Remarks on Synonyms of European Larinioides Species (Arachnida
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Arachnology (2015) 16 (8), 305–310 305 Remarks on synonyms of European Larinioides Only since the 1970s, in particular after publication of species (Arachnida: Araneae: Araneidae) Levi’s (1974) revision of the species (then placed in the genus Nuctenea), has the nomenclature of the three species been uniform worldwide. Rainer Breitling The situation was further complicated by the fact that, Faculty of Life Sciences, University of Manchester, for a long time, the existence of a vicariating sister species Manchester M1 7DN, UK of L. cornutus in southern and south-eastern Europe was not email: [email protected] widely realized. The difference between the northern and southern forms had been clearly established by Kulczyński Tobias Bauer (1901), but his work was published in a relatively obscure Hattenbachweg 12, series and neglected by most subsequent authors. This was 70599 Stuttgart, Germany probably also due to the fact that Kulczyński had used a email: [email protected] presumed junior synonym of L. cornutus, L. folium (Schrank, 1803), to refer to the southern species. The discovery of specimens of the southern form in Austria, and its careful Summary redescription by Thaler (1974), led to a considerable clar- ification and, in 1998, Levy determined that the earliest The five European species of the genus Larinioides are among available name for the species is L. suspicax, considering the most conspicuous spiders of the area, and some of them can be the dominant orb weavers of their habitat. Three of the species L. folium as a nomen dubium (Levy 1998). Since this work, were described by Clerck (1757), yet their nomenclatural history all five European species of Larinioides finally had widely since then has been marred by considerable confusion. Only in accepted valid names. the last 40 years has the nomenclature in the genus gradually However, recently it has been proposed that the names stabilized. Recently, however, it has been suggested to rename two of the species (Larinioides sclopetarius and L. suspicax) of two of the species be changed yet again. Mikhailov yet again. This note intends to show that these recent name (2013) suggested that Larinioides folium (Schrank, 1803) changes are not supported by the available evidence, and that is a valid species name and therefore the senior synonym of the established names L. sclopetarius and L. suspicax continue L. suspicax. This name change was accepted by Šestáková, to be valid. Marusik & Omelko (2014) in their careful revision of the entire genus. Šestáková, Marusik & Omelko (2014: 76) also suggested that “Araneus sclopetarius is confused with Introduction L. cornutus in the figures by Clerck (1757). Therefore the name Araneus sclopetarius Clerck, 1757 becomes a junior Five species of the orb-weaver genus Larinioides are synonym of L. cornutus and the senior synonym of A. sclo- known to occur in Europe. Three of them, L. cornutus, petarius, Araneus sericatus Clerck, 1757, is revalidated.” L. patagiatus, and L. sclopetarius, were described in the All European Larinioides species are large and conspic- founding document of spider nomenclature, Clerck’s (1757) uous orb weavers. Three of them have Holarctic distribu- Svenska Spindlar; a fourth one, L. ixobolus, was added tions and sometimes reach dramatic population densities in 1873 by Thorell (1873). The fifth species, L. suspicax in parasocial aggregations (L. sclopetarius) or can be the (O. Pickard-Cambridge, 1876), the Mediterranean sister dominant orb-weaver species in wetlands or field margins species of L. cornutus, was only relatively recently widely (L. cornutus). They have been the subject of a wide range accepted as an independent species (Thaler 1974). As with of biological studies, from morphology and behaviour to many of Clerck’s spiders, the nomenclature of Larinioides ecology and evolution (Ysnel 1990, 1993a,b; Ramousse species has been confused; for a long time, it was uncer- 1993; Ysnel, Lengwiler & Benz 1994; Sherman 1994; tain if Clerck’s names were actually available, as they were Heiling & Herberstein 1997, 1999a,b; Heiling 1999; Herber- published shortly before the 10th edition of Carl von Linné’s stein & Heiling 2001; Jeanson, Deneubourg & Theraulaz Systema Naturae, the official starting point of Zoological 2004; Schmitt 2004; Schmitt & Nioduschewski 2007a,b; nomenclature (ICZN 1959; Kronestedt 2010). Kleinteich & Schneider 2010, 2011; Sensenig et al. 2010; Moreover, the species are quite variable and widespread Jones et al. 2011a,b; Kralj-Fišer & Schneider 2012; Richter, and, even in Clerck’s original paper, five different names for Goebel & Schmitt 2012; Kralj-Fišer et al. 2013; Kleinteich, the three species were used. Many additional junior syno- Wilder & Schneider 2015; Shearer & Pruitt 2014). Changing nyms were introduced later by other early authors, repeat- the name of any of these species requires excellent argu- edly rediscovering the species across Europe. The identity ments, preferably based on new evidence. As the type spec- of the three Clerckian species was clarified by Westring imens of all affected species are lost, a detailed examination (1851) and especially by Thorell (1856, 1870), who also of the original descriptions and subsequent early revisions examined the available type material in Clerck’s collection is required in this case. in Stockholm (Thorell 1858), much of which is now lost (Kronestedt 2010). But, even afterwards, different names were often used, in particular due to disagreements about Status of Larinioides sclopetarius (Clerck, 1757) the priority of the simultaneously published synonyms in Clerck’s work (sclopetarius vs. sericatus; patagiatus vs. Šestáková, Marusik & Omelko (2014) suggested that ocellatus), sometimes without any justification (e.g. using Larinioides sclopetarius (Clerck, 1757) is a synonym of L. the junior synonyms foliatus or folium for L. cornutus). cornutus (Clerck, 1757), rather than of L. sericatus (Clerck, 306 Remarks on Larinioides Fig. 1: Habitus drawings of Araneus cornutus, A. sericatus and A. sclo- Fig. 2: Illustration of a male pedipalp referred to in the description of petarius in Clerck’s Svenska Spindlar. © The University of Man- Araneus sclopetarius in Clerck’s Svenska Spindlar. © The Univer- chester. sity of Manchester. 1757). They concluded that the species previously known as Šestáková, Marusik & Omelko (2014) are probably L. sclopetarius is now to be referred to as L. sericatus. They correct that the schematic, yet quite distinct, palpal illus- based this decision on the fact that the male pedipalp illus- tration (Fig. 2) is not showing the correct species, but this trated by Clerck, together with his description of Araneus figure was not intended as diagnostic. No comparative sclopetarius, shows characteristic features of L. cornutus. figures are shown for any of the other species of Araneidae, The text and figure of the habitus of Araneus sclope- nor are their male genitalia discussed. Clerck was aware of tarius, however, strongly suggest that the intended species the relevance of the male pedipalps as organs of reproduc- was indeed quite distinct from A. cornutus. For example, tion, and he even mentioned that their shape can be used to while the description of A. sericatus contains a cautionary discriminate species, but there is no indication that he ever note that this could turn out to be identical to A. cornutus, tried doing so for closely related species. The use of spider such a caveat is absent for A. sclopetarius. genitalia for species identification became widespread Clerck based his diagnosis of Araneus sclopetarius on its only in the 19th century. Clerck’s text also contains strong black abdomen with an off-white wavy margin and a char- internal evidence that the illustration of the pedipalp is not to acteristic pattern (1757: 43 “a figure explains better than a be considered an integral part of the description of Araneus description” [… som as ritningen bättre, än beskrivning, sclopetarius, but perhaps as a generic illustration (it is note- göres känbar …]; Fig. 1), as well as a white hairy margin of worthy that A. sclopetarius is the last of the large orb-web the carapace and a profusion of white fluffy hair in the eye weavers discussed in Clerck’s text). The text specifies that region. The relevance of this character is, however, not quite the name refers to the species illustrated in “Pl. 2. Tab. 3. clear, because Clerck drew his figures from living speci- Fig. 1.” (Clerck 1757: 43, i.e. the habitus drawing), and thus mens, which tend to show a much more prominent pilosity implicitly excludes the pedipalp drawing (Pl. 2. Tab. 3. Fig. than those preserved in alcohol. The associated figure shows 2). This differs strikingly from all other instances where the a (probably subadult) male but, from the text, there can be genitalia are illustrated; in those cases the name is always little doubt that it is intended to represent a general pattern applied to the entire Table, not a specified figure. Examples for both sexes. That the illustrated specimen is not adult from the same plate as A. sclopetarius are the illustrations is supported not only by the general body proportions and of A. segmentatus (Pl. 2. Tab. 6), A. pallidulus (Pl. 2. Tab. the lack of fully enlarged pedipalps but also by a compar- 7), A. labyrinthicus (Pl. 2. Tab. 8), and A. domesticus (Pl. 2. ison with the very similar illustration of a subadult male Tab. 9). of Araneus angulatus in Clerck’s work. In the latter case, It seems most likely that the description of the pedipalp, the description explicitly states that the depicted specimen although included in the description of A. sclopetarius, was not yet mature (1757: 22 “In mid-June I found a male, is intended to be applicable to all of the species of large which I fed until the 26th, hoping I would also find a female. orb-web weavers. We have found no indication in the text But, because it grew weak for two days, I illustrated it, that Clerck was aware that the species differed in the details considering it of ripe old age (see Pl.