To be or not to be a tibial comb: A discussion on the (past) use of tibial armature in tribal/subtribal organization in Cholevinae (Coleoptera: Leiodidae)




Coleoptera, Cholevinae, Ptomaphagini, Eucatopini, Tibial morphology


Detailed studies of microstructure has recently been shown to provide phylogenetic signals at several supraspecific levels within leiodid coleopterans, as well as in other insects. The tribe Ptomaphagini (Leiodidae: Cholevinae), with a Holarctic-Neotropical-Oriental distribution, has been characterized, among other things, by having a comb of equal-sized, flat spines around the apex of the tibiae of all legs, with a row of spines extending along the outer edge of the protibiae in the subtribes Baryodirina and Ptomaphaginina (but not in Ptomaphagina). A pattern similar to the one in Ptomaphaginina also occurs in the Neotropical cholevine tribe Eucatopini, and this has been used to indicate a phylogenetic relationship between the two tribes (but recent phylogenetic studies have not supported such a close relationship). We here review and revise the presence and structure of periapical (here called an ‘apical crown’) and marginal (here called an ‘external comb’) combs of spines on tibiae in Ptomaphagini, using other cholevines (with and without apical tibial combs) for comparison. We find a phylogenetic signal in an apical crown of tibial spines not interrupted at the outer spur, which seems to be an additional synapomorphy of Ptomaphagini, differing from the pattern in Eucatopini and remaining cholevines with an apical comb of spines, in which the comb is interrupted. We highlight differences not previously noticed between the apical protibial armature of Ptomaphaginina and Eucatopini.


Download data is not yet available.

Author Biographies

  • Pedro Gnaspini, Universidade de São Paulo (USP), Instituto de Biociências (IB‑USP), Departamento de Zoologia - São Paulo, SP, Brasil.

    Rua do Matão, travessa 14, 321, Butantã, CEP 05508‑090, São Paulo, SP, Brasil.

  • Stewart Blaine Peck, Canadian Museum of Nature, Research and Collections Division, Entomology - Ottawa, ON, Canada.

    PO Box 3443, Station D, Ottawa, ON, K1P 6P4, Canada.

  • Caio Antunes-Carvalho, Universidade Federal Fluminense (UFF), Instituto de Biologia (EGB), Departamento de Biologia Geral (GBG) - Niterói, RJ, Brasil.

    Outeiro São João Batista, s/nº, Centro, CEP 24020‑141, Niterói, RJ, Brasil.

  • Michel Perreau, Université de Paris, IUT Paris Diderot - Paris, France.

    20 quater Rue du Département 75018 Paris, France.

  • Eduardo Coutinho Gomyde, Universidade de São Paulo (USP), Instituto de Biociências (IB‑USP), Departamento de Zoologia - São Paulo, SP, Brasil.

    Rua do Matão, travessa 14, 321, Butantã, CEP 05508‑090, São Paulo, SP, Brasil.


Antunes-Carvalho, C. & Gnaspini, P. 2016. Pretarsus and distal margin of the terminal tarsomere as an unexplored character system for higher-level classification in Cholevinae (Coleoptera, Leiodidae). Systematic Entomology, 41: 392‑415.

Antunes-Carvalho, C.; Ribera, I.; Beutel, R.G. & Gnaspini, P. 2019. Morphology-based phylogenetic reconstruction of Cholevinae (Coleoptera: Leiodidae): a new view on higher-level relationships. Cladistics, 35: 1‑41.

Fresneda, J.; Grebennikov, V.V. & Ribera, I. 2011. The phylogenetic and geographic limits of Leptodirini (Insecta: Coleoptera: Leiodidae: Cholevinae), with a description of Sciaphyes shestakovi sp. n. from the Russian Far East. Arthropod Systematics and Phylogeny, 69: 99‑123.

Gnaspini, P. 1993. Brazilian Cholevidae (Coleoptera), with emphasis on cavernicolous species. II. Subgenus Iutururuca nov., subgenus of Adelopsis. Annales de la Société Entomologique de France, 29: 77‑87.

Gnaspini, P. 1994. The genus Eucatops (Coleoptera, Cholevidae, Eucatopinae) – description of new species and considerations on its systematic position. Iheringia (Zoologia), 76: 33‑42.

Gnaspini, P. 1996. Phylogenetic analysis of the tribe Ptomaphagini, with description of new Neotropical genera and species (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini). Papéis Avulsos de Zoologia, 39: 509‑556.

Gnaspini, P.; Antunes-Carvalho, C. & Leschen, R.A.B. 2017a. Pretarsal structures in Leiodidae and Agyrtidae (Coleoptera, Staphylinoidea). Journal of Morphology, 278: 1354‑1379.

Gnaspini, P.; Antunes-Carvalho, C.; Newton, A.F. & Leschen, R.A.B. 2017b. Show me your tenent setae and I tell you who you are – Telling the story of a neglected character complex with phylogenetic signals using Leiodidae (Coleoptera) as a case study. Arthropod Structure and Development, 46: 662‑685.

Jeannel, R. 1936. Monographie des Catopidae. Mémoires du Muséum National d’Histoire Naturelle (n.s.), 1: 1‑433.

Latella, L. 2010. Redescription of Eocatops ambiguous, Peyerimhoff, 1924 (Coleoptera, Cholevidae, Catopinae, Eucatopinae) from Libya. Bollettino del Museo Civico di Storia Naturale di Verona, Botanica, Zoologia, 34: 87‑91.

Newton, A.F. 1998. Phylogenetic problems, current classification and generic catalog of World Leiodidae (including Cholevidae). In: International Congress of Entomology, 20º. Proceedings. Phylogeny and evolution of subterranean and endogean Cholevidae (= Leiodidae Cholevinae) Torino, Museo Regionale di Scienze Naturali di Torino. p. 41‑177.

Newton, A.F. 2016. Leiodidae Fleming, 1821. In: Beutel, R.G. & Leschen, R.A.B. (Eds.). Coleoptera, Beetles, Volume 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2.ed. Berlin, Walter de Gruyter. v.1, p. 364‑376. (Handbook of Zoology: Arthropoda: Insecta)

Newton, A.F. 2019. StaphBase: Staphyliniformia world catalog database (version Nov 2018): Staphylinoidea, Hydrophiloidea, Synteliidae. In: Roskov, Y.; Ower, G.; Orrell, T.; Nicolson, D.; Bailly, N.; Kirk, P.; Bourgoin, T.; DeWalt, E.; Decock, W.; van Nieukerken, E.; Zarucchi, J. & Penev, L. (Eds.). Species 2000 & ITIS Catalogue of Life. Species 2000: Naturalis. Leiden, the Netherlands. Available at: Access in: 09/07/2019.

Perreau, M. 2000. Catalogue des Coléoptères Leiodidae Cholevinae et Platypsyllinae. Memoires de la Societé Entomologique de France, 4: 1‑460.

Perreau, M. 2019. Phylogeny of Bathysciotina Guéorguiev, 1974, based on morphology with a special emphasis to Italian genera and with the description of a new species of Halbherria (Coleoptera Leiodidae Cholevinae Leptodirini). Zootaxa, 4590: 367‑381.

Perreau, M. & Tronquet, M. 2002. Révision du genre Perriniella Jeannel et description d’une nouvelle espèce des Pyrénées françaises (Coleoptera: Leiodidae, Cholevinae, Leptodirini). Annales de la Societé Entomologique de France, 37: 481‑490.

Schilthuizen, M.; Perreau, M. & Njunjić, I. 2018. A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae). Zookeys, 777: 57‑108.

Wang, C.‑B. & Zhou, H.‑Z. 2015. Taxonomy of the genus Ptomaphaginus Portevin (Coleoptera: Leiodidae: Cholevinae: Ptomaphagini) from China, with description of eleven new species. Zootaxa, 3941: 301‑338.






Original Article

How to Cite

Gnaspini, P., Peck, S. B., Antunes-Carvalho, C., Perreau, M., & Gomyde, E. C. (2020). To be or not to be a tibial comb: A discussion on the (past) use of tibial armature in tribal/subtribal organization in Cholevinae (Coleoptera: Leiodidae). Papéis Avulsos De Zoologia, 60(special), e202060(s.i.).18.

Funding data