The cuckoo bee genus Sphecodes Latreille, 1804 (Hymenoptera, Halictidae) in Iran

An overview is provided of the Iranian fauna of cleptoparastic bee species in the genus Sphecodes Latreille (Halictinae: Halictini: Sphecodina). In total, 25 species are recorded from Iran, eight of which are newly recorded for the country: Sphecodes anatolicus Warncke, 1992, S. croaticus Meyer, 1922, S. haladai Warncke, 1992, S. pectoralis Morawitz, 1876, S. rubicundus Hagens, 1875, S. rufiventris (Panzer, 1798), S. saxicolus Warncke, 1992, and S. tadschicus Blüthgen, 1935. A new species S. ebmeri Astafurova & Proshchalykin, sp. n. (Mazandaran) is described and illustrated. For many species of Sphecodes the distribution data within Iran are enlarged.

The present paper is part of a series of works dealing with the bees of the genus Sphecodes of the territory of the Palaearctic region , 2015a, b, c, 2016a, b, 2017a, b, c, 2018, 2018a. Currently 77 species are known from this region, but the Sphecodes fauna of Iran is particularly under-recorded.
There are currently 862 species of bee known from Iran (Ascher and Pickering 2018), but new records and species continue to be added (e.g., Khodaparast and Monfared 2012, Khaghaninia et al. 2013, Nadimi et al. 2014Kuhlmann and Proshchalykin 2015, Safi et al. 2018, Proshchalykin and Kuhlmann 2018. The genus Sphecodes has been one of the more overlooked taxa partly due to the complexity of positively identifying material. Only one species -Sphecodes persicus Blüthgen, 1924(=S. pinguiculus Pérez, 1903 of this genus has been described from Iran (Blüthgen 1924) so far, and in total only 16 species have been recorded for the country (Warncke 1992, Ascher andPickering 2018). Records of Sphecodes from Iran in other published sources (Bogusch and Straka 2012, Özbek et al. 2015, Astafurova and Proshchalykin 2017b were not original and referenced from a previous paper (Warncke 1992).
Here we have attempted to build a survey of Iranian material available and to provide an overview of the 25 species found across Iran. Of these species, eight are recorded for the first time for the Iranian fauna and one species is described as new. For each species we provide a list of Iranian localities from the material examined and give previously published occurrences along with a summary of the wider distribution.

Materials and methods
The results presented in this paper are based on 324 specimens collected in Iran and currently housed in the Zoological Institute (St. Petersburg, Russia) and private collection of Maximilian Schwarz (Ansfelden, Austria). Acronyms for collections from which specimens were borrowed or are deposited are as follows: OÖLM -Oberösterreichisches Landesmuseum, Linz, Austria; PCMS -private collection of Maximilian Schwarz, Ansfelden, Austria; ZISP -Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia.
The taxonomy and distribution of species generally follows that of Warncke (1992), Bogusch and Straka (2012), Astafurova and Proshchalykin (2017b), and Özbek et al. (2015). A detailed current synonymy of the species has been given by Astafurova and Proshchalykin (2017b). Morphological terminology employed below follows that of Michener (2007) and Engel (2001). The ventral surfaces of some flagellomeres bear a distinctive patch or zone of sensilla trichodea A (sensu Årgent and Svensson 1982), and we refer to these as the "tyloids", easily observable under light microscopy. The abbreviations F, T, and S are used for flagellomere, metasomal tergum, and metasomal sternum, respectively. The density of integumental punctures is described using the following formula: puncture diameters (in μm) / ratio of distance between punctures to average puncture diameter, e.g., 15-20 μm / 0.5-1.5. Integumental sculpturing, aside from distinctive surface punctation, is described as follows: reticulate: superficially netlike or made up of a network of raised lines; rugose: irregular, nonparallel, wrinkled raised lines (rugae); tessellate: a regular network of shallow grooves with flat interspaces.
Specimens were studied with the use of a Leica M205A stereomicroscope and photographs taken with a combination of stereomicroscope Olympus SZX10 and digital camera Canon EOS70D. Final images representing a composite of several photographs taken at different focal planes and combined using Helicon Focus 6. All images were post-processed for contrast and brightness using Adobe Photoshop.
The species are presented alphabetically and those that could not be inspected in this paper are quoted from published sources. Provinces are presented in alphabetical order and the names of provinces are given in bold type. New distributional records are noted with an asterisk (*). Variation. Specimens from Iran differ from specimens from the rest of the range by having an entirely dark metasoma with distinctly punctate T2 in male. Diagnosis. The male is close to Sphecodes nomioidis Pesenko, 1979 owing to similar structure, sculpture of the body including distinctly punctate metasomal terga and similar small trapezoidal gonostylar shape (Fig. 1a, b). In gonostylar shape the new species is also close to S. geoffrellus (Kirby, 1802) ( Fig. 1 c), but differs from both species by having less developed tyloids covering about 1/2 ventral surface of flagellomeres ( Fig. 1d) (covering at least 4/5 ventral surface of flagellomeres in S. nomioidis and S. geoffrellus) (Fig. 1e).

Sphecodes croaticus
Description. (Male). (Fig. 1f ). Total body length 5.5 mm. Head ( Fig. 1g) black (except brown mouthparts and antenna); weakly transverse, at most 1.1 times as wide as long; vertex not elevated, distance from top of head to upper margin of a lateral ocellus about 2 lateral ocellar diameters as seen in dorsal view; antenna attain middle of mesoscutum, F1 transverse, 0.75 times as long as wide, F2 1.3 times as long as wide, remaining flagellomeres about 1.2 times as long as wide; tyloids (from F3 onwards) covering about 1/2 ventral surface of flagellomeres; clypeus, paraocular area and frons with confluent punctures (15-25 μm); area between ocellus and eye with punctures separated at most by 1.5 of a puncture diameter; vertex behind ocellus and gena rugose; face below antennal toruli with snow-white, plumose pubescence, obscuring the underlying integument.
Female unknown.
Etymology. This species is dedicated to the Austrian hymenopterist Andreas W. Ebmer in recognition of his outstanding contributions to bee researches.
General distribution. Only known from the type locality in Iran (Mazandaran).

Sphecodes pellucidus Smith, 1845
Published records. Ascher   Remark. We examined some specimens determined by K. Warncke, P. Bogusch, J. Starka and M. Schwarz from Iran deposited in PCMS and concluded that Sphecodes schenckii has been confused with S. tadschicus Blüthgen in Popov, 1935. The specimen recorded from Alborz Province (Ascher and Pickering 2018) probably also belongs to S. tadschicus Blüthgen in Popov, 1935. Nevertheless records of this species in north-western provinces of Iran are certain in the future as it is known from Caucasus and Turkey.

Discussion
Iran is a mostly mountainous country, with a landscape dominated by mountain ranges that separate various basins or plateaux from one another. A great variety of terrestrial ecosystems are situated on the territory of Iran: desert, semi-desert, steppe, forest steppe, forests and woodlands. However, dry biotopes occupy the majority of the Country and only 7% of the surface area is forested (most forest is found on the mountain slopes). Landscape and biotope diversity have a great effect on the diversity of bees, observable in the Sphecodes fauna of Iran, which includes forest species of temperate zoneas well as steppe and desert species of subtropics and endemic mountain species. The majority of the Sphecodes fauna of Iran is formed by 14 widespread species, distributed from Europe to West Siberia or even to the Russian Far East and Japan: These are S. albilabris, S. alternatus, S. crassus, S. ephippius, S. gibbus, S. longulus, S. monilicornis,  In total, 25 species of Sphecodes are recorded from Iran. This is distinctly less in comparison with the adjacent fauna of Turkey, Caucasus and Central Asia (Table 1). However this number will probably increase at least by a third and could be richer owing to eleven Western Palaearctic species which could be found in Iran: S. cristatus, S. hakkariensis, S. ferruginatus, S. nomioidis, S. dusmeti, S. niger, S. nurekensis, S. intermedius, S. armeniacus, S. crassanus, and S. geoffrellus. There are some species distributed in southern Europe, Turkey and the Caucasus that could be found in north-western Iran, whereas species mostly distributed in Central Asia could be recorded in central and eastern Iran. Because of these species the Iranian fauna of Sphecodes may be more diverse than those of Turkey or Central Asia.