Olivia Evangelista

A phylogeny of the treehopper subfamily Heteronotinae reveals convergent pronotal traits (Hemiptera: Auchenorrhyncha: Membracidae)

Even within an insect family famous for its morphological diversity, the treehopper subfamily Heteronotinae is a microcosm of pronotal variation, displaying remarkably dissimilar thoracic ornamentations among its ten included genera. Presented here is a reconstruction of heteronotine relationships based on DNA nucleotide sequence data from five nuclear and two mitochondrial genes from a comprehensive sample of ingroup taxa (including exemplars of all genera except for the monotypic Dysyncritus Fowler, 1895). Concordant phylogenetic estimates support the monophyly of Heteronotinae sensu stricto (i.e. excluding Darnoides Fairmaire, 1846), as well as the monophyly of the included genera Heteronotus Laporte, 1832, Nassunia Stål, 1862, Omolon Walker, 1862, Rhexia Stål, 1867, and Smiliorachis Fairmaire, 1846. Conversely, Anchistrotus Buckton, 1902, Allodrilus Evangelista, 2014 and Iria Stål, 1867 were not recovered as monophyletic, although topology comparison tests failed to reject the nonmonophyly of the latter two genera, nor Heteronotinae, as currently circumscribed. These results are interpreted in the context of available morphological evidence, which generally supports these findings. Also addressed here is the evolution of selected pronotal features representing major aspects of heteronotine morphology. Bayesian stochastic mapping of pronotal traits estimated multiple character state transitions, suggesting repeated acquisition of pronotal features, most of which have evolved independently within the diverse genus Heteronotus. These convergences may provide insight into the role of the pronotum in heteronotine groups that appear to serve different adaptive strategies, such as mimicry, crypsis and defence against predators. Furthermore, our results show a substantial diversity in need of formal description, including possible new genera, and several unique sexually dimorphic syndromes that may constitute a species complex within Heteronotus delineatus Walker, 1858. Nomenclatural changes are proposed for Nassunia nigromacula (Funkhouser, 1940) new combination and Smiliorachis inornata Stål, 1862 combination reinstated, which were found to be misplaced at the generic and subfamily levels according to our analyses.

An annotated catalogue of the Membracidae types in the Museu de Zoologia da Universidade de São Paulo, Brazil (Hemiptera: Auchenorrhyncha: Cicadomorpha)

Here presented is an annotated catalogue of the Membracidae types deposited at the Museu de Zoologia da Universidade de São Paulo (MZSP), with information on 106 primary types and 171 secondary types. These type specimens represent 82 nominal species currently classified in 8 treehopper subfamilies and 17 tribes. According to our estimates, at least 10 primary types and 44 secondary types are missing in the collection. A comprehensive list of literature sources, taxonomic remarks, label data and descriptions on the state of preservation of specimens are given.

Figures 95 – 100 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 95 – 100. Allodrilus deitzi Evangelista sp. nov. (paratype male, NCSU): 95. Abdomen, lateral view; 96. Abdomen, ventral view, detail of the VIII sternite (indicated); 97. Pygofer, lateral view; 98. Subgenital plate, ventral view. 99 – 100. Aedeagus and style, respectively, lateral view.

Figures 87 – 94 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 87 – 94. Allodrilus colombiensis Evangelista sp. nov. (holotype male, MPUJ): 87 – 88. Abdomen, lateral and ventral view (pygofer removed), respectively; 89. Pygofer, lateral view; 90 – 91. Pygofer (aedeagus, style, and subgenital plate removed), latero-caudal and caudal view, respectively; 92 – 93. Aedeagus and styles, lateral and ventral view, respectively; 94. Subgenital plate, ventral view.

Figures 79 – 86 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 79 – 86. Allodrilus alboferrugineus Evangelista sp. nov. (paratype male, MZSP): 79 – 80. Abdomen, lateral and ventral view, respectively. 81. Pygofer, lateral view; 82. Pygofer, caudal view (subgenital plate removed); 83. Subgenital plate, ventral view; 84 – 85. Aedeagus and style, respectively, lateral view; 86. Hind tibia, lateral view (setal rows indicated).

Figures 70 – 78 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 70 – 78. Allodrilus nitidipennis (Funkhouser) comb. nov.: 70 – 71. Abdomen, lateral and ventral view, respectively, abdominal segments indicated. 72. Pygofer, lateral view; 73. Pygofer, caudal view; 74. Subgenital plate and aedeagus, ventral view; 75 – 76. Aedeagus and style, respectively, lateral view; 77. VIII sternite, ventral view (indicated); 78. Hind tibia, lateral view (setal rows indicated); aa: aedeagal aphophysis.

Figures 62 – 69 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 62 – 69. Dysyncritus intectus Fowler: 62 – 63. Abdomen, lateral and ventral view, respectively (abdominal segments indicated); 64. Pygofer, lateral view; 65. Pygofer, caudal view (subgenital plate removed); 66. Subgenital plate, ventral view; 67. Style, lateral view; 68. Aedeagus, lateral view; 69. Hind tibia (male paralectotype, BMNH), lateral view, rotated clock-wise from left to right, (a) detail of the tibial surface, densely covered in minute setae.

Figures 56 – 61 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 56 – 61. Immatures and tending ants of A. nitidipennis (Funkhouser) comb. nov.: 56 – 58. 4 – 5 th instar nymph, frontal, lateral and dorsal view, respectively; 59. Camponotus femoratus (Fabricius, 1804), lateral view; 60. Crematogaster sp., lateral view; 61. Ectatomma tuberculatum (Oliver, 1792), lateral view. Scale bar = 1 mm.

Figures 51 – 55 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 51 – 55. Species of Allodrilus gen. nov., dorsal view: 51. A. deitzi Evangelista sp. nov., holotype male (NCSU); 52. A. granulatus Evangelista sp. nov., holotype female (USNM); 53. A. horizontalis Evangelista sp. nov., holotype male (NHRS); 54. A. intermedius Evangelista sp. nov., holotype male (USNM); 55. A. similis Evangelista sp. nov., holotype male (NCSU). Scale bar = 1 mm.

Figures 5 – 12 In The Identity Of The Treehopper Genus Dysyncritus Fowler, With Descriptions Of New Related Taxa (Hemiptera: Membracidae: Heteronotinae)

FIGURES 5 – 12. Species of Dysyncritus and Allodrilus gen. nov., lateral view: 5 – 6. D. intectus Fowler, lectotype female and paralectotype male, respectively (BMNH); 7 – 9. A. nitidipennis (Funkhouser) comb. nov., (7) holotype female (USNM), (8 – 9) color and pronotal variation: female (DZUP) and male (DNA voucher MEM 136, NCSU), respectively; 10 – 12. A. alboferrugineus Evangelista sp. nov., (10) holotype female (DZUP), color variation: (11) paratype male (USNM), (12) paratype female (MZSP). Scale bar = 1 mm.