Quaderno di Studi e Notizie di Storia Naturale della Romagna Quad. Studi Nat. Romagna, 52: 111-174 (dicembre 2020) ISSN 1123-6787 Marco Villani Proposal of a new arrangement of the Amelini genera Ameles Burmeister, 1838 and Parameles Saussure, 1869 status restauratus, with taxonomic remarks on some species (Insecta: Mantodea: Amelidae) Abstract The taxonomy of the current Amelini genera Ameles Burmeister, 1838 and Pseudoyersinia Kirby, 1904 is substantially revised using many morphological features described in the text. Excluding the well characterized genus Apteromantis Werner, 1931, Amelini are divided into 2 genera: Ameles Burmeister, 1838 and Parameles Saussure, 1869 restored as a valid genus. The genus Ameles Burmeister, 1838 includes 4 subgenera (2 new): 1) subg. Ameles (nominotypical) including Ameles spallanzania, Ameles maroccana, Ameles gracilis, Ameles occidentalis combinatio nova and Ameles battistoni nomen novum; 2) subg. Apterameles status restauratus including Ameles heldreichi, Ameles decolor, Ameles syriensis, Ameles arabica, Ameles kervillei, Ameles massai, Ameles wadisirhani, Ameles dumonti, Ameles moralesi and Ameles confusa status novus; 3) subg. Canariameles subgenus novum including Ameles limbata, Ameles teydeana combinatio nova, Ameles canariensis combinatio nova, Ameles subaptera combinatio nova, Ameles betancuriae combinatio nova and Ameles pilipes combinatio nova; and 4) subg, Pilosameles subgenus novum including Ameles persa and Ameles aegyptiaca. The genus Parameles Saussure, 1869 status restauratus includes 4 subgenera (2 new): 1) subg. Parameles (nominotypical) including Parameles picteti and Parameles acuta species nova; 2) subg. Pseudoyersinia combinatio nova including Parameles brevipennis combinatio nova, Parameles lagrecai combinatio nova, Parameles inaspectata combinatio nova, Parameles kabilica combinatio nova and Parameles salvinae combinatio nova; 3) subg. Stenameles subgenus novum including Parameles poggii combinatio nova and Parameles melillensis status restauratus et combinatio nova, and 4) subg. Leptameles subgenus novum including Parameles paui combinatio nova, Parameles paradecolor combinatio nova, Parameles andreae combinatio nova and Parameles insularis combinatio nova. The nomen novum Ameles battistoni is established for Pseudoyersinia maroccana after its new combination (homonymy with Ameles maroccana Uvarov, 1931). Taxonomy of some taxa is discussed: Mantis abjecta should be considered a nomen dubium; synonymy of Mantis nana and Mantis brevis with Ameles spallanzania is confirmed; synonymy of Ameles fasciipennis and Ameles spallanzania obscura with Ameles spallanzania is proposed; synonymy of Mantis assoi with Parameles picteti is confirmed. The new species Parameles acuta is described from Andalucía, 111 Morocco and Algeria. It differs from Parameles picteti for more distinctly acute eyes and for shape of male genitalia. Key words: new subgenera, new species, Amelidae, Amelini, Ameles, Pseudoyersinia, Canariameles, Apterameles, Leptameles, Pilosameles, Stenameles, supra-specific arrangement. Riassunto [Proposta di una ristrutturazione sopra-specifica degli Amelini dei generi Ameles Burmeister, 1838 e Parameles Saussure, 1869 status restauratus, con osservazioni tassonomiche su alcune specie (Insecta: Mantodea: Amelidae)] La tassonomia degli attuali generi di Amelini Ameles Burmeister, 1838 e Pseudoyersinia Kirby, 1904 è sostanzialmente revisionata mediante l’uso di numerosi caratteri morfologici descritti nel testo. Escludendo il ben caratterizzato genere Apteromantis Werner, 1931, gli Amelini sono divisi in due generi: Ameles Burmeister, 1838 e Parameles Saussure, 1869 restituito al rango di genere. Il genere Ameles Burmeister, 1838 include 4 sottogeneri (2 nuovi): 1) Ameles (s. str.) che comprende Ameles spallanzania, Ameles maroccana, Ameles gracilis, Ameles occidentalis combinatio nova and Ameles battistoni nomen novum; 2) subg. Apterameles status restauratus et combinatio nova che comprende Ameles heldreichi, Ameles decolor, Ameles syriensis, Ameles arabica, Ameles kervillei, Ameles massai, Ameles wadisirhani, Ameles dumonti, Ameles moralesi e Ameles confusa status novus; 3) subg. Canariameles subgenus novum che comprende Ameles limbata, Ameles teydeana combinatio nova, Ameles canariensis combinatio nova, Ameles subaptera combinatio nova, Ameles betancuriae combinatio nova and Ameles pilipes combinatio nova; 4) subg. Pilosameles subgenus novum che comprende Ameles persa e Ameles aegyptiaca. Il genere Parameles Saussure, 1869 status restauratus comprende 4 sottogeneri (2 nuovi): 1) subg. Parameles (s.str.) che comprende Parameles picteti e Parameles acuta species nova; 2) subg. Pseudoyersinia combinatio nova che comprende Parameles brevipennis combinatio nova, Parameles lagrecai combinatio nova, Parameles inaspectata combinatio nova, Parameles kabilica combinatio nova e Parameles salvinae combinatio nova; 3) subg. Stenameles subgenus novum che comprende Parameles poggii combinatio nova e Parameles melillensis status rest. et combinatio nova; 4) subg. Leptameles subgenus novum che comprende Parameles paui combinatio nova, Parameles paradecolor combinatio nova, Parameles andreae combinatio nova e Parameles insularis combinatio nova. Si stabilisce il nomen novum Ameles battistoni per la specie Pseudoyersinia maroccana a seguito della sua nuova combinazione (omonimia con Ameles maroccana Uvarov, 1931). Si discute la tassonomia di alcuni taxa: Mantis abjecta deve essere considerata un nomen dubium; si confermano le sinonimie di Mantis nana e Mantis brevis con Ameles spallanzania; si propongono le sinonimie di Ameles spallanzania obscura e Ameles fasciipennis con Ameles spallanzania; si conferma la sinonimia di Mantis assoi con Parameles picteti. Si descrive la nuova specie Parameles acuta dell’Andalucia, del Marocco e dell’Algeria. Essa differisce da Parameles picteti per gli occhi più distintamente acuti e per la morfologia dei genitali maschili. Parole chiave: nuovi sottogeneri, nuova specie, Amelidae, Amelini, Ameles, Parameles, Pseudoyersinia, Canariameles, Apterameles, Leptameles, Pilosameles, Stenameles, revisione sopra-specifica. 112 Introduction The family Amelidae Westwood, 1889, according to the last revision (Schwarz et Roy, 2019) includes two tribes: Litaneutrini Jantsch, 1999, with three genera widespread in the arid areas of North America, and Amelini Westwood, 1889, widespread in Western Palearctic. More recent studies (starting from Giglio- Tos, 1927, with the addition of one genus by Uvarov, 1931) subdivide the Palearctic Amelini into three genera: Ameles Burmeister, 1838, characterized by brachypterous females and winged males, Pseudoyersinia Kirby, 1904, with both sexes brachypterous, and Apteromantis Werner, 1931, with both sexes completely apterous. However, no Author has studied critically this subdivision into genera, therefore the wings length continues to be adopted as a discriminating feature. Personal observations, corroborated by many morphological examinations combined with biogeographical considerations, take me to reject the previous generic arrangement of Amelini, with the exception of Apteromantis, which is an uniform and well-characterized genus with only two described species. The flying organs length should not be considered a valid character for generic division of Amelini anymore, for its high variability that presumably depends on environmental factors. In the morphological analysis less macroscopical but more constant features have been used: shape of the vertex (important in the generic subdivision), shape of the eyes (useful in the subgeneric division of Parameles), shape of pronotum (important at subgeneric level in Ameles), shape of fore femora (important at subgeneric level in Ameles), colour and pubescence of walking legs (adopted in generic and subgeneric subdivision), shape of cerci (useful in generic division), morphology of male genitalia (important at both genus and subgenus level). In this first contribution to the knowledge of the taxonomy of Amelini I mainly propose a supra-specific rearrangement, taking stock of some misinterpreted species (Mantis abjecta, Mantis nana, Parameles picteti, Mantis assoi, see the dedicated chapters). I postpone to further studies the treatment of single species of some groups. History - Burmeister (1838) in his description of Mantis (Ameles) lists 4 species, namely (in the order as they are listed): Mantis nana Charpentier, 1825, Mantis minima Charpentier, 1825, Mantis aurantiaca Burmeister, 1838 and Mantis flavicincta Oliver, 1792. The last three species were later assigned to different genera (in the order, Iris Saussure, 1869, Acontista Saussure, 1872, Otomantis Saussure, 1871). The first one is therefore considered the type species of genus Ameles. - Saussure (1869) raises Ameles to genus level (within tribus “Mantii”, subtribus 113 “Mantites”) and describes two other genera, which he nevertheless places in a different tribe for their conical eyes (tribus “Acanthopsii”, subtribus “Harpagites”): Yersinia and Parameles. According to Saussure, Ameles includes the species Ameles spallanzania (Rossi, 1792), with Mantis nana as synonym, and Ameles decolor (Charpentier, 1825). Yersinia includes Yersinia brevipennis (Yersin, 1860) (previously Mantis brevipennis) and Yersinia mexicana (Saussure, 1859) (described as Acanthops mexicana). Parameles includes only one species, described in the same work, Parameles picteti Saussure, 1869. - Saussure (1871) downgrades Yersinia and Parameles to subgenera of Ameles. The species Mantis abjecta Cyrillo, 1787 is recognized as a probable synonym of Ameles spallanzania. That is incorrect because the latter has been described 5 years after abjecta, whose name has accordingly the priority over spallanzania. Other taxa have been synonymized with this species: Mantis nana (probable, “?”), Mantis brevis Rambur, 1939 and Mantis soror Serville, 1839. In addition the Author put Mantis wagneri Kittary, 1849 within genus Ameles, subgenus Yersinia. Later the species was transferred into genus Oxyothespis Saussure, 1870 (Uvarov, 1912). Saussure mentions also Mantis limbata Brullé, 1838 and Mantis gracilis Brullé, 1838 as probable members of the genus Ameles, only on the basis of the original description and figures (Brullé, 1838). - Jacobson and Bianki (1902) report the species Ameles spallanzania as synonym of Mantis abjecta, correcting the priority mistake made by Saussure (1871). - Kirby (1904) raises again Yersinia and Parameles to genus level. He reports the species Ameles abjecta with the confirmed synonym of Mantis nana (type species of Ameles). The Author reports also the following species within genus Ameles: Ameles alliberti (Guérin-Méneville, 1843) (previously in the genus Perlamantis Guérin-Méneville, 1843; later it will return into its own genus, as Perlamantis allibeti, in Giglio-Tos, 1913), Ameles minima (not yet transferred within genus Iris), Ameles pusilla (Eversmann, 1854) (described as Mantis, then placed within Gonypeta Saussure, 1869 (Saussure, 1871), later in Armene Stål, 1877), Ameles paui Bolívar, 1898 and Ameles gracilis. In the same work Kirby reports in the genus Yersinia the species Yersinia mexicana and Yersinia solitaria Scudder, 1898 (later placed in genus Yersiniops Hebard, 1931). The species Yersinia brevipennis is placed in a new genus, Pseudoyersinia, with the species wagneri and, with doubt, aptera Fuente, 1894 (described as Ameles). Kirby puts within Parameles the following species: Parameles picteti, Parameles limbata, Parameles assoi (Bolívar, 1873) (described as Mantis), Parameles brevis, Parameles heldreichi (Brunner, 1882) (described as Ameles) and Parameles 114 taurica Jakovleff, 1903. Therefore the Author places in Parameles the species with conical eyes, the feature used by Saussure (1869) to distinguish it from Ameles. - Giglio-Tos (1927) places Parameles as a synonym of Ameles. The synonymy is justified by the fact that Ameles picteti (type species of Parameles) is considered synonym of Ameles nana (type species of Ameles). Ameles nana is therefore interpreted as an elongate species with conical eyes, and not a more robust one with rounded/ovoid eyes. Ameles soror is removed from the synonymy with Ameles abjecta stated by Saussure (1871)1, and Kirby (1904). According to the Author, Ameles soror differs from Ameles abjecta by its shorter tegmina (less than 4 times the pronotum length) and for female’s not-folded abdomen. Ameles brevis appears in the keys as a species close to Ameles nana, and it means that it is another species far from Ameles abjecta, being part of a group with more elongate body and conical eyes. Also Ameles modesta (Bolívar, 1914) has been misinterpreted by Giglio-Tos (and by Kirby) because it is placed next to the species of Parameles (more slender, with conical eyes) while the syntypes of Ameles modesta (MNCN, MNHN) are clearly specimens of Ameles spallanzania. Giglio-Tos places in the genus Pseudoyersinia also the species aptera, resolving the doubt of Kirby (1904) about its generic assignment. The Author clearly distinguishes it from the other species of the genus for the lack of flying organs. - Werner (1931) describes the genus Apteromantis for the species Ameles aptera Fuente, 1894 and Pseudoyersinia bolivari Werner, 1929. - Uvarov (1948) notes that Ameles nana and Ameles brevis are both described as species with a short pronotum, while other Authors (Giglio-Tos, 1927; Morales Agacino, 1947) considered them similar to Ameles picteti or synonyms of it. He establishes the synonymy of Mantis brevis with Mantis nana. Moreover the Author stresses that Ameles abjecta is a species with long pronotum and distinct from Ameles nana. - Beier (1950) describes the new genus and species Apterameles rammei from Macedonia. The description is based on a nymph of Ameles heldreichi (Kaltenbach, 1963). - Kaltenbach (1963) synonymizes Apterameles rammei with Ameles heldreichi. He also synonymizes Mantis nana with Mantis spallanzania. - Agabiti (2002) in her thesis deals with some Mediterranean species of Ameles. 24 species are mentioned, including Ameles abjecta as species inquirenda. In this work Ameles nana returns to be considered, wrongly, a species with long pronotum. - Battiston, Picciau, Fontana et Marshall (2010) mention 17 species of 1 Saussure (1871) synonymized Ameles soror with A. spallanzania, at that time wrongly deemed to take priority over A. abjecta. 115 Ameles and 13 of Pseudoyersinia in the Euro-Mediterranean area. Treatment of Ameles nana and Ameles assoi are the same as in Agabiti (2002). Ameles limbata is synonymized with Ameles gracilis without giving any reason. - Agabiti, Salvatrice et Lombardo (2010) mention 24 species of Ameles (excluding Ameles abjecta). Ameles nana continues to be considered a valid species with long pronotum and Ameles assoi is resurrected from synonymy with Ameles picteti (Agabiti, 2002) without giving a reason. Ameles modesta Bolívar, 1914 and Ameles africana Bolívar, 1914 become synonyms of Ameles spallanzania without explanation. Materials and methods A total of 181 specimens of 17 species have been examined. In addition to directly examined material, I obtained photos of type specimens of some species from the Museums and I have seen some of them on the high definition photos of some web sites (mnhn.fr; mantodearesearch.com): Ameles fasciipennis, Ameles spallanzania obscura, Ameles confusa, Ameles moralesi, Ameles kervillei, Parameles paui, Parameles brevipennis, Parameles lagrecai. I read the description of all known species of Palearctic species of Ameles and Parameles and further descriptions provided by other Authors, especially Agabiti (2002), Agabiti, Salvatrice et Lombardo (2010) and Battiston et alii (2018). On the occasion of a species-level revision of these genera I will examine type material of almost all known taxa, but it is not the intent of this study, which is a merely supra-specific revision of Amelini, with only some resolutions on a few critical species. Dried specimens have been studied using a stereomicroscope with 20x and 40x enlargement. Male genitalia have been mounted on a transparent acetate label and included in Canada Balsam or Euparal. The specimens are preserved in Marco Villani private collection, in the collectors' private collections or in museums, where specified. Abbreviations: MHNG: Muséum d’Histoire naturelle, Geneva, Switzerland MNHN: Muséum national d’Histoire naturelle, Paris, France MSNG: Museo Civico di Storia Naturale, Genova, Italy MNCN: Museo Nacional de Ciencias Naturales, Madrid, Spain. NMW: Naturhistorisches Museum Wien, Austria. 116 Results The result of this study is an almost complete taxonomic revision of the current genera Ameles and Pseudoyersinia. The genus Parameles Saussure, 1869, previously considered a synonym of Ameles (Giglio-Tos, 1927), is reinstated, and Pseudoyersinia becomes a subgenus of it. The genus Ameles is divided into 4 subgenera (2 of which are new) and the genus Parameles in 4 (2 new). Apterameles is rehabilitated to become a subgenus of Ameles. Many new combinations are consequently proposed. The genus Apteromantis has not been involved in this taxonomic study for its homogeneity and for the presence of only two described species. A review of Apteromantis can be found in Battiston, Ortego, Correas et Cordero (2014). The new taxonomic system I propose, adopted for Ameles and Parameles in this work, is the following. Genus Ameles Burmeister, 1838 Ameles (Ameles) Burmeister, 1838 Ameles (Ameles) battistoni nomen novum = Pseudoyersinia maroccana Battiston, Correas, Lombardo, Mouna, Payne et Schütte, 2018 nec Ameles (Ameles) maroccana Uvarov, 1931 Ameles (Ameles) gracilis (Brullé, 1838) sub Mantis gracilis Brullé, 1838 Ameles (Ameles) maroccana Uvarov, 1931 Ameles (Ameles) occidentalis (Bolívar, 1914) combinatio nova sub Pseudoyersinia occidentalis Bolívar, 1914 Ameles (Ameles) spallanzania (Rossi, 1792) sub Mantis spallanzania Rossi, 1792 = Mantis nana Charpentier, 1825 synonymum novum = Mantis brevis Rambur, 1839 synonymum novum = Mantis soror Serville, 1839 = Ameles abjecta africana Bolívar, 1914 = Parameles modesta Bolívar, 1914 = Ameles fasciipennis Kaltenbach, 1963 synonymum novum = Ameles spallanzania obscura Battiston, Correas, Lombardo, Mouna, Payne et Schütte, 2018 synonymum novum 117 Ameles (Canariameles) subgenus novum Ameles (Canariameles) limbata (Brullé, 1838) sub Mantis limbata Brullé, 1838 = Ameles canaria Koçak & Kemal, 2008 Ameles (Canariameles) betancuriae (Wiemers, 1993) comb. nova sub Pseudoyersinia betancuriae Wiemers, 1993 Ameles (Canariameles) canariensis (Chopard, 1942) comb. nova sub Pseudoyersinia canariensis Chopard, 1942 Ameles (Canariameles) pilipes (Chopard, 1954) comb. nova sub Pseudoyersinia pilipes Chopard, 1954 Ameles (Canariameles) subaptera (Chopard, 1942) comb. nova sub Pseudoyersinia subaptera Chopard, 1942 = Pseudoyersinia lindbergi Chopard, 1954 Ameles (Canariameles) teydeana (Chopard, 1942) comb. nova sub Pseudoyersinia teydeana Chopard, 1942 Ameles (Pilosameles) subgenus novum Ameles (Pilosameles) aegyptiaca Werner, 1913 Ameles (Pilosameles) persa Bolívar, 1911 = Ameles crassinervis Dirsch, 1927 Ameles (Apterameles) Beier, 1950 status restauratus et status novus Ameles (Apterameles) arabica Uvarov, 1939 Ameles (Apterameles) confusa Morales Agacino, 1948 status novus sub Ameles moralesi confusa Morales Agacino, 1948 Ameles (Apterameles) decolor (Charpentier, 1825) sub Mantis decolor Charpentier, 1825 Ameles (Apterameles) dumonti Chopard, 1943 Ameles (Apterameles) heldreichi Brunner, 1882 = Ameles cypria Uvarov, 1936 = Parameles taurica (Jakovlev, 1903) 118 = Apterameles rammei Beier, 1950 = Parameles shelkovnikovi Bogatchov, 1946 Ameles (Apterameles) kervillei Bolívar, 1911 Ameles (Apterameles) massai Battiston et Fontana, 2005 Ameles (Apterameles) moralesi Bolívar, 1936 Ameles (Apterameles) syriensis Giglio-Tos, 1915 Ameles (Apterameles) wadisirhani Kaltenbach, 1982 Genus Parameles Saussure, 1869 status restauratus Parameles (Parameles) Saussure, 1869 Parameles (Parameles) acuta species nova Parameles (Parameles) picteti Saussure, 1869 = Mantis assoi (Bolívar, 1873) synonymia restaurata Parameles (Pseudoyersinia) Kirby, 1904 status novus Parameles (Pseudoyersinia) brevipennis (Yersin, 1860) comb. nova sub Mantis brevipennis Yersin, 1860 Parameles (Pseudoyersinia) inaspectata (Lombardo, 1986) comb. nova sub Pseudoyersinia inaspectata Lombardo, 1986 Parameles (Pseudoyersinia) kabilica (Lombardo, 1986) comb. nova sub Pseudoyersinia kabilica Lombardo, 1986 Parameles (Pseudoyersinia) lagrecai (Lombardo, 1984) comb. nova sub Pseudoyersinia lagrecai Lombardo, 1984 Parameles (Pseudoyersinia) salvinae (Lombardo, 1986) comb. nova sub Pseudoyersinia salvinae Lombardo, 1986 Parameles (Stenameles) subgenus novum Parameles (Stenameles) melillensis Bolívar, 1914 status novus sub Parameles assoi melillensis Bolívar, 1914 119 Parameles (Stenameles) poggii (Lombardo, 1986) combinatio nova sub Ameles poggii Lombardo, 1986 Parameles (Leptameles) subgenus novum Parameles (Leptameles) andreae (Galvagni, 1976) combinatio nova sub Pseudoyersinia andreae Galvagni, 1976 Parameles (Leptameles) insularis (Agabiti, Salvatrice et Lombardo, 2010) combinatio nova sub Ameles insularis Agabiti, Salvatrice et Lombardo, 2010 Parameles (Leptameles) paradecolor (Agabiti, Salvatrice et Lombardo, 2010) combinatio nova sub Ameles paradecolor Agabiti, Salvatrice et Lombardo, 2010 Parameles (Leptameles) paui (Bolívar, 1898) combinatio nova sub Ameles paui Bolívar, 1898 This is the taxonomic arrangement I will use in the following pages. The features I will describe below the principal traits useful in the study of Amelini and discuss their use in the taxonomic arrangement of the tribe. 1) Vertex. Vertex in Amelini is essentially of two types: either convex, that is elevated in the middle at least in a moderate median trait; or straight, or even concave, sometimes slightly elevated only in a short median trait. In the first case the convexity appears less accentuated in males for the high development of eyes that leads to a compression of lateral margins of vertex. Species of Ameles have a constantly convex vertex, which is a typical feature of this genus, but in some species with a “derived” habitus the convexity is less accentuated and limited to the ventral trait due to the very developed eyes (Ameles pilipes, Ameles canariensis). The straight vertex is a prerogative of genus Parameles, and also of Apteromantis. The real appearance of vertex can be appreciated only in a perfectly frontal view of the head. Neanids at first stage of Parameles (examined species: Parameles lagrecai) have a straight vertex, like in the adult stage. In Ameles (examined species: Ameles spallanzania, Ameles heldreichi, Ameles decolor) the convexity of vertex is clearly visible already in the first stage. 120