Bothriocerini

Cesne, Maxime Le, Huber, Elisabeth & Bourgoin, Thierry, 2025, Historical account and redescription of Bothriocera substigmatica (Lethierry, 1881) (Hemiptera, Fulgoromorpha, Cixiidae), an endemic species from the lesser Antilles, Zootaxa 5665 (1), pp. 85-98 : 86-87

Bothriocerini Muir , 1923

Historical taxonomical account. In 1835, Burmeister established the genus Bothriocera Burmeister, 1835 , and classified it under the group Cicadines Fulgorina, corresponding to Latreille's 1807 classification of Cicadariae Fulgorellae. Spinola (1839:145), noting the unique position of the antennae—situated within a notch on the head (referred to as "Néfliers de la maison Burdin (op. cite: 138)"—implicitly placed the genus within his group Cicadariae Fulgorellae Fulgorites Cixioides, though he did not name it explicitly. Spinola (1852: 52) confirmed this placement within the Fulgoritae, Cixioideae. Among these Fulgorites, Stål described the genus Andana Stål, 1856 , which he synonymized with Bothriocera a decade later, placing it within Fulgorida Cixiida ( Stål, 1866) and thereby reinforcing Spinola's initial classification. Since then, Bothriocera has remained within the family Cixiidae Spinola, 1839 .

In his foundational classification of planthopper families based on male genitalia morphology, Muir (1923) established the tribe Bothriocerini using a dual-character criterion: "Subantennal process present OR antennae sunk into pits”. The tribe initially included five genera— Bothriocera , Borysthenes Stål, 1866 , Kinnara Distant, 1906 , Euryphlepsia Muir, 1922 , and Stenophelpsia Muir , 1922 —designated "for convenience" (p. 98) in the first part of his 1925a paper (pp. 97–110). However, in the second part of this work (1925b: pp. 156–163), Muir reclassified Kinnara and the cixiid subfamily Meenoplinae Fieber, 1872 , removing them from Cixiidae to form the families Kinnaridae Muir, 1925 , and Meenoplidae Fieber, 1872 , respectively. It is interesting that Muir acknowledged that the tribe might be "only one of convenience," as he did not identify any unique characters uniting the "considerably different" of the four genera that remained within it (op. cit.: 158). Additionally, Euryphlepsia was originally described three years earlier as part of the Cixiidae tribe Oecleini ( Muir, 1922) , a tribe Muir did not revisit in subsequent work and which remained unaddressed until its revival by Emeljanov (1989).

In 1930, Muir , while publishing the foundation of the current planthopper classification, confirmed the tribe Bothriocerini . However, he mistakenly included the five genera previously mentioned in the first part of his 1925 work, despite Kinnara having already been reclassified as the type genus of the Kinnaridae . A few years later, Metcalf (1938) elevated Bothriocerini to subfamily rank and separated Bothriocerini , retaining only the genus Bothriocera , from Stenophlepsini Metcalf, 1938. Stenophlepsini was noted to include “at least four genera,” though only three were attributed: Borysthenes , Euryphlepsia , and Stenophelpsia ( Metcalf, 1938: 239).

Fifty years later, Emeljanov (1989) revisited the classification of the family Cixiidae , providing a fundamental reorganization that, together with his subsequent publications ( Emeljanov, 2000, 2002; Holzinger et al., 2002), still underpins our current classification although that framework is now in strong disagreement with recent phylogenetic analyses ( Le Cesne et al., 2022; Bucher et al., 2023; Luo et al., 2024). Within the Cixiidae and from the Cixiinae , Emeljanov separated a new subfamily: Borystheninae including the single genus Borysthenes and Bothriocerinae including Bothriocera and the genus Bothrioceretta Caldwell 1950 . The latter was distinguished based on differences in the vertex structure, male genitalia, and overlapping forewings ( Caldwell 1950). Emeljanov restricted Stenophepsini to Stenophelpsia and Euryphlepsia , moving them to the subfamily Cixiinae , where they were to be "compared with the Oecleini (= Myndini Metcalf, 1938), as Muir himself did (1922)" ( Emeljanov, 1989:94).

With the advent of molecular phylogenetics, the position of the Bothriocerinae has been further clarified. All recent studies ( Ceotto et al., 2008; Le Cesne et al., 2022; Bucher et al., 2023; Luo et al., 2024) place this subfamily within the Oecleini , rendering the latter paraphyletic. To ensure classification stability and prevent rapid successive revisions with premature conclusions, Luo et al. (2021, 2024) established the Oecleinian lineage (without assigning it a formal Linnaean rank), encompassing the tribes Duiliini, Stenophepsiini, Oecleini , and Bothriocerini . While the first three tribes remain within Cixiinae , Bothriocerini was downgraded to a tribe but remains formally distinct and separate from Cixiinae .

Although currently restricted in distribution to the Nearctic and Neotropical regions, the group has been enriched in recent years by a series of Eocene fossil genera from Western Europe ( Bothriobaltia Szwedo, 2002 ; Delwa Szwedo, 2019 ; Klugga Szwedo, 2019 ; and Liwakka Szwedo, 2019 ), indicating that Bothriocerini were much more widely distributed in the past.