North American Issidae
Thionia bullata (photo by Clyde Sorensen, North Carolina State University)
Issidae is the 5th largest family of planthoppers, consisting of 229 genera and 1,168 species (Bourgoin 2012; excluding Caliscelidae, including Hemisphaerinae). The higher classification of Issidae has been the subject of recent investigation. Fennah (1954) revised the classification of Issidae, in which he included Acanaloniidae as a subfamily, and in 1984, removed a series of North American issid genera (Bladinini subtribe Gaetuliina) to Nogodinidae, which subsequently were moved to Tropiduchidae (as tribe Gaetuliini) (Gnezdilov 2007). Several authors have since provided evidence that Issidae as defined by Fennah (1954, 1984) is not monophyletic (e.g., Emeljanov 1991, Yeh & Yang 1999). Emeljanov (1999) recognized Caliscelidae as a family, and revived Acanaloniidae (subsumed under Issidae by Fennah 1954), which was defined to include Euthiscia, and 2 Old World subfamilies (Trienopinae and Tonginae), subsequently moved to Nogodinidae and Tropiduchidae (Gnezdilov 2007, 2008). A series of additional changes to the higher taxonomy have been proposed, but for taxa north of Mexico the most important changes are the subtribal definitions proposed by Gnezdilov (2003b, 2009).
North American issids, as currently defined, can be recognized by the hind tibiae bearing 1-4 lateral spines (Acanaloniidae have none), forewings covering the abdomen and usually not reticulate (if reticulate then uniformly so, without patches of clear cells), and pronotum usually not extending anteriorly beyond the middle of the eyes. Like all higher planthoppers, Issidae have a pair of spines on the second tarsomere of the hind leg.
Hind tibia of Thiona elliptica showing lateral spines.
Issidae north of Mexico consist of 11 genera and 24 species. North American Issidae (including Caliscelidae and Gaetuliini) were revised by Doering (1936, 1938, 1940, 1941, 1958), and these works can be consulted for species identification. O’Brien (1988) provided a revised key to genus based on Doering (1938), later updated by Gnezdilov & O'Brien (2006b). Gnezdilov (2003a, 2004a) noted that North American Hysteropterum are not congeneric with the European species and 5 genera (Abolloptera, Balduza, Exortus, Kathleenum, Stilbometopius) were subsequently erected for North American Hysteropterum (Gnezdilov 2004a, Gnezdilov & O'Brien 2006b). Gnezdilov & O’Brien (2006a) report Hysteropterum severini and H. beameri as synonyms of Agalmatium bilobum (Fieber), a European species. Tylana ustulata was transferred to Hysteropterum by O’Brien (1988), subsequently to Tylanira by Gnezdilov & O’Brien (2006b), who also raised Paralixes from subgenus to genus status out of Ulixes Stål, 1861. Paralixes and Ulixes are rare in collections, and there appear to be undescribed species in these genera, but infraspecific variation is poorly understood.
In the U.S., Issidae is most species rich in the south, with most genera also occurring also in Central America, except the adventive Agalmatium. Thionia as currently defined includes 70 species, a few of which are Old World. Of the remaining genera, all are limited to North and Central America. The genera Paralixes (total of 4 species), Picumna (10 species) and Traxus (2 species) include Central American species not found in the U.S.
Distribution of Issidae in the U.S. (figure from Bartlett et al. in review).
Issidae is usually associated with woody plants both as adults and nymphs, and are found on the above-ground portion of plants. Most species are reported as monophagous (Wilson et al. 1994). Wheeler & Wilson (1987, 1988) describe the life history of Thionia elliptica, T. bullata and T. simplex. Thionia elliptica feeds on Quercus (in Pennsylvania mostly on Quercus ilicifolia Wangenh.), with a single generation per year, adults emerging in July and August, overwintering as eggs (Wheeler & Wilson 1987). Thionia bullata is univoltine on pines (including the introduced Pinus sylvestris L.), overwintering as eggs, with development requiring about 10 weeks (Wheeler & Wilson 1988, Wheeler 1996). Thionia simplex is polyphagous on a variety of herbaceous and woody plants (Wheeler & Wilson 1988). Schlinger (1958) describes the biology of the adventive Agalmatium bilobum (as Hysteropterum beameri) in California, which creates ‘mud cases’ for its eggs (see also Boulard 1987).
The higher taxonomy of Issidae north of Mexico can be summarized as follows:
(A key to genus will be added later)
Issinae Spinola, 1839
Colpopterini Gnezdilov, 2003b
Colpoptera Burmeister, 1835 (Type species Colpoptera sinuata Burmeister, 1835)
(note: Gnezdilov 2012d has moved this tribe to Nogodinidae; a change that will be reflected in this website at a later date).
Issini Spinola, 1839
Subtribe Agalmatiina Gnezdilov, 2002
Agalmatium Emeljanov 1971 (Type species Cercopis grylloides Fabricius, 1794; jr. syn. of Fulgora flavescens Olivier, 1791).
Subtribe Hysteropterina Melichar, 1906
Abolloptera Gnezdilov & O’Brien 2006b (Type species Hysteropterum bistriatum Caldwell, 1945).
Balduza Gnezdilov & O’Brien 2006b (Type species Hysteropterum unum Ball, 1910).
Exortus Gnezdilov 2004 (Type species Hysteropterum punctiferum Walker, 1851).
Kathleenum Gnezdilov 2004 (Type species Hysteropterum cornutum Melichar, 1906).
Paralixes Caldwell 1945 (Type species Issus scutatus Walker, 1851).
Stilbometopius Gnezdilov & O’Brien 2006b (Type species Issus auroreus Uhler, 1876).
Traxus Metcalf 1923 (Type species Traxus fulvus Metcalf, 1923).
Tylanira Ball 1936 (Type species Tylanira bifurca Ball, 1936).
Subtribe Thioniina Melichar, 1906
Picumna Stål 1864 (Type species Picumna varians Stål, 1864).
Thionia Stål 1859 (Type species Issus longipennis Spinola 1839).
Bliven, B. P. 1966. New genera and species of Issidae. Occidental Entomologist 1(9):103-107.
Boulard, M. 1987. Contribution a l'etude des Issidae. L'ootheque terreuse des 'Hysteropterum', un probleme evolutif (Hom. Fulgoroidea). Bulletin de la Societé Entomologique de France 92(1-2): 5-17.
Caldwell, J. S. 1945. Notes on Issidae from Mexico (Homoptera: Fulgoroidea). Annals of the Entomological Society of America 38: 89-120.
Caldwell, J. S. and D. M. DeLong. 1948. A new species of Issidae from California (Homoptera: Fulgoroidea). Ohio Journal of Science 48:176-177.
Doering, K. C. 1936. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part I. University of Kansas Science Bulletin 24(17): 421-467.
Doering. K. C. 1938. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part II. University of Kansas Science Bulletin 25(20): 447-575.
Doering. K. C. 1939. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part III. University of Kansas Science Bulletin 26(2): 83-167.
Doering, K. C. 1941. A contribution to the taxonomy of the subfamily Issinae in America north of Mexico (Fulgoroidea: Homoptera. Part IV. University of Kansas Science Bulletin 27(10):185-233.
Doering, K. C. 1958. A new species of Hysteropterum from grape (Issidae, Fulgoroidea, Homoptera). Journal of the Kansas Entomological Society 31: 101-103.
Emeljanov, A. F. 1991. An attempt to construct a phylogenetic tree for planthoppers (Homoptera, Cicadina). Entomological Review 70: 24-28 (Translation of Entomologicheskoye Obozreniye 1990, 69: 353-356, in Russian).
Emeljanov, A. F. 1999. Notes on the delimination of families of the Issidae group with description of a new species of Caliscellidae belonging to a new genus and tribe (Homoptera, Fulgoroidea). Zoosystematica Rossica 8(1): 61-72.
Fennah, R. G. 1954. The Higher classification of the family Issidae (Homoptera: Fulgoroidea) with descriptions of new species. Transactions of the Royal Entomological Society of London 105(19): 455-474. pdf
Fennah, R. G. 1955. Lanternflies of the family Issidae of the Lesser Antilles (Homoptera: Fulgoroidea). Proceedings of the United States Museum 105(3350): 23-47.
Gnezdilov, V. M. 2002. Morphology of the ovipositor in the subfamily Issinae (Homoptera, Cicadina, Issidae). Entomologicheskoe Obozrenie 81(3): 605-626, 780. [In Russian]
Gnezdilov, V. M. 2003a. Review of the family Issidae (Homoptera, Cicadina) of the European fauna, with notes on the structure of ovipositor in planthoppers. Chteniya Pamyati Nikolaya Aleksandrovicha Kholodkovskogo (Meetings in memory of N.A. Cholodkovsky) 56(1): 1-145. [Russian with English summary].
Gnezdilov, V. M. 2003b. A new tribe of the family Issidae with comments on the family as a whole (Homoptera: Cicadina). Zoosystematica Rossica 11(2): 305-309.
Gnezdilov, V. M. 2004a. Two new genera of the family Issidae (Homoptera: Cicadina: Fulgoroidea) from North America. Russian Entomological Journal 13 (1-2): 1-2.
Gnezdilov, V. M. 2004b. Morphology of the ovipositor in members of the subfamily Issinae (Homoptera, Cicadina, Issidae). Entomological Review 82(8): 957-974.
Gnezdilov, V. M. 2009. Revisionary notes on some tropical Issidae and Nogodinidae (Hemiptera: Fulgoroidea). Acta Entomologica Musei Nationalis Pragae 49(1): 75-92.
Gnezdilov, V. M. 2011 A new genus and new species of the family Issidae (Hemiptera, Fulgoroidea) from Venezuela. Deutsche Entomologische Zeitschrift 58(1): 119-121.
Gnezdilov, V. M. 2012d. Revision of the tribe Colpopterini Gnezdilov, 2003 (Homoptera, Fulgoroidea: Nogodinidae). Entomologicheskoe Obozrenie XCI(4): 757-774 [in Russian].
Gnezdilov, V. M. and L. B. O'Brien. 2006a. Hysteropterum severini Caldwell and DeLong, 1948, a synonym of Agalmatium bilobum (Fieber, 1877) (Hemiptera: Fulgoroidea: Issidae). Pan-Pacific Entomologist 82(1): 50-53.
Gnezdilov, V. M. and L. B. O'Brien. 2006b. Generic changes in United States Issini (Hemiptera, Fulgoroidea, Issidae). Insecta Mundi 20(3-4): 217-225.
Gnezdilov, V. M. and L. B. O'Brien. 2008. New taxa and combinations in Neotropical Issidae (Hemiptera: Fulgoroidea). Insecta Mundi 31: 1-26.
O'Brien, L. B. 1988. Taxonomic changes in North American Issidae (Homoptera: Fulgoroidea). Annals of the Entomological Society of America 81(6): 865-869.
Schlinger, E. I. 1958. Notes on the biology of a mud egg-case making fulgorid, Hysteropterum beameri Doering (Homoptera: Fulgoroidea). Journal of the Kansas Entomological Society 31(2): 104-106.
Stroinski, A. and J. Szwedo. 2008. Thionia douglundbergi sp nov from the Miocene Dominican amber (Hemiptera: Fulgoromorpha: Issidae) with notes on extinct higher planthoppers. Annales Zoologici 58(3) 529-536.
Wheeler, A. G., Jr. 1996. Use of an introduced host, scotch pine, by a native planthopper, Thionia bullata (Homoptera: Fulgoroidea: Issidae). Proceedings of the Entomological Society of Washington 98(2): 374-375.
Wheeler, A. G., Jr. and S. W. Wilson. 1987. Life history of the issid planthopper Thionia elliptica (Homoptera: Fulgoroidea) with description of a new Thionia species from Texas. Journal of the New York Entomological Society 95(3): 440-451.
Wheeler, A. G., Jr. and S. W. Wilson. 1988. Notes on the biology and immatures of the issid planthoppers Thionia bullata and T. simplex (Homoptera: Fulgoroidea). Journal of the New York Entomological Society 96(3): 266-273.
Wilson, S. W., C. Mitter, R. F. Denno, and M. R. Wilson.1994. Evolutionary patterns of host plant use by delphacid planthoppers and their relatives. In: R. F. Denno and T. J. Perfect, (eds.). Planthoppers: Their Ecology and Management. Chapman and Hall, New York. Pp. 7-45 & Appendix.