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A new fossil mosquito, with notes on the morphology and taxonomy of other species reported from Eocene Baltic amber (Diptera: Culicidae)


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A new fossil mosquito, with notes on the morphology and taxonomy of other species reported from Eocene Baltic amber (Diptera: Culicidae) Males of Culiseta gedanica sp. n. and Culex erikae Szadziewski & Szadziewska, 1985 are described, and an incomplete male of Coquillettidia is reported from Baltic amber for the first time. Aedes perkunas Podenas, 1999 is recognized as a junior synonym of Culex erikae , syn. n. Aedes serafini Szadziewski, 1998 is transferred to the genus Ochlerotatus , comb. n. Mosquitoes are rare fossils in Baltic amber. They represent six species of extant genera with a worldwide distribution: Culiseta, Ochlerotatus, Coquillettidia, Culex and Aedes. A key to the identification of males is also provided.
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VOL. 80: 765-777 Gdynia 31 December 2011
DOI: 10.2478/v10200-011-0057-5
A new fossil mosquito, with notes on the morphology and taxonomy
of other species reported from Eocene Baltic amber
(Diptera: Culicidae)
Department of Invertebrate Zoology, University of Gdańsk, Piłsudskiego 46, 81-378
Gdynia, Poland; e-mail: *
ABSTRACT. Males of Culiseta gedanica sp. n. and Culex erikae SZADZIEWSKI & SZADZIEWSKA,
1985 are described, and an incomplete male of Coquillettidia is reported from Baltic amber for the
first time. Aedes perkunas PODENAS, 1999 is recognized as a junior synonym of Culex erikae, syn. n.
Aedes serafini SZADZIEWSKI, 1998 is transferred to the genus Ochlerotatus, comb. n. Mosquitoes
are rare fossils in Baltic amber. They represent six species of extant genera with a worldwide
distribution: Culiseta, Ochlerotatus, Coquillettidia, Culex and Aedes. A key to the identification of
males is also provided.
KEY WORDS: Diptera, Culicidae, Eocene, Baltic amber, Gulf of Gdańsk.
The Culicidae is a well-known family of nematocerous flies. Over 3,200 species are known
in the extant world fauna. They are frequent parasites of mammals and birds as females
feed on their blood. Mosquito larvae and pupae inhabit exclusively shallow stagnant waters.
Despite the fact that the phylogenetic history of mosquitoes goes back to the Mesozoic
era, they are rarely preserved as fossils. The oldest true mosquito is reported from Upper
Cretaceous Canadian amber (POINAR et al. 2000). Fossil mosquitoes from the Cenozoic are
reviewed by EDWARDS (1923), STATZ (1944), EVENHUIS (1994) and SZADZIEWSKI (1998).
Mosquitoes are very rare in Eocene Baltic amber (SZADZIEWSKI 1998, PODENAS 1999),
while they are more common in younger Miocene Dominican amber (SZADZIEWSKI &
GROGAN 1994, POINAR 2005).
Polish Journal of Entomology 80 (4)
This paper presents the results of an examination of mosquitoes from the private
collection of Christel and Hans Werner Hoffeins of Hamburg and the Museum of Amber
Inclusions of the University of Gdańsk.
We are much indebted to Christel and Hans Hoffeins of Hamburg (Germany) for
arranging the loan of materials. This paper is a contribution to the research grant “Extinct
and extant genera in the palaeontological record of recent families of insects” from the
Ministry of Science and Higher Education of Poland NN 303 2979 37 awarded to RSz for
the years 2009-2012.
Four male mosquitoes from Baltic amber were examined. They are embedded in
artificial resin. For an explanation of general morphological terminology and abbreviations,
see DAHL (1997), SZADZIEWSKI (1998) and BECKER et al. (2010).
Subfamily Culicinae
Genus Culiseta FELT, 1904
Culiseta gedanica sp. n.
(Figs 1-3)
The male of the species is characteristic in having prespiracular setae, a larger claw on
the fore and mid legs with a basal and median tooth, and a smaller claw with a basal tooth.
The gonocoxite is simple, not modified, and the gonostylus bears an apical expansion and
distinct tooth (see key).
Male. Body length (without proboscis) 4.6 mm; thorax and abdomen 4.2 mm (Fig. 1A).
Wing length 4.17 mm. Eyes touching, flagellum distorted. Proboscis 2.06 mm long. Palpus
2.63 mm long (Fig. 1B,C); fourth palpomere stout, 0.59 mm long, fifth palpomere slender,
0.48 mm long. Scutellum trilobed. Prespiracular (=spiracular) setae present, postspiracular
absent. Wing vein R3 2.0 times longer than R2+3; veins with slender scales (Fig. 2A). Legs
incomplete: right fore and hind legs missing; left mid leg without tarsus; hind tarsi absent.
Fifth tarsomere of fore leg with strong basal horn-like expansion ventrally (Fig. 2B). Larger
SZADZIEWSKI R., GIŁKA W.: A new fossil mosquito, with notes on morphology and taxonomy
claw of fore and mid legs with basal and median tooth, smaller claw with basal tooth (Figs
2B,C). For length of leg segments, see Table 1.
Table 1. Lengths (in mm) of leg segments in male Culiseta gedanica sp. n.
fe ti ta1 ta2 ta3 ta4 ta5
p1 - - 1.03 0.37 0.24 0.12 0.20
p2 1.77 1.89 1.42 0.63 0.40 0.15 0.21
p3 - 1.89 - - - - -
Fig. 1. Culiseta gedanica sp. n., male: A – total habitus, B – palpus, C – head.
Polish Journal of Entomology 80 (4)
Fig. 2. Culiseta gedanica sp. n., male: A – wing, B – distal tarsomeres of fore leg, C – distal
tarsomeres of mid leg.
Genitalia (Fig. 3A, B) well preserved. Tergite IX with group of strong short
posteromedian spines. Gonocoxite simple, not modified, slender. Gonostylus distinctly bent
at apex, armed with apical expansion and distinct tooth (Fig. 3C). Claspettes curved with
serrated margins and pointed apices.
Female. Unknown.
Material examined
Holotype male. Hoffeins’ collection #981.3. Syninclusions: two collembolans. The
holotype will be deposited at the Senckenberg Deutsches Entomologisches Institut (SDEI,
The specific name refers to the Gulf of Gdańsk, where the largest deposits of Eocene Baltic
amber are to be found.
SZADZIEWSKI R., GIŁKA W.: A new fossil mosquito, with notes on morphology and taxonomy
The new species is included in the extant genus Culiseta as the adult has well visible
prespiracular setae. The new fossil species is distinctly smaller than extant Culiseta and has
unique gonostyli with an apical expansion. This is the first report of a fossil in the genus.
Fig. 3. Culiseta gedanica sp. n., male genitalia, photo (A) and drawing (B, C), C – apex of
Genus Ochlerotatus LYNCH ARRIBALZAGA, 1891
Ochlerotatus serafini (SZADZIEWSKI, 1998), comb. n.
(Fig. 4)
Aedes serafini SZADZIEWSKI, 1998: 240 (male, Baltic amber).
Notes on morphology
The present re-examination of the holotype resulted in new morphological details,
which were not shown in the original description of the species. They are: Fifth tarsomere
of fore leg with strong basal expansion ventrally. Larger claw of fore and mid legs with
Polish Journal of Entomology 80 (4)
median tooth, smaller claw with basal tooth, claws of hind leg small equal, each with basal
tooth. Tergite IX with group of four setae placed on a spatulate process in caudomedian
position. Base of gonocoxite with distinct ventral swelling; subtriangular expansion at mid
length of inner margin of gonocoxite bearing three stout spine-like setae directed ventrally,
and apical inner lobe (Fig. 4). Gonostyli invisible. Claspettes barely visible.
Fig. 4. Ochlerotatus serafini (SZADZIEWSKI), male genitalia (gonostyli not visible).
SZADZIEWSKI R., GIŁKA W.: A new fossil mosquito, with notes on morphology and taxonomy
Material examined
Holotype male of Aedes serafini, Museum of Amber Inclusions, University of Gdańsk,
MAI #112.
REINERT (2000) suggested dividing the composite genus Aedes MEIGEN into two genera
Aedes and Ochlerotatus; this proposition was widely accepted, and all subgenera
subsequently received generic status (HARBACH 2011).
The gonocoxite in Ochlerotatus is characteristic in having a distinct apical inner lobe.
Such a lobe, as observed in serafini, is very similar to that found in extant O. excrucians
(WALKER) (present examination), which indicates that this fossil species can be placed in
the extant genus Ochlerotatus.
Two other species of the genus Aedes, described by SZADZIEWSKI (1998) from Baltic
amber (A. hoffeinsorum and A. damzeni), are left in the genus of traditional concept, and
need further materials and studies. They were originally placed in the subgenus Finlaya
THEOBALD, which now is a distinct genus. However, A. hoffeinsorum and A. damzeni
cannot be placed in Finlaya, because in this genus the males have bigger claws on the fore
and mid legs armed with a basal and median tooth, whereas the smaller claw has a basal
tooth (HARBACH 2011). In A. hoffeinsorum all claws are simple, and in A. damzeni the
claws have no basal tooth (SZADZIEWSKI 1998).
Genus Coquillettidia DYAR, 1905
Coquillettidia sp. indet.
(Fig. 5)
Male (Fig. 5A). Hind legs, right wing and genitalia missing. Wing length 2.7 mm.
Fourth palpomere stout (0.47 mm), fifth palpomere slender (0.44 mm). Prespiracular and
postspiracular setae absent. Fifth tarsomere of fore leg with strong basal expansion
ventrally. Scales on wing veins broad (Fig. 5B). Larger claw of fore and mid legs with basal and
median tooth, smaller claw simple, without teeth. For length of legs segments, see Table 2.
Table 2. Length (in mm) of legs segments in male Coquillettidia sp. indet.
ta1 ta2 ta3 ta4 ta5
p1 0.92 0.29 0.20 0.09 0.13
p2 1.37 0.56 0.31 0.12 0.15
Polish Journal of Entomology 80 (4)
Fig. 5. Coquillettidia sp. indet., male: A – total habitus, B – wing veins with broad scales.
SZADZIEWSKI R., GIŁKA W.: A new fossil mosquito, with notes on morphology and taxonomy
Material examined
Hoffeins’ collection, male. #1719-1.
The subgenus Coquillettidia was recently removed from the genus Mansonia
BLANCHARD and elevated to a distinct genus (DAHL 1997, BECKER et al. 2010, HARBACH
2011). In the extant European fauna Mansonia is absent, while Coquillettidia is represented
by one species C. richiardii (FICALBI). Larvae and pupae of species from both genera
pierce water plants and take air from plant tissues. They are able to inhabit lakes and ponds
with fish, hiding from predators among plants in the littoral zone. This is the first record of
the genus in Eocene Baltic amber.
Genus Culex LINNAEUS, 1758
(Figs 6, 7)
Culex erikae SZADZIEWSKI & SZADZIEWSKA, 1985: 515 (female, Baltic amber, Eocene).
Aedes perkunas PODENAS, 1999: 113 (male, Baltic amber, Eocene), syn. n.
Male. Almost complete (Fig. 6A). Only distal portion of right flagellum missing. Left
side of thorax covered with milky fog. Body length about 5.0 mm (without proboscis).
Flagellum 1.77 mm long, plume well developed; distal flagellomeres 12 and 13 elongated,
each 0.48 mm. Terminal flagellomere armed with cylindrical apical prolongation. Proboscis
2.06 mm. Palpus 2.74 mm; fourth palpomere 0.53 mm long, fifth palpomere 0.45 mm long,
both slender, combined length of 4th and 5th palpomeres 1.56 times longer than 3rd
palpomere (Fig. 6B). Wing length measured from basal arculus 2.98 mm. Vein R2 4.1 times
longer than vein R2+3. Wing scales slender (Fig. 6C). Fifth tarsomere of fore leg with strong
basal expansion ventrally. Larger claw of fore and mid legs with median tooth, smaller
claw with basal tooth, claws of hind leg equal, simple (Fig. 7A-F). For length of legs
segments, see Table 3.
Table 3. Length (in mm) of leg segments in male Culex erikae.
Fe ti ta1 ta2 ta3 ta4 ta5
p1 - - 1.37 0.40 0.21 0.09 0.20
p2 - - 1.77 0.68 0.38 0.14 0.18
p3 1.71 1.94 2.19 1.14 0.78 0.43 0.25
Polish Journal of Entomology 80 (4)
Fig. 6. Culex erikae SZADZIEWSKI & SZADZIEWSKA, male: A – total habitus, B – palpus, C – wings.
SZADZIEWSKI R., GIŁKA W.: A new fossil mosquito, with notes on morphology and taxonomy
Fig. 7. Culex erikae SZADZIEWSKI & SZADZIEWSKA, male: A-C – distal tarsomeres of fore (A), mid
(B) and hind leg (C); D-F – tarsal claws of fore (D), mid (E) and hind leg (F); G, H – lateral aspect of
Genitalia (Fig. 7G, H). Observable in lateral aspect. Base of gonocoxite barely visible; a
ventromedian group of four long spatulate spines placed subapically, well visible.
Gonostylus evenly bent, tapering to pointed apex armed with short tooth.
Polish Journal of Entomology 80 (4)
Material examined
Hoffeins collection, #981-2a. male. Syninclusions in amber piece 2b: Mycetophilidae
and Hymenoptera.
The holotype male of Aedes perkunas is not complete: it lacks a hypopygium and hind
legs. According to the original description, the male of this mosquito has the claws of the fore
and mid legs unequal, and the larger one is armed with a median tooth. This character is
usually present in extant and fossil Culex, Ochlerotatus and Aedes. However, in the examined
males of these genera, the smaller claw has a basal tooth. We suspect that this character was
overlooked in the species described by PODENAS (1999), as was the basal-ventral tubercle on
the fifth tarsomere of the fore leg, which is usually present in male mosquitoes. The male of
Aedes perkunas, devoid of genitalia, is similar to O. serafini and Culex erikae. We took into
consideration the proportions of the palpomeres (3rd / 4th + 5th), which in A. perkunas and C.
erikae are the same (1.6), whereas they differ distinctly from that found in O. serafini (1.9).
Thus we propose to treat A. perkunas as a junior synonym of C. erikae.
Key to male mosquitoes from Baltic amber
1. Wing veins with broad scales (Fig. 5B) ........................................ Coquillettidia sp. indet.
-. Wing veins with slender scales (Fig. 2A) .......................................................................... 2
2. Prespiracular setae present. Larger claw of fore and mid leg with basal and median tooth
(Fig. 2B, C) ...............................................................................… Culiseta gedanica sp. n.
-. Prespiracular setae absent. Larger claw of fore and mid leg simple or with median tooth,
basal tooth absent (Fig. 7D, E) ......................................................................................… 3
3. Gonocoxite with a group of subapical spatulate spines on ventromedian surface (Fig. 7G, H)
…………………….………………… Culex erikae SZADZIEWSKI & SZADZIEWSKA, 1985
-. Gonocoxite simple, without modifications ........................................................................ 4
4. Palpus shorter than proboscis …………...……. Aedes hoffeinsorum SZADZIEWSKI, 1998
-. Palpus longer than proboscis ….…………………………………....…………………… 5
5. Fourth palpal segment slender, fifth one short .......... Aedes damzeni SZADZIEWSKI, 1998
-. Fourth palpal segment stout, fifth one long ... Ochlerotatus serafini (SZADZIEWSKI, 1998)
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Received: December 6, 2011
Accepted: December 18, 2011
... The great majority of the extant Holarctic species belongs to the genus Ochlerotatus. The oldest fossil species in the genus is Ochlerotatus serafini (Szadziewski 1998) from Eocene Baltic amber (Szadziewski & Giłka 2011). ...
... and C. varivestita (Statz 1944b), n. comb. The oldest report of the genus is from Eocene Baltic amber (Szadziewski & Giłka 2011 Description. Female and male. ...
The Diptera fauna from the late Eocene of the Isle of Wight (Bembridge Marls) is studied including redescriptions of formerly described material. The fauna includes the following
... In rock deposits they are barely preserved and details of their morphology usually are not visible. They are reported from the late Eocene Insects Limestone of the Isle of Wight (Ochlerotatus Lynch Arribálzaga, 1891; Coquillettidia Dyar, 1905;Culex Linnaeus, 1758;Edwards, 1923b;Szadziewski, 2019), from Miocene deposits of Rott (Coquillettidia, Anopheles Meigen, 1818;Statz, 1944;Szadziewski, 2019;Szadziewski & Giłka, 2011), Eocene deposits of Green River (Culex; Scudder, 1890), Eocene deposits in Colorado (Culex; Cockerell, 1919) and from Eocene shale deposits in Montana (Culiseta Felt, 1904;Harbach & Greenwalt 2012). Inclusions of mosquitoes from Cenozoic ambers are well preserved and allow more detailed studies. ...
... Inclusions of mosquitoes from Cenozoic ambers are well preserved and allow more detailed studies. They are very rare in Eocene Baltic amber (Culex Linnaeus, Coquillettidia Dyar, Culiseta, Ochlerotatus, Eoaedes Harbach & Greenwalt, 2012;Aetheapnomyia Harbach & Greenwalt, 2012;Szadziewski, 1998;Podenas, 1999;Szadziewski & Giłka, 2011;Harbach & Greenwalt, 2012), while they are more common in younger Miocene Dominican and Mexican ambers (Anopheles, Culex, Toxorhynchites Theobald, 1901;Szadziewski & Grogan, 1994;Poinar, 2005;, 2008. ...
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... Genitalia (Fig. 5I, J): Gonocoxite long (~360 μm), tapering toward rounded apex; gonostylus slender (~310 μm), hockey-stick-shaped, distinctly curved subapically, with apico-medial lobe and long apical tooth; claspette short, curved, with darkly pigmented, serrate margins and pointed apex (cf. Szadziewski & Giłka 2011: fig. 3). ...
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This study was conducted to know the common mosquito species in Amant Al-Asimah (Sana’a city) – Yemen, during the period from August 2018 to April 2019. Larvae mosquito were collected and identified to presence of two species in two genera. Culisetalongiareolata was more prevalence (98.1%) and well distributed in most districts of Amant Al-Asimah, while Culexpipiens was found with less prevalence (1.9%) and distributed in only four districts.Vectors of viruses and nematodes which caused different diseases were recorded in this study.
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The paper overviews the fossil record of insects with long mouthparts and rostra adapted to feeding on floral nectar and pollination drops of extinct gymnosperms. The presence of suctorial mouthparts is demonstrated for the first time for the Permian mecopterans Permochoristidae and Permotanyderidae. The long-proboscid scorpionflies Mesopsychidae are recorded for the first time from the Upper Jurassic of Kazakhstan. A new finding of a detached head of a long-proboscid nectar-feeding brachyceran fly is reported from the Lower Cretaceous of Transbaikalia. Three major radiations of long-proboscid nectar feeders are identified: the Paleozoic, the Mesozoic, and the Cenozoic one; they were related to the Paleozoic seed ferns, the Bennettitales and other Mesozoic entomophilous gymnosperms, and the flowering plants, respectively. The earliest long-proboscid nectar feeders, found in the Lower Permian deposits of the Cis-Urals, belong to Protomeropidae (stem-Amphiesmenoptera). The few other Paleozoic insects specialized to nectarivory probably also included some long-proboscid Permochoristidae. The diversity of long-proboscid nectar feeders shows a dramatic increase since the Middle Jurassic. About 70 Mesozoic species with preserved long mouthparts and rostra are known to date; they belong to 12 families and 3 orders (Mecoptera, Neuroptera, and Diptera) and can be clustered into three morphogroups. With the beginning of the Cenozoic the long-proboscid Mecoptera and Neuroptera were supplanted by Hymenoptera and Lepidoptera in the nectar-feeding niche, while Diptera on the whole retained this specialization. Considerable abundance of long-proboscid nectar feeders before the appearance of flowers with hidden nectar indicates that complex pollination systems first evolved in gymnosperms. Therefore, insect pollination cannot be considered the key novelty in flowering plants crucial for their evolutionary success.
Professor Ryszard Szadziewski’s adventure with entomology started in 1968 with his participation in the “Wielka Gra” (The Great Game) Polish TV 1 game show. With his mathematics and physics background and mind, he took a part in this contest as an amateur (it was one of the rules of the show). Even though he did not win this contest, he won entomology as a passion and profession.
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A new species of fossil mosquito, Culex malariager (Culicidae: Diptera), is described from Dominican amber. This species differs from extant members of the genus by the following combination of characters: 1) proboscis shorter than antennae; 2) wing scales both long and narrow and short and broad; 3) base of R vein (remigium) with two groups of dorsal setae; 4) clypeus as long as broad; 5) empodium on foretarsus; 6) vein 2A with rows of erect, short setae, and 7) postgenital lobe covering base of cerci. Inside the body cavity of C. malariensis are various stages (oocysts, sporozoites, ookinete, and microgametocyte) of a Plasmodium malarial parasite. This is the first record of a fossil mosquito vector of a vertebrate pathogen.
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Blood sucking gnats (Culicidae) are very rare in the Baltic amber (the Eocene). Only three males belonging to genus Aedes have been described so far. All three are regarded as separate species. No fully preserved females of that genus are known. A. perkunas sp. n. from the collection of Vilnius university is described. The diagnosis of the genus and the key to males of Aedes species from the Baltic amber are given.
Hennig’s (1966) concept of hierarchies of monophyletic taxons based on common ancestors (with plesiomorphies or retained primitive, ancestral characters) and shared synapomorphies (homologous shared characters inferred to have been present in the nearest common ancestor but not in earlier ancestors nor in the taxa outside this group), has provided the theoretical basis for taxa formation. During the first decades following its proposal, it resulted in the establishment of cladistic trees where recency of common ancestry is the sole criterion for grouping of the taxa. The problem of ranking taxa and tree formation in a Darwinian evolutionary context was the next scientific step (Eldridge and Cracraft 1980). Opinions about what different nodes represent, how to deal with branch lengths, and how to rank monophyletic entities, have become part of a scientific field of its own (Britton et al. 2007). Applying both morphologic and genetic taxonomic methods, and working with different groups of characters to reveal evolutionary relationships between insect orders or families as monophyletic groups, is now becoming standard. In the new millennium this work has resulted in several new hypotheses of phylogenetic trees of Insecta. Wheeler et al. (2001) established the relationship between Diptera and Strepsiptera. Grimaldi and Engel (2005) in their comprehensive work on extinct and extant Insecta, summarized different hypotheses for Diptera. They accepted five suborders of Lower Diptera (Nematocera): Tipulomorpha, Psychodomorpha, Culicomorpha, Blephariceromorpha and Bibionomorpha. They placed Anisopodidae as a sistergroup to Brachycera (all higher Diptera) and discarded Nematocera as a paraphyletic group (including a most recent common ancestor and some, but not all, of it descendants). This view has been strongly advocated by Amorim et al. (2006). They recognised seven suborders/infraorders instead of Nematocera and added Brachycera as the eighth suborder/infraorder for the rest of the Diptera.
Mosquitoes and Their Control presents a wealth of information on the bionomics, systematics, ecology, research techniques and control of both nuisance and disease vector mosquitoes in an easily readable style, providing practical guidelines and important information for professionals and laymen alike. Ninety-two European species and more than 100 globally important vector and nuisance species are included in the book. Most of them, including all European species, are described in the fully illustrated identification keys, followed by a detailed description of the morphology, biology, distribution and medical importance of each species, including over 700 detailed drawings. Mosquitoes and Their Control includes: Systematics and biology medical significance research techniques illustrated identification keys for larval and adult mosquito genera morphology, ecology, and distribution of the species identified in the keys biological, chemical, physical and genetic control of mosquitoes Mosquitoes and Their Control is a valuable tool for vector ecologists, entomologists, and all those involved with mosquito control, biology, ecology, and systematics world-wide. It will especially benefit those professionals, scientists and students dealing with mosquitoes and their control on a day-to-day basis. Society as a whole stands to gain from improved, environmentally responsible mosquito management programs designed on the basis of a broader understanding of mosquitoes and their control, as provided in this enlightening book. © Springer-Verlag Berlin Heidelberg 2003, 2010. All rights reserved.
In 1916 Prof. T. D. A. Cockerell described some fossil mosquitoes in the United States National Museum, from the Oligocene of the Isle of Wight. The material that he described consisted only of duplicates from the Brodie Collection, the main portion of which remained in the Geological Department of the British Museum (Natural History). When Prof. Cockerell came to England in 1920, he undertook the study of the main Brodie Collection of fossil insects, and found in it a considerable number of mosquitoes; at his suggestion I readily undertook to work out this material, and I wish to express my indebtedness to him for assistance and advice. My thanks are also due to Dr. A. Smith Woodward, F.E.S. and to Dr. F. A. Bather, F.E.S., for the facilities which they have afforded me. The descriptions and figures of Cockerell not being entirely clear, Dr. Bather obtained from Dr. R. S. Bassler, of Washington, photographs of the types of CockerelFs three species. These are reproduced here (PI. VII), and have been of great assistance in deciding upon the synonymy. As explained by Prof. Cockerell in a recent paper, the material examined belongs in part to the British Museum and in part to the late Mr. R. W. Hooley, F.G-.S. In the descriptions which follow, specimens belonging to the Museum are referred to by the letter I and their register-number, those belonging to Mr. Hooley (but deposited at the Museum) by the letter H. All the material is from G-urnet Bay (Isle