Checklist of the Sarcophagidae (Diptera) of Croatia, with new records from Croatia and other Mediterranean countries

Abstract An updated checklist of Croatian flesh flies is presented based on the literature, on material collected from 2004 to 2017, and on specimens in museum collections. The checklist comprises 22 genera and 148 species (two left unnamed), 105 of which are represented by new Croatian records. Twenty-five species are recorded from Croatia with certainty for the first time: Amobiapelopei (Rondani, 1859), Apodacraseriemaculata Macquart, 1854, Craticulinatabaniformis (Fabricius, 1805), Macronychiastriginervis (Zetterstedt, 1838), Metopiacampestris (Fallén, 1810), Miltogrammabrevipila Villeneuve, 1911, Miltogrammaiberica Villeneuve, 1912, Miltogrammaoestracea (Fallén, 1820), Miltogrammapunctata Meigen, 1824, Oebalia cylindrica (Fallén, 1810), Phyllotelespictipennis Loew, 1844, Senotainiaconica (Fallén, 1810), Taxigrammahilarella (Zetterstedt, 1844), Taxigrammastictica (Meigen, 1830), Agriamonachae (Kramer, 1908), Nyctialugubris (Macquart, 1843), Blaesoxipha (Blaesoxipha) aurulenta Rohdendorf, 1937, Blaesoxipha (Blaesoxipha) batilligera Séguy, 1941, Blaesoxipha (Blaesoxipha) plumicornis (Zetterstedt, 1859), Sarcophaga (Helicophagella) okaliana (Lehrer, 1975), Sarcophaga (Heteronychia) amita Rondani, 1860, Sarcophaga (Heteronychia) ancilla Rondani, 1865, Sarcophaga (Heteronychia) pseudobenaci (Baranov, 1942), Sarcophaga (Myorhina) lunigera Böttcher, 1914 and Sarcophaga (Stackelbergeola) mehadiensis Böttcher, 1912. Taxigrammahilarella, Nyctialugubris, Agriamonachae, Blaesoxipha (Blaesoxipha) aurulenta and Sarcophaga (Heteronychia) amita are recorded from Southeast Europe with certainty for the first time. The species Sarcophaga (Sarcophaga) hennigi Lehrer, 1978 is omitted from the list, as previous records from Croatia are shown to be based on an erroneous synonymy with Sarcophaganovaki Baranov, 1941 (= Sarcophaga (Sarcophaga) croatica Baranov, 1941). Blaesoxipha (Blaesoxipha) rufipes (Macquart, 1839) could not be confirmed from Croatia and is not included in the checklist. Three new synonymies are proposed: Golania Lehrer, 2000 = Thyrsocnema Enderlein, 1928, syn. nov., Parasarcophaga (Liosarcophaga) kovatschevitchi Strukan, 1970 = Sarcophaga (Liosarcophaga) marshalli Parker, 1923, syn. nov., and Sarcophagasubvicinassp.novaki Baranov, 1941 = Sarcophaga (Sarcophaga) croatica Baranov, 1941, syn. nov. As part of an effort to update the European distributions of all Croatian species, the following new national and regional records are also provided: Miltogrammabrevipila, Miltogrammataeniata Meigen, 1824 and Sarcophaga (Heteronychia) pandellei (Rohdendorf, 1937) new to Greece; Sarcophaga (Liosarcophaga) harpax Pandellé, 1896 and Sarcophaga (Sarcophaga) croatica new to Italy (respectively mainland and mainland and Sicily); Miltogrammaiberica new to Bulgaria and Sardinia; Pterellaconvergens (Pandellé, 1895) new to mainland Italy and Sicily; Nyctialugubris new to mainland Italy and Sardinia; Blaesoxipha (Blaesoxipha) litoralis (Villeneuve, 1911) new to Sardinia and thus confirmed for Italy; Apodacraseriemaculata, Macronychiastriginervis, Protomiltogrammafasciata (Meigen, 1824) and Blaesoxipha (Blaesoxipha) ungulata (Pandellé, 1896) new to Sardinia and Sicily; Macronychiadolini Verves & Khrokalo, 2006, Macronychiapolyodon (Meigen, 1824), Metopiaargyrocephala (Meigen, 1824), Senotainiaalbifrons (Rondani, 1859), Taxigrammamultipunctata (Rondani, 1859), Taxigrammastictica, Blaesoxipha (Blaesoxipha) unicolor (Villeneuve, 1912) and Sarcophaga (Helicophagella) agnata Rondani, 1860 new to Sardinia; Metopodiapilicornis (Pandellé, 1895), Miltogrammaoestracea, Miltogrammarutilans Meigen, 1824, Nyctiahalterata (Panzer, 1798), Blaesoxipha (Blaesoxipha) lapidosa Pape, 1994 and Blaesoxipha (Blaesoxipha) plumicornis new to Sicily.


Introduction
Sarcophagidae, commonly known as flesh flies because many species feed on the soft tissues of animals , comprise ca 2800 species worldwide (Pape 1996 and unpublished) and are currently divided into three subfamilies: Miltogramminae, Paramacronychiinae and Sarcophaginae . The flesh fly fauna of Croatia has been poorly studied. The earliest data were published between the mid-19 th century and the beginning of the 20 th century (Schiner 1862; Brauer and Bergenstamm 1891;Böttcher 1912Böttcher , 1913Rohdendorf 1937), whereas the first studies more specifically focused on Croatian flesh flies were undertaken before and during World War II by Baranov ( , 1929Baranov ( , 1930Baranov ( , 1931Baranov ( , 1939Baranov ( , 1940Baranov ( , 1941aBaranov ( , 1941bBaranov ( , 1943, who provided information on the general morphology and taxonomy of Croatian Sarcophaginae as well as descriptions and identification keys. Baranov's work was continued by Strukan ( , 1968, and further data were added after the breakup of Yugoslavia by Povolný and Znojil (1994), , , Znojil (1998, 1999), ,  and Whitmore et al. (2013). The primary aim of this paper is to summarize all available data on the flesh fly fauna of Croatia, based on a critical review of literature records, on data obtained from recent collecting (2004)(2005)(2006)(2007)(2008)(2009)(2010)(2011)(2012)(2013)(2014)(2015)(2016)(2017), and on older specimens in the collections of the Natural History Museum of Denmark (Copenhagen), the National Museum of Natural History, Smithsonian Institution (Washington, D.C.), the Natural History Museum (London) and Museum für Naturkunde (Berlin). We also report 33 new national and regional species records for Bulgaria, Greece, Italy, Sardinia and Sicily.

Study area
Croatia has an area of 56,542 square kilometers and borders with Slovenia, Hungary, Bosnia and Herzegovina, Serbia, Montenegro and Italy. The country is divided into three biogeographical regions: Pannonian-Peripannonian, Alpine, and Mediterranean (Bertić et al. 2001). The Pannonian-Peripannonian region extends between the rivers Drava, Sava and Danube, the Alpine region covers the area of the Dinaric Alps, Gorski Kotar and Lika, whereas the Mediterranean region extends along the Adriatic coast and includes a multitude of islands, cliffs and ridges. The study area includes 38 localities belonging to the Mediterranean, Alpine and Pannonian-Peripannonian regions of Croatia (Figs 1-6, Table 1).

Specimen sampling and identification
The recent sampling effort was carried out between 2004 and 2017. Most flies were collected with hand nets, with additional specimens collected by Malaise traps, plastic bottle traps and multipurpose traps used to collect biting flies. Specimens were preserved in 96% ethanol or killed in ethyl acetate and pinned shortly after sampling. Male terminalia were extended in loco when specimens were still fresh, or male and female terminalia were dissected at a later stage following the method described by Richet et al. (2011). Male abdomens were removed and soaked in a 10% KOH solution for 72 hours, immersed in 10% acetic acid for one minute and rinsed in water for another minute before being dehydrated in beech-wood creosote for four hours; the phallus, pre-and postgonites, sternite 5, cerci and surstyli were separated from the rest of the abdomen, placed in a drop of Canada balsam on a microscope slide and covered with a coverslip. Female abdomens were detached, washed in an ethanol solution and left to macerate in a 10% KOH solution for one hour; abdomen tips were then separated by cutting the membrane between the last two tergites, and rinsed in acetic acid and water before being left between slide and coverslip in beech-wood creosote for a period of four hours; they were then mounted in Canada balsam between two coverslips. Identifications were carried out using current keys for Sarcophagidae (Pape 1987;Povolný and Verves 1997;Richet et al. 2011) and descriptions and illustrations in , Whitmore et al. (2013) and Whitmore and Perry (2018).

Results
Altogether, 1534 specimens were examined from Croatia and other Mediterranean countries. New locality records are provided for 132 species, including new Croatian records for 105 species. In Croatia, the highest number of species (72) was recorded in the Mediterranean region, followed by the Pannonian-Peripannonian region with 48 species and the Alpine region with 39 species. The most common species in this study was Sarcophaga (Sarcophaga) croatica Baranov, 1941 with 233 specimens from 12 localities. The next most common species were S. (Sarcophaga) (Pape 1994; Verves and Khrokalo 2014) based on a misreading of . Records of Sarcophaga (Sarcophaga) hennigi Lehrer, 1978
Distribution: Croatia, France (mainland), Italy (mainland). Remarks: Unverified record.  published a list of 1323 species in 50 Diptera families from Croatia, but these data are partly outdated (Britvec 2000) and many species may have been misidentified. Miltogramma contarinii is a rare species so far recorded with certainty only from Italy and France.

Sphenometopa steinii (Schiner, 1862)
Literature records: Dubrovnik (Schiner 1862; Strobl 1900); . European distribution: Croatia, Cyprus, Greece (mainland).  . Distribution: Croatia, France (mainland). Remarks: This species was probably described from Croatia (Dalmatia), even though the type locality was originally given by Stein (1924) as "Europe" (see .  listed it for Croatia but considered its presence in France as doubtful.  NB: These Sardinian specimens do not all fall within the range of variation of T. multipunctata defined by Richet et al. (2013), particularly with regard to setation of the parafacialia. Further studies are required to determine whether T. multipunctata sensu auct. is a morphologically variable species or a species complex.
Remarks: Larvae recorded on sheep by Mikačić (1938) were identified as belonging to W. magnifica. However, these records should be treated with caution because the larvae of Wohlfahrtia Brauer & Bergenstamm are difficult to identify to species level.
Remarks: Unverified record. Although male and female terminalia provide important characters for distinguishing W. meigenii from other closely related species of Wohlfahrtia, misidentifications in older generalist publications cannot be ruled out. The taxonomy of the nominal taxa W. meigenii (Schiner) and W. vigil (Walker) is unsettled, as discussed by Hall et al. (2009) and Ge et al. (2018). We are here using W. meigenii due to its Palaearctic type locality and because this was the name used by  and .
Remarks: This species was not listed for Sardinia by , but has been known from the island at least since .

Sarcophaga (Sarcophaga) baranoffi Rohdendorf, 1937
New Until now, these eight species have been recorded in one to three European countries only, and four of them were described based on specimens collected in Croatia (Baranov 1939(Baranov , 1941bPovolný 1987;. The other species recorded in this study are widely distributed throughout Europe and beyond . The 57 species recorded by Baranov ( , 1929Baranov ( , 1930Baranov ( , 1931Baranov ( , 1939Baranov ( , 1940Baranov ( , 1941aBaranov ( , 1941bBaranov ( , 1943 and the 18 species recorded by Strukan ( , 1968 were confirmed in this study. Our recent identifications and examination of the literature have enabled us to update the checklist of Croatian flesh flies to 148 species (including two left unnamed). The several new country records indicate that further studies of the flesh fly fauna of Croatia are necessary.