Research Article |
Corresponding author: José Miguel Vela ( josem.vela@juntadeandalucia.es ) Academic editor: Ron Beenen
© 2020 José Miguel Vela, Miguel Ángel Alonso-Zarazaga, Mauro Daccordi.
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.
Citation:
Vela JM, Alonso-Zarazaga MÁ, Daccordi M (2020) The species of Timarcha Samouelle, 1819 described by Linnaeus (Coleoptera, Chrysomelidae). ZooKeys 986: 55-80. https://doi.org/10.3897/zookeys.986.57158
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Linnaeus described five species presently included in the genus Timarcha: Chrysomela goettingensis, Tenebrio caeruleus, Tenebrio laevigatus, Tenebrio latipes, and Tenebrio rugosus. After a study of the relevant material, the identity of these species has been established. The following synonyms are proposed or confirmed: Timarcha goettingensis (Linnaeus, 1758) = T. latipes (Linnaeus, 1767), syn. nov.; Timarcha caerulea (Linnaeus, 1758), comb. nov. = T. balearica Gory, 1833, syn. nov. = T. balearica Pérez Arcas, 1865, syn. nov.; Timarcha rugosa (Linnaeus, 1767) = T. scabra (Olivier, 1807), syn. conf. = T. generosa Erichson, 1841, syn. conf.; Timarcha laevigata (Linnaeus, 1767) = T. tenebricosa (Fabricius, 1775), syn. conf.. The type of Tenebrio caeruleus is a Chrysomelidae currently belonging to genus Timarcha and therefore can no longer be considered a Tenebrionidae (Helops caeruleus) nor the type species of genus Helops. For the sake of nomenclatural stability, an application to the International Commission on Zoological Nomenclature to change the relative precedence of Timarcha caerulea and retain usage of T. balearica will be made. An application to change the relative precedence of Timarcha laevigata has been submitted, which would lead to the conservation of usage of T. tenebricosa as valid. Lectotypes are designated for Chrysomela goettingensis, Tenebrio latipes, Tenebrio caeruleus, Timarcha balearica Gory, T. balearica Pérez Arcas, Tenebrio rugosus, Chrysomela scabra, Timarcha generosa, Tenebrio laevigatus, and Chrysomela tenebricosa. For each of the valid species the diagnosis, distribution, and host-plant data are reported.
Chrysomelinae, Europe, leaf beetles, nomenclature, North Africa, synonyms, taxonomy, Tenebrionidae
The taxonomy of the genus Timarcha Samouelle, 1819 (Coleoptera, Chrysomelidae) is among the most challenging of all Palaearctic chrysomelids because: i) the types were rarely consulted by authors, and ii) there exists a high variability in traits as size, sculpture and form of the pronotum (
As a starting point for the revision of the genus Timarcha, we have studied the species authored by Carl Linnaeus. He described five species presently belonging to this genus. They are Chrysomela goettingensis Linnaeus, 1758, Tenebrio caeruleus Linnaeus, 1758, Tenebrio laevigatus Linnaeus, 1767, Tenebrio latipes Linnaeus, 1767, and Tenebrio rugosus Linnaeus, 1767. One of us (MAAZ) studied and photographed the types of T. latipes and T. laevigatus in the collection of the Linnean Society of London. As well, consultation of photographs of Linnean types on the website of the Linnean Collections (http://linnean-online.org/) of the Linnean Society of London, together with a study of type specimens of other nominal species in other museums, led us to reconsider the availability and nomenclatural status of the five species of Timarcha described by Linnaeus.
Measurements of body length were made using the ocular grid of a Lomo MBS-10 binocular microscope at 10× magnification. Body size was considered the total length of the specimen from the anterior region of head to the apex of elytron. Photographs of type specimens of Tenebrio laevigatus and T. latipes were taken with a Canon EOS 7D camera attached to a MP-E 65 mm f/2.8 1–5× macro lens. Photographs of type specimens of Chrysomela goettingensis, Tenebrio caeruleus, and T. rugosus were kindly provided by Linnean collections staff (The Linnean Society of London) and of Timarcha balearica Gory, 1833 by Antoine Mantilleri (MNHN). Types, and their parts, of Timarcha balearica Pérez-Arcas, 1865, T. scabra (Olivier, 1807), and T. generosa Erichson, 1841 were photographed with an Olympus Stylus TG-3 digital compact camera. Photographs of other specimens or their parts were done with a Canon EOS 550D attached to a bellows with a Schneider Componon-S 50mm f/2.8 objective. Combine ZM was used for resolving the stack of photos.
The methodology to name the vestiture under of the tarsi I–III, to dissect the sclerites of endophallus, and to inflate the endophallus is explained by
In the treatment of type material from the collection of Carl Linnaeus, we have followed Recommendation 73F (
Host plants are given using their valid names. If a name, now a synonym, was originally mentioned, this follows the valid name between round brackets. Plant nomenclature follows
The material examined is housed in the following collections (curators mentioned between round brackets):
LSUK The Linnean Collections of the Linnean Society, London (Isabelle Charmantier, Suzanne Ryder)
MNHN Muséum National d’Histoire Naturelle, Paris (Antoine Mantilleri)
ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin (Johannes Frisch, Bernd Jäger)
ZMUK Zoologisches Museum, Universität Kiel, Kiel (Michael Kuhlmann).
The label data for all type specimens is cited as follows: a double slash (//) divides the texts on different labels, a single slash (/) divides the text in different rows. Type localities are cited with their original spellings. Comments and notes are cited in square brackets: [p] preceding data are printed, [h] preceding data are handwritten, [w] white label, [r] red label.
The webpage of the Linnean Collections (http://linnean-online.org/), of the Linnean Society of London, has been a critical source of information.
Chrysomela goettingensis
Tenebrio latipes
Chrysomela goettingensis: “Germania”. Tenebrio latipes: “Africa” [type locality wrong].
Chrysomela goettingensis: not examined. The images of the lectotype (♂, presently designated, Fig.
Tenebrio latipes: Lectotype (♀, presently designated, Fig.
In the Linnean collections there are two different species under Chrysomela goettingensis. One of them, specimen LINN 5537 labelled “goettingensis” [w, h, Linnaeus’ handwriting] (http://linnean-online.org/22922/), is the lectotype of Chrysomela goettingensis Linnaeus, 1758: 368 (presently in Timarcha, Fig.
Authors such as
Males: 7.4–12.6 mm (lectotype of T. goettingensis: 8.7 mm; Fig.
Timarcha goettingensis 1 lectotypus of Chrysomela goettingensis in dorsal (a) and side (b) view, and label (c) (photos provided by The Linnean Society, with permission to reproduce 2 lectotypus of Tenebrio latipes in dorsal (a) and side (b) view, lateral side of pronotum (c), underside of third metatarsomere (d), and label (e) (photos by Alonso Zarazaga and Ren Li, with permission to reproduce from The Linnean Society) 3 male habitus from Germany, Frankenhausen (a) and Germany, Erfurt (b) 4 pronota of males from France, Lozère (a), Germany, Bad Frankenhausen (b) and Germany, Erfurt (c).
Most of Europe, from northern Spain to European Russia, and reaching Great Britain and Sweden (
Scabiosa atropurpurea L. (= S. maritima L.) (Caprifoliaceae); Plantago lanceolata L., P. coronopus L. (Plantaginaceae), Cruciata laevipes Opiz, Galium aparine L., G. arenarium Loisel., G. mollugo L., G. odoratum (L.) Scop. (= Asperula odorata L.), G. saxatile L., G. uliginosum L., G. verum L., Rubia peregrina L. (Rubiaceae) (
Timarcha goettingensis 5 aedeagi of Germany, Erfurt in dorsal (a) and side view (b) and Germany, Bad Frankenhausen in dorsal (c) and side (d) view 6 sclerite of the endophallus from Germany, Bad Frankenhausen, in dorsolateral view (taken from
Tenebrio caeruleus
Tenebrio caeruleus
Timarcha caerulea
(
Timarcha balearica
Timarcha balearica
Timarcha balearica
Timarcha balearica var. violaceus
Timarcha balearica var. martini
Timarcha balearica ab. viridipennis
Timarcha balearica ab. coerulescens
Timarcha balearica ab. longicornis
Timarcha balearica ab. nigriventris
Timarcha balearica ab. olivacea
Timarcha balearica ab. semicoerulea
Timarcha balearica ab. discolor
Timarcha balearica ab. tricolor
Tenebrio caeruleus: “Hispania”. Timarcha balearica G.: “Les Iles Baléares”. Timarcha balearica P. A.: “Mahón (Menorca), Alcudia de Mallorca”.
Tenebrio caeruleus: not examined. The images of the lectotype (♂, presently designated, Fig.
Timarcha balearica G.: Lectotype (♂, presently designated, Fig.
Timarcha balearica P.-A.: Lectotype (♂, presently designated, Fig.
Timarcha balearica was described for the first time as figure 8 in planche 49 (
For the sake of stability (Art. 23.2,
Males: 12.0–14.5 mm (lectotype of Tenebrio caeruleus: 12.2 mm, Fig.
Timarcha balearica 8 lectotypus of Tenebrio caeruleus in dorsal (a) and side (b) view, and label (c) (photos provided by The Linnean Society, with permission to reproduce) 9 lectotypus of Timarcha balearica Gory in dorsal (a), ventral (b) and side (c) view, and label (d) (photos provided by Antoine Mantilleri (MNHN), with permission to reproduce) 10 lectotypus of Timarcha balearica Pérez Arcas, habitus (a), pronotum (b) and labels (c) 11 male habitus from Spain, Baleares, Palma de Mallorca 12 aedeagus from Spain, Baleares, Palma de Mallorca, in dorsal (a) and side (b) view 13 sclerite of the endophallus from Spain, Palma de Mallorca, in side view (taken from
Balearic Islands: Mallorca and Menorca (
Rubia peregrina L. (= R. angustifolia L.), Galium spp., Asperula spp. (Rubiaceae) (
Tenebrio rugosus
Chrysomela scabra
Timarcha generosa
Tenebrio rugosus: “Africa” [other localities mentioned in the original description as “Hispania” and “Gallia” are erroneous and should not be taken into consideration following Recommendation 76A.2 of the Code (
Tenebrio rugosus: not examined. The photographs of the lectotype (♀, presently designated, Fig.
Chrysomela scabra: Lectotype (♀, presently designated, Fig.
Timarcha generosa: Lectotype (♂, presently designated, Fig.
Timarcha rugosa 15 lectotypus of Tenebrio rugosus in dorsal (a) and side (b) view, and label in upper (c) and lower view (d) (photos provided by The Linnean Society, with permission to reproduce) 16 lectotypus of Timarcha scabra in dorsal (a), and ventral (b) view, pronotum (c) and label (d) 17 Lectotypus of Timarcha generosa in dorsal view (a), pronotum (b) and label (c).
Males: 11.8–19.0 mm (lectotype of T. generosa: 15.8 mm, Fig.
Timarcha rugosa 18 male habitus from Algery, Mandoura (a), Morocco, Debdou (b) and Morocco, Aguelmame Sidi Ali (c) 19 pronota of males from Algery, Mandoura (a), Morocco, Debdou (b) and Morocco, Aguelmam Sidi Ali (c) 20 aedeagi from Morocco, Oujda in dorsal (a) and side (b) view, Morocco, Aguelmame Sidi Ali in dorsal (c) and side (d) view and Morocco, Annual in dorsal (e) and side (f) view 21 sclerite of the endophallus from Algery, Batna in dorsal view (taken from
Tunisia, Algeria, and Morocco. Spain and France, as in the original description of T. rugosus, are wrong.
Plantago albicans L. (Plantaginaceae), Asperula sp. (Rubiaceae) (
Tenebrio laevigatus
Tenebrio coeruleus
Chrysomela tenebricosa
Chrysomela tenebriosa:
Chrysomela tenebrioides:
Chrysomela tenebricosa:
Tenebrio coeruleus:
Timarcha tenebricosa:
Chrysomela laevigata: Duftschmid 1825: 161 (combination).
Timarcha laevigata:
Tenebrio laevigatus: “Africa” [wrong type locality]. Chrysomela tenebricosa: “Europa australiori”.
Tenebrio laevigatus: Lectotype (♀, designated herein, Fig.
Chrysomela tenebricosa: not examined. Syntypes (1 ♂, 1♀, Fig.
Timarcha tenebricosa 23 lectotypus of Tenebrio laevigatus in dorsal (a) and side view (b), pronotum (c) and metatarsus in lower view (d) (photos by Alonso Zarazaga and Ren Li, with permission to reproduce from The Linnean Society) 24 typi of Chrysomela tenebricosa Fabricius, lectotypus male in dorsal (a) and ventral (b) view, and pronotum (c), paralectotypus female in dorsal (d) and ventral (e) view, and pronotum (f) (photographed by Michael Kuhlmann, Zoologischen Museum Kiel, reproduced with permission).
The lectotype of T. laevigata (L.), a female (Fig.
As the name Timarcha laevigata, in the sense here fixed as a synonym of T. tenebricosa, has at least one usage since 1899 (in
Males: 14.6–17.2 mm; females: 16.1–18.2 mm (lectotype of Tenebrio laevigata = 17.5 mm; Fig.
Timarcha tenebricosa 25 male habitus from France, Poitiers 26 pronota of males of males of Crimea (a), Austria, Vienna (b), France, Paimport (c) and France, Poitiers (d) 27 aedeagi from France, Alps Maritimes, Caussols in dorsal (a) and side (b) view, France, Normandie, Bréal in dorsal (c) and side (d) view, Crimea in dorsal (e) and side (f) view 28 sclerite of the endophallus from England, Launceston in dorsolateral view (taken from
Most of Europe, from northern Spain to Great Britain and Ireland, eastwards to Georgia, Azerbaijan, and Asiatic Turkey; not recorded in Scandinavia (
On Rubiaceae: Asperula cynanchica L., Cruciata laevipes Opiz, Galium aparine L., G. mollugo L., G. parisiense (L.), G. verum L., Rubia peregrina L. (
The difficult task of studying the types of old species has been greatly facilitated with the quick access to high-quality images. As a result, it is now easier for taxonomists to verify the status of old synonyms or interpretations which were based upon very short, generalized descriptions. The high-quality images of the types in the Linnaean collections have proven extremely useful for the zoological community. In addition, most museums and their curators are willing to help with search for and loan of types, which can facilitate taxonomic work. Incorrect species concepts, not based on the examination of name-bearing types, have sometimes been maintained despite the identity of extant type material, which has been carefully cared for and maintained for many years for the benefit of science.
In the case of genus Timarcha, whose revision is very necessary, the slow and sometimes difficult work of consulting types has become absolutely necessary, as various authors have made different interpretations for a long time. Although historical misinterpretations of T. laevigata and T. latipes have ascribed these to different North African species or even to both sexes of the same species, these two species are actually two very different European species. Also, T. caerulea is revealing, as it was considered a Tenebrionidae, when in actuality the type is clearly a male of the genus Timarcha, identical to T. balearica (Chrysomelidae).
The main synonyms presented here can be summarized as follows:
Timarcha goettingensis (Linnaeus, 1758)
= T. latipes (Linnaeus, 1767), syn. nov.
Timarcha balearica Gory, 1833 (to be proposed to ICZN as a nomen protectum)
= Timarcha caerulea (Linnaeus, 1758), syn. nov., comb. nov. (to be proposed to ICZN as a nomen oblitum)
Timarcha rugosa (Linnaeus, 1767)
= T. scabra (Olivier, 1807), syn. conf.
= T. generosa Erichson, 1841, syn. conf.
Timarcha tenebricosa (Fabricius, 1775) (proposed to ICZN as a nomen protectum)
= T. laevigata (Linnaeus, 1767), syn. conf. (proposed to ICZN as a nomen oblitum)
We thank Ren Li (Institute of Zoology, Chinese Academy of Sciences, Beijing, China) for the photographs of the types of T. laevigata and T. latipes. We also thank the following persons. Elaine Charwat and Mike Fitton, as former Deputy Librarian and Curator of Entomology, respectively, and Isabelle Charmantier and Suzanne Ryder, as present Head of Collections and Curator of Entomology, respectively (Linnean Collections, The Linnean Society of London), helped with the searching of Linnean types. Andrea Deneau (Digital Assets Manager in the Linnean Collections) facilitated photographs and permission to publish the Linnean types. Michael Kuhlmann (Zoologisches Museum Kiel) and Antoine Mantilleri (Muséum National d’Histoire Naturelle, Paris) kindly sent us to photographs of types of Timarcha tenebricosa and T. balearica, respectively. Mercedes París (Museo Nacional de Ciencias Naturales, Madrid) and Johannes Frisch and Bernd Jäger (Museum für Naturkunde der Humboldt-Universität, Berlin) were very helpful locating the types under their care. Patrice Bouchard (Canadian National Collection of Insects, Ottawa) made important suggestions in the text and revised and improved the English. The cooperation of Maurizio Bollino (Museo di Storia Naturale del Salento, Calimera, Lecce) was very important for the endophallus eversions. We are grateful to Lyubomir Penev for his support in the publication of our paper.