The jumping plant-lice (Hemiptera, Psylloidea) associated with Schinus (Anacardiaceae): systematics, biogeography and host plant relationships

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Abstract
The gall-forming psylloid fauna associated with the temperate Neotropical plant genus Schinus (Anacardiaceae) is revised, and keys are provided for adults, larvae and galls. Fourteen species are recognized in the genus Calophya (Calophyidae), 10 of which are described as new; C. gallifex and C. schini are reinstated from former synonymy with C. rubra. An additional closely related new species, C. clavuligera sp. n., is described from Lithrea spp. In the genus Tainarys (Psyllidae, Rhinocolinae), six species are recognized and, for five of these, Schinus species are confirmed hosts. Two of them are described as new and one is recombined from Leurolophus. A new Leurolophus species is described from Lithrea molleoides. Based on a cladistic analysis Pelmatobrachia, Microceropsylla and Neocalophya are synonymized with Calophya, producing five new and one revised species combinations as well as one nomen nov. A list of the 59 currently recognized species is provided with information on synonymy, distribution and host plants. The cladistic analyses suggest that the Schinus inhabiting calophyids form two monophyletic groups, the C. rubra and C. hermicitae groups, respectively. A discriminant analysis of 15 variables for both males and females provides morphometric characters for separating the morphologically very homogenous adults of the C. rubra group. The phylogenetic relationships of the Rhinocolinae are re-evaluated, and the genera Tainarys and Leurolophus are redefined. Notophyllura is transferred from the Euphyllurinae to the Rhinocolinae. Tainarys forms the sister group of Leurolophus + Notophyllura. Based on mapped distributions four areas of endemism (distribution) are defined: A (Eastern temperate South America), B (Central and Southern Chile, including adjacent parts of Argentina), C (Central Argentina) and D (Peru, Far Northern Chile, and Eastern Bolivia). Cladistic biogeographical analyses with TAS, COMPONENT and an algorithm by Enghoff yielded the general area cladogram (A +(C +(B +D))); the one calculated with BPA differed in the position of D: (D +(A +(B +C))). The reconstruction of vicariant events using a protocol by Hovenkamp, corroborates the former general area cladogram, which is compared to competing hypotheses. The history of dispersal and vicariance events is reconstructed with the programme DIVA. Evidence for cospeciation of Schinus with its associated psylloids is evaluated with COMPONENT and TREE MAP. The likelihood for cospeciation is high in the Calophya hermicitae group, but low in the Calophya rubra group and in Tainarys. The gall shape depends partly from the psylloid, partly from the Schinus phylogeny.