mid legs and one on hind legs, latter spur sometimes weak. Mid coxa divided into
anterodorsal portion, basicoxite, and posteroventral one, disticoxite. Hind coxa bearing
polished condyle on anteroventral corner and sometimes haired behind on posteromedial
margin.
Abdomen. Densely pollinose, integument rarely bare and shining. Elongate, tapering,
with six or seven visible segments. Hypopygium (Figs. 6-10) on ninth segment enlarged
and conspicuous, tilted slightly dorsad; ninth tergum variously modified; dististylus simple,
elongate or elbowed, or with ventral flanges; cerci flattened and leaf-like, or elongate and
narrow. Hairs moderate to dense.
Female. Head (Figs. 1B-5B). Shape similar to male; frontal width above antennae
one-fourth to one-half head width, widening above. Eye facets uniform. Face strongly
widened beneath antennae and as in male, bare or with few hairs. Occipital bristling short
and dense, longer on front and parafacial areas, latter sometimes bare. Antenna as described
for male except first segment usually shorter and bristling less dense. Proboscis elongate,
length subequal to height of face or head, directed down and slightly forward; mouthparts
fitted for biting with well developed mandibles, labellum fleshy. Palpus similar to male except
less bristled and second segment usually thickened medially and curved, rarely thin.
Thorax, wing, and legs. As described for male except hair and bristling shorter and
groups of bristle-like hairs on coxae lacking.
Abdomen. Simple in structure, tapering evenly to apex, shorter than in male, segments
frequently telescoped. External genitalia simple with variously shaped, 2-segmented cerci
on tenth abdominal segment.
KEY TO SUBGENERA
AND SPECIES-GROUPS
OF Symphoromyia
1. First antennal segment short, scarcely longer than second segment (Fig. 1 ) ; upper
portion of sternopleuron with, at most, few scattered hairs; first sternum bare
toward base; male ninth tergum nearly transverse on posterior margin (Fig. 6A);
female palpus elongate and slender. Europe . . . . . . - . . . Symphoromyia s. str.
First antennal segment much longer than second, often strongly swollen; upper
sternopleuron with few to many hairs; first sternum haired basally; male ninth
tergum transverse o r not, often produced apically; female palpus at least somewhat swollen medially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Third antennal segment concave (rarely straight) along anterior margin beneath
arista, outline hatchet-shaped; parafacials usually haired laterally o n upper half,
infrequently bare (Fig. 3 ) ; male dististylus strongly elbowed and usually without
strong ventral flange(s) (Fig. 8 C ) . (Axinicera n. subgen.) . . . . . . . . . . . . 3
Third antennal segment convex beneath arista, outline nearly round o r kidneyshaped; parafacials without hair or, if present, arranged otherwise, male dististylus not elbowed or, if so, with strong flange(s) ventrally . . . . . . . . . . . . 4
3. Abdomen dark brown, gray, o r black but never marked with orange; female abdomen usually densely gray pollinose; male middle coxa bearing black, bristle-like
hairs ("hair pencil") on anteroventral surface (Fig. 11) ; dististylus without
ventral, inner process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . johnsoni group
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857
6-10. Male terminalia (A, dorsal view of ninth tergum; B, lateral view of left side of
ninth tergum; C, left dististylus). 6, S. (Symphoromyia) immaculata (Meigen); 7, S. (Paraphoromyia) fulvipes Bigot; 8, S. (Axinicera) johnsoni Coquillett; 9, S. (Pogonaria) pachyceras
Williston; 10, S. (Ochleromyia) kinkaidi Aldrich.
FIGS.
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Abdomen completely yellowish orange in female and black with orange markings
dorsally and laterally in male (rarely entirely black); mid coxa of male haired
anteriorly but hair scattered and not grouped into definite "hair pencil"; dististylus with ventral, anterior-directed process . . . . . . . . . . . . . . . . . . sackeni group
Hind coxa haired behind on posterior margin (Fig. 12) (except varicornis Loew
and trucis Coquillett, ninth tergum of male is broadly cleft apically and female
first antennal segment is pale); male hypopygium wi'th ninth tergum nearly
transverse along posterior margin and not or scarcely produced apically
(Fig. 7A); dististylus short, almost straight, usually with 'broadened preapical
region but lacking deep ventral lobe or process (Fig. 7C) (Paraphoromyia
Becker) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.
.. . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Hind coxa bare behind; ninth tergum of male variously produced apically; dististylus
long and slender or ventrally with definite process or broad flange . . . . . . . . 8
First antennal segment of male strongly swollen and deeply convex beneath; female
front broad above antennal bases, about one-half head width when viewed from
front . . . . . . . . . . . . . . . . . . . . . . . ..
.
.
. . . . . . . . . . . . . . . . . . . crasdcornis group
First antennal segment of male slender or only slightly more convex below than
above; female front narrow, usually smaller than eye width, abdomen pale
orange on apical segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Hind coxa bare behind on posterior margin; sternopleuron above with only few
medial hairs; firs't antennal segment of female pale; male hypopygium with
ninth tergum slightly produced and distinctly notched. Coastal California and
.
group
associated Channel Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . varicornis
Hind coxa haired behind; upper portion of sternopleuron usually with many hairs;
male ninth tergum not or scarcely produced, posterior margin entire. Lower
elevation and montane forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Abdomen glossy-black or brown, pollen (if present) sparse; third antennal segment
yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . plagens group
Abdomen more or less densely gray pollinose; third antennal segment reddish brown
to black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fulvipes group
Parafacials densely haired; third antennal segment subequally rounded above and
below (Fig. 4 ) ; male eyes distinctly separated, upper and lower facets strongly
differentiated; female abdomen typically densely gray pollinose (Pogonaria
n. subgen.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
.
.. . . . . . . . . . . . . . . . . . . . 9
Parafacials usually bare or (at most) with few hairs on upper half (females) or
lower half (males); third antennal segment less rounded above than below
(Fig. 5); male eyes broadly contiguous above, facets weakly differentiated;
female abdomen usually glossy, rarely pollinose (Ochleromyia n. subgen.) . 12
Proboscis elongate, subequal to head height; third antennal segment almost circular
in outline; female front one-third to one-half head width. California . . . . . . . .
........................................
cruenta group
Proboscis length less than head height, at most slightly greater 'than clypeal height;
third antennal segment kidney-shaped; female front less than one-third head
width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
~ostorbitalbristles of male long and readily visible from front; body usually densely
pollinose black in male and gray in female, rarely subshiny, never glossy;
female halter and palpus black or dark brown . . . . . . . . . . . . pachyceras group
Postorbital bristles of male shorter, scarcely visible from front; both sexes usually
densely gray pollinose (males sometimes black); female abdomen infrequently
glossy; female halter and palpus sometimes yellow . . . . . . . . . . . . . . . . . . . . 11
Wing membrane clear or evenly infuscated; male dististylus deeply lobed ventrally.
Washington, Idaho, and Colorado . . . . . . . . . . . . . . . . . . . . . . . . . . plumbea group
Wing infuscated apically but never evenly over entire membrane; male dististylus
narrow (sometimes twisted), never with pronounced ventral lobe or deep
ventral flange. California . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .inconspicua group
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THE CANADIAN ENTOMOLOGIST
859
Hind tibia of male bearing long, erect posteroventral hairs that are distinctly longer
than tibial diameter; female with black halter and palpus; abdomen usually
glossy (rarely pollinose gray). Montane species, infrequently occurring near
sea level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alripes group
Hind tibia of male with short, erect hairs on basal half, scarcely longer than tibial
diameter; female halter and palpus yellow (latter may sometimes appear black
ventrally). Pacific Coastal species, rarely from montane areas . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . kinkaidi group
Subgenus Symphoromyia Frauenfeld sensu strict0
Figs. 1, 6
Type-species: Atherix rnelaena Meigen, 1820, Syst. Beschr. 2: 109 (original designation).
The typical subgenus is apparently restricted to Europe and includes the
species melaena (Meigen) and immaculata (Meigen). There are no records of
these species biting man or animals although immaculata has well developed
mouthparts.
This subgenus is unique as the first antennal segment is less than twice as long
as the second and not or only scarcely swollen. Also, the pleural areas are weakly
haired, the first abdominal segment is bare beneath, and the upper sternopleuron
is typically bare. The bare sternopleuron is found outside the group only in
varicornis (Loew), but this species is included in the subgenus Paraphoromyia
based on its enlarged first antennal segment and other features to be discussed. In
females of Symphoromyia s. str. the front appears very narrow just above the
antennal bases, resembling similar development in the Nearctic species.
Subgenus Paraphoromyia Becker
Figs. 2, 7, 12
Parapheromyia Becker (as genus) 1921: 59; Szilady, 1932: 48. Type-species:
Atherix crassicornis Panzer, 1809, Faunae insectorum germanicae initiae oder
Deutschlands Insecten, Hft. 105, No. 10 (original designation).
Paraphoromyia, Becker, 1922: 185; SCguy, 1926: 114; Steyskal, 1953: 239.
Pwapheromyia (as subgenus) Szilady, 1934: 254.
Paraphoromyia (as subgenus) RozkoSn? and Spitzer, 1965: 363.
When Becker ( 192 1 ) first proposed Paraphoromyia, his name appeared
instead as Parapheromyia in the key (p. 42), the generic description (p. 59), and
subsequent discussion (p. 72). Although the stem "pher" and "phoro" have
essentially the same meaning ("to carry or bear"), as a coordinate taxon of Symphoromyia, not to mention the euphony, the name properly formed should have
been Paraphoromyia. Becker evidently was quick to recognize this error, for
Paraphoromyia was used exclusively throughout his subsequent work (1922) which
followed the first within a year. Unfortunately no mention of emendation is made
in this later work. Between Becker's first and second publications, Lindner
(1922), in commenting on the status of Paraphoromyia, used the emended form
but similarly offered no comment. All recent workers have accepted Paraphoromyia except Szilady (1932, 1934), who, possibly working from the first
paper, perpetuated the use of Parapheromyia.
After careful consideration of the matter, I have assumed that the subsequent
emendation of the original spelling is justified and reflects a case of either lapsus by
Becker or perhaps copier's or printer's error. (One is reminded that lapsus
includes slips of the mind as well as slips of the pen.) In addition to maintaining
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August 1974
stability of the name in use, further support for this position lies in the fact that
neither Becker nor Lindner, for that matter, considered the misspelled name significant enough to mention the emendation in subsequent papers. Therefore thz
name accepted here is Paraphoromyia. Even though the original name, Parapheromyia, has been included among the list of synonyms, it is recommended that
the emended form be accepted with the original author and date.
This subgenus is widespread throughout the northern temperate region and
appears to be the only subgenus with species occurring in both the Nearctic and
Palaearctic rgion. Paraphoromyia has several species-groups which show a range
of forms. The crassicornis group, which includes the species algens Leonard,
crassicornis (Panzer), and montana Aldrich, is recognized by the strongly swollcn
first antennal segment in both sexes and very broad female front. There is but a
single record of biting for this group. Luigioni ( 1921) reported S. grisea (Meigen)
(= crassicornis) as biting man in Italy, but this record has not been subsequently
verified. In contrast, the fulvipes group of the biting species fulvipes Bigot,
trivitatta Bigot, and hirta Johnson, has only a moderately swollen first antennal
segment and narrow female front. Included in this group also are cinerea Johnson,
currani Leonard, and pleuralis Curran, all uncommon species of the eastern United
States. The plagens group is similar in both morphology and biting habits to the
previous group with plagens Williston and limata Coquillett differing only in the
swollen palpi and very glossy abdomens. Two last species, varicornis (Loew) and
trucis Coquillett, are restricted to California and its off-shore islands and almost
deserve a subgenus of their own. Unlike all other species of this subgenus, the
hind coxae of varicornis and rrucis are not haired behind, the upper sternopleuron
appears almost bare, and the male ninth tergum is broadly produced and apically
notched. Nonetheless they appear more closely related to Paraphoromyia than to
any other based on gross overall similarities in head, antennae, and abdominal
features but especially in the male hypopygium. For these reasons both varicornis
and trucis are included in Paraphoromyia.
FIGS. 11-12. 11, Lateral view of middle coxa of S. (Axinicera) johnsoni Coquillett male
showing group of bristle-like hairs. 12, Posterolateral view of right hind coxa of S. (Paraphoromyia) fulvipes Bigot with hairs behind on posterior margin. ex, coxa; fe, femur;
hp, bristle-like hairs ("hair pencil").
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86 1
In general the subgenus can be recognized by the characters used in the key,
as other features have proved too variable as diagnostic features.
Axinicera new subgenus
Figs. 3, 8, 1 1
Type-species: Symphoromyia johnsoni Coquillett, 1894, J1 N.Y. ent. SOC.2: 5'4.
The subgenus Axinicera is restricted almost entirely to the State of California
with the exception of two species which extend northward into British Columbia
and three other isolated species occurring in Utah, Colorado, and Wyoming.
Nearly all the species appear to bite and some show a degree of host specificity.
As a number of undescribed species are involved, the nature of this specificity will
be discussed in subsequent treatment of the species of this subgenus.
This subgenus can easily be recognized by the hatchet-shaped third antenna1
segment. Other features include the narrow front and parafacials in the females,
outwardly and densely haired parafacials in all males and many females, and thz
strongly elbowed male dististylus. In most species the male ninth tergum is produced into a long, attenuated process.
Two distinct species-groups are recognized within the subgenus. The first
and most conspicuous is typified by sackeni AIdrich, the females of which are
yellow-bodied and the males dark. The male abdomen is usually marked with
yellow on the lateral and dorsal aspects, and the dististylus always bears a ventral,
anterior-directed process. At present there are a number of undescribed species
referred to the name sackeni. The second group is characterized by johnsoni
Coquillett. The males are gray to brown and females appear somewhat uniformly
gray pollinose on the abdomen. Males lack the inner ventral process on the
dististylus of the sackeni species-group, but they usually have a well defined group
of bristle-like hairs on the ventral portion of the middle coxae. There are several
yet undescribed species in addition to barbata Aldrich, inurbana Aldrich, securifern
Coquillett, and the type-species, johnsoni Coquillett.
Pogonaria new subgenus
Figs. 4, 9
Type-species: Symphoromyia pachyceras Williston, 1886, Trans. Am. ent. SOC.13:
287.
This subgenus appears restricted to western North America with the greatest
number of species occurring in California. Three species extend north into
Washington and two additional ones are known from Colorado and Arizona. All
apparently bite with some evidence of distinct host specificity.
Species of this subgenus typically have dense and evenly haired parafacial
areas, narrowly separated eyes in males, and strongly flanged dististyli. 'The
females are densely gray pollinose and their front is about one-third to two-fifths
head width. In the males, the abdomen is often enlarged and laterally bristled on
the fourth and fifth sternites, the following segment noticeably narrower than either
the preceding segments or the hypopygium.
The species are divided into four main groups. The first includes pachyceras
Williston and cervivora Turner and Chillcott, with long postorbital bristles in the
black colored male and with darkened wings in both sexes; the second group
includes inquisitor Aldrich and plumbea Aldrich with short male postorbital bristles
and clear wings; among the third group are the recently described truncata, nana,
and inconspicua (Turner and Chillcott 1973) and several undescribed species with
862
THE CANADIAN ENTOMOLOGIST
August 1974
short postorbital bristles, bicolored wings, and slender dististyli bearing little or no
ventral lobe. The final group is represented by cruenta Coquillett and includes
one or more undescribed forms, with very long probosces and nearly circular third
antennal segments.
It should be noted that plumbea Aldrich, to some extent, shows intermediate
characters between this subgenus and Ochleromyia. It has a male distyle structure
similar to the latter but the female head is characteristic of Pogomria.
Ochleromyia new subgenus
Figs. 5, 10
Type-species: Symphoromyia kinkaidi Aldrich, 1915, Proc. U.S. natn. MUS. 49 :
129.
The systematics of this subgenus will be discussed in a later paper (in prep.).
This subgenus consists of nine species, six of them undescribed, which are
restricted to western North America from the mountainous areas of southern California to Alaska. They may be segregated into two groups. The first, typified
by kinkaidi Aldrich, contains three species, two restricted to coastal areas of the
west and one species found all along the Sierra Nevada of California. Females of
this group have yellow halteres and palpi and very narrow front. A second group
of six species is found in the Cordilleran and Coast Ranges. Species of this group,
which includes atripes Bigot, are widespread, except pullata Coquillett which
appears endemic to Colorado, and an undescribed species restricted to the Californian Sierra Nevada. Within this group females are recognized by black halteres
and palpi and a somewhat wider front than found in the previous subgroup.
The species of Ochleromyia are all biters of man and other vertebrates and
can be recognized, with few exceptions, by the almost entirely bare parafacial areas
( a few dorsal hairs in females or ventral hairs in males), the reduced upper half of
the third antennal segment, narrow front in the female, and contiguous eyes of the
male. As mentioned above, this subgenus is most closely related to Pogonaria
with plumbea Aldrich occupying an intermediate position.
Biology
Knowledge of habits and life stages of Symphoromyia is fragmentary at best
and based on occasional observations, often anecdotal notes, which have contributed little. Immature forms have been described for several species, but
information about specific larval habits or their habitats is lacking for most.
Female biting behavior is perhaps best known, but even here only few hosts have
been recorded, while the feeding habits of most species remain unknown. Male
swarming behavior has been reported for only one North American species
(Shemanchuk and Weintraub 1961) and possibly one other.
LIFEHISTORY.The following generalized account of the life history of Symphoromyia has been reconstructed from field observations by the author and from scattered
reports in the literature. In spring or early summer, after apparently overwintering as
either egg or lama, the maggot-like larva becomes active and begins to feed on detritus
or on other organisms in the surrounding soil. The lama molts, probably three times,
and when fully grown, ceases feeding, moves close to the soil surface and pupates.
The imago emerges from its puparium after about 2 weeks in the pupal stage, usually in
late spring or summer. Males of many species then move to aggregation sites (e.g.,
groves of trees, slopes and crests of hills or mountain tops-Turner 1971). On warm,
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THE CANADIAN ENTOMOLOGIST
sunny days at these sites they engage in a characteristic sustained flight termed "swarming", which differs among species in appearance, location, timing, and other salient
features. Females are believed to pass quickly through these stationary swarms, are
recognized by the males, pursued, and eventually are mated. This supposed sequence
of events remains to be verified. There are reports of females taking part in the normally all-male swarms (Shemanchuk and Weintraub 1961). Only after insemination does
the female seek out its appropriate vertebrate host, take a full blood meal, and retire to
nearby vegetation to digest the blood and convert the proteinaceous material into egg
tissue. When her complement of eggs has matured, she moves to the proper oviposition
site and deposits them. Oviposition occurs on or in vegetation (Sommerman 1962) or
into moist soil. She then searches out another host and begins again the gonadotrophic
cycle of vitellogenesis and deposition which may be repeated several times before her
death (Anderson, pers. comm.). Most, if not all, Syrnphoromyia species are probably
univoltine as suggested by the typically short, well defined seasonality and definite
sequences of species at any one locality. Hoy and Anderson (1966) using these criteria
of seasonal occurrence and sequence have determined that several Californian species
of the subgenus Pogonaria are univoltine.
IMMATURES.Beling (1882) first recorded and described an immature Symphoromyia based upon the larva and pupa of the European crassicornis (Panzer). The
larva was found during mid-May in the upper layer of soil in a grass-covered roadway
at the edge of a beech forest. It pupated after 10 days and the adult emerged 13 days
later. Similar developmental timing for several undescribed Alaskan species is reported
by Solmmerman (1962). Although Beling collected additional Iarvae, none was reared
successfully. Subsequent descriptions of Symphoromyia larvae by Lundbeck (1907),
Malloch (19 17), Hennig (1952), and Brauns (1954) merely repeat the description by
Beling. Sommerman (1962, 1963) has contributed most to our current knowledge
about the ecology of and rearing techniques for immature stages through her work with
Alaskan Symphoromyia. She discovered and described both larval and pupal stages of
four species taken during May and June in Alaska from damp soil in mountain meadows
near and above timberline and from sheltered areas on s t e e ~ .well drained sloms. Also
described for the first time were eggs deposited in the laboratory by one species. Sommerman found both larvae and pupae located very near to the surface of damp soil,
often within or just beneath a surface layer of moss. In most cases, these immatures
were close to roots of certain plants, particularly Veratrum eschscholtzii, false hellebore,
and Heracleum lanatum, cow parsnip. Frohne (1957, 1959) had suggested earlier
that areas near Juneau, Alaska, might be breeding sites for the particularly abundant
species atripes Bigot and kinkaidi Aldrich encountered there. Two prominent plants
in these areas are V. eschscholtzii and H. tanatum. Near Corvallis, Ore., Levburn F.
Lewis (pers. comm.) extracted many Symphoromyia larvae from moist but not wet soil
near these same plants and reared imagos of several species. Cow parsnip was also
adjacent to a principally coniferous area in Alberta frequented by hirta Johnson (Shemanchuk and Weintraub 1961). This insect-plant association is probably not trophic
but rather that the insect immatures and the "indicator" plants probably require the
same soil type and moisture.
Sommerman (1962) suggested that the Alaskan larvae probably were not predaceous, ". . . at least not on any very active animals because they are slow-moving". She
reared numerous immatures in vials containing only soil, decaying vegetable matter, and
moss. Whether different species of Symphoromyia are obligate or facultative predators,
herbivores, or detritus feeders, or perhaps all of these, is not definitely known.
NATURALENEMIES. While the natural enemies of immature Symphoromyia are
unknown, several species of wasps, flies, and spiders prey on adults. Most appear
opportunistic rather than specific as Symphoromyia represents only a portion of total
prey. No predator took large numbers of Symphoromyia although they may capture
several individuals of a single species.
x ,
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THE CANADIAN ENTOMOLOGIST
August 1974
Several families of predaceous flies prey upon rhagionids and, on occasion, Symphoromyia. Powell and Stage (1962) reported that the asilid Stenopogon engelharti
Bromley, collected in the White Mountains of eastern California (Mono County), had
captured a Symphoromyia. I examined that prey and found it to be a female of an
undescribed species closely related to montana Aldrich. Foster (1970) cited a report
on the predatory behavior of the yellow dung fly, Scatophaga stercoraria (Anthomyiidae),
normally a predator of coprophilic flies, preying on unidentified species of Symphoromyia
in tall grass, some 0.25 to 0.5 km from the nearest sheep dung where males of the dung
fly were aggregating. Males of the family Empididae (especially Empis, Rhamphomyia,
and Hilara) are also known to take any appropriately-sized insect that can be captured
on the wing. In northern California the author observed males and mating pairs of
Rhamphomyia sudigeronis Coquillett with Symphoromyia as prey as they perched on
flowers and clung to tops of tall grasses in open meadows. Of the single empidids or
mating pairs subsequently sampled seven had female pachyceras Williston as prey, two
had males, and the remaining prey were other species of small flies. It could not be
determined at the time whether the empidids were capturing Syrnphoromyia resting on
vegetation, from swarm aggregations, or from other sources.
Among the widely distributed crabronine wasps (Sphecidae, Crabroniinae), several
genera, including Crabro, Crosserus, and Ectemnius, reportedly stock their nest cells
with rhagionid flies (LeClercq 1954). Certain wasps prey on nearly any insect they
can subdue and manage physically. Among their nest booty, rhagionids form only a
portion of the total prey stock. Other wasps tend to restrict their predation to a particular life stage or group of organisms. In Norway and Sweden, for example, Adlerz
( 1903) and Sparre-Schneider ( 1906) have reported Crabro ( Anothy reus) lapponicus
(Zett.) collecting crassicornis (Panzer), while Soot-Ryen (1924) noted that both sexes
of this species were equally represented in provisions of nests he examined. Near
Hopland in northern California, several Symphoromyia species were discovered among
the assortment of dipterous prey from a wasp nest in a rotting log. The wasp, Ecternnius (Hypocrabro) species near spiniferus (Fox), had collected, in addition to two
species of Rhagio, the following Symphoromyia: pachyceras Williston (four males, two
females) ; cervivora Turner and Chillcott (two males, two females) ; truncata T. and C .
(two females) ; and plagens Williston (one male, one female). Since the log containing
the nest was lying in an open meadow the wasps must have gathered the flies either in
the open area or beneath and about oak trees or other vegetation some 30 to 50 yd away.
Bechtel and Schlinger (1957) reported that the twig-boring E. spiniferus takes principally Ogcodes eugonatus Loew (Acroceridae) as prey, but no acrocerids were found
among these nest provisions.
Spiders are often common in foliage near where Symphoromyia females tend to
feed, males swarm, and both sexes occasionally alight. However, the author observed
only two instances of predation by spiders. In each case, the flies were ensnared in leaf
webs of Dictynus sp. (Dictynidae) and both were females of the pachyceras group.
Their abdomens appeared dark and swollen with blood from recent engorgement. 'This
suggests that they were probably entrapped as they sought a place to digest the blood
meal and mature their eggs after feeding nearby.
FEMALE
BITINGACTIVITY. While adult feeding habits of many rhagionids are not
definitely established, females of Symphoromyia and Spaniopsis apparently suck blood.
White (1914) reports an account by G . H. Hardy who was bitten while strolling through
a very large swarm of Spaniopsis tabaniformis. Several species of Spania were listed
by Paramonov (1962) as biting, but as he synonymized the Australian Spaniopsis with
the Palaearctic Spania, his biting records no doubt refer to species of the former than
to the more northern Spania. Three additional closely related genera, now considered
as belonging to the recently established family Athericidae (Stuckenberg 1973), are
also documented blood suckers. These genera are: Dasyornma (Philippi 1865; Malloch
1932), Atrichops (Desportes 1942), and Suragina (Knab 1912; Nagatomi 1962).
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THE CANADIAN ENTOMOLOGIST
865
Humans and other vertebrates apparently serve as hosts for most haematophagous
rhagionids and athericids, except for three species of Atrichops which evidently suck
blood only from frogs (Desportes 1942; Nagatomi 1962).
Symphoromyia translated from Greek means "accompanying fly. This suggests
that Frauenfeld in forming the name may have observed, or at least realized, that
females of certain species form biting aggregations that often trail after the host. This
is quite extraordinary as biting records of European species with which he was familiar
are quite rare. Becher (1882) first described and illustrated the female mouthparts of
the European melaena Meig. and immaculata Meig. Ross (1940) did likewise for the
Nearctic atripes Bigot, but it was Snodgrass (1943) who noted the similarity in between
homologous structures of Symphoromyia mouthparts and those of female tabanids.
The first report of human haematophagy was by Osten Sacken (1877) who experienced
bites from several unidentified Symphoromyia species in the Sierra Nevada of California
and noted that the female of one species bites in a manner similar to Tabanus. Subsequent to this observation, numerous reports of blood-sucking in Symphoromyia were
published. These reports were summarized in part by Aldrich (1915), and Hoy and
Anderson (1966), the latter with particular regard to Californian species. The species
involved include: atripes Bigot; crassicornis (Panzer); fulvipes Bigot; hirta Johnson;
inquisitor Aldrich; pachyceras Williston; sackeni Aldrich; limata Coquillett, and unidentified species in the johnsoni and atripes groups.
Nearly all of the above probably represent groups of related forms rather than
individual species. For example, atripes Bigot and kinkaidi A l d r i ~ hare only two of
nine very similar species belonging to the subgenus Ochleromyia. A n almost identical
situation exists for pachyceras Williston and sackeni Aldrich of the subgenera Pogonaria
and Axinicera respectively. Therefore, the wide host spectrum attributed in the past
to these "species" may actually represent biting records of several similar species under
a single name. Only when each species cian be biologically defined will it be possible
to determine precisely the host preferences of Symphoromyia. It may be found, of
course, that host biting records will provide diagnostic clues for recognition of species,
particularly in those groups where species are at present very difficult to separate.
While Symphoromyia species are often locally abundant in certain years and can
annoy man, livestock, and game animals, their medical and veterinary importance
remains undetermined. There is no evidence of any species vectoring pathogens o r
parasites, but their potential for doing so remains an important possibility.
MALE SWARMING
BEHAVIOR.Swarming is widespread among insects and well
documented throughout the Diptera (McAlpine and Munroe 1968; Downes 1969).
The first well known account of swarming in Synlphoromyia was by Shemanchuk
and Weintraub (1961), who observed large swarms of hirta Johnson (or a closely
related species) along certain forest pathways and among trees in southern Alberta.
These aggregations, some up to 100 yd long, were comprised primarily of males but
females were abundant in the vicinity and frequently entered swarms (at most in a ratio
of one female to five males), While it was possible to establish the association of sexes
by sampling the swarms, no mating pairs were observed or captured. Individual males
oriented "predominantly to a slight wind and at right angles to the sun", although
conflict between these two orientations is conceivable. Swarm acivity was reduced
when clouds obscured the sun and was absent entirely on cool, cloudy days. There
was progressive upward movement of the swarm during the day.
An earlier report by Knowlton and Maddock (1944) of swarming of hirta Johnson
has been generally overlooked. This account is based on observations made in the
Wasatch Mountains, Utah, at about 9600 ft, and is as follows: "The most interesting
and conspicuous behavior noted was the great abundance of these flies on and around
one of the two foot high cast-iron water hydrants, standing in a cleared spot. A rock
base and a wet area surrounded each hydrant. On one of the hydrants approximately
350 S. hirta rested, while an estimated 2000 flies or more swarmed around the warm
metal hydrant in a column which usually varied from three and one-hlalf feet in
866
THE CANADIAN ENTOMOLOGIST
August 1974
diameter and extended f r o m a few inches above the ground t o 6 o r 8 inches above the
hydrant". While the sex of the swarming and resting flies was not indicated, it was
assumed f r o m the context of the report that they were females (Shemanchuk and Weintraub 1961). However, the physical setting of the area (i.e., a n open, sunny spot and
a vertical metal hydrant) as well as the nature of the swarm (i.e., columnar aggregation)
suggests that the authors observed a male swarm.
T h e purpose of swarming i n Diptera is a matter of considerable conjecture, and
it is n o less clear f o r species of Symphoromyia. Swarming is nevertheless widespread
among a large number of species (Turner 197 1) . F r o m extensive field observations
made by the author, he concludes that this activity is species-specific ( o r a t least different
between species a t the same locality) and like biting behavior may eventually prove t o
be taxonomically applicable.
Acknowledgments
The a u t h o r wishes t o t h a n k Drs. Maurice T. James, Lynda L. Richards, and
H o r a c e S. Telford, Washington S t a t e University, f o r their critical review of the
m a n u s c r i p t and f o r o t h e r c o m m e n t s . I a m grateful to Drs. Willis W. W i r t h
National M u s e u m of N a t u r a l History, Washington, D.C.) and H e r b e r t J. Teskey
(Canadian N a t i o n a l Collection, O t t a w a ) f o r loan of European specimens f o r comparison. Dr. T e s k e y also made available notes and papers of t h e late J. G. Chillc o t t and f o r this kindness I am m o s t appreciative. Finally I am i n d e b t e d to
Mr. M i c h a e l M. Bentzien, Drs. Frank R. Cole and E v e r t I. Schlinger f o r assistance
in determining spiders, empidids, and sphecid wasps respectively.
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(Received 25 February 1974)