New members of the genera Neanura MacGillivray, 1893 and Deutonura Cassagnau, 1979 (Collembola: Neanuridae) from the Middle East

New species of Neanura MacGillivray, 1893 and Deutonura Cassagnau, 1979 are described from northern Iran. Both taxa are characterized by unusual features that place them in isolated positions within the genera. Neanura deharvengi sp. nov. differs from congeners by the extreme reduction of head chaetotaxy and fused lateral tubercles on the head. These characteristics of the new species broaden the existing diagnosis of the genus Neanura. An updated diagnosis is provided herein. Deutonura persica sp. nov. is most similar to D. plena (Stach, 1951), known from the Carpathians. The new species can be distinguished by the strong reduction of its head, labial, and labral chaetotaxy as well as the relative length of chaetae De2 and De3 on abdominal segments I–III. The Iranian records of D. decolorata (Gama & Gisin, 1964) are questioned. Brief remarks on the importance of the newly described species for the knowledge of both genera are also provided.


Introduction
In recent years, there have been a series of publications about Iranian species in the subfamily Neanurinae (Shayanmehr et al. 2013;Mayvan et al. 2015;Smolis et al. 2012Smolis et al. , 2016aSmolis et al. , 2016bSmolis et al. , 2017)).To date, the local fauna of this subfamily encompasses 13 species classifi ed into 8 genera.Due to the size and diversity of Iran these numbers are certainly an underestimate.Confi rming this, we supply the descriptions of two new Iranian species of this group.
The genus Neanura MacGillivray, 1893 is the oldest taxonomic unit in the subfamily.Cassagnau (1979) redefi ned the genus and split it into four subgenera: Neanura sensu stricto, Cryptonura, Deutonura and Endonura.Shortly after, this was expanded by Deharveng (1982b).As currently understood, Neanura includes only 7 species (Deharveng 1982b;Buşmachiu & Deharveng 2008;Smolis & Deharveng 2017).The distribution of most of them, excluding the cosmopolitan N. muscorum (Templeton, 1835), is restricted to Europe, especially to the western and central parts of the continent.Taxonomically, Neanura is probably most closely related to the Siberian genus Kalanura Smolis, 2007, from which it differs in the arrangement of lateral tubercles and the relative length of chaetae A and B on the head (Smolis 2007).
In contrast to the genus Neanura, Deutonura Cassagnau, 1979 is nowadays one of the largest genera of Neanurinae, comprising 57 valid species (e.g., Deharveng 1982a;Deharveng & Weiner 1984;Porco et al. 2010;Deharveng et al. 2015).The highest diversity of the genus is documented for western Europe; for example, 19 species have been recorded in France (Deharveng et al. 2015).Due to the fusion of tubercles Di and De on the head and tubercles Di on the penultimate abdominal segment, Deutonura strongly resembles the monotypic genus Albanura Deharveng, 1982, although it differs from the latter by having separate tubercles Di on abdominal tergite IV (fused in Albanura) and by the number of chaetae Di on abdomen V (3+3 or very rarely 2+2 chaetae in Deutonura, 2+2 in Albanura) (Deharveng 1982b).
In the present contribution, we describe Neanura deharvengi sp.nov.and Deutonura persica sp.nov.from northern Iran.The former species is especially interesting since it possesses two unusual characters: extreme reduction of cephalic chaetotaxy and fused lateral tubercles on the head.Nevertheless, this new species is also characterised by several features (i.e., 3+3 eyes, the cross-type arrangement of chaetae on the head, separateness of cephalic chaetae Dl and presence of male ventral organ) which are characteristic of the genus Neanura.It convinced us that a new genus shouldn't be erected for it at the moment and that the diagnosis of Neanura can be modifi ed to accommodate the new species.We additionally include short remarks on the published record of D. decolorata (Gama & Gisin, 1964) from Iran and on the global distribution of the genera Neanura and Deutonura.

Material and methods
Specimens were collected from soil and litter samples.The samples were extracted using a Berlese-Tullgren apparatus.Specimens were cleared in Nesbitt's fl uid, subsequently mounted on slides in Swan's medium and studied using a Nikon Eclipse E600 phase contrast microscope.Figures were drawn with a camera lucida and prepared for publication using Adobe Photoshop CS3.The material is deposited in the Department of Invertebrate Biology, Evolution and Conservation, Institute of Environmental Biology, University of Wrocław, Poland (DIBEC).
The following abbreviations are used in the text, fi gures and tables:
Since the new species described below is characterized by a number of characters unknown in other members of the genus, e.g., the absence of cephalic chaetae A and the fusion of lateral tubercles on the head into one mass, the currently accepted diagnosis of the genus as proposed by Deharveng (1982) is emended here.Moreover, three recently erected genera (Kalanura, Xylanura Smolis, 2011 andPersanura Mayvan et al., 2015;see Smolis 2007, 2011and Mayvan et al. 2015) morphologically strongly resemble the genus Neanura.As a result, its diagnosis requires extension based on new morphological characters (e.g., relative length of cephalic chaetae A and B, presence of tubercles Di on Abd., arrangement of tubercles on Abd.V, presence of male ventral organ) diagnostic and useful for generic characterization within the tribe Neanurini.

Emended diagnosis
Body colour blue to grey or whitish.

Etymology
The new species is dedicated to our colleague and friend Louis Deharveng for his magnifi cent contribution to our knowledge of Collembola.
LEGS.Claw without internal tooth.On tibiotarsi, chaeta M present and chaetae B4 and B5 relatively short and pointed.

Remarks
Neanura deharvengi sp.nov. is most similar to N. pallida Deharveng, 1979 andN. minuta Gisin, 1963, both of which are also characterized by the presence of chaetae L' on Abd.V, the absence of chaetae Ocp on the head and tubercles Di on Th.I.Besides the unique characters of the new species, a reduction

Etymology
The specifi c epithet 'persica' refers to the historical name of Iran, the terra typica of the new species.
LEGS.Claw without internal tooth.On tibiotarsi, chaeta M present and chaetae B4 and B5 relatively short and pointed.

Variability
A complete fusion of the cephalic tubercles Af and Cl has been observed in a single specimen.Nevertheless, based on the available material it is hard to assess defi nitively whether it is just an aberrant form or an example of morphological variation within the species.

Remarks
Using the recent published key of the genus (Deharveng et al. 2015), Deutonura persica sp.nov.seems to be most similar to D. plena (Stach, 1951), a species described and only known from the Western and Eastern Carpathians.These two taxa, however, can easily be distinguished using a number of characters: chaetae C and E on the head (absent in persica; present in plena), elementary tubercles DE and EE on the head (absent in persica; present in plena), number of lateral chaetae (L+So) on the head (8 in European Journal of Taxonomy 406: 1-16 ( 2018

Discussion
The two new taxa described in this paper broaden and enrich our knowledge of the genera Neanura and Deutonura, especially with regard to their morphology and biogeography.For example, as mentioned in the Introduction, the genus Neanura, excluding the cosmopolitan N. muscorum, is primarily restricted to western and central Europe.The discovery of N. deharvengi sp.nov.has shown that other species of the genus can be expected outside Europe.In addition, the new species is characterized by a peculiar set of characters, including the absence of chaetae A on the head.Although many genera and species of Neanurinae are characterized by a more or less advanced degree of reduction of cephalic chaetotaxy, these processes usually do not include the mentioned chaetae.For instance, within the Western Palaearctic a similar reduction of chaetae A is observed exclusively in the genus Bilobella Caroli, 1912, belonging to Paleonurini, another tribe of Neanurinae (e.g., Deharveng 1981;Smolis & Kaprus' 2008).
In contrast to the previous genus, the range of Deutonura is notably broader as its members are distributed both in the Western Palaearctic (western, southwestern, and central Europe, northern Africa; 46 species) and in the Eastern Palaearctic (Korea, Japan and Russian Far East; 11 species).Two species are exceptions to this general distribution pattern: D. frigida (Yosii, 1969), which ranges from central Siberia (the basin of the Yenisei river) to north-eastern America (Deharveng & Weiner 1984;Babenko & Fjellberg 2006) and D. gibbosa Porco, Bedos & Deharveng, 2010, which was probably introduced by humans to South Africa (Deharveng et al. 2015).In spite of the fact that more than 80% of the known members of the genus occur in the Western Palaearctic, the range of Deutonura in the region is highly concentrated and localized.For example, there are no species of Deutonura in such areas as the British Isles, Scandinavia, the Balkan Peninsula, Crimea, Asia Minor, or the islands of the East Mediterranean Basin (e.g., Rhodes, Crete, Cyprus).The absence of Deutonura in northern Europe is easy to understand, as the Neanurinae fauna of these areas is generally impoverished.The rest of the present picture can probably be explained by incompleteness of our knowledge.Nevertheless, some areas like Greece (including Crete and Rhodes), Albania, Bulgaria, Turkey, or the Caucasus have been more or less well investigated for Neanurinae diversity and many species belonging to the subfamily have been recorded there (e.g., Cassagnau & Péja 1979;Ellis 1976;Deharveng 1982aDeharveng , 1982b;;Smolis & Kuznetsova 2016).The observed gap in the known distribution of Deutonura is striking, and the answer to this question can undoubtedly be pivotal to understanding its history and evolution.
on Ant.IV iv = ordinary chaetae on ventral Ant.IV L' = ordinary lateral chaeta on Abd.V Mc = short macrochaeta Mcc = very short macrochaeta SMOLIS A. et al., New species of Neanura and Deutonura from the Middle
head separate.Tubercles Di on Th.I present or absent.Th.II-III with 3 chaetae Di.Tubercles Di on Abd.I-IV present.Tubercles Di never fused on abdominal segment IV, but fused, partially fused or separate on Abd.V.Each half of Abd.V with 3 or 2 chaetae Di.Cryptopygy absent or slightly developed.Abd.VI bilobed.Furcal remnant without microchaetae.Chaeta L' present or absent.Adult males usually with European Journal of Taxonomy 406: 1-16 (2018) modifi ed chaetae on ventral side of abdomen (male ventral organ).Tibiotarsi with 19, 19, 18 chaetae, clavate chaetae absent.Claw without inner tooth.

Table 2a .
(Deharveng 1979)o et al. 2010)13)p.nov.: cephalic chaetotaxy-dorsal side.Only one species of Deutonura, D. decolorata(Gama & Gisin, 1964), has previously been recorded(Cox 1982;Shayanmehr et al. 2013)from Iran.This species was described and is otherwise known only from the French Alps(Gisin 1964;Porco et al. 2010).Because of the fused cephalic tubercles Af and Cl, D. decolorata belongs to the small conjuncta species group, containing only 4 European species(Deharveng 1979).We have detected a similar fusion of cephalic tubercles in a single individual of D. persica sp.nov.(see Variability).In light of the known distribution of D. decolorata and the above observation, the record of D. decolorata from Iran should probably be treated as highly uncertain.