North American Dictyopharidae
Scolops sulcipes. Photo by Dave Funk of Stroud Water Research Center, Avondale, PA.
Dictyopharidae is the 6th largest family of planthoppers worldwide, with 164 genera and 730 species at present (Bourgoin 2012). Dictyopharidae is represented north of Mexico by 14 genera and 85 species. The southwest includes nearly all species known from the contiguous United States (14 genera, 80 species), with the rest of the regional fauna consisting mostly of widespread taxa. Nine genera and 39 species are Orgeriinae, an unusual, brachypterous, arid-adapted subfamily, all of which are endemic to the southwest U.S. and adjacent Mexico. Among Dictyopharinae, Scolops is the most species-rich genus, including 36 taxa found in nearly all states and Mexico; Phylloscelis (4 species) is found only north of Mexico, whereas Mitrops (3 species), Nersia (12 species) and Rhynchomitra (5 species) are also found elsewhere in the Neotropics. Tribes of Dictyopharinae were recently revised by Emeljanov (2008, 2011).
Dictyopharidae and Fulgoridae are sister groups, with problematic family-level delineation, although the enigmatic taxa are mostly Old World (Urban & Cryan 2009). The families are similar in several respects, including the tendency to have an elongated head, and both having a row of teeth on the apex of the second hind tarsomere (a plesiomorphic feature for the superfamily). North of Mexico, the families can be separated by Dictyopharidae not having reticulate venation in the hind wing (see below). North American dictyopharids are usually smaller than fulgorids, but there is overlap (although U.S. fulgorids always have patterned forewings whereas the larger U.S. dictyopharids have clear wings). Many dictyopharids have 3 carinae on the frons in addition to the lateral carinae (separating the functional frons from the genal and temple region). Orgeriinae is brachypterous, as are most Scolops and Phylloscelis (although the brachypterous wings of the latter 2 genera will cover the abdomen, whereas in Orgeriinae the forewings leave several tergites exposed). Dictyopharidae is characteristically long-legged and have a long beak, and stand in a curious upright position to feed.
Wings of Diacira sp. (Cladodipterini, from Brazil)
Since Metcalf (1946), Phylloscelis has been revised (McPherson 1994, McPherson & Wilson 1995), and Doering (1955) revised Acinaca and Deserta, providing a key to genera of North American Orgeriinae. Since few species of Dictyopharidae have been described from North America since Metcalf (1946), earlier taxonomic works are still useful, including Doering & Darby (1943) for Orgerius. Breakey (1929) provides a key to Scolops, but 12 taxa have been described since that time, and a synthetic work is badly needed. Fennah (1944) presented keys to most of the New World dictyopharid genera. The key is revised from Doering & Darby (1943) and Fennah (1944), with Loxophora and Timodema treated as subgenera of Ticida following Emeljanov (2006).
Dictyopharid distribution in the United States (from Bartlett et al in press)
Dictyopharids feed on a variety of plants with adults and nymphs both found on above-ground portions of plants. Most species are associated with dicots in a variety of plant families, and are most often reported as monophagous (Wilson et al. 1994). The biology of Nersia florens was described by Wilson and McPherson (1981a). This species (in Illinois) is apparently univoltine, and feeds on Rumex crispus L. (Polygonaceae) and Ageratina altissima (L.) King & H. Rob. (Asteraceae), with adults emerging in August and September. The biology of Phylloscelis was examined by McPherson (1994) and McPherson & Wilson (1995, 1996). Phylloscelis is also univoltine (except Florida populations of P. atra), and overwinters as eggs. Phylloscelis pallescens was found exclusively on Pycnanthemum tenuifolium Schrad. (Lamiacae) with nymphs producing abundant wax. Phylloscelis rubra (the ‘cranberry toad bug’) is reported only from cranberry (Vaccinium macrocarpon Aiton; Ericaceae) and has been implicated in wilting and dieback (Sirrine & Fulton 1914, Wilson & O’Brien 1987). Taosa inexacta (Walker 1858) is being considered as a biological control for water hyacinth (Eichhornia crassipes (Mart.) Solms, Pontederiaceae; e.g., Cordo 1999). Rhynchomitra microrhina was recently reported from the exotic grass Eragrostis curvata (Schrad.) Nees and appears to be bivoltine (Wilson & Wheeler 2005).
The genera of Dictyopharidae found north of Mexico are as follows:
Dictyopharinae Onuki, 1901
Nersiini Emeljanov 1983 (sensu Emeljanov 2011)
Mitrops Fennah, 1944 (Type species Fulgora noctivida Linnaeus, 1767).
Nersia Stål, 1862 (Type species Nersia haedina Stål, 1862).
Rhynchomitra Fennah, 1944 (Type species Dictyophara microrhina (Walker, 1851).
Phylloscelini Emeljanov, 1983
Phylloscelis Germar, 1839 (Type species Phylloscelis pallescens Germar, 1839).
Scoloptini Emeljanov, 1983
Scolops Schaum, 1850 (Type species Fulgora sulcipes Say 1825)
= Ornithissus Fowler, 1904 (Type species Ornithissus cockerelli Fowler, 1904); syn. by Kirkaldy 1907: 248.
Subgenus Belonocharis Uhler, 1891 (Type species Belonocharis fumida Uhler, 1891); status (subgenus) Van Duzee 1916: 78.
Subgenus Scolops Schaum 1850 (Type species Fulgora sulcipes Say 1825).
Orgeriinae Fieber, 1872
Orgeriini Fieber, 1872
Acinaca Ball & Hartzell, 1922 (Type species Acinaca lurida Ball & Hartzell, 1922).
Aridia Ball & Hartzell, 1922 (Type species Orgerius compressus Ball, 1909).
Deserta Ball & Hartzell, 1922 (Type species Orgamara bipunctata Ball, 1909).
Orgamara Ball, 1909 (Type species Orgamara acuta Ball, 1909)
Orgerius Stål, 1859 (Type species Orgerius rhyparus Stål, 1859).
= Ranissus Fieber, 1866 (Type species Ranissus leptopus Fieber, 1866); syn. by Fieber 1872: 4.
Subgenus Orgerius Stål, 1859
Subgenus Opsigonus Emeljanov, 2006 (Type species Orgerius minor Ball, 1909).
Ticida Uhler, 1891 (Type species Ticida cingulata Uhler, 1891).
= Loxophora Van Duzee, 1908 (Type species Loxophora transversa Van Duzee, 1908); syn. by Emeljanov 2006: 73.
Subgenus Heicophora Emeljanov, 2006a (Type species Loxophora dammersi Van Duzee, 1934).
Subgenus Ticida Uhler, 1891 (Type species Ticida cingulata Uhler, 1891).
Subgenus Timodema Ball, 1909 (Type species Timodema miracula Ball, 1909); generic status revised by Emeljanov 2006: 74 (reduced to subgenus of Ticida).
Ticrania Emeljanov, 2006 (Type species Ticida chamberlini Van Duzee, 1923).
Timonidia Ball & Hartzell, 1922 (Type species Timonidia solitaria Ball & Hartzell, 1922).
Yucanda Ball & Hartzell, 1922 (Type species Orgamara albida Ball 1909).
Key to the genera of Dictyopharidae of North America, North of Mexico.
1. Wings as long as abdomen in dorsal view in both macropters and brachypters, often transparent (brachypters with wings usually extending to end of abdomen); claval fold present; tegulae present ... (Dictyopharinae) 2
1.’ Wings always brachypterous, shorter than abdomen, leaving several segments exposed when viewed from above; claval fold absent; forewing opaque; tegulae hidden ... (Orgeriinae) 6
Left to right: Orgerius concordus (Orgeriinae), Rhynchomitra microrhina, and Scolops sulcipes (Dictyopharinae)
2. Tibiae and femora of forelegs greatly foliaceous; vertex not produced anteriorly ... Phylloscelis Germar
2.’ Tibiae and femora of forelegs not greatly foliaceous (if tibiae flattened, then head produced); vertex produced anteriorly ... 3
Phyllocelis rubra (left) and Phylloscelis pallescens (right); note expanded legs.
3. Forewings leathery, opaque, brownish or grayish, brachypterous or macropterous; body brown, head projection long and thin ... Scolops Schaum
3.’ Forewings membranous, translucent, greenish, macropterous; body green ... 4
4. Tegulae distinctly carinate; vertex short-triangular ... Nersia Stål
4.’ Tegulae not carinate; vertex elongate-triangular ... 5
Top to bottom: Mitrops dioxys, Nersia florens, and Rhynchomitra microrhina
Head and thorax of Nersia florens showing carinate tegula.
5. Six or seven areoles adjoining nodal line, latter distinct, pronotum notched, but shallowly ... Mitrops Fennah
5.’ No regular areoles on nodal line, latter distorted by distal reticulation; pronotum with a deep narrow notch on hind margin basally ... Rhynchomitra Fennah
Pronotal notch of Nersia (left) and Rhynchomitra (right) .
6. Callosity behind eye evident from lateral view; vertex elongate or angulate ... 7
6.’ No callosity behind eye; vertex rounding or, if elongate, broad if seen from side ... 10
Callosity behind eye: Acinaca lurida (left, callosity absent) and Orgerius concordus (right, callosity present).
7. Vertex short, less than twice the length of the eyes ... Orgerius Stål
7.’ Vertex elongate, more than twice the length of the eyes ... 8
Deserta obscura (left), Orgamara argentia (middle) and Yucanda albida (right) (Orgeriinae), dorsal view showing elongated head.
8. Cephalic process, as seen from side, beak-like, the apex obliquely rounding from above, the lower angle slightly produced ... Deserta Ball
8.’ Cephalic process, as seen from the side, truncate at the extremity, five angled ... 9
9. Cephalic process gradually tapering, as seen from top and side ... Orgamara Ball
9.’ Cephalic process parallel margined; apex, as seen from the side, slightly enlarged, projecting at an angle with the vertex ...Yucanda Ball & Hartzell
10. Head angulate in lateral view, produced in front of eyes for distance greater than 2/3 width of eyes; head widening toward apex ... Acinaca Ball & Hartzell
10.’ Head rounded or angulate, produced in front of eyes for distance less than 2.3 width of eyes... 11
Acinaca lurida (Orgeriinae); Photo by Rick Donovall, University of Delaware, Department of Entomology)
11. Vertex broad and short; apical cell of vertex (areolet) absent; front usually with horizontal black band above frontoclypeal suture; fore and middle tibiae sometimes foliaceous ... 12
11.’ Vertex longer, apical cell of vertex (areolet) present; front without horizontal black band above frontoclypeal suture; tibiae not foliaceous ... 13
12. Front with horizontal black band above frontoclypeal suture; pronotum without lateral carinae, posterior margin shallowly concave; forewing with median and claval veins as evident carinae ... Ticida Uhler
12.’ Front without horizontal black band above frontoclypeal suture; pronotum with lateral carinae, posterior margin deeply U-shaped; forewing with uniform net of veins ... Ticrania Emeljanov
Ticida cingulata (top) and Ticida (Timodema) miracula (bottom) (Orgeriinae); Photos by Rick Donovall, University of Delaware, Department of Entomology)
Aridia compressa (left), Ticrania chamberlini (middle, holotype), and Timonidia solitania (right); photo of Aridia compressa by Rick Donovall, University of Delaware, Department of Entomology); photo of holotype of Ticrania chamberlini courtesy Norm Penny, Department of Entomology, California Academy of Sciences); photo of Timonidia solitania by Kimberley Shropshire, Department of Entomology, University of Delaware.
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