Conocephalinae

Mugleston, Joseph D., Naegle, Michael, Song, Hojun & Whiting, Michael F., 2018, A Comprehensive Phylogeny of Tettigoniidae (Orthoptera: Ensifera) Reveals Extensive Ecomorph Convergence and Widespread Taxonomic Incongruence, Insect Systematics and Diversity (AIFB) 2 (4), No. 5, pp. 1-27 : 22

Conocephalinae

The conehead katydids ( Conocephalinae ), so named for the hypognathous faces giving the head a cone-like appearance, form a large and diverse lineage that is well supported as a monophyletic group. This subfamily is further split into two subclades. The first subclade ( Fig. 12 ) consists of the monophyletic tribe Conocephalini (meadow katydids). In agreement with our prior work ( Mugleston et al. 2013, 2016), the Agraeciini and Copiphorini tribes ( Fig. 13 ) are paraphyletic. The monophyly of these tribes has been a difficult topic for more than a century, as the characters that separate the tribes are not clear. Caudell (1911) following Redtenbacher (1891) separated the two by the fastigium (point) of the vertex being noticeably narrower than the first antennal segment ( Agraeciini ) rather than wider than the first segment ( Copiphorini ). The difficulty of placing taxa within these similar tribes was recognized early on ( Caudell 1918, Zeuner 1936). Walker and Gurney (1972) provided a table with five characters used to differentiate the two tribes, but four of the five characters included the qualifiers ‘usually’, ‘seldom’, or ‘often’. One character on Walker and Gurney’s table was fixed in both tribes: the ventral tooth of the vertex. The lack of characters to distinguish the two tribes has led to a number of taxa being difficult to place. For example, Sphyrometopa (Carl, 1908) has a broad fastigium typical of Copiphorini , but a curved ovipositor and no tooth on the ventral surface of the vertex. The latter set of characters has led to Sphyrometopa being placed within Agraeciini , although our results show it is more closely related to the Neotropical Copiphora (Serville, 1831) . Overall, the differences between these tribes are limited to a minor difference in the projection from the vertex and this does not appear to be phylogenetically informative. From our results it is clear in some cases biogeographic regions are a better indicator of relationships within this conehead subclade (e.g., Indomalayan Agraeciini, Australasian Agraeciini , and Neotropical Agraeciini + Copiphorini ). An obvious exception to this is the clade of slender Copiphorini ( Neoconocephalus , Euconocephalus (Karny, 1907) , Ruspolia , Belocephalus (Scudder, 1875) , Pseudorhynchus , etc.). These katydids are widespread and represent multiple oversea dispersals leading to their current worldwide distribution. The overlap in morphology between the two tribes has made the differences between them difficult to ascertain. The single character used to distinguish the tribes is not useful, and there appears to be no real support or justification for the continued use of both tribes Agraeciini and Copiphorini . To alleviate further confusion, Copiphorini should no longer be viewed as a valid tribe and the species currently within this tribe should be placed within Agraeciini , the senior listing.

Figures

Fig. 12. Conocephalinae group: Conocephalini (Conocephalinae). Posterior probability values over 90 are marked with a circle at the node.Vertical bars indicate subtribes, tribes, and subfamilies. Paraphyletic groups are marked with an asterisk. Photo credit:Arthur Anker.

Fig. 13. Conocephalinae group: Euconchophorini, Agraeciini, and Copiphorini.Vertical bars indicate subtribes, tribes, and subfamilies. Paraphyletic groups are marked with an asterisk. Posterior probability values over 90 are marked with a circle at the node. Photo credit: Joseph Mugleston.