Lipotactinae, Ingrisch, 1995

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 : 23-24

Lipotactinae , Terpandrini , Austrosaginae , and Holarctic Tettigoniinae

Lipotactinae is monophyletic and sister to Neobarrettia + (( Austrosaginae + Chlorobalius ) + Holarctic Tettigoniinae ) ( Fig. 15 ). Lipotactinae was originally described within Tympanophorinae ( Zeuner 1936) but elevated to a subfamily by Ingrisch (1995) due to

Necessary changes to paraphyletic subfamilies:

• Meconematinae should no longer be used

• Listroscelidinae should no longer be used

• Mecopodinae should no longer be used unless changes are made to exclude Aprosphylini and include the tribes Ischnomelini and Phrictini (formerly Pseudophyllinae ) and the subfamily Phyllophorinae

• Pseudophyllinae should no longer be used unless Simoderini, Phrictini , and Ischnomelini are removed

• Tettigoniinae should only include the Holarctic shield back tribes and Ephippigerini (formerly Bradyporinae ).

• Phaneropterinae should include Zichyini (formerly Bradyporinae )

Paraphyletic tribes and genus groups that should no longer be used unless further revised

• Nedubini • Poreuomenini

• Copiphorini • Ducetiini

• Agraeciini • Ephippithytae

• Terpandrini • Elimaeini

• Tettigoniini • Mirolliini

• Platycleidini • Holochlorini

• Sexavaini • Steirodontini

• Phyllomimini • Trigonocoryphini

• Cymatomerini • Pycnopalpini

• Platyphyllini • Pycnopalpina

• Pleminiini • Phyllopterini

• Cocconotini • Phyllopterina

• Anaulacomerina • Sudderiini

• Teleutiini • Microcentrini

• Phaneropterini • Amblycoryphini

• Odonturini • Insarini

• Barbitistini • Terpnistrini

• Acrometopini

Necessary changes to the recently named supertribes:

• Pseudophylliti o Remove genera within Phrictini

• Pleminiiti o Remove genera within Ischnomelini

Genera rearranged as a result of this study

Arachnoscelis should no longer be included within Phisidini differences in thoracic sterna, thoracic auditory spiracle, compressed tibia, etc. While emphasis was placed on the extant taxa when these subfamilies were divided, the fossil katydids were not addressed, leaving Eomortoniellus spp. under Tympanophorinae instead of moving them along with their modern counterparts Lipotactes (Brunner von Wattenwyl, 1898) to Lipotactinae . Gorochov referred to Lipotactinae as a tribe (presumably under Tympanophorinae) and included Eomortoniellus with the other Lipotactinae ( Gorochov 2010) . Tympanophorinae and Lipotactinae are not closely related and similarity between these two groups is likely due to ecomorphic convergence as in the other distantly related katydid subfamilies. Listroscelidinae was shown to be paraphyletic ( Mugleston et al. 2013) and the distantly related taxa ( Meiophisis (Jin, 1992) and Arachnoscelis ) have since been removed from Listroscelidinae . However, with our additional sampling, the ‘taxonomic dump’ of Listroscelidinae was again verified ( Mugleston et al. 2016). In addition to the Requenini mentioned above, samples from North American and Australian taxa within the tribe Terpandrini were included in this study. Australian Chlorobalius are sister to Australian Austrosaginae and not North American Neobarrettia . Terpandrini historically has been included in both Saginae ( Gorochov 2007) and Listroscelidinae ( Rentz 2001, Naskrecki and Rentz 2010, Fialho et al. 2014). Likewise, Austrosaginae genera were also included within Saginae and only elevated to subfamily rank in the last few decades ( Rentz 1993). Austrosaginae , Saginae, and the taxa previously described under the paraphyletic Listroscelidinae are all predatory katydids with similar habitus. The taxonomic confusion is another apparent case where convergence in ecomorphs has led to invalid taxonomic groupings. Saginae is only a distant relative to the Austrosaginae and species traditionally described under Listroscelidinae . Additionally, Listroscelidinae are more closely related to biogeographically close taxa in separate subfamilies than to other Listroscelidinae once again verifying that this subfamily does not represent a monophyletic group and should no longer be considered valid.

The nominate subfamily Tettigoniinae is paraphyletic as it is currently defined. The Holarctic Tettigoniinae is not a monophyletic group because the Bradyporinae genus Ephippiger sp. (Berthold, 1827) is nested within this group. As with the other large katydid subfamilies, widespread tribes are not monophyletic and similar morphology may have more to do with similarity in habitat and independent selective pressures than with phylogeny. Two relatively recent dispersals to Nearctic regions occurred. The first gave rise to Anabrus simplex (Haldeman, 1852) , the Mormon cricket. The second is a more recent transition that gave rise to the more apical North American shieldbacks. Tettigoniinae are largely recognized by features associated with spines, plantula, and ovipositor but the characters that are used to define this group do not account for the various southern hemisphere taxa that are currently described as Tettigoniinae but only distantly related. If Tettigoniinae is to continue being used, it should only include the Holarctic taxa + Ephippiger sp. and exclude the taxa currently described in the tribe Nedubini .

Figures

Fig. 15. Tettigoniinae group:Terpandrini,Austrosaginae,and HolarcticTettigoniinae.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.