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Accepted by G.W. Courtney: 5 Oct. 2012; published: 30 Oct. 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3530: 25–34 (2012)
www.mapress.com/zootaxa/Article
25
urn:lsid:zoobank.org:pub:31CC9B59-447E-4005-99B1-84D9232DB97E
Two Eocene species of Culiseta (Diptera: Culicidae) from the Kishenehn
Formation in Montana
RALPH E. HARBACH1 & DALE GREENWALT2
1Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, U.K. E-mail: r.harbach@nhm.ac.uk
2Paleobiology Department, National Museum of Natural History, Washington, DC, 20013, USA. E-mail: GreenwaltD@si.edu
Abstract
Culiseta kishenehn, sp. n. and Cs. lemniscata, sp. n. (Diptera: Culicidae: Culisetini) are described from compression fos-
sils from the 46 million year old Kishenehn shale deposits in Montana, USA. The new species appear to share features
with extant species of subgenera Climacura and Culicella, respectively. The antiquity of Culiseta is examined and previ-
ously described Eocene fossil species are discussed. Eoaedes gen. n. and Aetheapnomyia gen. n. are established for Aedes
damzeni Pod nas and Ae. hoffeinsorum Szadziewski, two Eocene fossil species in Baltic amber.
Key words: Aetheapnomyia gen. n., compression fossils, Culiseta kishenehn sp. n., Cs. lemniscata sp. n., Eoaedes gen.
n., mosquitoes
Introduction
Family Culicidae (Diptera) includes 3,523 extant species classified in 111 genera (including the 80 genera of tribe
Aedini recognized in the phylogenetic classification of Reinert et al., 2009) (Harbach, 2012). Fossil Culicidae have
been described from amber inclusions (11 species) and compression/impressions (12 species) from localities in
Canada, Dominican Republic, England, Germany, France, Mexico, Myanmar, Poland and the United States.
Twenty-one of the fossil species are from the Tertiary, including species of Anopheles Meigen, Culex Linnaeus,
Mansonia Blanchard, Toxorhynchites Theobald and two extinct genera, and two are from the Cretaceous. The
discovery of the two Cretaceous species confirms Edwards’ (1923) view that Culicidae must have originated in the
Mesozoic. The oldest fossil, Burmaculex antiquus Borkent & Grimaldi, is in Burmese amber from the mid-
Cretaceous (89.3–99.6 million years ago, Mya) and the second oldest fossil, Paleoculicis minutus Poinar,
Zavortink, Pike & Johnston, is in Canadian amber from the Late (Upper) Cretaceous (66.0–100.5 Mya). Ten of the
Tertiary species are from the Oligocene (23.0–33.9 Mya; 5 Late Oligocene 23.0–28.1 Mya), nine from the Eocene
(33.9–58.0 Mya; 2 Late Eocene 33.9–41.3 Mya, 2 Early Eocene 41.3–56.0 Mya), one from the Early Miocene to
Late Eocene (13.8–41.3 Mya) and one from the Early Miocene to Late Oligocene (13.8–33.9 Mya). The Eocene
fossil species are placed in two or three extant genera, Anopheles (1 species), Culex (4 species) and perhaps
Toxorhynchites (1 species), and four species that are doubtfully assigned to the extant genus Aedes Meigen
(Harbach, 2012).
Two Eocene species of genus Culiseta Felt are described here from compression fossils discovered in shale
deposits of the Kishenehn Basin (46.2 ±0.4 Mya) in North America. The depositional environment of the
Kishenehn Basin 46 Mya was a large shallow body of water, both lacustrine and paludal. Data from several studies
suggest that the climate of the Kishenehn Basin 46 Mya was wet subtropical to tropical: (1) Approximately 10
million years after the higher temperatures of the Paleocene-Eocene Thermal Maximum, the mean annual
temperature for what is now northern Montana was approximately 15°C higher than it is now (Wolfe, 1995). (2)
The early arboreal primate Tarkadectes montanensis, originally described from the Coal Creek member (McKenna,
1990), was recently assigned to the extinct family Omomyidae (Ni et al., 2010). Most closely related to the
Omomyidae are the insectivorous Tarsiidae, extant species of which are restricted to islands of Southeast Asia
HARBACH & GREENWALT
26 · Zootaxa 3530 © 2012 Magnolia Press
(Groves & Shekelle, 2010). (3) The molluscan fauna of the Kishenehn Formation is divided into four different
successional groups, the first exemplified by Gastrocopta miniscula. Gastrocopta pellucida, selected as the extant
analog of G. miniscula, lives in an environment characterized by a mean annual temperature of 25–27°C (Pierce &
Constenius, 2001). Thus, mosquitoes that existed in the Nearctic Region during the Eocene flourished in a climate
that was considerably warmer than it is today, and that fauna is likely to be more closely related to species now
distributed mostly in the tropics and subtropics.
As Huber & Greenwalt (2011) pointed out, specimens preserved as compression fossils are generally less
informative morphologically than specimens preserved in amber. Nevertheless, as is evident from the Kishenehn
fossils described here, “they can be diagnosed moderately well and differentiated with reasonable certainty from
each other and from other extinct and extant genera”. The two Eocene species described below are the first fossil
species of Culiseta discovered thus far.
Materials and methods
Ten specimens of fossil Culicidae (5 females; 5 males) were collected in August, 2009 (USNM numbers 546534
and 546535 [Kishenehn numbers 20093.1 and 30294.5 respectively]) and August, 2011 (USNM numbers
546528–546533, 547065 and 547066 [Kishenehn numbers 505.1, 610.4, 624.1, 635.3, 686.1, 693.1, 30542.1 and
30624.1 respectively]) at three sites: Dakin (W 113° 42.173¢, N 48° 23.476¢) (USNM numbers 546535, 547065
and 547066 [Kishenehn numbers 30294.5, 30542.1 and 30624.1]), Disbrow Creek (W 113° 40.999¢, N 48°
22.572¢) (USNM number 546534 [Kishenehn number 20093.1]) and Constenius Park (W 113° 42.085¢, N 48°
23.517¢) (USNM numbers 546528–546533 [Kishenehn numbers 505.1, 610.4, 624.1, 635.3, 686.1 and 693.1
respectively]). All three collection sites contain exposures of the middle sequence of the Coal Creek member of the
Kishenehn Formation exposed along the Middle Fork of the Flathead River in northwestern Montana between
Paola and Coal Creeks. The middle sequence of the Coal Creek member has been estimated to be 46.2 ±0.4 million
years old (Lutetian) by 40Ar/39Ar analysis and 43.5 ±4.9 million years old by fission-track analysis (Constenius,
1996). These dates are supported by biostratigraphic correlations with mammalian and molluscan fossils as well as
palynology (Constenius et al., 1989; Pierce & Constenius, 2001).
Fossils were immersed in 95% ethanol for examination and photography. Specimens were photographed with
an Olympus SZX12 microscope equipped with a Q-Color5 Olympus camera. Image-Pro Plus 7.0 software (Media
Cybernetics, Inc., Bethesda, Maryland) was used to capture and record the images and measurements. The
anatomical terminology and abbreviations of Harbach & Knight (1980, 1982), revised and updated by Harbach
(2012), are used for the descriptions. The descriptions are based on examination of all available specimens, all of
which have various missing or obscured body parts and structures. The fossils are deposited in the National
Museum of Natural History, Washington, D.C. The ages indicated for mosquito fossils follow the geological
timescale determined by the International Commission on Stratigraphy (http://www.stratigraphy.org). These ages
generally differ slightly from those reported in the literature on mosquito fossils.
Taxonomy
Culiseta kishenehn, sp. n.
(Figs 1, 2, 4A)
Adults. Darkly ornamented mosquitoes, maxillary palpus, proboscis, legs (except ventral area of femora), wing and
abdominal terga dark-scaled; proboscis slender, longer than forefemur; cell R2 longer than vein R2+3, tibiae and tarsi
dark-scaled; base of hindcoxa below base of mesomeron; pulvilli absent or inconspicuous.
Female. Head: Ornamentation of head capsule not evident; antenna 1.6–1.7 mm, about 0.7 length of proboscis;
maxillary palpus 0.3–0.5 mm, 0.15−0.20 length of proboscis, with 4 palpomeres; proboscis 2.2−2.4 mm, 1.3−1.5
length of forefemur, slightly to moderately bent ventrad (appearing straight in ventral view, specimen no. USNM
547066). Thorax: Black, ornamentation obscured. Legs: Femora without knee spots; forefemur 1.6–1.9 mm,
0.7–0.8 length of proboscis; tibiae slightly swollen distally; ungues simple, fore- and midungues relatively large,
Zootaxa 3530 © 2012 Magnolia Press · 27
TWO EOCENE SPECIES OF CULISETA (DIPTERA: CULICIDAE)
equal, hindungues smaller, equal. Wing (Fig. 4A): Length 3.1–3.7 mm (mean 3.3 mm); base of subcosta with
sparse(?) setae (radial setae of Maslov, 1989); distal half of wing with outstanding linear scales on most veins; vein
R2 longer than vein R2+3 (R2/R2+3 = 1.7–2.0); base of vein M3+4 intersects M1+2 proximal to intersection of
mediocubital crossvein with M1+2; anal vein (vein 1A) terminates well beyond intersection of mediocubital
crossvein and cubitus. Abdomen: Apically rounded; terga entirely dark-scaled, no evidence of basolateral pale
patches. Genitalia: Cerci short, rounded and scarcely visible; 3 spermathecal capsules.
Male. Essentially as female. Head: Antenna about 0.7 length of proboscis; flagellomeres 1–11 each with whorl
of very long dense setae, flagellomeres 12 and 13 very long, their combined length greater than total length of
preceding flagellomeres; maxillary palpus longer than proboscis (1.1 times as long), comprised of 5 palpomeres,
palpomeres 2 and 3 ankylosed, long, about 0.75 length of proboscis, palpomere 3 slightly swollen, bearing setae
apically, palpomeres 4 and 5 not noticeably swollen, generally projecting straight forwards or turned slightly
upwards, palpomere 4 and base of palpomere 5 with numerous long setae, remainder of palpomere 5 with some
relatively short setae; proboscis about 1.4 length of forefemur. Legs: Ungues of fore- and midlegs enlarged, larger
anterior unguis with proximal tooth; hindungues small, equal. Abdomen: Terga with well-developed setae along
posterior margin. Genitalia: Prominent; gonocoxite elongate, relatively stout basally, gradually narrower toward
apex, with crescentiform (ridge-like) basal mesal lobe (basal lobe of authors, gonofurca of Maslov, 1989)
apparently bearing row(s) of setae on crest, no distinct apicodorsal lobe (apical lobe of authors); gonostylus borne
at apex of gonocoxite, rather long and narrow, undivided; paraprocts of proctiger elongate, heavily sclerotized,
with strong apical denticle.
Egg, larva and pupa. Unknown.
Etymology. The Kishenehn Formation is named for Kishenehn Creek. Kishenehn (American spelling) is a
Kutenai Native American word meaning “no good”. Why the Kutenai adopted this name for the creek is unknown.
The specific name kishenehn is understood to be masculine, but the evidence for usage as either a noun or an
adjective is not decisive; consequently, it is regarded here to be a noun in apposition to the generic name of
Culiseta, which is feminine.
Systematics. In view of the importance of larval characters for recognizing the subgenera of Culiseta
(Edwards, 1932; Dobrotworsky, 1954, 1960, 1971), it is not possible to place Cs. kishenehn in an extant subgenus.
The weakly produced basal mesal lobe of the male genitalia is shared with species of subgenera Culiseta,
Climacura Howard, Dyar & Knab and Culicella Felt. The origin of vein M3+4 proximal to the radiomedial crossvein
(Fig. 4A,B), the entirely dark-scaled abdominal terga and the absence of an apicodorsal lobe on the gonocoxite of
the male preclude the species from being placed in subgenus Culiseta, species which have the base of M3+4 more or
less in line with the radiomedial crossvein, banded abdominal terga and an apicodorsal lobe on the gonocoxite. The
distal position of the radiomedial crossvein relative to the origin of vein M3+4 and the absence of an apicodorsal lobe
on the gonocoxite are characteristics of subgenera Climacura and Culicella. Subgenus Climacura includes five
species in different parts of the world: Cs. antipodea Dobrotworsky in southeastern Australia, the type species Cs.
melanura (Coquillett) in eastern North America, Cs. tonnoiri (Edwards) and Cs. novaezealandiae Pillai in New
Zealand, and Cs. marchettei Garcia, Jeffery & Rudnick in Southeast Asia. Culiseta kishenehn resembles these
species, except the last one, in having entirely dark-scaled tarsi and abdominal terga, and also resembles Cs.
melanura in having dark-scaled maxillary palpi, which are longer than the proboscis in males. It is not certain, but
Cs. kishenehn seems to further resemble species of subgenus Climacura in having few subcostal setae
(Dobrotworsky, 1971). Subgenus Culicella includes 14 species with distributions in Australia (seven species),
North America (one species) and the Palaearctic Region (six species). Most of the species have banded tarsi and
abdominal terga (three Australian species have unbanded terga and two of these have unbanded tarsi), but in most
respects Cs. kishenehn seems to be more similar to species of subgenus Climacura. Considering the North
American provenance of Cs. kishenehn and Cs. melanura, their close similarity in adult habitus is unlikely to be
due to convergence.
Type series. Holotype male (USNM no. 546528; Kishenehn no. 505.1), allotype female (USNM no. 546529,
Kishenehn no. 610.4), paratypes males (USNM nos. 546530 546533, Kishenehn nos. 624.1, 635.3, 686.1 and
693.1 respectively), Constenius Park (W 113° 42.085¢, N 48° 23.517¢), paratype female (USNM no. 546534,
Kishenehn nos. 20093.1), Disbrow Creek (W 113° 40.999¢, N 48° 22.572¢) and paratype female (USNM no.
547066, Kishenehn no. 30624.1), Dakin (W 113° 42.173¢, N 48° 23.476¢), Kishenehn Formation, Montana,
USA.
HARBACH & GREENWALT
28 · Zootaxa 3530 © 2012 Magnolia Press
FIGURE 1. Culiseta kishenehn, sp. n. A,B. Holotype male (USNM 546528), (A) head and its appendages, (B) genitalia (tergal
aspect). C. Paratype male (USNM 546530), body, maxillary palpi, proboscis and abdominal terga entirely dark-scaled. D. Paratype
female (USNM 546534), head with short maxillary palpus. E. Allotype female (USNM 546529), distal part of left wing (dorsal)
showing the projecting linear scales and long cell R2. BML = basal dorsomesal lobe; Ppr = paraproct; R2 = cell R2; R2+3 = vein R2+3.
Zootaxa 3530 © 2012 Magnolia Press · 29
TWO EOCENE SPECIES OF CULISETA (DIPTERA: CULICIDAE)
FIGURE 2. Culiseta kishenehn, sp. n. A. Allotype female (USNM 546529), habitus, appendages of head and thorax entirely
dark-scaled (abdomen poorly preserved). B. Paratype female (USNM 547066), abdomen (left side) and right wing (ventral),
terga and wing entirely dark-scaled, long cell R2, apex of abdomen blunt. C. Paratype female (USNM 546534), apex of
abdomen showing the short cercus (partially obstructed by tergum VIII) and three spermathecal capsules. Ce = cercus; R2 = cell
R2; SCa = spermathecal capsule; VIII-S = sternum VIII.
HARBACH & GREENWALT
30 · Zootaxa 3530 © 2012 Magnolia Press
FIGURE 3. Culiseta lemniscata, sp. n. A,B. Holotype female (USNM 547065) and paratype female (USNM 546535)
respectively, habitus, proximal portion of proboscis pale, legs and wings dark-scaled, abdominal terga with basal pale bands
(arrows). C. Same as A, apex of abdomen showing the short cerci and three (indistinct) spermathecal capsules. Ce = cercus;
SCa = spermathecal capsule; VIII-S = sternum VIII.
Zootaxa 3530 © 2012 Magnolia Press · 31
TWO EOCENE SPECIES OF CULISETA (DIPTERA: CULICIDAE)
FIGURE 4. A. Culiseta kishenehn, sp. n. Left wing of allotype female (USNM 546529). B–D. Culiseta lemniscata, sp. n. B.
Right wing (dorsal) of holotype female ((USNM 547065). C,D. Same as B, (C) base of wing showing subcostal setae (actually
on ventral surface), (D) distal portion of wing showing narrow semi-erect fusiform scales on post-costal veins. M2+3 = vein M2+3;
rm = radiomedial crossvein; R2 = cell R2; R2+3 = vein R2+3; Sc = subcosta; SuS = subcostal setae.
Culiseta lemniscata, sp. n.
(Figs 3, 4B,C)
Female. Similar to Cs. kishenehn but generally paler in overall habitus; maxillary palpus, wing, tibiae and tarsi
entirely dark-scaled; differing as follows. Head: Ornamentation of head capsule not evident; antenna 2.0–2.2 mm,
about 0.8 length of proboscis, flagellomeres 1–3 slightly thicker and gradually tapering to width of distal
HARBACH & GREENWALT
32 · Zootaxa 3530 © 2012 Magnolia Press
flagellomeres; maxillary palpus with 4 palpomeres and possibly a fifth minute palpomere, length 0.5 mm, 0.20
length of proboscis; proboscis slender, longer than forefemur, length 2.5–2.7 mm, 1.3−1.5 length of forefemur, bent
ventrad, dark-scaled with lighter scaling on proximal 0.6, slightly swollen before labella. Thorax: Dark,
ornamentation obscured; mesonotum rather strongly arched. Legs: Base of hindcoxa below base of mesomeron;
femora without knee spots; forefemur 1.8−2.0 mm, 0.7−0.8 length of proboscis; hindfemur pale ventrally to near
apex; tibiae slightly swollen distally; foretibia and foretarsomere 1 with complete anteroventral and posteroventral
rows of prominent setae (evident from large alveoli); foretarsomere 1 nearly as long as foretarsomeres 2–5
combined; ungues of all legs equal, simple; pulvilli absent or inconspicuous. Wing (Fig. 4B–D): Length 3.9 mm;
base of subcosta with numerous conspicuous setae, distal half of wing with semi-erect narrow fusiform scales on
all veins except costa; cell R2 longer than vein R2+3 (R2/R2+3 = 1.6–1.7); base of vein M3+4 proximal to mediocubital
crossvein; vein CuP distinct from vein CuA; anal vein (vein 1A) terminates well beyond intersection of
mediocubital crossvein and cubitus. Abdomen: Apex rounded; terga with basal pale bands. Genitalia: Cerci short
and rounded; 3 spermathecal capsules.
Male, egg, larva and pupa. Unknown.
Etymology. The specific name lemniscata is a Latin adjective (feminine form of the masculine lemniscatus)
meaning “ribboned” or “adorned with ribbons”. The name refers to the pale banding of the abdominal terga.
Systematics. Culiseta lemniscata, like Cs. kishenehn, cannot be placed in an extant subgenus of Culiseta based
solely on adult characters. The distal position of the radiomedial crossvein relative to the origin of vein M3+4 (Fig.
4B), as noted above, is characteristic of subgenera Climacura and Culicella. The numerous subcostal setae (Fig.
4C) and banded abdominal terga (Fig. 3) indicate a closer relationship with species of subgenus Culicella than to
species of Climacura. In fact, the female of Cs. lemniscata keys to Culicella in Dobrotworsky’s (1971) key to the
world subgenera of Culiseta.
Type series. Holotype female (USNM no. 547065, Kishenehn no. 30542.1) and paratype female (USNM no.
546535, Kishenehn no. 30294.5), Dakin (W 113° 42.173¢, N 48° 23.476¢), Kishenehn Formation, Montana, USA.
Discussion
Fossil records can provide insights into anatomical diversification, historical biogeography and the antiquity of
taxa. Edwards (1923) surmized that “The origin and phylogenetic history of the Culicidae must go back to well into
the Mesozoic Era” (~66.0 Mya). From divergence times based on sequence data for nuclear protein-coding genes
and fossil calibration points, it appears that major culicid lineages date to the Early (Lower) Cretaceous
(100.5–145.0 Mya), and genus Culiseta may have appeared as early as 191.0 Mya (CL = 154.7–218.8 Mya) during
the Middle Jurassic (Reidenbach et al., 2009). Extant species of Culiseta exhibit generalized features that indicate
the genus is a primitive lineage of subfamily Culicinae (Edwards, 1932; Belkin, 1962; Marks, 1968). Culiseta may
be what paleontologists refer to as a “stem group” (Smith, 1994), a paraphyletic or polyphyletic assemblage of
species that share features of extinct taxa. The spotted distribution of the “living fossil” species of Culiseta
suggests that their extinct relatives (primitive lineage) may have existed before the separation of Pangaea during
the Late Paleozoic and Early Mesozoic Eras from about 300–200 Mya.
A number of fossil species have been assigned to Culicidae since the beginning of binomial nomenclature, but
only 23 can be placed in the family with confidence. The oldest fossil, Burmaculex antiquus from the mid-
Cretaceous (89.3–99.6 Mya), bears several plesiomorphic features, including a relatively short proboscis, which
suggest it is a stem-group mosquito that is intermediate between extant mosquitoes and other midges. In fact, the
phylogenetic analysis of morphological data conducted by Borkent & Grimaldi (2004) indicates that Burmaculex is
the sister group of all other fossil and modern mosquitoes. Morphological features of the second oldest fossil,
Paleoculicis minutus from the Late Cretaceous (66.0–100.5 Mya) indicate that Paleoculicis shares a closer affinity
with culicine than anopheline mosquitoes, which suggests that this ancestral lineage is younger than the lineage
that gave rise to subfamily Anophelinae. Anopheles dominicanus Zavortink & Poinar and Anopheles? rottensis
Statz are the only fossil anopheline mosquitoes. The former is in Dominican amber from the Late Eocene
(33.9–41.3 Mya) and the latter is a compression fossil from the Late Oligocene of Germany (13.8–33.9 Mya).
The new fossil species described above are from the Middle Eocene (41.3–47.8 Mya); hence, they are older
than the two previously described species from the Late Eocene (33.9–41.3 Mya): Anopheles dominicanus and
Zootaxa 3530 © 2012 Magnolia Press · 33
TWO EOCENE SPECIES OF CULISETA (DIPTERA: CULICIDAE)
Culex erikae Szadziewski & Szadziewska (Baltic amber). They are also older than Culex malariager Poinar
(Dominican amber) from the Early Miocene to Late Eocene (13.8–41.3 Mya) and Toxorhynchites mexicanus
Zavortink & Poinar (Mexican amber) from the Early Miocene to Late Oligocene (13.8–33.9 Mya). Seven other
Eocene species have been described. These include Culex damnatorum Scudder (Green River, Wyoming, Early
Eocene, compression/impression), Culex pipiens Linnaeus (an extant species in Baltic amber), Culex winchesteri
Cockerell (Cathedral Bluffs, Colorado, Early Eocene, compression/impression) and four species placed in the
extant genus Aedes (broad traditional sense).
According to Edwards (1923), Culex damnatorum is “Evidently a Culicine [sic] mosquito; perhaps a true
Culex, but it cannot be definitely assigned to that genus on the information available”. Cockerell (1919) named
Culex winchesteri from “a female with short maxillary palpi (about 0.4 mm), distinctly curved proboscis (3 mm)
and an abdomen “like that of true Culex, obtuse at the end, not tapering as in Aedes”. No further information is
given, but it is possible, perhaps likely, that this fossil species belongs to genus Culiseta, even though it is older
(49.0–54.8 million years old) than the two new species described here.
The four Eocene mosquitoes described as species of Aedes are in Baltic amber. Three of these species, Ae.
damzeni Szadziewski, Ae. hoffeinsorum Szadziewski and Ae. serafini Szadziewski, were described as species of
the extant subgenus Finlaya Theobald (broad traditional sense) (Szadziewski, 1998) and the fourth, Ae. perkunas
Pod nas, was regarded as “most probably” belonging to that subgenus (Pod nas, 1999). These species are certainly
culicine mosquitoes, but they cannot be assigned to an extant genus of Aedini based on the available information.
The absence of spotted wings in these species precludes them from being included in genus Finlaya (restricted
sense of Reinert et al., 2009).
The claspettes of the holotype male of Ae. damzeni are elongate with an apical sickle-like appendage as in
extant Ochlerotatus Lynch Arribálzaga, but the proctiger is said to be “barely visible” and what appears to be the
aedeagus (Szadziewski, 1998: fig. 3b,c) bears a closer resemblance to the aedeagus of members of the Aedes group
of genera, e.g. Aedimorphus Theobald, than to members of the Ochlerotatus group. Szadziewski (1998) did not
describe the ornamentation of this species, but his photograph of the holotype (fig. 5b) shows that it is a darkly
ornamented species. However, in addition to features of the male genitalia, the species differs markedly from extant
Aedini in having a short fifth palpomere (about 0.4 the length of palpomere 4) and a prominent setose lobe borne
ventrally at the base of foretarsomere 5. Based on these distinctions and other information contained in the
descriptions and illustrations of Szadziewski (1998) and Pod nas (1999), Ae. damzeni cannot be assigned to a
currently recognized genus (extinct or extant) of Aedini; hence, Eoaedes gen. n. is hereby proposed for this fossil
species. The name is derived from the Greek eos (dawn, morning, early, east) and Aedes, an extant genus. The
gender is masculine.
The type specimen of Ae. hoffeinsorum is a male with short maxillary palpi, shorter than the proboscis, a
greatly reduced palpomere 5 (about 0.3 length of palpomere 4), proboscis noticeably swollen distally, vein R2 of the
wing much shorter than vein R2+3 and the fore- and midungues borne well before the apex of the fifth tarsomere,
unique features which suggest that this species does not belong to a currently recognized genus of Aedini. The
genitalia of the specimen (illustrated but not described) bear some resemblance to those of certain members of the
Ochlerotatus group of genera, e.g. Acartomyia mariae (Sergent & Sergent), but this only supports the tribal
placement of the species. Based on the combination of unusual characters noted here and the likelihood that re-
examination of the specimen will reveal additional distinctive characters not mentioned in Szadziewski’s (1998)
brief description, Aetheapnomyia gen. n. is proposed here for the fossil species originally named Aedes
hoffeinsorum. The name is derived from the Greek aethes (unusual, strange), apnoos (dead) and myia (fly). The
gender is feminine.
The presence of postspiracular setae and claspettes with blade-like apical filaments (Szadziewski, 1998) clearly
place the holotype male of Ae. serafini within Aedini. Other than these features, the description of the specimen only
includes some measurements of head and thoracic appendages and characteristics of the ungues. In the absence of
more detailed information, it is not possible to ascertain the affinities and generic placement of this species.
Pod nas (1999) described Ae. perkunas from a male that is missing its genitalia. The specimen is darkly
ornamented, the maxillary palpi are developed as they are in many Aedini, the larger unguis of the fore- and
midlegs bears a median tooth and midtarsomere 4 is expanded ventrally toward the apex. The thoracic pleura are
partially obscured and distal parts of the wings are twisted. The specimen could be allied with extant species of
Ochlerotatus, but we are loath to relegate it to that genus in the absence of genital and wing characters.
HARBACH & GREENWALT
34 · Zootaxa 3530 © 2012 Magnolia Press
Acknowledgements
We gratefully thank two anonymous reviewers for their insightful comments on an earlier draft of this manuscript.
The second author thanks Conrad Labandeira, Department of Paleobiology, National Museum of Natural History
(NMNH), Washington, DC, for sponsorship and administrative support. This is contribution number 280 of the
Evolution of Terrestrial Ecosystems Consortium of the NMNH.
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