Journal of the American Control Association, 2O(2):2Og_214,2O04 Copyright A 20M by the American Mosquito Control Association. Inc.

SCIENTIFICOPINION

THE AND SUBGENUS CATEGORIES WITHIN CULICIDAE AND PLACEMENT OF OCHLEROTATUS AS A SUBGENUS OF

HARRY M. SAVAGE' ,q,NoDANIEL STRICKMANa3

ABSTRACT Many - species of Culicidae are of major medical, veterinary, and economic importance. To facilitate discussion among taxonomists, medical entomologists, ecologists, and vector control specialists, it is essential that culicidologists be able to readily recognize individual genera. Adult female mosquiioes, the stage most often encountered in surveys, should be identifiable to genus *itho,rt dissection with the aid of a good- quality dissecting microscope. Female adult specimens of Ochlerotatus and Aedes as defined by Reinert c'annot be identified morphologically without dissection, and no distinct differences in biology, behavior, and ecology distinguish these 2 taxa as currently defined. Use of these names as genera complicatis mosquito identificatron and interferes with information retrieval and communication among taxonomisis, medical entomologists, and vector control specialists. Therefore, it is our opinion that Ochlerotafzs Lynch Arribalzaga should be-placed as a subgenus of Aedes Meigen, Aedes (Ochlerotatus). We believe that the usage of thJgenus Aedei and the subgenus Ae. (Ochlerotaras) should be restored to the traditional usage during ihe intervai 1906-2000.

KEY WORDS Aedes, Ochlerotalus, Culicidae, genus and subgenus categories, mosquito control, oviposition behavior

Unlike the species category, the genus and sub- of interest to many professional groups such as ep- genus categories are not based on a biological con- idemiologists, veterinarians, physicians, politicians, cept nor are they strictly defined. Usage and defi- public health workers, urban planners, vector con- nition of generic and subgeneric taxa may vary trol specialists, and entomologists, the use of well- among groups and even between authors defined genera facilitates communication and infor- working on the same group of . Tradition- mation exchange. Such communication has seldom ally, phylogenetic taxonomists defined genera as a been more important in the United States and group of related species sharing a common phylo- worldwide. Mosquito-borne diseases draw sincere genetic origin that differ from species of other re- attention from those charged with responsibility of lated genera by a decided gap in morphological public health at all levels. The professional and characters (Mayr 1969). Entomologists have tended nonprofessional public relies heavily on simple dis- to group species within genera such that the size of tinctions between mosquito genera to make deci- the morphological gap was in inverse ratio to the sions about risk and remediation. Therefore, the number of species within the genus (Michener distinctions between medically important mosquito 1957). Following these general concepts, genera genera should be as clear as possible morphologi- with many species, such as Culex Linnaeus and cally and useful operationally. Anopheles Meigen, should be well defined and rec- Work on the and classiflcation of mos- ognizable, and separated by distinct morphological quitoes developed slowly. In the l0th edition of gaps from other related genera. Genera with few Linnaeus's (1758) Systema naturae, 2 mosquito species, for example Toxorhynchites Theobald, species were placed in the single genus Culex. In would be separated from other genera by larger, 1818, Meigen (1818) established 2 new genera, more exclusive morphological gaps. In many ani- Anopheles and Aedes Meigen, bringing the total mal groups, particularly vertebrates, morphological number of genera to 3. During the remainder of the gaps were associated with adaptation to particular l9th century, 9 new generic names were published ecological zones or niches (Inger 1958), and genera including 4 genera that remain in use today: Flae- differed in morphology and biology. The generic magogus Williston, Psorophora Robineau-Des- concept employed within a family often is a func- voidy, Sabethes Robineau-Desvoidy, and Urano- tion of the professional groups who use the classi- taenia Lynch Anlbalzaga (Thompson and Gaimari fication. For families such as the Culicidae that are 2OOO).In 19O0, experimental transmission of hu- man pathogens by mosquitoes was demonstrated. Patrick Manson experimentally infected 2 human rCenters for Disease Control and Prevention, PO Box volunteers with malaria by bites of mosquitoes 2087. Ft. Collins. CO 80522. in the genus Anopheles 2 Department of , Walter Reed Army Insti- brought from Italy, where they tute of Research, 503 Robert Grant Avenue, Silver Spring, had fed upon a malaria patient, and Walter Reed MD 20910. and colleagues working in Havana, Cuba, demon- 3 Present address: Santa Clara County Vector Control strated the transmission of virus to District, 976 Lenzen Avenue, San Jose. CA 95126. healthy volunteers by Ae. (Stegomyia) aegypti L. JuNe2004 ScrENTrFrc OPINIONS 209 infected by feeding on hospitalized patients (Philip smaller genera will result from future taxonomlc and Rozeboom 1973). These events of 1900 study and phylogenetic analysis. We agree that phy- spurred work on the taxonomy and biology of mos- logenetic analysis is desperately needed within the quitoes. Just l0 years later, Theobald (1910) rec- family Culicidae, but suggest that the best way to ognized 1,050 species in 149 genera (Edwards advance toward a natural classification is to focus 1932). When Edwards (1932) began work on the on the use of subgenera. Whenever possible, sub- Genera Insectorum he faced taxonomic chaos and genera should be deflned as monophyletic groups the task of reviewing 327 geneic and subgeneric of species. We encourage all mosquito workers to names (Thompson and Gaimari 20O0). Edwards report the subgeneric name upon I st usage of a spe- (1932) placed 1,400 species in 30 genera and 89 cies name in tables or text, for example Aedes subgenera and developed a classification ofthe Cu- (Stegomyia) albopictus (Skuse) or Culex (Neoculex) licidae that forms the basis for the present classi- tenitans Walker. If the monophyletic nature of sub- fication. Edwards (1932) pointed out that the use of genera becomes firmly established, then analysis of more inclusive generic concepts made the relation- the evolutionary relationships among the subgenera ships among species clearer and the generic limits will eventually lead to a natural classification of more easily defined. Edwards (1932) also pointed genera and other higher level taxa. As subgenera out that smaller groups of species can be usefully become well defined, taxonomists may choose to designated by subgeneric names. The broad defi- elevate selected subgenera to genera based on firm nitions of genera developed by Edwards (1932) al- phylogenetic evidence. lowed adult females to be identified to genus in the In 1906, Dyar and Knab defined Aedesbasedpri- laboratory with a good-quality dissecting micro- marily on larval morphology. This broad usage of scope and allowed mosquito genera to be recog- the genus Aedes was accepted by Edwards (1917) nized in the field as morphological and ecological and by Barraud (1928), and the name Aedes was units (Service 1993). universally employed in this inclusive sense during We believe that recognition of genera by medical the interval 1906-2000. In 1918, Dyar pointed out entomologists is essential in order to use taxa for that species of Aedes could be placed in 2 groups the generalization of biological characteristics and based on differences in the claspette of the male to facilitate subsequent species identification. Adult genitalia ("harpago" ofDyar t19181). The claspette female mosquitoes, the stage most often encoun- was developed into a columnar stem bearing a ter- tered in surveys, should be identifiable to genus minal bladelike filament in Ae. (Ochlerotatus) without dissection with the aid of a good-quality Lynch Arribalzaga and related subgenera, whereas dissecting microscope. Field personnel should be in Ae. (Aedes) Meigen and Ae. (Aedimorphus) able to recognize the common genera (e.9., Culex, Theobald the claspette is rudimentary and repre- Anopheles, and Aedes) of Culicidae in the field by sented by setae on a conical base. Edwards (192'l) morphological characters, general appearance, and confirmed Dyar's finding and noted that the 2 biological and behavioral characteristics. groups also could be defined by distinct differences With the advent of cladistics (Hennig 1966), tax- in the shape of the phallosome of the aedeagus onomists began to emphasize that genera should be ("mesosome" of Edwards t192ll). One group, rep- monophyletic assemblages of species and that gap resented by the subgenera Ae. (Ochlerotatus) and criteria were inappropriate (Wiley 1981). The es- Ae. (Finlnya) Theobald, has a simple tubelike phal- tablishment of monophyletic genera and subgenera losome, whereas the phallosome in the 2nd group, is a laudable goal, and progress toward this goal represented by Ae. (Aedes), Ae. (Aedimorphus), and will facilitate information retrieval and communi- Ae. (Stegomyla) Theobald, is a paired structure with cation among culicidologists. However, for most spiny, brushlike structures apically (Edwards 1921). groups of including the Culicidae, the rig- This dichotomy was so distinctive that Edwards "Both orous phylogenetic studies required to demonstrate (1921) stated these distinctions are so well the monophyletic nature of genera are in their in- marked that one would expect to find correspond- fancy or lacking. It is important to note that even ing distinctions in the body characters or in the lar- though some mosquito genera inchtding Aedes vae." Edwards (1941) reiterated that the subgenera (sensu lato) appear heterogeneous, no author has of Aedes could be divided into 2 groups based on presented any data or argument that Aedes (sensu the 2 characters of the male genitalia mentioned lato) is polyphyletic. Although only the manuscript above, the phallosome and claspette. In subgeneric of Isoe (2O00) provided data to suggest that Aedes descriptions, Edwards (1941) pointed out that the is paraphyletic (that is that some or perhaps all of shape of the insula of the female genitalia differed the nominal genera in the tribe may have among subgenera, but we are unable to find a clear evolved from within Aedes), many mosquito tax- statement in Edwards's publications indicating that onomists believe that Aedes is paraphyletic (Zavor- he realized that differences in the shape of the in- tink, personal communication). sula also corresponded to the dichotomy in male Zavortink (1990) suggested that the classification genitalia. Belkin (1962) added 2 additional char- of genera within the Culicidae is unnatural based acters to those supporting the dichotomy of sub- on observed species/genus ratios, and that more, genera wtthin Aedes, the presence or absence of 2to Jouruel op rHE AuenrceN Mosqurro CoNrnol AssocrA'r'IoN Vor-.20, No. 2 setae on the proctiger of the male genitalia, and the Ludlow, U day a Thurman, Ve rral lina Theobald, and presence or absence of seta l2-I in the larvae. Mat- Zeugnomyia Leicester appear to have evolved from tingly (1958) and Belkin (1962) knew of the pub- within lineage 2b (Fig. l), whereas the genera Hae- lications of Edwards and the obvious dichotomy magogus, Optfex Hutton, and Psorophora appear to within the genus Aedes, yet both cautioned against have evolved from within lineage 2a (Zavorlink, formalizing this dichotomy with taxonomic names. personal communication). We have little if any in- "The Mattingly (1958) stated genusAedes is a very formation on the phylogenetic relationships among large one, comprising more than five hundred cur- the subgenera of Aedes (sensu lato) and the other rently recognized species. To split it would involve Aedini genera, and it will likely be many years be- using various familiar names in unfamiliar combi- fore rigorous phylogenetic studies are conducted. nations and this is not to be undertaken lightly in Interpretation of the phylogenetic relationships a group of medical importance. It is not felt that it among genera and subgenera of the Aedini will can be justified at the present time." A few years likely vary considerably in the foreseeable future. "suggest later Belkin (1962) concurred, stating cau- We believe that it is unwise to modify the generic tion in proposing a reclassification of the genus or and subgeneric classiflcation within the Aedini until its dismemberment." phylogenetic relationships become stable and well Reinert (2000) recently elevated the subgenus documented. Ochlerotatus to generic rank and divided the sub- At least 3 aspects of the inclusive gents Aedes genera formerly placed in Aedes (sensu lato) among are unifying (lineage I in Fig. 1). First, all members the elevated genus Aedes and the elevated genus of the genus deposit an egg with a characteristic Ochlerotatus. Reinert (2000) based his decision to structure (Horsfall and Craig 1956; Horsfall et al. elevate Ochlerotatus on characters of the insula of 197O, 1973; Linley 1989; Linley and Clark 1989). the female genitalia that are not visible without dis- One common element is the nature of the exocho- section and a refined statement ofdifferences in the rion (consisting of the maternal follicular epitheli- male genitalia and larval seta 12-I following Ed- um). In Aedes, the exochorion forms a temporary wards (1921, 1941) and Belkin (1962'). Reinert pellicle that lasts for only a few weeks after the egg (2000) also noted exceptions to the characters he is deposited. Another consistent feature of the eggs used to separate these 2 new putative genera. As a is that a sagittal cross section is not symmetrical; result of Reinert's action (2000), medical entomol- the dorsal side is more arched than the ventral with ogists were forced to identify specimens I st to spe- respect to the final orientation of the . Finally, cies, and then to select a generic name associated on hatching, the larva cuts the chorion about one with the subgenus in which the species currently is quarter of the way from the anterior end, forming placed. We resisted initial pleas from medical en- a neatly separated cap. Although this line of dehis- tomologists and vector control specialists to correct cence has no external sign, the internal structure of this situation because the elevation of Ochlerotarus the serosal cuticle (secreted by the developing lar- resulted in 2 groups of subgenera that corresponded va) appears to include a line of separation corre- to the division of subgenera based on male genitalia sponding to the eventual hatching line (Clements discussed by Edwards (1941), and because we 1992). The presence of a preformed hatching line hoped that the elevation of Ochlerotatzs would lead also is supported by the common observation that to new characters that would allow for the identi- an embryonated egg of Aedes will pop open at the fication of these genera. Unfortunately, no addition- hatching line when exposed to gentle pressure from al support for the elevation of Ochlerotalas has a small probe. been published. The eggs of Psorophora, Haemagogus, Opifex, Although not mentioned by Reinert (2000), the and Eretmapodites are probably the most similar to most important aspect of his paper is that he defined those of Aedes. The spinose pattern of the exocho- characters that support the basal furcation (lineages rion of Psorophora is distinctive and typical of the 2a and 2b in Fig. l) within Aedes (sensu lato). entire genus (Horsfall et al. 1952, Bosworth et al. Based on his work and that of Dyar (1918), Ed- 1983). Horsfall et al. (1970) further distinguished wards (1921, 1941), and Belkin (1962), there is eggs of Psorophora from those of Aedes based on strong support for a basal furcation within Aedes the carinate shape of the chorionic pattern and on (sensu lato), with I branch leading to Aedimorphus, the more strongly arched shape of the eggs. Hae- Stegomyia, and related subgenera and the other magogus have fibers projecting from the dorsal side leading to Ochlerotatzs and related subgenera (Fig. of the chorion forming attachments to the substrate 1). However, not all furcations within phylogenetic on which the female oviposits. Although this char- trees need to be associated with formal zoological acter is different from the vast majority of Aedes, nomenclature. In fact, nearly all furcations are not several Asian species of Aedes are reported to have associated with formal taxonomic names. The basal eggs that are very similar to those of Haemagogus furcation within Aedes (Fig. 1) may actually rep- (Linley and Chadee 1991, Linley et al. 1991). The resent the basal furcation within the tribe Aedini. eggs of Eretmapodites are elongated relative to The Aedini genera Armigeres Theobald, Ayurakitia most Aedes and possess a more complicated cho- Thurman, Eretmapodites Theobald, Heizmannia rionic structure (Linley and Service 1994). Mat- Juue 2004 ScrENTrFrcOptr.ttoNs 2rl

Aedes Aedimorphus Alanstonea Albuginosus Belkinius Bothaella Cancraedes Christophersiornyia Diceronyia Edwardsaedes Fredwardsius Huaedes Indusius Isoaedes Leptosomqtomyia Lorrainea Neomelaniconion Paraedes Pseudarmigeres Scutomyia Skusea Stegomyia

Chaetocruiomyia Finlaya Geoshsea Halaedes Kmknightia Levua Macleaya Molpemyia Macidus Nothoskusea Ochlerotdus Prctomacleoya Pseudoskusea Rhinoskusea Rusticoidus Zavortinkius

Abraedes Aztecaedes Gymnometopa Howardina Kompia

Fig. l. Cladogram of subgenera of genus Aedes (sensu lato) as inferred from data presented by Reinert (2000). 1, Iineage 1, Aedes(sensulato);2a, lineage2a,Ochlerotatus sensuReinert2OOO;2b, lineage2b, AedessensuReinert 2000; 3, lineages Ieading to subgenera of Aedes (sensu lato).

tingly (1970b) drew parallels between the eggs of rather than as adults or larvae. Horsfall etal. (1973) Optf", and Aedes, but the basic difference that Op- reviewed all Culicidae in establishing a categori- fex deposits eggs below the water surface (a char- zation of hatching stimuli for mosquito eggs. The acteristic shared only with Mansonia Blanchard) is eggs of m:rny mosquito species do not require the significant. stimulus of decreasing oxygen tension for hatching. Second, and of great biological significance, the These eggs (e.g., those of Culex and Anopheles) eggs of Aedes are not deposited on the surface of hatch as soon as the embryo completes develop- the water. Speciflc morphological and physiological ment and are called type I. Hatching of type II eggs adaptations of the eggs allow hatching to delay for occurs after oxygen levels are depressed, the situ- weeks, months, or, in some cases, years. This char- ation for eggs of Aedes vexans (Meigen) lying in acteristic of the eggs probably is the cause of the the soil during a favorable season. Type III eggs adaptation of the genus to either container or flood- require conditioning by light or temperature or both water habitats with latent periods spent as eggs before hatching in water with decreasing oxygen 2t2 Jount.tlr- oF THE AMERTcIN Mosqurro CoNrnol AssocrarroN Vor-.20, No. 2 tension, corresponding to an egg in diapause. So Edwards, and Or. alba Baker\ have been observed far as is known, the development of biological to deposit eggs above the waterline, but because strategies dependent on type II and type III eggs eggs of Or. signifera hatch as soon as embryona- are unique to the genera Aedes and P.sorophora. tion is complete (Zavortink 1968), it seems likely Although it seems likely that eggs of Haemagogus that larvae make their way to the water rather than and, possibly, Armigeres would fit into the type II wait for inundation. Mattingly (1970c) reviewed the category, no information is available in the litera- bionomics of eggs of Orthopodomyia and fotnd ture suggesting that the eggs of these genera un- references to overwintering as an egg. Members of dergo a latency period characteristic of type III the genus Eretmapodites also deposit eggs above eggs. The almost exclusively tropical distribution the waterline (in plant axils) or in dry snail shells, of Haemagogus and Armigeres may be associated although these eggs apparently are not very resis- with the lack of capability to withstand cold tem- tant to drying (Mattingly l970a). The eggs of peratures in the egg stage. Anopheles dirus Peyton and Harrison (Rosenberg Horsfall's (1972) review of the bionomics of Cu- 1982), An. gambiae Giles, and An. arabiensis Pat- licidae remains the most thorough treatment of the ton (Beier et al. 1990) are capable of survival under entire family. This book is arranged by genera and dry conditions for I or 2 wk. Barr and Barr (1969) provides a convenient overview of the basic eco- reviewed the basic structure and function of sa- logical strategies for egg deposition and survival. bethine genera, and briefly mentioned that several Horsfall (1972) summarized the information on species of Wyeomyia and Malaya Leicester appar- eggs available at the time, with the following re- ently deposit eggs on dry substrates. Although sults. Genera that deposit their eggs singly on the Tripteroides bambusa (Yamada) deposits eggs di- surface of the water include Anopheles, Toxorhyn- rectly on the surface of the water, it reportedly chites, Trichoprosopon Theobald (in part, Tr. dig- overwinters as an egg in the northern part of its itatum lRondanil eggs are deposited as a maternally range (Hong and Kim 1995). Under colony condi- protected raft [Lounibos 1983]), Wyeomyia'[heo- tions, females of Tp. bambusa were observed to bald (eggs sometimes stranded on sides of plants deposit most of their eggs above the waterline on and subsequently hatch when flooded), Orthopo- a piece of balsa wood (Zavortink, personal com- domyia Theobald, Optfex (eggs deposited below the munication). water surface [Mattingly 1970b]), and Mimomyia The final unifuing characteristic of Aedes is that Theobald (Clements 1999). Genera that deposit the larvae swim with sinusoidal motion (Strickman their eggs in structured assemblages on the surface 1989) wherein the curvature of the anterior flexion of the water (rafts) include Coquillettidia Dyar, Cu- is similar in extent to the posterior flexion. Al- lex (vast majority of species [Clements 1999D, Cu- though this behavior is not unique to the genus, its liseta Felt, and Uranotaenia. Genera that cement consistency within the genus is striking given the their eggs to the leaves of aquatic plants include wide range of larval habitats occupied by the genus. Mansonia and Ficalbia Theobald, and some species The suite of morphological (egg structure), physi- of Cx. (Melanoconion) Theobald, Cx. (Neoculex') ological (resistant egg), and behavioral (oviposition Dyar, and Cx. (Lophoceraomyia) Theobald (Cle- habits and swimming motions) characters may have ments 1999). Genera that deposit their eggs singly arisen independently, but it seems more likely that above the waterline or in soil, waiting for inunda- these characters support the monophyletic origin of tion to hatch, include Aedes, Armigeres (in part, 6 Aedes (sensu lato) or lineage I (Fig. 1). species in the subgenus Leicesteria Theobald glue The only formal names available to name line- eggs to their own legs, with the female allowing ages are subgenera and genera. Systematists often larvae to hatch into water after embryonation use additional names, species groups, lineages, and [Lounibos 1983]), H aemagogus, and P sorophora. others, but these names are not recognized formally The ability to deposit eggs that resist drying by the International Code of Zoological Nomencla- above the waterline is most completely developed ture (ICZN 2000). Taxonomists must be very se- in the generaAedes, Haemagogus, and Psorophora. lective in assigning names to only the most impor- The great number of species in the genus Aedes tant and well-defined lineages. It seems prudent not suggests that the strategy of a resistant egg depos- to formalize these 2 lineages associated with the ited above the water is highly adaptive under geo- basal furcation within Aedes (sensu lato) (lineages Iogically recent ecological conditions. However, a 2a and 2b in Fig. l) with formal taxonomic names few other species and genera of Culicidae display because they are not supported by morphological these attributes. Although details were not given, characters in the female adult that are visible with- eggs of Optfex apparently are resistant to drying out dissection and by biological differences. In- (Clements 1999), presumably when eggs deposited deed, Reinert (2000) noted several exceptions to his below the waterline are later stranded. Orthopodo- new classification, and examination of recent mo- myia signifera (Coquillett) apparently glues its lecular data indicates that Aedes as defined by Rein- morphologically distinctive egg ("shaped like an ert (2000) is paraphyletic (Isoe 2000). We believe elm seed") above the water (Pratt and Kidwell that the wisest route at this juncture is to treat Aedes 1969). Other species of Orthopodomyia (Or. kummi (sensu lato) (lineage I in Fig. 1) as a genus and to Juxt 2004 ScrENTrFrcOpnqroNs zt3 focus on establishing the subgenera ofAedes (sensu of natural affinities. Insecutor Inscitiae Menstruus 6: lato) as monophyletic units (lineages at the level of 71-86. larvae of Culicidae classified arrow 3 in Fig. 1). Dyar HG, Knab F. 1906. The as independent organisms. J N Y Entomol Soc 14:169- Female adult specimens of Ochlerotatus and Ae- 230. des as defined by Reinert (2000) cannot be identi- Edwards FW. 1917. Notes on Culicidae, with descriptions and no fied morphologically without dissection, of new species. Bull Entomol Res 6:357-364. distinct differences in biology, behavioq and ecol- Edwards F"W. 1921. A revision of the mosquitoes of the ogy distinguish these 2 taxa as curently defined. Palaearctic region. Bull Entomol Res 12:263-351. Use of these names as genera complicates mosquito Edwards FW. 1932. Diptera fam. Culicidae. In: Wytsman identification and interferes with information re- P, ed. Genera Insectorum Fascicle 194. Brussels, Bel- trieval and communication among taxonomists, gium: Desmet-Verteneuil. p l-258. medical entomologists, and vector control special- Edwards FW. 1941. Mosquitoes of the Ethiopian region. ists. Therefore, it is our opinion tl:.at Ochlerotatus III.-CuIicine adults and pupae London, United King- (Natural Lynch Arribalzaga should be placed as a subgenus dom: British Museum History). W. 1966. Phylogenetic systematics Urbana, IL: of Aedes Meigen, Aedes (Ochlerotatus). We believe Hennig University of Illinois Press. that the usage of the genus Aedes and the subgenus Hong HK, Kim CM. 1995. Hiber:nation study on Aedes (Ochlerold/rs) should be restored to the tradi- Ae. dorsalis. Korean J Entomol 25:185-187. tional usage during the interval 1906-2000. Horsfall WR. 1972. Mosquitoes: their bionomics and re- lation to disease New York: Hafner Publishing Co. Horsfall WR, Craig GB Jr. 1956. Eggs of floodwater mos- ACI(NOWLEDGMENTS quitoes. IV. Species of Aedes common in Illinois (Dip- tera: Culicidae). Ann Entomol Soc Am 49:368-374. We thank Boris Kondratieff (Department of Horsfall WR. Fowler HW Jr, Moretti LJ, Larsen JR. 1973. Sciences and Pest Management, Bioagricultural Bionomics and embryology of the inland fioodwater Colorado State University, Fort Collins), F Chris- mosquito, Aedes vexans. Urbana, IL: University of 11- tian Thompson (Systematic Entomology Laborato- linois Press. ry, U.S. Department of Agriculture, Smithsonian Horsfall WR, Miles RC, Sokatch lT. 1952. Eggs of flood- Institution, Washington, DC), and Tom Zavortink water mosquitoes. I. Species of Psorophora. Ann En- (Bohart Museum, Department of Entomology, Uni- tomol Soc Am 45:618-624. versity of California, Davis) for valuable comments Horsfall WR, Voorhees FR, Cupp EW. 1970. Eggs of on a draft of this manuscript, although the authors floodwater mosquitoes. XIII. 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