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新热带区竹节虫研究XXVI:Cladomorformia类群(分类新属)的分类学综述,一种枝翅竹节虫亚科的谱系,并描述七个新属和41个新物种(竹节虫目:隐匿竹节虫亚目:枝翅竹节虫亚科)

Studies on Neotropical Phasmatodea XXVI: Taxonomic review of Cladomorformia tax. n., a lineage of Diapheromerinae stick insects, with the descriptions of seven new genera and 41 new species (Phasmatodea: Occidophasmata: Diapheromerinae).

作者信息

Hennemann Frank H, Conle Oskar V

机构信息

Tannenwaldallee 53; 61348 Bad Homburg; Germany.

Am Freischütz 16; 47058 Duisburg; Germany.

出版信息

Zootaxa. 2024 Apr 26;5444(1):1-454. doi: 10.11646/zootaxa.5444.1.1.

Abstract

Stick and leaf insects (Phasmatodea) are a moderately diverse order that comprises almost 3,500 extant species of large to very large often impressively camouflaged nocturnal herbivores. The order also stands out as one of the few insect orders that have until lately lacked a robust higher-level phylogeny and still the relationships between many New World taxa in particular remain unknown. The Diapheromerinae is one of the main lineages of the Occidophasmata and comprises a bulk of the diversity of New World stick insects. The clade is endemic to the Americas and includes the longest insects of that region. During the past 25 years Diapheromerinae has experienced various often inconsistent attempts of a classification, which have resulted in a complicated history that has meanwhile constituted a heterogenous mixture of historical and contemporary views. To counter these challenges workers have made use of rank-free taxa to provisionally group supposedly monophyletic clades. Currently the Diapheromerinae are sub-divided into the two tribes Diapheromerini and Oreophoetini and recent molecular analyses have shown taxa of the Cladomorphinae Günther, 1953 to belong to Diapheromerinae. Certainly, the clade still deserves much work to delimit meaningful sub-groups within Diapheromerinae reflective of their evolutionary history and the results of the latest comprehensive molecular-based phylogenetic studies already imply sub-divisions within the Diapheromerini. The rank-free taxon Cladomorformia tax. n. is established to comprise former Cladomorphini, Cranidiini, Otocrania Redtenbacher, 1908 as well as sections of Diapheromerini sensu Robertson et al., 2018, which are the genera that have previously been placed in the "Phanocles group" sensu Zompro, 2001 as well as two genera of the "Bacteria group" and one genus of the "Clonistria group": Alienobostra, Bostriana, Calynda, Globocalynda, Laciphorus, Phanocles, Phanocloidea and Trychopeplus. The genus Cranidium and all genera of Cladomorphini sensu Hennemann et al., 2016 are here formally transferred to Diapheromerinae, this is Aplopocranidium, Cladomorphus, Jeremia, Jeremiodes, Otocraniella and Xylodus. A holistic review of Cladomorformia at the genus level is conducted herein and new diagnoses, differentiations and keys to all 25 genera that are now contained in that clade are presented. Lists of species are provided for all genera, which include detailed type data, synonymies and distributional records. Moreover, identification keys to all 163 known valid species are provided to render identification of species within the 25 known genera possible. In total, 232 species are covered if the 69 synonymous taxa are included. The subfamily Haplopodinae is introduced to comprise all genera of the tribes Haplopodini, Hesperophasmatini and Pterinoxylini as classified by Hennemann et al. (2016) as well as the recently described Teruelphasmini. Renaming former Cladomorphinae sensu Robertson et al. (2018) into Haplopodinae is necessary, because the type-genus Cladomorphus can no longer be considered a member of that lineage and belongs in Diapheromerinae. Haplopodinae is the only New World clade that belongs into the Old World Oriophasmata. Bostranova Villet, 2023 has been introduced to replace the preoccupied Bostra Stål, 1875. The type-species of Bostranova, Bacteria turgida Westwood, 1859, however is here shown to belong in Phanocloidea Zompro, 2001. Thus, Bostranova is synonymised under Phanocloidea (n. syn.) and all species are transferred to Phanocloidea and other genera. Paraphanocles Zompro, 2001 (Type-species: Mantis keratosqueleton Olivier, 1792) is synonymised with Phanocles Stål, 1875 (syn. n.). Seven new genera are described: Globocrania gen. n. (Type-species: Bacteria emesa Westwood, 1859), Hirtuleiodes gen. n. (Type-species: Phibalosoma gibbosa Chopard, 1911), Lanceobostra gen. n. (Type-species: Bacteria aetolus Westwood, 1859), Ocreatophasma gen. n. (Type-species: Ocreatophasma elegans sp. n.), Parotocrania gen. n. (Type-species: Parotocrania panamae sp. n.), Phanoclocrania gen. n. (Type-species: Bostra dorsuaria Stål, 1875) and Spinocloidea gen. n. (Type-species: Spinocloidea panamaense sp. n.). All seven genera are described from both sexes and the eggs, with the exception of Ocreatophasma gen. n., which is known from the females only. Forty-one new species are described: Globocalynda cornuta sp. n. from Ecuador, Globocalynda marcapatae sp. n. from Peru, Globocalynda ruficollis sp. n. from Bolivia, Hirtuleiodes peruanus sp. n. from Peru, Jeremiodes costaricensis sp. n. from Costa Rica, Jeremiodes ecuadoricus sp. n. from Ecuador, Jeremiodes peruanus sp. n. from Peru, Lanceobostra chapalaense sp. n. from Mexico, Lanceobostra glabra sp. n. from Mexico, Lanceobostra oaxacaee sp. n. from Mexico, Lanceobostra ornata sp. n. from Mexico, Lanceobostra torquata sp. n. from Mexico, Lanceobostra tuckerae sp. n. from Mexico, Ocreatophasma elegans gen. n., sp. n. from Peru, Ocreatophasma fragile gen. n., sp. n. from Peru, Ocreatophasma modestum gen. n., sp. n. from Peru, Parotocrania acutilobata gen. n., sp. n. from Ecuador, Parotocrania curvata gen. n., sp. n. from Peru, Parotocrania panamae gen. n., sp. n. from Panama, Phanocles acutecornutus sp. n. from Ecuador, Phanocles barbadosense sp. n. from Barbados, Phanocles berezini sp. n. from Mexico, Phanocles brevipes sp. n. from Peru, Phanocles chiapasense sp. n. from Mexico, Phanocles cuzcoense sp. n. from Peru, Phanocles ecuadoricus sp. n. from Ecuador, Phanocles falcatus sp. n. from Ecuador, Phanocles maximus sp. n. from Panama, Phanocles mexicanus sp. n. from Mexico, Phanocles pleurospinosus sp. n. from Costa Rica, Phanocles rehni sp. n. from Honduras, Phanocles solidus sp. n. from Ecuador, Phanocles spectabilis sp. n. from Costa Rica, Phanocles superbus sp. n. from Ecuador, Phanocloidea sanguinea sp. n. from Ecuador, Phanocloidea semiptera sp. n. from Venezuela, Phanocloidea venezuelica sp. n. from Venezuela, Spinocloidea panamaense gen. n., sp. n. from Panama, Spinocloidea splendida gen. n., sp. n. from Colombia and Spinocloidea tumescens gen. n., sp. n. from Costa Rica. With a maximum recorded body length of 285.0 mm the new species Phanocles maximus sp. n. from Panama is the longest extant insect of the Americas and the Occidophasmata clade. A total of 132 taxonomic changes are conducted: 64 species are transferred to other genera and 43 new synonyms are established. Lectotypes are designated for 25 taxa to ensure stability of the concerned names or new synonymies here established. Moreover, the previously unknown females of five and males of twelve species are described and illustrated for the first time. Colour illustrations are presented of the eggs of 39 species, of which those of 26 species are formally described and illustrated for the first time.

摘要

竹节虫目昆虫是一个多样性适中的目,包含近3500种现存的大型至超大型夜行性食草动物,它们通常具有令人印象深刻的伪装。该目也是少数几个直到最近仍缺乏可靠高级系统发育关系的昆虫目之一,特别是许多新大陆类群之间的关系仍然未知。枝背竹节虫亚科是隐竹节虫类的主要谱系之一,包含了新大陆竹节虫的大部分多样性。该分支是美洲特有的,包括该地区最长的昆虫。在过去的25年里,枝背竹节虫亚科经历了各种常常不一致的分类尝试,导致了一段复杂的历史,同时形成了历史观点和当代观点的异质混合。为应对这些挑战,研究人员利用无等级分类单元来临时划分推测为单系的分支。目前,枝背竹节虫亚科被分为两个族,即枝背竹节虫族和山竹节虫族,最近的分子分析表明,1953年的枝形竹节虫亚科的类群属于枝背竹节虫亚科。当然,该分支仍需要大量工作来界定枝背竹节虫亚科中有意义的亚群,以反映其进化历史,最新的基于全面分子的系统发育研究结果已经暗示了枝背竹节虫族内的细分。建立了无等级分类单元枝形竹节虫形态类,以包括以前的枝形竹节虫族、头角竹节虫族、1908年的耳角竹节虫属,以及2018年罗伯逊等人所定义的枝背竹节虫族的部分类群,这些类群包括以前被置于2001年佐姆普罗所定义的“幻角竹节虫类群”中的属,以及“菌形竹节虫类群”的两个属和“克隆竹节虫类群”的一个属:异菌形竹节虫属、菌形竹节虫属、卡琳达竹节虫属、球卡琳达竹节虫属、刺缘竹节虫属、幻角竹节虫属、幻角竹节蛉属和曲角竹节虫属。2016年亨内曼等人所定义的头角竹节虫属和所有枝形竹节虫族的属在此正式转移到枝背竹节虫亚科,即尖头角竹节虫属、枝形竹节虫属、耶利米竹节虫属、耶利米蛉竹节虫属、小耳角竹节虫属和木竹节虫属。本文对枝形竹节虫形态类在属级水平上进行了全面综述,并给出了该分支中现有的25个属的新诊断、鉴别特征和检索表。提供了所有属的物种列表,包括详细的模式数据、同义词和分布记录。此外,还提供了所有163个已知有效物种的检索表,以便能够识别25个已知属内的物种。如果包括69个同物异名分类单元,总共涵盖232个物种。引入了单枝竹节虫亚科,以包括亨内曼等人(2016年)分类的单枝竹节虫族、西方竹节虫族和翼木竹节虫族的所有属,以及最近描述的特鲁埃尔竹节虫族。将2018年罗伯逊等人所定义的原枝形竹节虫亚科重新命名为单枝竹节虫亚科是必要的,因为模式属枝形竹节虫属不能再被视为该谱系的成员,而应属于枝背竹节虫亚科。单枝竹节虫亚科是唯一属于旧世界东方竹节虫类的新大陆分支。2023年引入了菌形竹节虫新属以取代已被占用的1875年的菌形竹节虫属。然而,菌形竹节虫新属的模式种,1859年韦斯特伍德命名的肿菌形竹节虫,在此被证明属于2

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