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An annotated checklist of the horse flies, deer flies, and yellow flies (Diptera: Tabanidae) of Florida.

The family Tabanidae includes horse flies, deer flies, and yellow flies, which are considered significant pests of livestock in the United States (Hansens 1979; Goodwin et al. 1985). Most females require a blood meal for egg production, although autogeny has been documented in several species (Rockel 1969; Anderson 1971; Burger & Lake 1980). In Florida, there is 1 known species, Asaphomyiafloridensis Pechuman, that does not feed on blood at all (Pechuman 1974), and the reduced mouthparts of the genus Merycomyia suggest that this genus also is not hematophagous although data on feeding habits and behavior is still lacking (Jones & Anthony 1964). Anautogenous tabanid females ingest blood by lacerating the skin with serrated mouthparts and lapping up the pooled blood, which can cause significant irritation to the host. Several commercial adult tabanid traps are available along with blueprints for homemade traps on the internet, though studies focusing on population reduction in the environment are lacking.

Eggs are laid in a variety of habitats, often on vegetation along the perimeter of permanent or temporary bodies of water (Jones & Anthony 1964). Most Tabanidae are thought of as having aquatic larval stages, with larvae inhabiting moist or saturated soils around lakes, streams, ponds, and even roadside ditches; however, Wilson (1969) collected several larval specimens of serious pest species from the soil and debris of mostly dry hardwood hammocks in an alluvial forest in Louisiana. The larval and pupal ecology of many species is still unknown, perhaps due to a sampling bias towards aquatic environments. Many larvae are predacious, feeding on macroinvertebrates in the environment. Larval and pupal descriptions, habitats, and life histories are still undocumented for several tabanid species. Life cycles can range from a few weeks to several years (Jones & Anthony 1964). Larval control methods have been attempted such as water impoundment and chemical controls with mediocre results (Anderson & Kneen 1969; Anderson 1985). Long-term control is difficult to achieve due to ecological patchiness of larval habitats, large population numbers, varying life histories, different seasonal distributions, and extensive life cycles.

Tabanids can easily become a major pest of man, especially salt marsh species that are known to readily feed on humans and often inhabit coastal tourist areas, golf courses, campgrounds, etc. (Hansens 1979). In extreme cases, tabanid infestations can cause a decrease in property values (Gerhardt et al. 1973). Tabanids are known to be mechanical vectors of several animal diseases such as equine infectious anemia virus, bovine leukemia virus, hog cholera virus, anaplasmosis, anthrax, tularemia, and several other serious diseases of veterinary concern (Krinsky 1976; Foil 1989). Perhaps even more significant than livestock pathogen transmission are the economic losses farmers experience during large tabanid infestations. Laceration of the skin by feeding females causes significant irritation to the host. Many livestock hosts respond with an attempt to dislodge the flies, but tabanids are persistent biters and will relocate until fully engorged (Foil 1983). During heavy infestations, livestock may decrease grazing and lose body weight due to the amount of time spent trying to dislodge the flies (Perich et al. 1986), which can also decrease milk production (Hansens 1979). Cattle that have been fed on by 66 to 90 horse flies per day may suffer from decreased feeding efficiency by up to 16.9% (Perich et al. 1986). Further studies on economic and veterinary impacts of tabanid feeding are still needed. In order to proceed with veterinary, ecological, and economic studies of the Tabanidae in Florida, we have developed a taxonomically organized and up-to-date record of species for the state. Currently, we have identified 124 species/subspecies documented from Florida, with an additional 15 possible species, representing 15 genera. In comparison, 138 species have been recorded from Georgia at the time of Burger's (1995) catalog. Sampling in Alabama has not extensively been undertaken, but it is likely that species numbers are similar to neighboring Georgia and Florida.

Materials and Methods

The last complete compilation of the Tabanidae of Florida was conducted in 1964 by Calvin Jones and Darrell Anthony (Jones & Anthony 1964). Since then, several taxonomic and distributional changes have been made by various authors. The included annotations following species names are those records that have been published since Jones and Anthony's 1964 publication or personal notes. If no annotations are present after a species name, no records have been updated since 1964. For annotations prior to 1964, see Jones & Anthony (1964) and Bargren (1961).

Nomenclature for our list follows that of Burger (1995). Burger (1995) noted that many species had designated variations or subspecies based solely on color. After studying several of these species, he determined that southern specimens often exhibited a melanistic variation (with intermediates) of the color form fairly consistently, and therefore determined that these are not varieties or subspecies but simply melanistic forms of the corresponding northern species.

Following Fairchild & French (1999), we have included 15 species (in parentheses) that have distribution records in surrounding states but no official records from Florida. It is possible, based on habitats and distribution, that these species are present in Florida though not yet documented. We removed Tabanus catenatus Walker, 1848 from that list due to the possible misidentification of the Georgia specimens. Burger (1995) lists the southern distribution of T. catenatus to only North Carolina; therefore it is not included here. To date, Florida has 124 species/subspecies of Tabanidae, with an additional 15 possible species representing 15 genera lacking a current Florida record.

Additionally, we have not included any misidentifications in the synonymy of species names unless they directly relate to Florida records. To see synonymous misidentifications, see Burger (1995). Subspecies status was assigned following Burger's (1995) catalog. A notation of "FSCA" after the Florida record indicates CMZN personally examined specimens housed at the Florida State Collection of Arthropods (FSCA) at the Department of Plant Industry in Gainesville, Florida. Notations on several species were compiled from literature and collection records, though much remains unknown, especially regarding larval ecology. In many cases, observations are based on 1 or few specimens or records. The FSCA houses many personal Tabanidae collections, and a thorough examination of individual specimens may yield extended flight times and distributions for the state. The majority of Florida species have a distribution of "North" or "North and Central" Florida. It is true that North Florida represents the southernmost range extension for many species; however, sampling has historically been biased towards these locations. Very little sampling has been done in Central and South Florida, wherefore a species identification should never be ruled out simply based on our locality data. It is also possible that there may be unidentified or undocumented species from that region, including Caribbean species that have established populations in South Florida.

A new state record is presented for Tabanus reinwardtii Wiedemann, 1828. Four specimens were caught in Bay County, Florida, in 1999 in a French 2-tier box trap by James Cilek and Mary Ann Olson of the Public Health Entomology Research and Education Center, Florida A&M University, Panama City, Florida. Identifications were confirmed by John Burger. Previously, T. reinwardtii was known only from Canada, south to Georgia and Louisiana, and west to Colorado (Burger 1995). A new state record is presented for Chlorotabanus mexicanus (L., 1758), previously recorded from Panama to Mexico. Three females and 1 male were caught and identified on Big Pine Key, Florida, by Gunter Muller of the Institute for Medical Research Israel-Canada, Hebrew University, Jerusalem, Israel, in 1999. Finally, a new state record also is presented for Tabanus yucatanus Townsend, 1897, previously documented from the Yucatan Peninsula. Two specimens were taken in a malaise trap on Cedar Key, Florida, by Muller and Revay, of the Institute for Medical Research Israel-Canada, Hebrew University, Jerusalem, Israel.

Results

Supplementary material for this article is online in Florida Entomologist 98(2) (June 2015) at http://purl.fcla.edu/fcla/entomologist/ browse. Each supplementary table and figure is referred to herein as either Suppl. Table or as Suppl. Fig.

Annotated checklist of Florida Tabanidae

Agkistrocerus finitimus (Stone), 1938 Florida Record: Stone 1938 (as Dicladocera finitima), FSCA Synonym: Dicladocera finitimus Stone, 1938

Agkistrocerus megerlei (Wiedemann), 1828 Florida Record: Osten Sacken 1878 (as T. megerlei), FSCA Synonymy: Tabanus megerlei Wiedemann, 1828, Dicladocera megerlei (Wiedemann)

Anacimas limbellatus Enderlein, 1923 Florida Record: Stone 1938 (as A. geropogon), FSCA Synonymy: Anacimas geropogon Philip, 1936 Fairchild 1978. Notes on classification.

Asaphomyia floridensis Pechuman, 1974 Florida Record: Pechuman 1974, FSCA Pechuman 1974. New species. Note: Listed as a "Species of Concern". Reviewed for threatened or endangered status by the Florida Fish and Wildlife Commission in 1994.

Chlorotabanus crepuscularis (Bequaert), 1926 (Suppl. Fig. 7) Florida Record: Osten Sacken 1876 (as T mexicanus), FSCA Synonymy: Chlorotabanus flavus Macquart, 1834; Chlorotabanus mexicanus, authors, not Linnaeus; Chlorotabanus sulphureus, authors, not Palisot de Beauvois

Chlorotabanus mexicanus (Linnaeus, 1758)

NEW RECORD

Florida Record: G. Muller & E. Revay 1998 (unpubl.). Three females and 1 male were collected in a malaise trap on the Big Pine Key in Apr 1999. Previously, this species is known from northern South America, throughout Mexico, and in the Caribbean.

The specimens are deposited in Col. G.C. Muller and in the Florida State Collection of Arthropods, Department of Plant Industry, Gainesville, Florida.

Chrysops abatus Philip, 1941 Florida Record: Philip 1941 Note: Species pair with C. dorsovittatus (Fairchild 1978).

Chrysops amazon Daecke, 1905 Florida Record: Philip 1955, FSCA Synonymy: Chrysops amazon hubbelli Philip 1955 Burger 1995. Notes on classification.

Chrysops atlanticus Pechuman, 1949 (Suppl. Table 2) Florida Record: Walker 1848 (as C. canifrons), FSCA Synonymy: Chrysops canifrons Walker, 1848 Note: Member of the C. flavidus species group (Baier 1999).

Chrysops beameri Brennan, 1935 Florida Record: Fairchild 1937 (misidentified as C. hinei)

Chrysops bistellatus Daecke, 1905 Florida Record: Fairchild 1937

Chrysops brimleyi Hine, 1904 Florida Record: Fairchild 1937, FSCA

Chrysops brunneus Hine, 1903 Florida Record: Brennan 1935, FSCA Synonymy: Chrysops brunnea (Hine 1903) Note: Member of the C. flavidus species group (Baier 1999).

Chrysops callidus Osten Sacken, 1875 (Suppl. Table 2) Florida Record: Johnson and Coquillett 1895 Synonymy: Chrysops callidulus Philip, 1941

Chrysops calvus (Pechuman & Teskey, 1967) Florida Record: Pechuman & Teskey 1967, FSCA Pechuman & Teskey 1967. New species. Note: Species pair with Chrysops niger (Drees et al. 1980).

Chrysops carbonarius Walker, 1848 Florida Record: Philip 1955 (as C. carbonaria) Synonymy: Chrysops provocans Walker, 1850

Chrysops celatus Pechuman, 1949 Florida Record: Tidwell 1973, FSCA Synonymy: Chrysops flavidus celata Pechuman, 1949 Note: Member of the C. flavidus species group (Baier 1999).

Chrysops cincticornis Walker, 1848 (Suppl. Table 2) Florida Record: Fairchild 1937, FSCA Synonymy: Chrysops celer Osten Sacken, 1875

Chrysops cincticornis nigropterus Fairchild, 1937 Florida Record: Fairchild 1937, FSCA Synonymy: Chrysops celer var. nigroptera var. nov. Fairchild, 1937

Chrysops cursim Whitney, 1879 (Suppl. Fig. 1) Florida Record: Brennan, 1935, FSCA

Chrysops dacne Philip, 1955 Florida Record: Philip 1955

Note: Species pair with C. parvulus as noted in collections in the Florida State Collection of Arthropods, Department of Plant Industry, Gainesville, Florida.

Chrysops dimmocki Hine, 1905 Florida Record: Philip, 1947 (mentioned as part of the C. pudicus species group), FSCA

Chrysops divisus Walker, 1848 Florida Record: Walker 1848 Synonymy: Chrysops atropos Osten Sacken, 1875

Chrysops dixianus Pechuman, 1974 Florida Record: Pechuman 1974, FSCA Pechuman 1974. New species. Note: Member of the C. flavidus species group (Baier 1999).

Chrysops dorsopunctus Fairchild, 1937 Florida Record: Fairchild 1937 (as C. fulvistigma var. dorsopuncta var. nov.) Synonymy: Chrysops fulvistigmus dorsopunctus Fairchild, 1937

Chrysops dorsovittatus Hine, 1907 Florida Record: Hine 1907 Note: Species pair with C. abatus (Fairchild 1978).

Chrysops flavidus Wiedemann, 1821 (Suppl. Table 2; Suppl. Fig. 2) Florida Record: Johnson & Coquillett 1895, FSCA Synonymy: Chrysops pallida Macquart, 1838; Chrysops pallidus Bellardi, 1859

Note: The C. flavidus species group contains: C. atlanticus, C. brunneus, C. dixianus, C. flavidus, C. reicherti, (C. sandyi), C. tumidicornis, and C. celatus (Baier 1999).

Chrysops floridanus Johnson, 1913 Florida Record: Johnson 1913 (as C. vittatus var. floridanus), FSCA Synonymy: Chrysops vittatus floridanus Johnson, 1913 Burger, 1995. Note on classification.

Note: Species pair with C. vittatus. Chrysops vittatus is thought to be an inland species and C. floridanus is thought to be a coastal species (Burger 1995).

Chrysops fuliginosus Wiedemann, 1821 (Suppl. Fig. 3) Florida Record: Walker, 1848, FSCA Synonymy: Chrysops plangens Wiedemann, 1828; Chrysops confuses Harris, 1835

Note: There is a larger inland variation of C. fuliginosus with distinct yellow abdominal markings. Ecology of the inland variation is unknown.

Chrysops fulvistigma Hine, 1904 Florida Record: Philip 1950 Chrysops geminatus Wiedemann, 1828 Florida Record: Fairchild 1937

Chrysops hinei Daecke, 1907 (Suppl. Fig. 4) Florida Record: Johnson 1913

Chrysops hyalinus Shannon, 1924 Florida Record: Philip & Jones 1962 Synonymy: Chrysops vitripennis Shannon, 1916; Chrysops claripennis Krober, 1926

Chrysops ifasi Fairchild, 1978 Florida Record: Fairchild 1978 Fairchild, 1978. New species.

Chrysops macquarti Philip, 1961 Florida Record: Philip 1961 Synonymy: Chrysops univittatus, authors, not Macquart Philip 1961. Note on classification, name change.

Chrysops moechus Osten Sacken, 1875 Florida Record: Bargren 1961

Chrysops montanus Osten Sacken, 1875 (Suppl. Table 2) Florida Record: Fairchild 1937 Synonymy: Chrysops montanus perplexa Philip, 1955 Burger 1995. Note on classification.

Chrysops niger Macquart, 1838 Florida Record: Philip 1950 (as C. nigra) Synonymy: Chrysops nigra taylori Philip, 1955 Burger 1995. Note on classification.

Note: Species pair with Chrysops calvus (Drees et al. 1980).

Chrysops nigribimbo Whitney, 1879 Florida Record: Philip 1947, FSCA

Chrysops obsoletus Wiedemann, 1821 (Suppl. Table 2) Florida Record: Osten Sacken 1875 (as C. morosus), FSCA Synonymy: Chrysops obsoletus lugens Wiedemann, 1821; Chrysops trinotatus Macquart, 1838; Chrysops morosus Osten Sacken, 1875; Chrysops ultimus Whitney, 1914

Chrysops parvulus Daecke, 1907 Florida Record: Johnson 1913

Note: Species pair with C. dacne as noted in collections in the Florida State Collection of Arthropods, Department of Plant Industry, Gainesville, Florida.

Chrysops pikei Whitney, 1904 Florida Record: Fairchild 1937 Synonymy: Chrysops harmani Tidwell, 1973

Chrysops pudicus Osten Sacken, 1875 (Suppl. Table 1; Suppl. Fig. 5) Florida Record: Osten Sacken 1875, FSCA

Note: Member of the C. flavidus species group (Baier 1999).

Chrysops reicherti Fairchild, 1937 Florida Record: Fairchild 1937, FSCA Synonymy: Chrysops flavidus reicherti Fairchild, 1937

Note: Member of the C. flavidus species group (Baier 1999).

(Chrysops sandyi Baier, 1999) Baier 1999. New species.

Chrysops tidwelli Philip & Jones, 1962 Florida Record: Philip & Jones 1962 Philip & Jones 1962. New species.

Chrysops tumidicornis Baier, 1999 Florida Record: Baier 1999, FSCA Baier 1999. New species.

Note: Member of the C. flavidus species group (Baier 1999).

Chrysops univittatus Macquart, 1855 Florida Record: Johnson & Coquillett 1895 Synonymy: Chrysops fraternus Krober, 1926; Chrysops wiede manni Krober, 1926

Chrysops upsilon Philip, 1950 Florida Record: Fairchild 1978 Fairchild 1978. New record.

Chrysops vittatus Wiedemann, 1821 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 6) Florida Record: Johnson & Coquillett 1895, FSCA Synonymy: Chrysops areolatus Walker, 1848; Chrysops lineatus Jaennicke 1867; Chrysops vittatus floridana Johnson, 1913; Chrysops ornatus Krober, 1926

Diachlorus ferrugatus (Fabricius), 1805 (Suppl. Table 1; Suppl. Fig. 8) Florida Record: Walker 1848 (as Chrysops approximans), FSCA

Synonymy: Diachlorus americanus Palisot de Beauvois, 1819; Diachlorus ataenia Macquart, 1838; Diachlorus approximans Walker, 1848

Haematopota punctulata (Macquart), 1838 Florida Record: Johnson & Coquillett 1895 (as H. americana; corrected by Johnson 1912), FSCA Synonymy: Chrysozona punctulata (Macquart), 1838

Hamatabanus annularis (Hine), 1917 Florida Record: Stone 1935 (as Dicladocera sexfasciata Stone), FSCA Synonymy: Hamatabanus sexfasciatus (Stone), 1935 Burger 1995. Note on classification.

Hamatabanus carolinensis (Macquart), 1838 Florida Record: Philip 1947 (as H. scitus), FSCA Synonymy: Hamatabanus scitus (Walker), 1848; Tabanus hirtioculatus Macquart, 1855; Tabanus cerastes Osten Sacken, 1876; Tabanus fraterna Krober, 1931; Tabanus frater Krober, 1934

Hamatabanus exilipalpis (Stone), 1938 Florida Record: Stone 1938 (as T. exilipalpis), FSCA

Hamatabanus floridensis (Hine), 1912 Florida Record: Hine 1912 (as T. floridensis), FSCA Synonymy: Stenotabanus floridensis (Hine), 1912; Tabanus floridensis Hine

Burger 1995. Note on classification. Hybomitra cincta (Fabricius), 1794 Florida Record: Jones & Anthony 1964, FSCA Synonymy: Tabanus cincta (F.), 1794

Hybomitra difficilis (Wiedemann), 1828 Florida Record: Philip 1961, FSCA Synonymy: Tabanus carolinensis, authors, not Macquart; Tabanus difficilis (Wiedemann), 1828

Hybomitra hinei (Johnson), 1904 Florida Record: Stone 1938 (as T. hinei), FSCA Synonymy: Tabanus politus Johnson, 1900; Tabanus hinei (Johnson), 1904; Hybomitra hinei wrighti (Whitney), 1915 Burger 1995. Note on classification.

Hybomitra trispila (Wiedemann), 1828 (Suppl. Table 2) Florida Record: Goodwin 1976, FSCA Synonymy: Tabanus trispila (Wiedemann), 1828 Goodwin, 1976. New record.

Leucotabanus annulatus (Say), 1823 Florida Record: Fairchild 1937 (as T. annulatus), FSCA Synonymy: Tabanus annulatus (Say), 1823; Tabanus argenteus Wiedemann in Philip, 1950

Merycomyia microcera (Walker), 1848 Florida Record: Stone 1953 (as M. brunnea), FSCA Synonymy; Merycomyia brunnea Stone, 1953

Note: Listed as a "Species of Concern". Reviewed for threatened or endangered status by the Florida Fish and Wildlife Commission in 1994.

Merycomyia whitneyi (Johnson), 1904 Florida Record: Philip 1954, FSCA Synonymy; Merycomyia geminata Hine, 1912; Merycomyia mixta Hine, 1912

Microtabanus pygmaeus (Williston), 1885 Florida Record: Williston 1885 (as T. pygmaeus), FSCA Synonymy: Tabanus pygmaeus (Williston), 1885; Atylotus pygmaeus (Williston)

Stenotabanus (Aegialomyia) magnicallus (Stone) 1935 Florida Record: Tidwell 1973, FSCA Synonymy: Tabanus magnicallus Stone, 1935; Tabanus maritimus Townsend, 1898; Tabanus nanus Macquart, 1846

Stenotabanus (Aegialomyia) psammophilus (Osten Sacken) 1876 Florida Record: Osten Sacken 1875 (as T. psammophilus), FSCA Synonymy: Tabanus psammophilus Osten Sacken, 1876

Tabanus aar Philip, 1941 Florida Record: Philip 1941, FSCA

Tabanus abdominalis Fabricius, 1805 Florida Record: Walker 1848, FSCA

Tabanus acutus (Bigot), 1892 Florida Record: Fairchild 1937, FSCA Synonymy: Atylotus acutus Bigot, 1892

Tabanus americanus Forster, 1771 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 9) Florida Record: Walker 1848 (as T ruficornis), FSCA Synonymy: Tabanus plumbeus Drury, 1773; Tabanus ruficornis Fabrucius, 1775; Tabanus limbatus Palisot De Beauvois 1806

Tabanus aranti Hays, 1961 Florida Record: Philip & Jones 1962; Recorded by Fairchild (1937) as T. nigrescens atripennis, though not reassigned to T. aranti until 1980 (Fairchild 1980), FSCA Goodwin 1976. New record.

Tabanus atratus Fabricius, 1775 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 10) Florida Record: Osten Sacken 1875, FSCA

Synonymy: Tabanus americanus Drury, 1773; Tabanusniger Palisot De Beauvois, 1806; Tabanus validus Wiedemann, 1828; Tabanus atratus nantuckensis Hine, 1917; Tabanus atratusfulvopilosis Johnson, 1919 Burger 1995. Note on classification

Tabanus birdiei Whitney, 1914 Florida Record: Whitney 1914, FSCA

Tabanus bishoppi Stone, 1933 Florida Record: Stone 1935, FSCA

(Tabanus calens Linnaeus, 1767) Florida Record: Williston 1885 (as T. giganteus)

Synonymy: Tabanus giganteus De Geer, 1776; Tabanus lineatus Fabricius, 1781; Tabanus pallidus Palisot De Beauvois, 1809; Tabanus bicolor Macquart, 1847; Tabanus coesiofasciatus Macquart, 1855

Note: Williston (1885) and Johnson & Coquillett (1895) reported this species from an unknown location in Florida as T. giganteus. Stone (1938) also reported this species from Florida and Fairchild (1950) lists Florida within the range, but T. calens has not been recorded, because Burger (1995) does not consider Florida within the distribution range and there is much debate about the identification of the original T. giganteus specimens (Philip 1952b).

Tabanus cayensis Fairchild, 1935 Florida Record: Fairchild 1935, FSCA

Tabanus cheliopterus Rondani, 1850 Florida Record: Philip 1936 (as T. subfronto) Synonymy: Tabanus cheliopterus subfronto Philip, 1936 Burger 1995. Note on classification.

Tabanus coarctatus Stone, 1935 Florida Record: Stone 1935, FSCA

Tabanus colon Thunberg, 1827 Florida Record: Fairchild 1937, FSCA Synonymy: Tabanus nigrescens atripennis Stone, 1935

Note: Both names are still used in the literature, but refer to the same species.

Tabanus conterminus Walker, 1850 Florida Record: Johnson 1913, FSCA

Note: Fairchild & French (1999) note that T. nigrovittatus and T. conterminus will key out the same. In Florida, there likely is 1 or more undescribed species in this complex (Sutton & Carlson 1997) and one can realistically expect only to identify specimens to the T. nigrovittatus complex, not to species (B. D. Sutton, Florida Department of Agriculture and Consumer Services, Department of Plant Industry, unpublished data.)

(Tabanus cymatophorus Osten Sacken, 1876) Florida Record: Johnson 1913

Note: Johnson (1913) notes specimens taken in Biscayne Bay and West Palm Beach, Florida. Muller caught 2 specimens near St. Augustine, Florida in the late 1990s (unpublished data). Burger (1995) lists the southern distribution of T. cymatophorus as Georgia.

Tabanus daedalus (Stone), 1938 Florida Record: Stone 1938 (as S. daedalus) Synonymy: Stenotabanus daedalus Stone, 1938

Tabanus endymion Osten Sacken, 1878 Florida Record: Fairchild 1937

(Tabanus equalis Hine, 1923) (Suppl. Table 2) Synonymy: Tabanus uniformis Hine, 1917

Tabanus fairchildi Stone, 1938 Florida Record: Stone 1938 Synonymy: Tabanus vivax, authors, not Osten Sacken

Tabanus fronto Osten Sacken, 1876 Florida Record: Williston 1885 Synonymy: T. cheliopterus var. fronto Philip, 1936 Burger 1995. Note on classification.

Tabanus fulvilineis Philip, 1957 Florida Record: Philip 1957 (as T. nigrovittatus fulvilineis), FSCA Synonymy: Tabanus nigrovittatus fulvilineis Philip, 1957 Burger 1995. Note on classification.

Note: Member of the Tabanus nigrovittatus complex. See note under T. nigrovittatus.

Tabanus fulvulus Wiedemann, 1828 Florida Record: Johnson 1913, FSCA Synonymy: Tabanus fulvofrater Walker, 1848; Tabanus mutates Walker, 1850

Tabanus fumipennis Wiedemann, 1828 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 11) Florida Record: Osten Sacken 1875 (as T. rufus), FSCA Synonymy: Tabanus rufus Palisot De Beauvois, 1809; Tabanus flammans Walker, 1848; Tabanus formosus Walker, 1848

(Tabanusfuscicostatus Hine, 1906) (Suppl. Table 2) Florida Record: Pechuman 1949, FSCA

Note: The presence of T. fuscicostatus in Florida is still unclear. Several Florida specimens exist at the FSCA labeled T. fuscicostatus; however, a note left by Sandy Fairchild sugg ests that based on the dark femurs and dark costal cells, these were misidentifications and T. fuscicostatus does not have a Florida distribution.

Tabanus fusconervosus Macquart, 1838 Florida Record: Walker 1848 (2 records by Walker, as T. recedens and T fusco-nervosus) Synonymy: Tabanus confusus Walker, 1848; Tabanus recedens Walker, 1848; Tabanus fur Williston, 1885, Tabanus turbidus Wiedemann, 1828

Tabanus fuscopunctatus Macquart, 1850 Florida Record: Osten Sacken 1876, FSCA

Tabanus fuscopunctatus pechumani Philip, 1960 Florida Record: Philip 1960 (as T. imitans pechumani), FSCA Synonymy: Tabanus imitans pechumani Philip, 1960 Burger 1995. Note on classification.

Tabanus gladiator Stone, 1935 Florida Record: Fairchild 1937, FSCA Tabanus gracilis Wiedemann, 1828 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 12) Florida Record: Williston 1885, FSCA

Tabanus hinellus Philip, 1960 (Suppl. Table 1; Suppl. Fig. 13) Florida Record: Fairchild 1983, FSCA Synonymy: Tabanus lineola hinellus Philip, 1960

Tabanus imitans Walker, 1848 Florida Record: Fairchild 1937, FSCA

Tabanus johnsoni Hine, 1907 (Suppl. Table 1; Suppl. Fig. 14) Florida Record: Hine 1907, FSCA

Tabanus kisliuki Stone, 1935 Florida Record: Philip 1950 (as T. quirinus) Synonymy: Tabanus quirinus Philip, 1950

Tabanus lineola complex Fabricius, 1794 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 15) Florida Record: Walker, 1848, FSCA

Note: The T. lineola complex has extensive morphological variation and further work needs to be done on this group. Several variations and subspecies have been proposed, though the taxonomy is still cloudy. This species is thought to be the inland variation of T. hinellus.

(Tabanus limbatinevris Macquart, 1847) Tabanus longiusculus Hine, 1907

Florida Record: Fairchild 1937, FSCA Tabanus maculipennis Wiedemann, 1928 Florida Record: Stone 1938 (as T. imitans var. excessus), FSCA Synonymy: Tabanus imitans var. excessus Stone, 1938

Tabanus melanocerus Wiedemann, 1828 Florida Record: Osten Sacken 1875, FSCA Synonymy: Tabanus lacustris Stone, 1935; Tabanus melanocerus var. lacustris Stone, 1935 Burger 1995. Note on classification.

Note: Burger (1995) places T. melanocerus and T. melanocerus lacustris as a synonymous group under the name T. melanocerus. After studying several specimens, it should be noted there is a distinct spur on vein R4 in the former T. melanocerus lacustris that is clearly absent in T. melanocerus. Following Fairchild & French (1999), we are labeling lacustris as a variation of T. melanocerus.

Tabanus mixis Philip, 1950 Florida Record: Philip 1950b, FSCA Synonymy: Tabanus molestus mixis Philip, 1950

Tabanus moderator Stone, 1938 Florida Record: Fairchild & French 1999, FSCA

Tabanus molestus Say, 1823 Florida Record: Johnson & Coquillett 1895, FSCA Synonymy: Atylotus tennessensis Bigot, 1892 Burger 1995. Note on classification.

Tabanus mularis Stone, 1935 (Suppl. Table 2) Florida Record: Stone 1935, FSCA

Note: Member of the Tabanus nigrovittatus complex. See note under T nigrovittatus.

(Tabanus nefarius Hine, 1907)

(Tabanus nigrescens Palisot de Beauvois, 1809) Florida Record: Stone 1938. No records since.

Tabanus nigrescens atripennis Stone, 1935 See Tabanus colon Thunberg, 1827

Tabanus nigripes Wiedemann, 1821 (Suppl. Table 1; Suppl. Fig. 16) Florida Record: Osten Sacken 1875 (as T. coffeatus), FSCA Synonymy: Tabanus coffeatus Macquart, 1847; Tabanus winthemi Krober, 1931

Tabanus nigrovittatus Macquart, 1847 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 17) Florida Record: Johnson & Coquillett 1895, FSCA Synonymy: Tabanus simulans Walker, 1848; Tabanus vicarious Walker, 1848 ex parte

Note: Tabanus nigrovittatus belongs to the T. nigrovittatus complex which contains several lined species. Fairchild & French (1999) note that T. nigrovittatus and T. conteminus will key out the same. Jones & Anthony (1964) list T. quinquevittatus as an inland Florida species, but note that some previous T. quinquevittatus specimens were misidentified and were actually T. nigrovittatus. Burger (1995) and Fairchild & French (1999) do not consider T. quinquevittatus sensu stricto to have a Florida distribution. In Florida, there likely is 1 or more undescribed species in this complex (Sutton & Carlson 1997) and one can realistically expect to identify specimens only to the T. nigrovittatus complex, not to species (B. D. Sutton, Florida Department of Agriculture and Consumer Services, Department of Plant Industry).

Tabanuspallidescens Philip, 1936 (Suppl. Table 1; Suppl. Table 2; Suppl. Fig. 18) Florida Record: Stone 1938 (as T fulvulus var. pallidescens), FSCA

Tabanus petiolatus Hine, 1917 (Suppl. Table 1; Suppl. Fig. 19) Florida Record: Fairchild 1937, FSCA Synonymy: Tabanus yulensus Philip, 1950

Tabanus proximus Walker, 1848 Florida Record: Walker 1848 Synonymy: Tabanus benedictus Whitney, 1904

Tabanus pumilus Macquart, 1838 (Suppl. Table 1; Suppl. Fig. 20) Florida Record: Osten Sacken 1875, FSCA

Tabanus quinquevittatus Wiedemann, 1821 (Suppl. Table 1; Suppl. Fig. 21) Florida Record: Osten Sacken 1875 (as T. costalis), FSCA

Synonymy: Tabanus baitimorensis Macquart, 1855; Tabanus costalis Wiedemann, 1828; Tabanus manifestus Walker, 1850; Tabanus vicarious Walker, 1848

Note: Member of the T nigrovittatus complex. Jones & Anthony (1964) and Bargren (1961) list T. quinquevittatus as an inland Florida species, but Jones & Anthony (1964) note that some previous T. quinquevittatus specimens were misidentified and were actually T. nigrovittatus. Burger (1995) and Fairchild & French (1999) do not consider T. quinquevittatus sensu stricto to have a Florida distribution. In Florida, there likely is 1 or more undescribed species in this complex (Sutton & Carlson 1997) and one can realistically expect to identify specimens only to the T. nigrovittatus complex, not to species (B. D. Sutton, Florida Department of Agriculture and Consumer Services, Department of Plant Industry).

Tabanus reinwardtii Wiedemann, 1828

NEW RECORD

Florida Record: Cilek & Olson 1999 (unpublished). Four specimens were collected in a 2-tier box trap in Bay County, Florida, in 1999 by Jim Cilek and Mary Ann Olson. Specimens confirmed by J. F. Burger. Synonymy: Tabanus erythrotelus Walker, 1850

Tabanus rufofrater Walker, 1850

Florida Record: Osten Sacken 1875 (as T. tener), FSCA Synonymy: Tabanus unicolor Macquart, 1847; Tabanus lateritius Rondani, 1863; Tabanus tener Osten Sacken, 1876

(Tabanus sackeni Fairchild, 1934)

(Tabanus sagax Osten Sacken, 1876) Synonymy: Atylotus baal Townsend, 1895; Tabanus dawsoni Philip, 1931

Tabanus sparus sparus Whitney, 1879 Florida Record: Johnson & Coquillett 1895, FSCA

Tabanus sparus milleri Whitney, 1914 (Suppl. Table 1; Suppl. Fig. 22) Florida Record: Whitney 1914 (as T. milleri), FSCA Synonymy: Tabanus sparus var. milleri Whitney, 1914

Tabanus stygius Say, 1823 Florida Record: Walker 1848

(Tabanus sublongus Stone, 1938)

Tabanus subsimilis Bellardi, 1859

Florida Record: Philip 1952a (as T. vittigerschwarti); originally recorded by Philip (1941), but Fairchild (1978) states it was most likely a misidentification. FSCA

Synonymy: Tabanus nippontucki Philip, 1942; Tabanus vittiger schwardti Philip, 1943

Tabanus sulcifrons Macquart, 1855 (Suppl. Table 2) Florida Record: Johnson 1913, FSCA Synonymy; Tabanus variegatus Fabricius, 1805; Tabanus tectus Osten Sacken, 1876; Tabanus exul Osten Sacken, 1878

(Tabanus superjumentarius Whitney, 1879) (Suppl. Table 2)

(Tabanus texanus Hine, 1907) Tabanus trijunctus Walker, 1854 (Suppl. Table 1; Suppl. Fig. 23) Florida Record: Osten Sacken 1858

Tabanus trimaculatus Palisot De Beauvois, 1806 Florida Record: Fairchild 1937, FSCA

Synonymy: Tabanus quinquelineatus Macquart, 1834; Tabanus apicalis Walker, 1848

Tabanus turbidus Wiedemann, 1828 Florida Record: Johnson & Coquillett 1895, FSCA

(Tabanus venustus Osten Sacken, 1876) (Suppl. Table 2)

Tabanus vittiger guatemalanus Hine, 1906 Florida Record: Fairchild 1978 (Florida forms of T. vittiger mentioned by Philip, 1957)

Synonymy: Tabanus bellardi Szilady, 1926; Tabanus appendiculatus Bequaert, 1940 (2 only); Tabanus carneus Bequaert, 1940 (2 only); Tabanus caymanicus Fairchild. 1942; Tabanus angustivitta Bequaert and Renjifo-Salcedo, 1946

Tabanus wiedemanni Osten Sacken, 1876 Florida Record: Osten Sacken 1875-76 Synonymy: Tabanus ater Palisot de Beauvois of Wiedemann, 1828 (Tabanus wilsoni Pechuman, 1962)

Tabanus yucatanus Townsend, 1897

NEW RECORD

Florida Record: Muller & Revay 2003 (unpublished). Two specimens were collected in a malaise trap in Cedar Key, Florida.

Note: Previously documented from the Yucatan Peninsula. The 2 Florida specimens are deposited in Col. G.C. Muller and in the Florida State Collection of Arthropods, Department of Plant Industry, Gainesville, Florida.

Tabanus zythicolor Philip, 1936 (Suppl. Table 1; Suppl. Fig. 24) Florida Record: Stone 1938

Whitneyomyia beatifica (Whitney), 1914 Florida Record: Whitney 1914 (as Tabanus beatificus), FSCA

Synonymy: Snowiellus styguis Enderlein, 1925; Tabanus ater Pallisot De Beauvois, 1811; Tabanus beatifica Whitney, 1914; Tabanus lugubris Macquart, 1838

Whitneyomyia beatifica var. atricorpus Philip, 1950 Florida Record: Philip 1950a, FSCA Synonymy: Whitneyomyia atricorpus Philip, 1950

Discussion

PEST SPECIES

We have listed 24 tabanid pest species of Florida (Suppl. Table 1) with their respective locations, habitats, biting preferences, and additional notes. Data were compiled through literature and collection records; however, the majority of collections throughout the years have been based in North or Central Florida. Many previous Florida tabanid researchers were based out of North Florida universities and other in stitutions, and therefore sampling in North Florida was more feasible and cost effective. Very few collection records exist for South Florida, and therefore species data may be biased towards North/Central Florida. In the case of C. pudicus, C. flavidus, C. vittatus, and T. nigrovittatus, we have only included representatives of the species groups. Different species within the species groups may be dominant in different regions of the state or the USA.

DISEASE TRANSMISSION

Tabanids are competent disease vectors in many parts of the world. In Africa, Chrysops species are the primary vectors of Loa loa (Cobbold, 1864) (Spirurida: Onchocercidae) filariasis, and several tabanid species have been implicated in the mechanical transmission of African trypanosomiasis, commonly known as nagana (Krinsky 1976; Foil 1989). In North America, tabanids are considered minor disease vectors but have the potential to become a serious problem if conditions are right. Tularemia, after L. loa, is the second principal human disease vectored by tabanids (Minter 2009). In western states, tularemia is often referred to as "deer fly fever." However, it is also spread through other means such as ticks and infected animal tissue, specifically rodents and rabbits (Jellison & Parker 1945; Hopla 1960; Cooney & Burgdorfer 1974). Interestingly, no cases of "deer fly fever" are reported from eastern states of the USA with a high incidence of tularemia, suggesting the principle tabanid vector is a western species, likely Chrysops discalis (Jellison 1950). Tularemia is a bacterial infection that causes flu-like symptoms in humans, often with the presence of a skin ulcer. In animals, infection can lead to death if untreated. Tularemia does not present with skin ulcers in animals, and is difficult to culture in the laboratory unless infection is suspected (Kahn 2005), wherefore detection in animals is difficult, but tularemia is not a major threat to the livestock industry in the USA. Although the suspected vector species are not present in Florida, Florida tabanids may be capable of disseminating the disease.

Anaplasmosis is a rickettsial infection that is common in livestock. In the USA, anaplasmosis is caused by Anaplasma marginale and mechanical transmission is the primary route of dissemination via flies (Foil 1989). Several species inhabiting or potentially inhabiting Florida have been found to be viable rickettsial vectors including T sulcifrons, T. lineola, (T. fuscicostatus), T. mularis, and T. pallidescens (Lotze & Yiengist 1941; Hawkins et al. 1982). Symptoms in hoofstock include anemia, haematuria, diarrhea, and anorexia. If detected, anaplasmosis can be treated with antibiotics.

The 3rd bacterial disease of concern to livestock owners is infection with Bacillus anthracis, commonly called anthrax. Anthrax spores are incredibly resilient and can persist dormant in the environment for years. Cutaneous anthrax outbreaks in hoofstock are often attributed to tabanids (Foil 1989), and there is evidence that tabanids are capable of transmitting the infection to humans. Although anthrax is no longer a common disease in the USA, outbreaks are severe and can take down entire herds in short periods of time. Prior to a mass-produced vaccine in the 1950s, anthrax was a major concern for livestock producers throughout North America and still is a concern in wild animal populations throughout the USA (Blackburn et al. 2007), though primarily centered in the Midwest and western states.

A greater concern to Florida is the transmission of the equine infectious anemia virus (EIAV) or "swamp fever." This retrovirus is spread through mechanical transmission and is present worldwide. EIAV-capable Florida tabanid vectors include T. fuscicostatus and C. flavidus (Foil 1983; Issel et al. 1988). Often, infected horses do not exhibit symptoms of the disease but are still capable of transmission. When horses do exhibit symptoms, they often present with recurring fever, anemia, and swelling of the abdomen and legs, and EIAV may cause abortion in pregnant mares. In extreme cases, the disease can be fatal. Infection is persistent for life once contracted, and positive individuals should be quarantined from the herd (Kahn 2005).

Bovine leukemia virus is a retrovirus of cattle that causes malignant tumors of the lymph nodes. The virus can be transmitted by flies, specifically T. fuscicostatus (Foil 1989) but is also transmitted through infected milk consumed by calves. Sheep and goats serve as secondary hosts to the virus. Most infected cattle do not exhibit symptoms of the virus but are capable of transmission. The virus is usually fatal for those that do exhibit symptoms and affects multiple organs within the body leading to organ failure and cardiac arrest. Dairy cows are generally more susceptible than beef cattle, and cattle between 4 and 8 yr old are most commonly affected (Kahn 2005).

Hog cholera virus (classical swine fever) is a mechanically transmitted Pestivirus, a genus of viruses that belong to the family Flaviviridae. Viruses in the genus Pestivirus infect mammals, including members of the Bovidae and the Suidae. Hog cholera is endemic to Florida and the USA, and it can affect both domestic and feral hogs. Swine may contract acute or chronic forms of the virus, both of which cause poor reproductive performance. Swine with severe infection present with fever, depression, constipation followed by diarrhea and hemorrhaging under the skin. Survival with chronic infections is usually less than 30 d (Kahn 2005). The USA has experienced several outbreaks in the past leading to an eradication program starting in 1961. The United States Department of Agriculture declared a national emergency in 1972 after an especially widespread epidemic, and in 1978, the Secretary of Agriculture declared the USA hog cholera-free (Martin 1978). In Florida, farmers are still urged to watch for resurgence as feral hogs are widespread and difficult to keep away from farms. Florida tabanid species implicated in hog cholera transmission include T. lineola, T. quinquevittatus, T. americanus, T. molestus, and T. atratus (Tidwell et al. 1972; Foil 1989).

Several other parasites and diseases can be transmitted by tabanids, though usually tabanids are not the primary vector. Borrelia burgdorferi (Lyme disease) has been detected in bloodfed female tabanids, though the competence for transmitting the spirochetes is unknown. There have been reports of erythema migrans forming at the site of deer fly bites, but transmission studies have not been done (Magnarelli et al. 1986; Foil 1989). We have compiled a list of Florida tabanid species with known disease transmission capabilities, and associated pathogens (Suppl. Table 2) (Tidwell et al. 1972; Krinsky 1976; Magnarelli et al. 1986; Foil 1989). For a comprehensive review of potential transmittable agents, see Krinsky (1976) and Foil (1989).

Acknowledgments

The authors would like to thank staff at the Florida Department of Agriculture and Consumer Services, Department of Plant Industry for the open use of the Florida State Collection of Arthropods, as well as help in locating specimens and literature in the collection archives. Thank you to Dr. John F. Burger for sending CMZN a personal copy of the Catalog of Tabanidae (Diptera) of North America North of Mexico. Thank you to Mr. Mark Dykes, Chief Apiary Inspector, Texas, and Dr. Larry Hribar, Research Director, Florida Keys Mosquito Control District for reviewing and editing this manuscript.

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Catherine M. Zettel Nalen (1) *, Daniel L. Kline (1), Bruce D. Sutton (2), Gunter Muller (3), and James E. Cilek (4)

(1) Center for Medical, Agricultural, and Veterinary Entomology, USDA-ARS, Gainesville, FL 32608, USA

(2) Department of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville, FL 32607, USA

(3) Department of Microbiology and Molecular Genetics, Institute for Medical Research Israel-Canada, Kuvin Centre for the Study of Infectious and Tropical Diseases, Hebrew University, Jerusalem, Israel

(4) Department of Testing and Evaluation, Navy Entomology Center of Excellence, Jacksonville, FL 32212, USA

* Corresponding author; E-mail: cnalen@wcchd.org

Supplementary material for this article in Florida Entomologist 98(2) (June 2015) is online at http://purl.fcla.edu/fcla/entomologist/browse
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