Superfamily Fulgoroidea: the planthoppers

Current Phylogenetic Hypothesis

Overview

Fulgoridae (aka, the lanternflies), and their closest relatives, Dictyopharidae, are two of 20 insect families that are included in the hemipteran superfamily Fulgoroidea (commonly referred to as “planthoppers”). Collectively, planthoppers are a diverse insect group including >9,000 described species, all of which are sap-feeding herbivores. Different researchers have proposed alternative hypotheses regarding how many planthopper families there are and how those families are related. The phylogeny shown here is based on the results of a molecular phylogenetic analysis of Fulgoroidea (using DNA nucleotide sequence data from four genes; Urban & Cryan, 2007). These results support the recognition of at least 19 planthopper families (but note that exemplars of Hypochthonellidae and Gengidae were unavailable for that study). Relationships among some families are depicted here as unresolved because different phylogenetic reconstruction methods produced conflicting results (Urban & Cryan, 2007); we plan to further explore the evolution of the planthopper superfamily in an expanded analysis including more taxa and more data.

These molecular data-based results support the previous hypotheses proposed by Asche (1987; evolutionary relationships based on morphology) and Wilson et al. (1994; host-plant associations): the most ancient planthopper families (Cixiidae and Delphacidae) have ovipositors shaped for piercing and sawing vegetation, and insert eggs into the tissues of monocots or in debris at the base of the host plant. The families of intermediate divergence age (Kinnaridae, Meenoplidae, Derbidae, Achilidae, Achilixiidae, Fulgoridae, and Dictyopharidae) have spade-like ovipositors for burying or covering eggs with soil or wax on monocot or dicot host plants. The most recently diverged families have piercing-excavating ovipositors used to excavate the woody tissue of dicot hosts for egg laying.

Fulgoroidea Phylogeny

Focal Group 1: Family Fulgoridae

Pyrops Phenax Phrictus Zanna Penthacodes

Overview

Fulgorid planthoppers (often called “lanternflies”) are typically arboreal, most often associated with large tropical trees. Fulgoridae is the only family in this intermediate group that deposits eggs higher up on the host plant (rather than in the soil) and covers them with a waxy substance. Fulgoridae are also unique among the planthoppers in that many species are large-bodied (as long as 95 mm), have pigmented, often brightly colored wings, and some produce cuticular waxes in a variety of forms, including plumes that extend well beyond the length of the abdomen. Although at least 10 other planthopper families have elongate head processes, many species of Fulgoridae exhibit comparatively exaggerated morphological diversity in head shape, such as the peanut-headed bug (Fulgora laternaria) and the dragon-headed bug (Phrictus quinquepartitus). The ~550 species of Fulgoridae are primarily distributed throughout the world’s tropical and subtropical regions, with only 17 species recorded from the Nearctic region, and no species recorded from the Palearctic (Metcalf, 1947).

Urban & Cryan (2009) conducted the first phylogenetic investigation of Fulgoridae (based on DNA nucleotide sequence data from five genes generated from a worldwide sampling of fulgorid species representing all 8 recognized subfamilies and 10 of 11 tribes). Results suggested that there have been multiple losses of the extended head process across fulgorid evolution, with what appears to be convergence (in shape and/or loss) in distantly related lineages. The higher classification of Fulgoridae, which is based primarily on characters associated with head morphology, was not well supported by this analysis, suggesting the need for a revised classification of Fulgoridae.

Focal Group 2: Family Dictyopharidae

Dictyopharidae Dictyopharidae Dictyopharidae Dictyopharidae Dictyopharidae

Overview

Dictyopharid planthoppers tend to be associated with non-tree host plants (i.e., grasses, sedges, and shrubs), oviposit in the soil at the base of their host plants, and cover their eggs with debris (Emeljanov, 1979). Dictyopharidae are typically smaller (< 30 mm) and cryptic, often with green or brown bodies and clear wings. Although most species generally do not exhibit the same degree of morphological diversity (in head shape, wing venation, wing shape, etc.) observed in Fulgoridae, some species of Dictyopharidae do rival the lanternflies in their bizarre morphology. The ~500 described species of Dictyopharidae are most diverse in tropical and subtropical regions worldwide. However, Dictyopharidae show greater diversity than Fulgoridae in temperate regions, with 35 species occurring in the Nearctic, and more than 200 in the Palearctic. It is estimated that 50-80% of dictyopharid species are presently undescribed and undiscovered. The phylogeny of Dictyopharidae has never been investigated.