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Manual of Praying Mantis Morphology, Nomenclature, and Practices (Insecta, Mantodea)

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Manual of Praying Mantis Morphology, Nomenclature, and Practices (Insecta, Mantodea) A peer-reviewed open-access journal ZooKeys 696:Manual 1–100 of(2017) praying mantis morphology, nomenclature, and practices (Insecta, Mantodea) 1 doi: 10.3897/zookeys.696.12542 MONOGRAPH http://zookeys.pensoft.net Launched to accelerate biodiversity research Manual of praying mantis morphology, nomenclature, and practices (Insecta, Mantodea) Sydney K. Brannoch1,2, Frank Wieland3, Julio Rivera4, Klaus-Dieter Klass5, Olivier Béthoux6, Gavin J. Svenson1,2 1 Department of Invertebrate Zoology, Cleveland Museum of Natural History, 1 Wade Oval Drive, Cleveland, Ohio, USA 2 Department of Biology, Case Western Reserve University, 10900 Euclid Avenue, Cleveland, Ohio, USA 3 Pfalzmuseum für Naturkunde - POLLICHIA-Museum, Hermann-Schäfer-Str. 17, 67098 Bad Dürkheim, Germany 4 Universidad San Ignacio de Loyola, Perú 5 Senckenberg Natural History Collections Dresden, Königsbrücker Landstrasse 159, D-01109 Dresden, Germany 6 Centre de Recherche sur la Paleobio- diversite et les Paleoenvironnements (CR2P, UMR 7207), Sorbonne Universites, MNHN, CNRS, UPMC- Paris6, Museum National d’Histoire Naturelle, 57 Rue Cuvier, CP 38, 75005 Paris, France Corresponding author: Sydney K. Brannoch ([email protected]) Academic editor: P. Stoev | Received 6 March 2016 | Accepted 19 June 2017 | Published 13 September 2017 http://zoobank.org/C7B4D1FF-AD2C-4E30-9679-8673D4AB8219 Citation: Brannoch SK, Wieland F, Rivera J, Klass K-D, Béthoux O, Svenson GJ (2017) Manual of praying mantis morphology, nomenclature, and practices (Insecta, Mantodea). ZooKeys 696: 1–100. https://doi.org/10.3897/ zookeys.696.12542 Abstract This study provides a comprehensive review of historical morphological nomenclature used for praying mantis (Mantodea) morphology, which includes citations, original use, and assignment of homology. All referenced structures across historical works correspond to a proposed standard term for use in all subse- quent works pertaining to praying mantis morphology and systematics. The new standards are presented with a verbal description in a glossary as well as indicated on illustrations and images. In the vast majority of cases, originally used terms were adopted as the new standard. In addition, historical morphological topographical homology conjectures are considered with discussion on modern interpretations. A new standardized formulation to present foreleg femoral and tibial spines is proposed for clarity based on previous works. In addition, descriptions for methods of collection, curation, genital complex dissection, and labeling are provided to aid in the proper preservation and storage of specimens for longevity and ease of study. Due to the lack of consistent linear morphometric measurement practices in the literature, we have proposed a series of measurements for taxonomic and morphological research. These measurements are presented with figures to provide visual aids with homologous landmarks to ensure compatibility and Copyright Sydney K. Brannoch et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Sydney K. Brannoch et al. / ZooKeys 696: 1–100 (2017) comparability across the Order. Finally, our proposed method of pinning mantises is presented with a photographical example as well as a video tutorial available at http://mantodearesearch.com. Keywords Mantodea, measurement, morphology, praying mantis, terminology Table of contents 1. Introduction ......................................................................................................... 3 2. Methods ............................................................................................................... 4 3. Results .................................................................................................................. 4 3.1. Head ........................................................................................................... 5 3.2. Wings and wing venation ............................................................................ 8 3.3. Pro-, meso-, and metathoracic legs ............................................................ 16 3.4. Thoracic structures .................................................................................... 24 3.5. Abdominal and genitalic structures ........................................................... 25 3.6. Oothecae ................................................................................................... 50 3.7. Specimen collection .................................................................................. 56 3.8. Specimen preparation ................................................................................ 57 3.9. Specimen mounting .................................................................................. 58 3.10. Male and female genitalia preparation ..................................................... 60 3.11. Specimen labels ....................................................................................... 61 3.12. Standards of measurement ....................................................................... 62 3.13. Glossary .................................................................................................. 73 4. Conclusion ......................................................................................................... 86 5. Acknowledgments .............................................................................................. 87 6. References .......................................................................................................... 87 Supplementary material 1 ....................................................................................... 96 Supplementary material 2 ....................................................................................... 96 Supplementary material 3 ....................................................................................... 97 Supplementary material 4 ....................................................................................... 97 Supplementary material 5 ....................................................................................... 98 Supplementary material 6 ....................................................................................... 98 Supplementary material 7 ....................................................................................... 99 Supplementary material 8 ....................................................................................... 99 Supplementary material 9 ..................................................................................... 100 Supplementary material 10 ................................................................................... 100 Manual of praying mantis morphology, nomenclature, and practices (Insecta, Mantodea) 3 1. Introduction The central motivation for this work is to produce an updated standard for morpho- logical nomenclature, specimen preparation, and measurement data capture. As there is currently a lack of standardization, some level of confusion exists about term use and ap- plication to features as well as the optimal method to measure features that retain highly variable or ambiguous boundaries. We believe this to be an important time to propose a set of standards due to a growth in taxonomic interest in Mantodea, the application of new technologies, and to improve lab workflow efficiency. In addition, we outline and propose new methodological standards to improve the ability to research specimens, which includes specimen preparation and pinning, genital dissections, and labeling. Justification to standardize: Coding of morphology: Congruence of terminology ensures accurate interpretations and future use of characters and their states in morphological analyses and deposi- tion into morphology databases such as MorphBank. The current lack of a system ensures barriers derived from language and chosen reference material. Morpho- logical terminology is suggested using topographical correspondances, which does not in all cases correspond to a hypothesis of homology. Formulaic descriptions: Telegraph style descriptions speed taxonomic work, but term standards ensure longevity, direct comparisons with other studies, reuse of descrip- tions, and extracting coded characters from descriptions. Imaging: Access to high resolution images of specimens are of great importance to taxo- nomic work by improving how we gather data, compare specimens, and identify spe- cies. However, the way a specimen is dry pinned will have great influence on how many images are needed of the same specimen in order to adequately provide access to the relevant features. Minimizing the number of images captured by standardizing the way a specimen is mounted will greatly increase digitization efficiency and access to feature information. If imaging equipment is not readily available, scientific illustration, when performed with high precision and heeding symmetry in bilaterally symmetric struc- tures, can capture important characters for taxonomic and morphological study. Morphometric analysis: Capture of measurement data requires standardization for broader future use in other analyses based on phylogenetics or species delimitation. The measurements described for suggested use are for linear morphometrics, to be used for taxonomic or morphological purposes, like distinguishing between
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