Cicadellidae, Latreille, 1825

Dietrich, Christopher H., Allen, Julie M., Lemmon, Alan R., Lemmon, Emily Moriarty, Takiya, Daniela M., Evangelista, Olivia, Walden, Kimberly K. O., Grady, Patrick G. S. & Johnson, Kevin P., 2017, Anchored Hybrid Enrichment-Based Phylogenomics of Leafhoppers and Treehoppers (Hemiptera: Cicadomorpha: Membracoidea), Insect Systematics and Diversity 1 (1), pp. 57-72 : 67-69

Cicadellidae : a Paraphyletic Family

Results of our analyses provide strong confirmation of the hypothesis of some previous authors (Hamilton 1983, Rakitov 1998, Dietrich et al.2001a) that treehoppers are a derived lineage of leafhoppers and, hence, that the family Cicadellidae as presently defined is paraphyletic. Previous analyses indicated that one or both of the leafhopper subfamilies Megophthalminae and Ulopinae are sister to the treehopper lineage comprising the families Aetalionidae , Melizoderidae ,

and Membracidae . Our results provide strong support for this relationship, although the status of Ulopinae and Megophthalminae as sister groups (also suggested by a previous analysis; Dietrich et al. 2001a) remains less certain. The sister-group relationship of treehoppers with these leafhopper subfamilies is consistent with behavioral observations by Rakitov (1996, 1998) that megophthalmine and ulopine leafhoppers share a unique form of anointing behavior called ‘bathing,’ by which the insects wallow in fluid containing secretory products of the Malpighian tubules that they deposit on the substrate. In other leafhoppers, the fluid is collected and spread over the body using the hind legs. Interestingly, in both leafhopper groups, the Malpighian secretion contains brochosomes, which consist of minute granules that form an extremely hydrophobic coating when the fluid dries. Although some treehoppers do exhibit bathing behavior, there is currently no evidence that they produce brochosomes ( Rakitov 1996, 1998). Thus, our results are consistent with Rakitov’s (1998) hypothesis that the derivation of treehoppers from a specialized clade of leafhoppers coincided with the loss of brochosome production. Presence of rows of enlarged setae on the hind tibia (used by modern leafhoppers to spread brochosomes over the body; Fig. 1C ) in at least some members of the extinct Mesozoic membracoid family Archijassidae ( Shcherbakov 2012) suggests that brochosome production originated in Membracoidea prior to the origin of Cicadellidae , the earliest fossils of which are recorded from the lower Cretaceous.

The lack of clear resolution of relationships among the major lineages of Membracoidea has, so far, hindered attempts to redefine Cicadellidae as a monophyletic group. While the results of our concatenated ML analysis did not identify a well-supported sister group to the (strongly supported) megophthalmine–ulopine–tree-hopper lineage, the coalescent gene tree (ASTRAL) analysis grouped the remaining Cicadellidae in a clade that was sister to this lineage with 96% posterior probability. If the latter result can be confirmed by further analysis, then the family Cicadellidae could be rendered monophyletic by excluding Ulopinae and Megophthalminae from Cicadellidae and elevating these subfamilies to separate family status.

Figures

Fig. 1. Representatives of Membracoidea: (A) Mapuchea chilensis (Nielson) (Myerslopiidae), Chile; (B) Mesargus sp. (Cicadellidae: Ulopinae), China; (C) Evacanthus sp. (Cicadellidae: Evacanthinae), China, using hind legs with rows of enlarged setae to spread brochosomes over wing surfaces; (D) Aetalion reticulatum (Linnaeus), (Aetalionidae), adult female guarding egg mass; (E) Melizoderes osborni (Funkhouser); (F) Notocera sp. (Membracidae), Brazil.