Updated Checklist of Vespidae (Hymenoptera: Vespoidea) in Iran

. 231 species of the family Vespidae (Hymenoptera, Vespoidea) of Iran, in 55 genera belonging to 4 subfamilies Eumeninae (45 genera, 184 species), Masarinae (5 genera, 24 species), Polistinae (2 genera, 17 species) and Vespinae (3 genera, 6 species) are listed. An overall assessment of the distribution pattern of the vespid species in Iran indicates a complex fauna of different biogeographic regions. 111 species are found in both Eastern and Western Palaearctic regions, while 67 species were found only in the Eastern Palaearctic region. Few species (14 species – 6.1%) of various genera are known as elements of central and western Asian area and their area of distribution is not known in Europe (West Palaearctic) and in the Far East. The species that were found both in the Oriental and Afrotropical Regions comprises 11.7 and 15.6% the Iranian vespid fauna, respectively. Many species (48, 20.8%) are exclusively recorded from Iran and as yet there is no record of these species from other countries. The highest percentage of the vespid species are recorded from Sistan-o Baluchestan (42 species, 18.2%), Alborz (42 species, 18.2%), Fars (39 species, 16.9%) and Tehran provinces (38 Species 16.5%), representing the fauna of the Southeastern, North- and South Central of the country.


Introduction
Vespidae is one of the largest and most diverse families of the order Hymenoptera with more than 5000 species worldwide in six subfamilies: Euparagiinae (single genus, 10 species), Masarinae (14 genera, 374 species), Eumeninae (204 genera, 3844 species), Stenogastrinae (7 genera, 72 species), Polistinae (25 genera, 1024 species), and Vespinae (4 genera, 69 species) (Pickett & Carpenter, 2010;Carpenter, unpubl.). Species of the family Vespidae play important roles in ecosystems, as biological control agents, since they feed their larvae with caterpillars and other insects (Ross & Matthews, 1991). They also serve as pollinators of various plants (Fateryga, 2010). Adult vespids are commonly black or brown in color, but often marked with yellow or white patterns. Most species are solitary, but many species are social (Goulet & Huber, 1993). The social species belong to the subfamilies Stenogastrinae, Polistinae and Vespinae, while all members of the Euparagiinae, Masarinae and Eumeninae live solitarily (Ross & Matthews, 1991). In solitary species, the larvae feed normally on larvae of other insects, especially caterpillars, within a cell constructed and provisioned by the adult female, but in the Masarinae the larva is supplied with a mixture of pollen and nectar instead. In social species the larva is progressively fed by the adult females on masticated insects (Ross & Matthews, 1991;Goulet & Huber, 1993).
In the last catalogue of Vespidae of Iran (Ebrahimi & Carpenter, 2008), 51 genera and 182 species were recorded with relevant data about distribution in various parts of the country. Since then, many other species have been recorded from Iran or their distribution in other parts of the country is documented. On the other hand, the status of some recorded species was changed from that considered in Ebrahimi & Carpenter (2008). Here we present an updated checklist of Vespidae from Iran with details based on the original references indicating the recent advances of the knowledge on the fauna of Vespidae in various parts of the country.

Material and methods
The known vespid species recorded from Iran were compiled and listed. Data about general distribution of the recorded species were compiled from the selected relevant literature. The provincial distribution of each species was also based on the locality records in the published data. Classification and nomenclature followed Carpenter (1996Carpenter ( , 2001, , Gusenleitner (2013Gusenleitner ( , 2019 and Schmid-Egger et al. (2017). The occurrence of species in Iran as originally indicated the junior synonyms under which the species were previously recorded from Iran are ignored. The valid name of each species is presented according to the last taxonomic modification. Other taxonomic changes and occurrence of the subspecies were also highlighted as separate remarks. Within the subfamilies the species are listed and presented alphabetically.

Remarks: According to Schmid-Egger et al. (2017) this species might be P. mongolicus du
Remarks: This species has been recorded from Iran as Paravespula germanica (Fabricius, 1793) .

Irrelevant records
A few vespid species are recorded from Iran on the basis of specimens which have never gone through the relevant identification by the experts mentioned as co-authors. The masarine species listed by Abd-Rabou et al. (2005) are obviously elements of the Afrotropical region (Gess & Gess, 2010) or have restricted distribution areas in southwestern Europe (Mauss & Castro, 2000, Mauss in litt. 2019, so that their occurrence in Iran is particularly unlikely. Considering the unreliable data that appeared in those publications, all the species records from these surveys are excluded from the fauna of Iran, until they may be found and confirmed in future reputable studies (Table 1). Gusenleitner, 2016;Celonites iranus Gusenleitner, 2018;Pseudosymmorphus angulatus Gusenleitner & Ockermüller, 2018; Pseudosymmorphus capillus Gusenleitner & Ockermüller, 2018 and Masaris agnolii Selis, 2019. None of these species are recorded from other countries yet.

Leptochilus simurgh
The genera Euodynerus, Polistes and Eumenes are the most species-rich groups in the vespid fauna of Iran with 18, 15 and 12 species respectively (Fig. 2) Giordani Soika, 1970;Gusenleitner, 1973b;Ebrahimi & Carpenter, 2008), represented by Sistan-o Baluchestan (26 genera, 42 species), Alborz (23 genera, 42 species), Fars (22,39) and Tehran (16,38) provinces, respectively. Some other provinces in the south-west (Khuzestan: 21, 34) and south-east (Kerman: 21, 31) were also explored considerably (Table 2). In his long series of taxonomic papers, Gusenleitner (1973bGusenleitner ( , 1985bGusenleitner ( , 1986bGusenleitner ( , 1995aGusenleitner ( , 1998cGusenleitner ( , 2000cGusenleitner ( , 2001Gusenleitner ( , 2006aGusenleitner ( , 2010aGusenleitner ( , 2011aGusenleitner ( , 2012aGusenleitner ( , 2012bGusenleitner ( , 2014Gusenleitner ( , 2016Gusenleitner ( , 2018 described 20 new species from 10 provinces, mainly from south and south-eastern areas in Fars, Hormozgan and Kerman provinces, with 6, 4 and 3 new species, respectively. The provincial distribution for many species is unclear, since the locality data are not recorded properly, or local names were used. Some of these species were recovered by the subsequent studies which provide the locality data. Furthermore, there were not sufficient data to assign the (five) species (Kohl & Handlirsch, 1889;Kostylev, 1935; recorded from the former Khorasan province, until it was subdivided into three separate provinces in 2004: Khorasan-e Razavi, North Khorasan and South Khorasan. More than 47 subspecies belonging mainly to the subfamily Eumeninae (43) (Fig. 1) are recorded from various regions of Iran Giordani Soika, 1970;Kurzenko, 1977;Gusenleitner, 2000b;2004;Ebrahimi & Carpenter, 2008;Gusenleitner, 2010a;Pashaei Rad et al., 2010;Gusenleitner, 2013). Some of these subspecies were later raised to separate species or synonymized (Kostylev, 1937(Kostylev, , 1940Giordani Soika, 1961;Ebrahimi & Carpenter, 2008). Further discussion about validity of these subspecies attributed to the knowledge about evolutionary patterns of the vespid lineages (Cumming, 1989;Carpenter, 2003). Additional investigations on both zoogeography and habitat preference, supplemented with molecular data are needed to define the taxonomic situation of these subspecies. A similar situation was also suggested for the sphecid fauna of Iran with many subspecies (Jahantigh et al., 2017). An overall analysis of the vespid fauna of Iran (Table 3) indicates the majority of species (77.1%) are distributed only in the (Eastern and Western) Palaearctic regions. The rest include the species that have a broader area of distribution, expanded into the Afrotropical Region, Oriental Region or both. A higher percentage of the species (11.3%) are found in Afrotropics compared to the Oriental Region (4.8%). There are few Palaearctic species that are found also in the Nearctic (1 species -Pseudepipona herrichii) or Neotropical region (1 species -Eumenes dubius). On the other hand, there are some species (Polistes olivaceus, Polistes dominula, Vespula germanica), that are widely distributed in various biogeographical regions both in the northern and southern Hemisphere. These species expanded their distribution through the association with human global transportation (Liebert et al., 2006;Masciocchi & Corley, 2013;Hammer & Jensen, 2019). While many areas remained unexplored, the known Vespidae of Iran indicate a rich fauna that needs further studies in different habitats. No comparison was done to determine the faunal similarity with the neighbouring countries, but the number of species (Turkey: 298 (species and subspecies) (Yildirim & Gusenleitner, 2012), Russia: 197 (Antropov & Fateryga, 2017), Pakistan: 79 (Rafi et al., 2017)) indicates a moderately well developed knowledge about the vespid fauna of Iran.