FORUM Classification of Mosquitoes in (Diptera: Culicidae): Paraphylyphobia, and Classification Versus Cladistic Analysis

HARRY M. SAVAGE

Centers for Disease Control and Prevention, P.O. Box 2087, Fort Collins, CO 80522

J. Med. Entomol. 42(6): 923Ð927 (2005) Downloaded from https://academic.oup.com/jme/article/42/6/923/886357 by guest on 29 September 2021 ABSTRACT Many are important vectors of human and diseases, and others are important nuisance species. To facilitate communication and information exchange among pro- fessional groups interested in vector-borne diseases, it is essential that a stable nomenclature be maintained. For the Culicidae, easily identiÞable genera based on morphology are an asset. Major changes in generic concept, the elevation of 32 subgenera within to generic status, and changes in hundreds of species names proposed in a recent article demand consideration by all parties interested in mosquito-borne diseases. The entire approach to Aedini systematics of these authors was ßawed by an inordinate fear of paraphyletic taxa or Paraphylyphobia, and their inability to distinguish between classiÞcation and cladistic analysis. Taxonomists should refrain from making taxonomic changes based on preliminary data, and they should be very selective in assigning generic names to only the most important and well-deÞned groups of species.

KEY WORDS Aedini classiÞcation, Aedes, , paraphylyphobia, mosquito classiÞcation

MANY MOSQUITO SPECIES IN the tribe Aedini, a cosmo- In this communication, I brießy review taxonomic politan group represented by 11 genera and Ϸ1,239 categories in a zoological classiÞcation, the Interna- species, are important vectors of human and animal tional Code of Zoological Nomenclature (the Code), diseases, and many others are of considerable eco- the development of generic concept within the Cu- nomic importance as nuisance or pest species. To licidae, classiÞcation within the tribe Aedini and facilitate communication and information exchange Aedes, and explain how confusion between classiÞca- among epidemiologists, physicians, veterinarians, vi- tion and cladistic analysis and paraphylyphobia lead to rologists, parasitologists, public health workers, and the classiÞcation of Aedini proposed by Reinert et al. medical entomologists, it is essential that a stable no- (2004). menclature or system of names be maintained. Such communication has seldom been more important Taxonomic Categories and the International Code worldwide, because mosquito-borne diseases draw of Zoological Nomenclature sincere attention from those charged with the respon- sibility of public health at all levels. Mosquito taxon- Unlike the species category, which is deÞned by the biological species concept (Mayr 1969, 1970), omists strive to use new methods of analysis and new the genus and subgenus categories are not based on data sets to address the phylogeny and classiÞcation of a biological concept nor are they strictly deÞned. mosquitoes, and future change in classiÞcation and Usage and deÞnition of generic and subgeneric taxa may nomenclature is inevitable. However, major changes vary among animal groups and even between authors in generic concept, the elevation of 32 subgenera working on the same group of . Traditionally, within Aedes (sensu Edwards 1932, 1941) to generic phylogenetic taxonomists deÞned genera as a group of status, and the resultant spelling changes in hundreds related species sharing a common phylogenetic origin of species names by Reinert et al. (2004) demand that differ from species of other related genera by a consideration by all parties interested in mosquito- decided gap in morphological characters (Mayr 1969). borne diseases. Although there are many technical For families such as the Culicidae that are of interest to issues associated with the manuscript of Reinert et al. many professional groups, well-deÞned, easily identiÞ- (2004), visit the following Web-based forum to access able genera based on morphology are an asset. the discussion, http://wrbu.si.edu/forums, I believe The Code (ICZN 1999) provides rules for forming that their entire approach to Aedini systematics was names in the species, subgenus, genus, , and ßawed by an inordinate fear of paraphyletic taxa or categories and requires that adjectival species paraphylyphobia, and their inability to distinguish be- names correspond to the gender of the genus in which tween classiÞcation and cladistic analysis. they are placed (Table 1). For example (Table 1), if 924 JOURNAL OF MEDICAL Vol. 42, no. 6

Table 1. Classificiation of Ae. (Stegomyia) albopictus tema Naturae (Linnaeus 1758), two mosquito species were placed in the single genus L. In 1818, Category Meigen (1818) established two new genera, Family Culicidae Meigen and Aedes Meigen, bringing the total number Subfamily of genera to three. During the remainder of the Tribe Aedini Genus Aedes 19th century, nine new generic names were pub- Lineage Aedes Lineage lished, including four genera that remain in use today: Subgenus (Stegomyia) Hemagogus Williston, Robineau-Desvoidy, Species Group Scutellaris Group Robineau-Desvoidy, and Lynch Species Ae. (Stegomyia) albopictus (Skuse) Arribalzaga (Savage and Strickman 2004). In 1900, experimental transmission of human pathogens by The formation of names in the family, subfamily, genus, subgenus, mosquitoes was demonstrated. Patrick Manson ex-

and species categories is treated by Articles in the International Code perimentally infected two human volunteers with Downloaded from https://academic.oup.com/jme/article/42/6/923/886357 by guest on 29 September 2021 of Zoological Nomenclature (the Code); above these categories are malaria by bites of Anopheles mosquitoes brought in boldface. The category Tribe is addressed by Recommendations in the Code. Other hierarchical levels sometimes used by biologist such from Italy where they had fed upon a malaria patient; as Lineage, Species Group, and Species Complex are not addressed by and Walter Reed and colleagues working in Havana the Code. demonstrated the transmission of virus to healthy volunteers by Aedes (Stegomyia) aegypti L. the species albopictus (Skuse) is placed in the mas- infected on hospitalized patients (Philip and Roze- culine genus Aedes, the correct species name is Aedes boom 1973). These events of 1900 spurred work on the albopictus (Skuse). However, if the feminine sub- and biology of mosquitoes. Just 10 yr later, genus Stegomyia was elevated to generic status, and if Theobald (1910) recognized 1,050 species in the species albopictus was placed in the genus Stego- 149 genera (Edwards 1932). In 1906, Dyar and Knab myia, the species name would have to be amended to (1906) developed a classiÞcation of the mosquitoes the feminine and the correct name would be based on larval morphology and deÞned genera, in- Stegomyia albopicta (Skuse). cluding Aedes, broadly. Howard et al. (1915, 1917), in Some public health workers falsely assume that their classiÞcation of the mosquitoes of North and the Code requires them to accept the results of new Central America and the West Indies, furthered de- taxonomic papers and new classiÞcations. However, veloped the concept that genera should be broadly this is absolutely false. The purpose of the Code is deÞned and easily identiÞable. This broad usage of to promote stability and universality of taxonomic genera within the mosquitoes, including Aedes, was names and to ensure that the valid name of each accepted by Edwards (1917) and by Barraud (1928), species is unique within each genus, according to the and the name Aedes was universally used in this rules of priority. The Code does not provide rules or inclusive sense during the interval 1906Ð2000 (Savage guidance on assigning rank. ClassiÞcations above the and Strickman 2004). Dyar (1928), in his comprehensive species level depend on subjective interpretation of classiÞcation of the mosquitoes of the Americas, deÞned taxonomic data and the philosophy adopted by the genera broadly and generally as they are used today, with investigator. The Code does not restrict freedom of well deÞned internal groups placed as subgenera. In taxonomic thought or action. The Preamble and the 1932, Edwards extended the generic and subgeneric con- Introduction to the Code make this very clear: The cepts employed by Dyar (Dyar and Knab 1906, Dyar Preamble states, “The objects of the Code are to pro- 1928) and placed 1,400 species in 30 genera and 89 sub- mote stability and universality in the scientiÞc names genera and offered a classiÞcation of the Culicidae of the of animals and to ensure that the name of each taxon World that forms the basis for the present classiÞcation. is unique and distinct. All its provisions and recommen- Edwards (1932) pointed out that the use of more inclu- dations are subservient to those ends and none restricts sive generic concepts made the relationships among spe- the freedom of taxonomic thought or actions”; and the cies clearer and the generic limits more easily deÞned. Introduction further states, “Nomenclature does not de- Edwards (1932) also pointed out that smaller groups of termine the inclusiveness or exclusiveness of any taxon, species can be usefully designated by subgeneric names. nor the rank to be accorded to any assemblage of animals, The broad deÞnitions of genera developed by Edwards but rather, provides the name that is to be used for a (1932) allowed adult female mosquitoes to be identiÞed taxon whatever taxonomic limits and rank are given to it” to genus in the laboratory with a good-quality dissecting (ICZN 1999). The only formal names available to name microscope and allowed mosquito genera to be recog- groups of species or lineages above species and below nized in the Þeld as morphological and ecological units. subfamily are subgenera and genera (see Table 1 for an example). Taxonomists should be very selective in as- Tribe Aedini signing generic and subgeneric names to only the most important and well deÞned lineages or groups of species. The traditional classiÞcation of the tribe Aedini is presented in Table 2. Eleven genera and Ϸ1,239 spe- cies are included. All genera except Aedes are small to Generic Concept with Culicidae moderate in species number. All genera were estab- Work on the taxonomy and classiÞcation of mos- lished based on distinct morphological characters be- quitoes developed slowly. In the 10th edition of Sys- tween 1818 and 1908, with the exception of two mor- November 2005 SAVAGE:CLASSIFICATION OF MOSQUITOES IN TRIBE AEDINI 925

Table 2. Traditional classification of genera within the Tribe Cladistics Analysis and Paraphyly Aedini (after Edwards 1932, Belkin 1962, Knight and Stone 1977, Reinert 1999) With the advent of cladistics (Hennig 1966), tax- onomists began to emphasize that genera should be Approximate monophyletic assemblages of species and that gap Genus no. of species criteria, or morphological distinction, were inappro- priate (Wiley 1981). A monophyletic group is a group Aedes Meigen 1818 928 that includes an ancestor and all of its descendants: for Theobald 1901 51 Ayurakitia Thurman 1954 2 example, Psorophora or Hemagogus in Fig. 1. The ac- Theobald 1901 48 ceptance of cladistic methods of phylogenetic analysis Williston 1896 28 lead to a search for monophyletic groups or taxa by a Heizmannia Ludlow 1905 34 Hutton 1902 1 generation of taxonomists. A polyphyletic group is a Psorophora Robineau-Desvoidy 1827 47 group that includes descendants from different an- Downloaded from https://academic.oup.com/jme/article/42/6/923/886357 by guest on 29 September 2021 Udaya Thurman 1954 3 cestors. Nearly all taxonomists agree that polyphyletic Verrallina Theobald 1903 93 taxa are to be avoided (Diggs and Lipscomb 2002). Zeugnomyia Leicester 1908 4 Total 1,239 However, cladists failed to realize that the removal of a monophyletic group from a phylogenetic tree re- The number of species in each genus was obtained by multiple sulted in the formation of a paraphyletic group (Brum- searches of the Water Reed Biosystematics Unit Web site in April, mitt 2002, 2003). A paraphyletic group is a group of 2005 (http://wrbu.si.edu). species that share a common ancestor but does not include all the descendants of the common ancestor: for example, the group containing the remaining spe- phologically unusual taxa with very few species cies in Fig. 1 after the monophyletic taxa Psorophora, established by Thurman in 1954 (Thurman 1954a, b; Hemagogus and Eretmapodites are removed. Although Mattingly 1958): Ayurakitia Thurman and Udaya the search for monophyletic groups deÞned by syna- Thurman. Verrallina Theobald was established in pomorphies, or shared derived character states, re- 1903, but redeÞned by Reinert (1999). ceived a conceptual basis from cladistic theory, it is The genus Aedes includes 44 subgenera (Table 3) important to note that 10 of the 11 Aedini genera in that can be divided into two lineages, the Aedes Lin- Table 2 were established before 1908 based on unusual eage and the Ochlerotatus Lineage, based primarily on morphological characters and that these 10 genera are characters of the male and female genitalia discovered monophyletic in the analysis of Reinert et al. (2004). by Dyar (1918), Edwards (1921), and Belkin (1962) These genera, or lineages, were cut out of the Aedini and discussed in detail by Reinert (2000) and Savage evolutionary tree, and the less morphologically well- and Strickman (2004). deÞned lineages representing the 44 subgenera of Aedes (Table 3) were left in the paraphyletic genus Aedes by early taxonomists. Although cladistic termi- Table 3. Classification of the subgenera of Aedes Meigen into nology was not in use, early workers recognized that two lineages after the arguments of Dyar (1918), Edwards (1921, 1941), Belkin (1962), Reinert (2000), and Savage and Strickman the distinct genera of Aedini, for example Psorophora (2004) and Hemagogus (Dyar 1928, Hendrickson and Sokal 1968), evolved from within Aedes and that Aedes was Aedes lineage Ochlerotatus lineage therefore paraphyletic. To avoid use of the resultant (Aedes) Meigen (Abraedes) Zavortink paraphyletic taxon Aedes, Reinert et al. (2004) felt (Aedimorphus) Theobald (Aztecaedes) Zavortink obligated to dismember Aedes by elevating all mono- (Alanstonea) Mattingly (Bruceharrisonius) Reinert phyletic groups at the corresponding level within the (Albuginosus) Reinert (Chaetocruiomyia) Theobald (Belkinius) Reinert (Finlaya) Theobald tree to genera (Kristensen 1982). For example, all taxa (Bothaella) Reinert (Geoskusea) Edwards represented by the unnamed lines in Fig. 1, which (Cancraedes) Edwards (Gymnometopa) Coquillett have common ancestors denoted by squares at ap- (Christophersiomyia) Barraud (Halaedes) Belkin (Diceromyia) Theobald (Howardina) Theobald proximately the same level as the common ancestors (Edwardsaedes) Belkin (Kenknightia) Reinert of Psorophora, Hemagogus, and Eretmapodites, also () Reinert (Kompia) Aitken would be elevated to generic status. If all of the mono- (Huaedes) Huang (Levua) Stone and Bohart phyletic groups in Fig. 1 can be deÞned by morpho- (Indusius) Edwards () Theobald (Isoaedes) Reinert (Molpemyia) Theobald logical characters that allow them to be identiÞed and (Leptosomatomyia) Theobald (Mucidus) Theobald recognized, this leads to the generation of numerous (Lorrainea) Belkin (Nothoskusea) Dumbleton new well-deÞned genera and would be considered (Neomelaniconion) Newstead (Ochlerotatus) Lynch Arribalzaga beneÞcial by many taxonomists. However, if the new (Paraedes) Edwards (Protomacleaya) Theobald (Pseudarmigeres) Stone & (Pseudoskusea) Theobald monophyletic groups are poorly deÞned and cannot Knight be recognized as morphological or biological groups, the (Scutomyia) Theobald (Rhinoskusea) Edwards destruction of well-established and deÞned paraphyletic () Theobald (Rusticoidus) Shevchenko and taxa into many poorly deÞned monophyletic taxa results Prudkina (Stegomyia) Theobald (Zavortinkius) Reinert in poor taxonomy and a useless classiÞcation (Michener 2000, Brummitt 2002, Diggs and Lipscomb 2002, Nordal Note subgeneric names are enclosed in parentheses. and Stedje 2005). Because most of these new genera are 926 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 42, no. 6 Downloaded from https://academic.oup.com/jme/article/42/6/923/886357 by guest on 29 September 2021

Fig. 1. Hypothetical phylogenetic tree. Black circles represent extant species; open circles and open squares represent postulated ancestor species. Tree is Ϸ3% of the size of the Aedini tree. Number of slash marks on basal lineages is indicative of degree of evolutionary change, either unusual or distinct morphological changes as noted by early taxonomists, or potential synapomorphies in the sense of cladistics. Heavy lines above extant species indicate limits of taxa, either the monophyletic genera Psorophora, Hemogogus, and Eretmapodites, or selected subgenera of Aedes from Table 3. poorly deÞned and difÞcult or impossible to identify, subgeneric names for fragments will signiÞcantly re- medical entomologists and other Þeldworkers would be duce confusion. If the monophyletic nature of sub- forced to identify a specimen Þrst to species by using a genera of Aedes becomes Þrmly established, then anal- broad Aedini key then look in an appendix or table to ysis of the evolutionary relationships among the determine what genus it was assigned to. ReinertÕs fear subgenera of Aedes and genera of Aedini will even- of paraphyletic taxa (Reinert et al. 2004), admitted on tually lead to a reliable phylogeny and stable classiÞ- page 327, “we stress that our study removes the evidently cation. As subgenera become well deÞned, taxono- paraphyletic ÔlargeÕ genera, with a necessary fragmenta- mists may choose to elevate selected subgenera to tion of traditional taxa and a subjective decision that the genera based on Þrm phylogenetic evidence and mor- fragments be treated as genera rather than subgenera,” phological characters that allow these new genera to results in the treatment of numerous poorly deÞned be reliably identiÞed. “fragments” as genera. A conservative and rational ap- It is unfortunate that Reinert et al. (2004) felt com- proach would have been to place these “fragments” as pelled to change the classiÞcation of Aedini to reßect subgenera within the paraphyletic genus Aedes as they the topology of their preferred tree based on study of were in the traditional classiÞcation of Edwards 1932, less that 10% (119/1,239) of the species in the tribe. If 1941 following the argument of Savage and Strickman these authors would have presented their preliminary (2004); see Table 3. The treatment of these fragments as results as the Þrst major attempt to understand the subgenera maintains all the information associated with phylogeny of the Aedini, their article would have been the subgeneric name, for example subgenus Ae. (Stego- hailed as an important contribution to mosquito sys- myia) and the genus Stegomyia, convey exactly the same tematics. These authors amassed a large morphologi- information, but use of the former subgeneric name cal data set and presented a framework for future avoids the need to amend the species names of included analyses. However, because these authors tried to species. The article by Reinert et al. (2004) was the Þrst translate the results of their cladistic analysis directly major cladistic analysis of the Aedini, and future studies into a reclassiÞcation of genera, their contribution using different sets of morphological characters or/and should be rejected. Other cladists, recognizing that molecular data will certainly produce different frag- more names are needed for clades within a tree than ments, and different relationships among the fragments are available in the Code, are developing an alterna- and well-deÞned monophyletic groups. If these frag- tive code of names, PhyloCode, for naming the many ments are continually treated as genera, the names of clades and recognizing hierarchy within phylogenetic common vector and nuisance species also will continu- trees (Cantino and de Queiroz 2004). These codes ally change. This will lead to disruption of communica- could be used simultaneously to discuss the evolution tion among those interested in mosquito-borne diseases, and biology of lineages or groups of species. However, the destruction of the information system associated only the most well-deÞned and hence stable lineages with a stable classiÞcation, and difÞculties in literature should be assigned the rank of genera. search and database management. Phylogenetic analysis is desperately needed within the family Culicidae. However, I am not hopeful that Acknowledgments the phylogeny, classiÞcation, and nomenclature of the I thank Richard K. Brummitt (The Herbarium, Royal Aedini will stabilize in the near future. The use of Botanic Gardens, Kew, Richmond, United ), Tom November 2005 SAVAGE:CLASSIFICATION OF MOSQUITOES IN TRIBE AEDINI 927

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