C. R. Biologies 339 (2016) 427–436 Contents lists available at ScienceDirect Comptes Rendus Biologies ww w.sciencedirect.com Biodiversity/Biodiversite´ The geographical pattern of distribution of the genus Tityobuthus Pocock, 1890, a typical Ananterinae element endemic to Madagascar (Scorpiones: Buthidae) Sche´ma de distribution ge´ographique du genre Tityobuthus Pocock, 1890, Ananterinae ende´mique typique de Madagascar (Scorpiones: Buthidae) a, b c Wilson R. Lourenc¸o *, Patrick O. Waeber , Lucienne Wilme´ a Muse´um national d’Histoire naturelle, Sorbonne Universite´s, Institut de syste´matique, e´volution, biodiversite´ (ISYEB), UMR 7205 – CNRS, MNHN, UPMC, EPHE, CP 53, 57, rue Cuvier, 75005 Paris, France b ETH Zurich, Department of ITES, Forest Management and Development (ForDev) Group, Universita¨tstrasse 16, 8092 Zurich, Switzerland c Missouri Botanical Garden, Madagascar Research & Conservation Program, BP 3391, Antananarivo 101, Madagascar A R T I C L E I N F O A B S T R A C T Article history: New comments are proposed for the Ananterinae (sensu Pocock) or the ‘Ananteris Group’. Received 28 April 2016 The worldwide pattern of distribution of the elements associated with the Ananterinae, as Accepted after revision 6 June 2016 well as aspects of their ecology, is discussed. The biogeographic patterns presented by Available online 22 July 2016 extant and fossil elements of this group confirm not only the characteristics of a lineage representing a typical Gondwanian distribution, but correspond also to older Pangean Keywords: patterns. One new species is described in the genus Tityobuthus Pocock. This new species is Scorpion also a possible endemic element to the Island of Nosy-Be or at least to the Sambirano Ananterinae/Ananteris Group region. Generally, the Madagascar pattern of Tityobuthus is following the Neogrosphus rule, Biogeographic pattern showing typical high species richness with low dispersal when the ancestral population Ecology had a large niche breadth. Tityobuthus ß 2016 Acade´mie des sciences. Published by Elsevier Masson SAS. This is an open access New species article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/ 4.0/). R E´ S U M E´ Mots cle´s : De nouvelles conside´rations sur les Ananterinae (sensu Pocock) ou le « groupe Ananteris » Scorpion sont propose´es. Le mode`le tre`s e´largi de distribution des e´le´ments associe´s aux Ananterinae/groupe Ananteris Ananterinae, ainsi que des aspects de leur e´cologie, sont discute´s. Les mode`les Mode`le bioge´ographique bioge´ographiques pre´sente´s par des e´le´ments, aussi bien fossiles que modernes, associe´s E´ cologie a` ce groupe confirment non seulement les caracte´ristiques d’une ligne´e gondwanienne, Tityobuthus mais aussi celles d’un groupe avec des traits datant de la Pange´e. Une nouvelle espe`ce est Nouvelle espe`ce de´crite pour le genre Tityobuthus Pocock. Cette nouvelle espe`ce est e´galement un possible e´le´ment ende´mique de l’ıˆle de Nosy-Be ou pour le moins de la re´gion du Sambirano. Le genre Tityobuthus de Madagascar se conforme globalement a` la « Neogrosphus rule » en * Corresponding author. E-mail addresses: [email protected] (W.R. Lourenc¸o), [email protected] (P.O. Waeber), [email protected] (L. Wilme´). http://dx.doi.org/10.1016/j.crvi.2016.06.003 1631-0691/ß 2016 Acade´mie des sciences. Published by Elsevier Masson SAS. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/). 428 W.R. Lourenc¸o et al. / C. R. Biologies 339 (2016) 427–436 montrant une richesse spe´cifique e´leve´e, avec une dispersion limite´e lorsque la niche de la population ancestrale e´tait e´tendue. ß 2016 Acade´mie des sciences. Publie´ par Elsevier Masson SAS. Cet article est publie´ en Open Access sous licence CC BY-NC-ND (http://creativecommons.org/ licenses/by-nc-nd/4.0/). presented by this genus in Madagascar [5]. For further 1. Introduction historical details about this genus and its increasing species richness, see Lourenc¸o [5]. The genus Tityobuthus appears as a particular element within the Malagasy scorpion fauna. Despite being an endemic element to Madagascar, Tityobuthus has a close 2. The Ananterinae or ‘Ananteris Group’; history, relationship with other genera of the Ananterinae, or composition and known distribution ‘Ananteris Group’ [1], which are only present in quite distinct geographical regions of the world. The most As outlined in recent papers [1,6], the position of marked example is the genus Ananteris Thorell, 1891, an subfamilies within the family Buthidae has always been a element of the Neotropical scorpion fauna, which, howev- controversial matter [7,8]. The subfamily Ananterinae was er, clearly appears as quite distinct from the other buthid proposed by Pocock [9] to accommodate the genus elements of this fauna. Ananteris Thorell. Pocock wrote: ‘I propose to eliminate Tityobuthus was established by Pocock [2] for Tityobu- from this subfamily (Buthinae) the isolated Neotropical thus baroni (Pocock, 1890) previously included in the genus genus Ananteris, which differs strikingly from the rest of Rhoptrurus Karsch, 1886, because this name was already the family in the structure of the pectines. The subfamily preoccupied by that of a snake genus, Rhoptrura Peters, Ananterinae may be created for its reception’. Kraepelin 1858 [3]. Confusion existed about the genera Tityobuthus [10] added Ananterinae, previously described by Pocock Pocock, Pseudobuthus Pocock and Odonturus Karsch until [9] and the Tityinae, within the Buthidae family. The they were revised by Vachon [3], who finally included only definition of a subfamily Ananterinae, or of one ‘Ananteris two species in the genus Tityobuthus: Tityobuthus baroni group’, remains to be decided. For more details, see (Pocock, 1890) and Tityobuthus gracilis (Fage, 1946). The Lourenc¸o [1,6]. latter was originally described in the genus Babycurus A number of genera may be accommodated within the Karsch 1886. T. gracilis was finally accommodated in its Ananterinae or ‘Ananteris Group’. Some are strongly own genus by Lourenc¸o [4] as Troglotityobuthus speciose, as Ananteris, Tityobuthus or Lychas C.L. Koch, gracilis. Subsequently, several other species of Tityobuthus and can be distributed over broad geographical ranges, have been described from the island [5]. whereas others may be represented by only one or two In this current paper, we bring some new insights into species such as Lychasioides Vachon or Troglotityobuthus the biogeographic patterns presented by the genus Lourenc¸o and are endemic to rather narrow ranges. The Tityobuthus in Madagascar. One new species is also global extant distribution of the Ananteris group is tropical described, attesting to the high levels of diversity in South America, tropical and subtropical in Asia, Oceania Fig. 1. Distribution of the nine genera in the Ananteris group, as well as the three fossil Cenozoic genera on the global map of climate variability modified from Chan et al. [11]. (STR: seasonal temperature range; MAP: mean annual precipitation; DTR: diurnal temperature range). W.R. Lourenc¸o et al. / C. R. Biologies 339 (2016) 427–436 429 and Sub-Sahara Africa, in climatic conditions encompass- are given in millimetres. The trichobothrial notations ing humid, dry and arid ones [11] (Fig. 1). follow Vachon [23], and the morphological terminology mostly follows Vachon [24] and Hjelle [25]. Ananteris Thorell, 1891 – Almost all the Neotropical region. 4. Taxonomic treatment Ananteroides Borelli, 1911 – Africa; Guinea Bissau, Guinea, Ivory Coast, Mauritania, Sierra Leone. Family Buthidae C.L. Koch, 1837 Himalayotityobuthus Lourenc¸o, 1997 – India and Nepal. Subfamily Ananterinae Pocock, 1900 Lychas C.L. Koch, 1845 – Africa, India, Southeast Asia, Genus Tityobuthus Pocock, 1890 Australia, Indian Ocean and Western Pacific Ocean islands. Tityobuthus lokobe sp. n. (Fig. 2) Lychasioides Vachon, 1974 – Africa; Cameroun. Microananteris Lourenc¸o, 2003 – French Guiana. Type material. Madagascar, Nosy-Be, R.N. Lokobe. Microananteroides Rossi & Lourenc¸o, 2015 – Africa; 1 female holotype with 5 juvenile paratypes, 4 males & Ghana. 1 female (one juvenile was removed by M. Vachon), IX/ Tityobuthus Pocock, 1893 – Madagascar. 1947 (J. Millot leg.), battage (RS-1428), deposited in the Troglotityobuthus Lourenc¸o, 2000 – Madagascar (cave Muse´um national d’Histoire naturelle, Paris. species). Etymology. The specific name is a noun in apposition to yPalaeolychas Lourenc¸o & Weitschat, 1996 – Cenozoic the generic name and refers to the locality where the new fossil, Samland Peninsula, Baltic coast. species was collected. yPalaeoananteris Lourenc¸o & Weitschat, 2001 – Cenozoic Diagnosis. Scorpion of moderate size, with respect to fossil, Samland Peninsula, Baltic coast. the genus, and with 31–32 mm in total length. General yPalaeotityobuthus Lourenc¸o & Weitschat, 2000 – Cenozoic coloration pale yellow throughout, both the body and the fossil, Samland Peninsula, Baltic coast. appendages with some spots on the carapace, tergites, legs, and partially on pedipalps and metasomal segments. Among the several genera of Cenozoic fossil scorpions Carapace with a moderate concavity. Pectines large and described from the Samland Peninsula along the Baltic less separate than in females of other species with 14 to coast (today Poland and Kaliningrad in Russia), most are 15 teeth; fulcra reduced. Telson moderately elongated, related to the Ananterinae [12]. Some of these fossil with a short and curved aculeus; subaculear tooth strong, specimens remain rather incomplete and their morpholo- between rhomboid and spinoid, with two subaculear gy cannot be globally known. Nevertheless, their affinities granules. Internal face of the patella with 9–10 spinoid with the extant Ananterinae genera are well supported by granules. Tibial spurs strongly reduced on leg IV, absent on a number of characters [12–15]. leg III. Pedipalp fixed and movable fingers with 8–9 slightly Several recent studies [4,5,16,17] attest that Tityobuthus oblique rows of granules. Trichobothrial pattern of type A- shows closer affinities with the genus Ananteris than to the a (alpha) – orthobothriotaxic.