Orthoptera

Skejo, Josip, Rebrina, Fran, Szövényi, Gergely, Puskás, Gellért & Tvrtković, Nikola, 2018, The first annotated checklist of Croatian crickets and grasshoppers (Orthoptera: Ensifera, Caelifera), Zootaxa 4533 (1), pp. 1-95 : 9-16

Orthoptera research in Croatia through history

(refer to Fig. 2A, B, C )

The pioneers (1811–1846): Orthoptera research in Croatia started on the eastern Adriatic coast with Ernst Friedrich Germar— director of the Mineralogical Museum and professor at University of Halle in Germany. During a trip in 1811 he collected various insects (mostly beetles) in Fiume (= Rijeka), Veglia (= Krk Island), Cherso (= Cres Island), Dalmatia (part of the French Kingdom of Italy), Dubrovnik area (until 1808 Respublica Ragusa) and Cattaro (= Kotor). Germar (1817) reported nine Orthoptera species from Croatia and described a new species from Krk Island— Gryllus hystrix (currently in the genus Prionotropis ). French entomologist baron Pierre Francǫis Dejean (former division general in Napoleon's Army) has collected various insects (also predominantly beetles) in the coastal belt of the Austrian Empire during his exile in 1817. Dejean travelled on foot accompanied by an Istrian local (Kollar in Carrara 1846, Nonveiller 1999). Parts of his collection have, just as Germar’s, ended in different European museums and have been revised later. Following Germar's (1817) publication, Toussaint von Charpentier (1825) enlisted four grasshopper species from Dalmatia in his monography (Horae entomologicae: De Orthopteris Europaeis). Joseph Ullrich collected Orthoptera in Pola (= Pula) and surroundings in 1820 and sent specimens to Franz Xaver Fieber, a Czech entomologist ( Krauss 1879). Baron Ferenc Ocskay von Ocskó from Sopron (Ödenburg), visited Adriatic coast of the Austrian Empire (Croatian littoral) and collected Orthoptera ( Nonveiller 1999) . From Dalmatia, he described a new cricket species— Acheta dalmatina (today in the genus Gryllomorpha ) ( Ocskay 1833). Between Fiume (= Rijeka) and Carlopago (= Karlobag), Ocskay collected four specimens of undescribed bush-cricket species and sent them to Charpentier for examination ( Ocskay et al. 1850, Ocskay 1863). The species was named Barbitistes ocskayi and described first time in the study of Ocskay written in 1847, but published in 1863 ( Ocskay 1863), later than in the last Charpentier’s paper ( Ocskay et al. 1850), published after his death by his assistents and Ocskay personally, accompanied by originally Sturm’s drawings. From Germar's collection, German entomologist Karl Hermann Konrad Burmeister from Halle-Wittenberg described Phaneroptera macropoda (today Acrometopa servillea macropoda ) from Dalmatia, and Phalangopsis araneiformis (currently in the genus Dolichopoda ) from Ragusa (= Dubrovnik) (Burmeister 1838). Carl Theodor Ernst von Siebold, professor in Erlangen ( Germany), collected Orthoptera around Pola (= Pula) in Istria in 1842 ( Krauss 1879). In Carrara’s monography on Dalmatia (1846), Austrian entomologist Vincenz Kollar, curator of the Royal Zoological Cabinet in Vienna, published the first review of Dalmatian orthopterans, reporting 14 species. From the material collected during this period, a German zoologist and mineralogist Leopold Heinrich Fischer described Gryllus squamiger (currently in the genus Pseudomogoplistes ) on a female collected near Ragusa (= Dubrovnik) ( Fischer 1853). Fischer also reported Pezotettix pedestris (currently in the genus Podisma ) from specimens collected near Fiume (= Rijeka) and deposited in Rosenheim’s collection. In his review of European Orthoptera (1853) , Czech entomologist and botanist Franz Xaver Fieber recorded seven species from Dalmatia, including Saga serrata Chp. (today Saga pedo ), and described Ephippigera discoidalis (currently in the genus Ephippiger ) on a specimen collected by baron Ocskay.

Golden research period in Austria-Hungary (1853–1882): In 1851, a scientific society named Zoologisch- Botanische Verein (eng. Zoological-Botanical Society) was founded in Vienna. Together with the University in Vienna and the Royal Zoological Cabinet, it supported scientific expeditions within the Monarchy of Austria- Hungary and in other parts of the world. The society brought together scientists from different branches of natural

sciences, organized lectures and published presented data in the society journal. A member of the society Josef Mann—taxidermist and insect collector—who worked as a technician at the Imperial and Royal Zoological Court Cabinet (Vienna), collected in 1853 five Orthoptera species around Fiume (= Rijeka) ( Krauss 1879) and recorded in 1866 Thamnotrizon littoralis (currently in the genus Pholidoptera ) in Joseptahl (= Josipdol) on Velika Kapela Mt. ( Krauss 1879, Nonveiller 1999). Swiss physician Karl (Carolus) Brunner von Wattenwyl, telegraph-office director and orthopterist, later vice-president of Zoological-Botanical Society, traveled several times to Austrian Istria, Hungarian-Croatian litoral and Austrian Dalmatia to collect Orthoptera ( Schulthess 1915) . He also visited Adriatic islands—Lussin (= Lošinj), Cherso (= Cres), Veglia (= Krk) and Lesina (= Hvar). Brunner von Wattenwyl's collection in Vienna was one of the largest Orthoptera collections from that period ( Kaltenbach 2001). Brunner von Wattenwyl (1861) described Stetheophyma variegatum var. brevipennis (currently valid name = Arcyptera brevipennis brevipennis ) from Zengg (= Senj) and Fiume (= Rijeka). He also published a record of Rhaphidophora cavicola (currently in the genus Troglophilus ), after specimens collected in 1860 in Lesina (= Hvar) by a professional cave fauna collector Josef Erber (Brunner von Wattenwyl 1861). Georg Ritter von Frauenfeld, Austrian malacologist, entomologist and curator at the Vienna Museum (from 1851 to 1873 secretary of Zoologico-Botanical Society in Vienna), in 1854 and 1856 collected Orthoptera in Austrian Dalmatia. With Brunner von Wattenwyl’s help, he published data collected during his excursions, with comments on Germar’s records, and Brunner von Wattenwyl's descriptions of three new species ( Frauenfeld 1861), all later synonimized ( Cigliano et al. 2018). Frauenfeld recorded 44 Orthoptera species in Dalmatia, 17 for the first time in the region. In 1865, Johann Zelebor, Viennese painter and taxidermist, collected Gryllus miniatus (today Stenobothrus rubicundulus ) on Velebit Mt. (Visočica peak) ( Krauss 1879) ( Anonymous 1890). Veit Graber, Austrian biologist, was employed as a high school zoology teacher in Vinkovci (Slavonia) from 1868 to 1870 ( Jaworowski 1892). He was the first to start an inventory of Orthoptera in Slavonia along the Sava River from Brod (= Slavonski Brod) and Syrmia (= Srijem, Srem, Szerém) eastwards ( Graber 1868, 1870). In eastern Posavina and what is today Croatian Srijem (= Podunavlje) he found 39 species, out of which 24 were recorded for the first time in Croatia. While preparing taxonomic revision of bush cricket genera then included in Decticinae, Ottó Herman , a Hungarian entomologist and curator at the Natural History Museums in Kolozsvár (Cluj), Vienna and Budapest, received a manuscript from Brunner von Wattenwyl with taxonomic illustrations of two new species from Croatia, to be included in the paper ( Herman 1874). Those were Gampsocleis abbreviata from Sign (= Sinj), collected by Täubel, and Rhacocleis buchichii from Lesina (= Hvar Isl.) collected by Grgur Bučić. Hence, the author of these descriptions is Brunner von Wattenwyl, not Herman ( Cigliano et al. 2018). Giambattista Cubich (Ivan Krstitelj Kubić, Johann Kubik), a physician from Veglia (= Krk Isl.) and a member of Zoological-Botanical Society in Vienna, reported eight Orthoptera species from Krk Isl. in his monography ( Cubich 1875). Furthermore, Brunner von Wattewyl (1878) described Barbitistes yersini (lectotype from Obrovazzo—Obrovac). In the same paper, he included Fieber's description of Barbitistes elegans , assigning it to the genus Poecilimon (lectotype from Monte Maggiore—Učka Mt.) ( Ingrisch & Pavićević 2010). In the monograph (1882) on European Orthoptera, Brunner von Wattenwyl reported 51 species from Croatia, noteworthy Orphania denticauda (currently in the genus Polysarcus ), Isophya kraussi (reported as I. camptoxipha ), Platycleis roeselii (currently in the genus Roseliana) and Gryllus frontalis (currently in the genus Modicogryllus ). He also described Pachytrachelus frater (currently in the genus Pachytrachis ) from Austrian Dalmatia (syntypes from Ragusa—Dubrovnik, and Sabioncello—Pelješac) (Brunner von Wattenwyl 1882). Later, he reported further 11 species new to Croatian fauna. Swiss mineralogist and entomologist Henri Louis Frédéric de Saussure (1877) described Arachnocephalus dalmatinus (synonym of A. vestitus ) on a specimen from Dalmatia collected by Brunner von Wattenwyl. In this period, the most comprehensive overview of Croatian Orthoptera was published by Hermann Krauss, a German physician from Thübingen and an orthopterist closely colaborating with Brunner von Wattenwyl. In 1874 and 1877 he visited Austrian Istria, Hungarian-Croatian litoral (Kvarner), later Hvar Isl. and other parts of Dalmatia. In the paper on Istrian Orthoptera, Krauss (1879) reported 80 species altogether, of which 33 were recorded for the first time in Croatia. He also gave remarks on microhabitats and phenology. He was the first othopterist in Croatia to make remarks on bioacoustics of the studied species. In the paper ( Krauss 1879), he described Troglophilus neglectus from Clana (= Klana) near Rijeka, collected by Brunner von Wattenwyl, and Thamnotrizon dalmaticus (currently in the genus Pholidoptera ) from Tschitscheria (= Ćićarija Mt.), Grobnik above Rijeka, and Draga near Bakar, collected by Krauss. Ten years later, Krauss (1888) described Mogoplistes novaki (currently in the genus Paramogoplistes ) and Chorthippus lesinensis (currently regarded a subspecies of Chorthippus mollis ) from the material sent to him by Ivan Krstitelj Novak from Lesina (= Hvar Isl.).

First Croats taking part in the research (1886–1914): Grgur Bučić (Gregorio Bucchich) from Hvar on Hvar Isl. was the first Croatian entomologist to publish papers on Orthoptera . Working as head of post and telegraph office in Hvar, he was also a versatile naturalist and a correspodent member of Zoological-Botanical Society in Vienna. He kept contact with Brunner von Wattenwyl. Bučić published a review of Orthoptera collected on Hvar Isl. and Curzola (= Korčula Isl.) ( Bucchich 1886). He reported altogether 45 species from Korčula Isl. and 40 species from Hvar Isl., with notes on microhabitats for some species. Giam Battista Novak (Ivan Krstitelj Novak), a naturalist and school teacher in Brusje, Vrbanje (= Verbagno) and Hvar on Hvar Isl., collected insects in Dalmatia from 1875. He was a member of Zoological-Botanical Society in Vienna from 1884 and kept contact with Krauss, who identified Novak's Orthoptera material collected on the island. One year after retirement, Novak published a contribution to orthopteroid insect fauna of Hvar Isl. ( Novak 1888). He found five orthopteran species previously not recorded on the island, including the first records of Myrmecophilus species in Croatia. In the paper, Novak (1888) included observations about microhabitats and phenology of the species. In his third and final contribution on insects, Novak (1891) reported 11 orthopteran species for Spalato (= Split) and one for Zara (= Zadar). Eugen Adolf Jurinac, high school teacher in Varaždin and a collaborator of Zoological Museum in Zagreb, in summer 1883 investigated cave and terrestrial fauna in karst between Velika and Mala Kapela Mts. (Dinaric Alps region). He was the first to record Troglophilus neglectus outside a cave ( Jurinac 1887a). He recorded seven orthopteran species, including T. cavicola (in two caves), near Krapina ( Jurinac 1886), was the first to record three Orthoptera species on Ivanščica Mt. ( Jurinac 1887a), and reported 10 species from the surroundings of Oriovac (Slavonia) ( Jurinac 1887b). He identified the species after Brunner von Wattenwyl (1882). Anton Lodes, a forester on Krk Isl., reported Barbitistes ocskayi causing damage on trees ( Lodes 1897). Roman Puschnig, Austrian zoologist reported altogether 12 species of Orthoptera collected during his journey in Bosnia and Herzegovina and Dalmatia in 1895 ( Puschnig 1896), of which two frequent ones, Acridium aegyptium (today Anacridium aegyptium ) and Locusta viridissima (today Tettigonia viridissima ) recorded in Spalato and Salona (=Solin) in Croatia. The oldest Orthoptera specimens deposited in the Croatian Natural History Museum in Zagreb originated from 1880s, when first entomological collections were formed with encouragement from its director and curator Spiridion Brusina. Franjo Dobijaš (alias M. Padewieth, after his wife Matilda Padewieth), natural goods trader from Senj, between 1895 and 1898 collected Orthoptera along the Croatian littoral, and on the southern slopes of Velika Kapela Mt. and Velebit Mt. He reported altogether 104 species, of which 8 were recorded for the first time in the country, and provided locations and remarks about their microhabitats. Padewieth was in contact with Krauss, who helped him with identification. With Krauss' help, Padewieth (1900) described Platycleis kraussi (currently in the genus Bicolorana ) from Senj surroundings. A year prior to Padewieth's (1900) contribution, a Hungarian orthopterist Gyula Pungur published a checklist of Orthoptera of the Hungarian Kingdom ( Pungur 1899) based on literature and museum data (collection of the Budapest Natural History Museum). In 1868, a semi-autonomous Kingdom of Croatia-Slavonia was created by merging Kingdom of Croatia and Kingdom of Slavonia within the Hungarian Kingdom. Pungur's (1899) data relevant for Croatia were quoted as either Regio croatica (Pannonian region) or Regio adriatica (Croatian littoral—coastal part between Rijeka and Karlobag). Pungur reported altogether 100 species for the Kingdom of Croatia-Slavonia (60 Ensifera , 40 Caelifera ) of which two represented first records for the country— Psorodonotus Fieberi (currently Psorodonotus illyricus ) and Acrotylus insubricus . In this period, Austrian orthopterists took several scientific excursions along the Adriatic coast and islands. Researchers from the Trieste Zoological Station, supported by the Kaiserliche Akademie der Wissenschaften in Vienna, took an excursion from May to June 1901 across the Adriatic islands. Among the researchers were August Ginzberger, a botanist and research assistant at the University of Vienna and Egon Galvagni, an entomologist. They visited Lošinj, Susak, Rab, Kornati, Vis, Biševo, Sveti Andrija, Lastovo and Palagruža Islands ( Galvagni 1902, Werner in Ginzberger 1916). In 1906, a physician and entomologist Hugo Karny took part in an expedition to Meleda (= Mljet) (Karny 1907a, 1907b, 1908). In 1912, the University of Vienna organized an expedition from Spalato (= Split) to Brazza (= Brač Isl.) and Sabioncello (= Pelješac peninsula). Professor Franz Josef Maria Werner, a herpetologist and entomologist, published full account of this excursion with additional comments on his data from the 1880s expeditions to Adriatic islands ( Werner 1920). After Ebner's (1908) short review of findings in Austrian Southern Dalmatia (mostly Montenegro, only three species collected in Dalmatia), Hungarian orthopterist Dezső Kuthy (1908) published records from the Hungarian part ( Croatia, Slavonia, Dalmatia) of Austria-Hungary, but mostly providing the same data as Pungur (1899) and Karny (1907a). A record of Poecilimon affinis from Károlyváros (= Karlovac) is dubious. It could be but a misidentification of another Poecilimon species ( Szövényi & Puskás 2012) or erroneous toponym usage (Károlyváros could also stand for Karlovci in Srem, Serbia). Willy Adolf Theodor Ramme, curator of the Berlin Natural History Museum and a well-known orthopterist, visited Istria and Rijeka region in 1912 ( Ramme 1913). From the visited area he described a new variety— Oedipoda miniata var. meridionalis Ramme, 1913 (currently O. meridionalis ). Austrian forester Victor Apfelbeck, appointed curator of the Entomological Department of the National Museum in Sarajevo, and his assistant technician Adolf Winneguth, collected cave crickets on Korčula Isl. for the museum collection ( Karaman 1958). Umberto Girometta, high school biology teacher, mountaineer and speleologist from Split, collected cave crickets in Dalmatian caves ( Girometta 1913, 1914) and later donated the material to the newly founded Natural History Museum in Split.

Two World wars and the period between them (1914–1944): August Langhoffer, an entomologist of Hungarian origin, who worked as a high school teacher in Senj, collected numerous Orthoptera species from the surrounding area before 1892. His collection was unfortunately destroyed ( Padewieth 1900). While he was director of the Zoological Museum in Zagreb (1901–1927) and Entomology professor at the University of Zagreb, he organized cave fauna surveys, but without interesting results ( Langhoffer 1915a, 1915b). In 1926, Langhoffer wrote on pest grasshopper and bush cricket species causing tree damage along the southern slopes of Velebit Mt. ( Langhoffer 1928). Croatian entomologist Franjo pl. Košćec collected orthopterans in the surroundings of Varaždin (NW Croatia) between 1923 and 1956. His collection was deposited in the Varaždin City Museum. Petar Novak from Hvar, son of Ivan Krstitelj Novak, worked as a viticulture inspector and entomologist at the Department of Phytopathology (Fitopatološki odsjek) of the Experimental and Control Station for Agriculture (Poljoprivredna ogledna i kontrolna stanica) in Split, and as a director of the Natural History Museum in Split from 1946 to 1952. He had a large collection of various insects, mostly beetles. Giussepe Müller from Trieste, his high school friend and fellow entomologist who accompanied Novak on several field trips across Dalmatia, helped him identify orthopterans. Orthoptera material is currently deposited in the collection of Museo Civico in Trieste (revised by Roy Kleukers) and in the Natural History Museum in Belgrade. After the Second World War, Müller published records of Orthoptera collected during scientific excursions to Dugi otok Isl. and Kornati Archipelago, organized by the Yugoslav Academy of Sciences and Arts from 1925 to 1927 ( Müller 1957). He reported 20 species from Dugi otok Isl. and 2 from Kornat Isl. His record of Chorthippus mollis was the first faunistic record of this species in Dalmatia. He reported cave cricket specimens found in Strašna peć cave as an unknown subspecies of Troglophilus neglectus (after Karaman et al. (2011) they likely belong to T. ovuliformis ). Novak (1928) was the first who reported high density of Dociostaurus maroccanus populations around Sinj (Dalmatia) in 1923, causing crop damage. Richard Ebner, teacher, herpetologist and entomologist from Austria, described Psorodonotus fieberi illyricus (currently regarded a distinct species) from Mte Maggiore (= Učka Mt.) ( Ebner 1923). The species was reported from Croatia earlier as P. fieberi ( Pungur 1899) . In 1929, Ramme visited area around Senj (in search of Platycleis kraussi ) and the surroundings of Zagreb (to collect Isophya kraussi from the type locality). He described Isophya brevicauda , a new species from the surroundings of Samobor and Krapina and presented the first records of Poecilimon gracilis and P. schmidtii from Croatia ( Ramme 1931). Ramme sent grasshopper specimens collected above Senj and identified as Stenobothrus istrianus ( Ramme 1931) to a well-known orthopterist Sir Boris Petrovich Uvarov, specialised in Acridomorpha, who worked at the Imperial Bureau of Entomology in London. Uvarov compared those specimens with S. nigromaculatus istrianus paratypes, concluding they were not conspecific. Hence, Ramme described a new species— Stenobothrus croaticus ( Ramme 1933) . Before his emigration to England, German orthopterist and eminent paleontologist Frederick Everard Zeuner described Pholidoptera dalmatica maritima (in this study a synonym of P. dalmatica ) from Lesina (= Hvar Isl.) ( Zeuner 1931). Zeuner was employed at the Anti-Locust Research Centre in London, founded by Uvarov, where he earned his PhD in 1942 on Orthoptera related topic. Again, Ramme visited Dalmatia (Split, Hvar Isl.) and Dubrovnik area in 1939, collecting rare and not well-known species of Orthoptera from the region ( R. buchichii , P. frater ). Ramme (1951) published data from the expedition after the Second World War. Benno Wolf, a well-known German speleologist, gathered literature data and provided new records of cave crickets ( Dolichopoda araneiformis , Troglophilus neglectus , T. cavicola ) from Istria, Dalmatia, Dubrovnik area and Brač Isl. in the third volume of his Animalium Cavernarum Catalogus ( Wolf 1938). Pëtr A. Us, a Russian emigrant from Ukraine and highschool teacher in Pirot, Serbia, published records of 24 Orthoptera species from Papuk Mt. and Ravna gora Mt. („Slavonisches Waldgebirge“), collected by Austrian botanist August Ginzberger ( Us 1938). Us’ paper contained first faunistic records of Chrysochraon dispar and C. brachypterus (currently in the genus Euthystira ). Sándor Pongrácz (1944), an entomologist, director general of the Hungarian Natural History Museum in Budapest, published a short review of orthoptheran fauna of Kopački rit—area where the mouth of Drava River joins Danube („Drauwinkel“) in southern Baranja (part of the Kingdom of Hungary at the time). He provided the first faunistic records of Dociostaurus brevicollis , Chorthippus brunneus and Mecostethus parapleurus .

Decades under Tito's Yugoslavia (1945–1992): Imre von Igalffy, a Hungarian hunter and natural goods collector, donated his entomological collection (including many orthopterans) to Croatian National Zoological Museum in Zagreb. Most specimens were likely collected in the surroundings of his castle Trnovec in Gornja Pačetina, NW Croatia. Božidar Antolić, a tehnician at the Entomological collection of the Croatian National Zoological Museum, collected orthopterans in excursions on Pag Isl. from 1955 to 1962. Boža Pokopac (married name Tvrtković), high school teacher in Bjelovar, graduated from the Faculty of Science in 1956, with a thesis on Orthoptera from the surroundings of Bjelovar. The remains of her material are currently deposited in the Entomological Collections of the Division of Zoology (Department of Biology, Faculty of Science in Zagreb). Her collection has been revised recently (Šerić-Jelaska & Skejo 2014 after Pokopac 1956). Pokopac’ finding of Stenobothrus crassipes was the first record of the species in the country. Of originally reported 27 species and 286 specimens, the remaining collection includes 18 species and only 99 specimens, while the rest was lost or destroyed. Zora Karaman, an entomologist born in Istria to a Slovenian family (and wife of the well-known Yugoslav zoologist Stanko Luka Karaman), who worked for a short time as a curator at the Entomological Collection of the City Museum in Dubrovnik, published her and her husband’s findings of Troglophilus and Dolichopoda species from Dalmatia and Dubrovnik area ( Karaman 1958). The list also contained new speleological data from Vladimir Redenšek (Croatian entomologist and speleologist), Beatrica Đulić (Croatian mammalogist) and Sonja Mikšić (Bosnian orthopterist at the National Museum in Sarajevo), together with comments on previous records by Apfelbeck and Winneguth ( Karaman 1958). Reviewing firstly Macedonian ( Karaman 1960), then Palearctic Tetrigidae (1965) , Mladen Karaman reported Tetrix ceperoi from Cres Isl. as a new subspecies (T. c. balcanicus ), not valid anymore ( Harz 1969). Czech orthopterist Josef Mařan received grasshoppers from V. Švarc, collected in 1961 on Lastovo and Korčula Islands. From the material, Mařan described two new subspecies of Chorthippus lesinensis (at the time regarded as a species distinct from both C. vagans and C. mollis ) —C. l. lastovensis (from Lastovo Isl.) and C. l. svarci (from Korčula Isl.) ( Mařan 1965). The subspecies were later synonymized with the nominal one— C. mollis lesinensis , by Willemse et al. (2009). Russian orthopterist Oleg Grebenščikov and Serbian orthopterist Živko R. Adamović jointly visited Dubrovnik area in August 1946 and found D. araneiformis in Močiljska cave, as well as Locusta migratoria in the alluvial plain of Konavli ( Adamović 1964). In the same paper, Adamović reported 45 species for Dubrovnik area, with first faunistic records of Trigonidium cicindeloides (from Ombla) and Saga natoliae („in the surrouding country“ of Dubrovnik). From 1959 to 1964, Us collected orthopterans during his holidays (in June/July) at Cres Isl. and Lošinj Isl., reporting altogether 37 species ( Us 1964). Us (1970) published cave crickets findings from Croatia by Slovenian speleologist and coleopterogist Egon Pretner. He erroneously identified some Troglophilus neglectus specimens from caves northern of Plitvice as T. brevicauda ( Karaman et al. 2011). Adamović (1972) described Eupholidoptera chabrieri usi (currently a synonym of E. schmidti ) from Lošinj on the specimens from Us' collection deposited in the Slovenian Museum of Natural History in Ljubljana. With the help of ornithologist and orthopterist Sergej Matvejev, Us (1967) (often cited as Us & Matvejev 1967) prepared the first catalogue (checklist) of Yugoslav Orthopteroidea (Catalogus faunae Jugoslaviae—Orthopteroidea), reporting data for each constituent republic. Us (1967) divided Croatia into the following units, based on the available data: Istria, Kvarner, Dalmatia, and Slavonia. Us listed 94 Ensifera and 67 Caelifera , i.e. 161 orthopteran species for Croatia. However, 11 species were synonyms of other species in the list, and five were recorded in Austrian Dalmatia outside current Croatian borders. Thus, altogether 145 species were reported for Croatia. Voucher specimens collected for the catalogue were deposited in London, because Us sent them to Uvarov for the help with identification ( Nonveiller 1999). From the material collected earlier (see paragraphs above), Ramme prepared a monograph published in 1951. In this comprehensive overview, Ramme (1951) described Conocephalus dilatatus (currently synonymous with C. fuscus ) from the Ombla mouth (Dubrovnik area), Pholidoptera brachynota (in this study synonymized with P. dalmatica ) from the hills above Senj, Odontopodisma fallax from Veprinaz (= Veprinac) from Mte. Maggiore (= Učka Mt.), and O. decipiens from Krapina (NW Croatia). German orthopterist Kurt Harz published two comprehensive monographs (Die Orthopteren Europas), fundamental books on European Orthoptera—the first volume on Ensifera ( Harz 1969) and the second on Caelifera ( Harz 1975) . Furthermore, he described Barbitistes kaltenbachi on Brunner von Wattenwyl's material from Lesina (= Hvar) ( Harz 1965), Ephippiger ephippiger vicheti (currently synonymous with E. persicarius ) from Pisino (= Pazin) ( Harz 1966), Chorthippus bornhalmi (likely synonymous with C. maritimus ) on Bornhalm's specimens from Dubrovnik ( Harz 1971). In Brunner von Wattenwyl's Odontopodisma material, Harz (1973) found specimens from the surroundings of Rijeka, labeled ' O. fraudatrix sp.nov. ' by Ramme, and assigned them to a new species— O. rammei ( Harz 1973) . In the current checklist we provide evidence that O. rammei is but a synonym of O. fallax (see comments under O. fallax and O. rammei syn.nov.). Eminent Dutch orthopterist Ferdinand M. H. (Fer) Willemse (1973) revised the genus Paracaloptenus and assigned specimens from Učka Mt. to a new species— P. cristatus . Previously published P. caloptenoides records from Croatia also belonged to this species. Austrian entomologist Astrid Kohlich (1975) published the first faunistic record of Platycleis romana in Croatia, from the material collected during (the Institute of Zoology at the University of Vienna) students' excursion to Istria. Orthopterist Alfred Peter Kaltenbach from the Natural History Museum in Vienna helped her with identification. A well-known German orthopterist Sigfrid Ingrisch visited Istria twice and found numerous rare orthopteran species, including Andreiinimon nuptialis and Epacromius coerulipes ( Ingrisch 1981) . Italian orthopterist Antonio Galvagni (1986, 1987) revised the genus Miramella in Europe. He concluded that the records of the long-winged M. alpina from the Balkans (including old records from Croatia) belong to M. irena . Italian orthopterist Baccio Baccetti (1992) was the first to report Myrmecophilus myrmecophilus (from Split, Stalling pers. comm.) and Omocestus minutus (from Biševo Isl.) from Croatia. Of 13 Gryllotalpa species native to Europe, Baccetti described 7 of them (Baccetti & Capra 1978, Baccetti 1991, 1992) on the basis of chromosome numbers. Cytotaxonomic definition of south European species makes the systematics and identification of mole crickets rather difficult.

Systematic research in the last decades (1993–2018): Two Austrian excursions to Cres Isl. ( Kühling et al. 1998, Schuster et al. 1998) resulted in some surprising, but doubtful findings (e.g. Isophya sp., Pteronemobius lineolatus ), since they are not in accordance with the known distribution of these species in Europe ( Hochkirch et al. 2016) and our results. Bulgarian orthopterist Dragan Petrov Chobanov revised numerous endemic genera of the Balkans and Europe. While revising Bulgarian Orthoptera, Chobanov (2003) published a record of Stenonemobius bicolor ponticus from Zengg (=Senj), collected by Padewieth and correctly identified by Russian cricket specialist Andrey Vasilyevich Gorochov. Hungarian assistant lecturer at the University of Debrecen Antal Nagy surveyed orthopterans in the Žumberak and Samoborsko gorje Mts. Nature Park in 2003. Nagy (2006) reported 37 species, among them the first faunistic records of Metrioptera brachyptera and Chorthippus eisentrauti in Croatia. Furthermore, he found Mediterranean elements, e.g. Eupholidoptera schmidti (reported as E. chabrieri ) and Pholidoptera dalmatica , but did not provide exact localities. German biologists (not specialized in Orthoptera ) Andy Sombke and Matthias Schlegel studied Orthoptera of Istria and Šipan Isl. (near Dubrovnik) from 2001 to 2006. They published records of 36 species—23 for Šipan and 30 for Istria ( Sombke & Schlegel 2007). Some records are to be checked, especially those of Omocestus haemorrhoidalis (could represent O. minutus ), Acrotylus insubricus on Šipan Isl. and Chorthippus vagans in Vrsar (could represent C. mollis ignifer ). In the revision of the genus Gryllomorpha, Gorochov (2009) designated G. d. dalmatina neotype from Trsteno near Dubrovnik. Ingrisch revised numerous ensiferan taxa from the Balkans together with Serbian coleopterologist and orthopterist Dragan Pavićević. They described Poecilmon albolineatus from Durmitor Mt. in Montenegro, a species within the elegans group, numerous taxa from the ornatus group, and Leptophyes intermedia , a species from the punctatissima group ( Ingrisch & Pavićević 2010). Serbian entomologist Ivo Mladen Karaman, grandson of Stanko Luka and Zora Karaman, and son of Mladen Karaman, revised the genus Troglophilus in the western Balkans ( Karaman et al. 2011). They concluded that T. ovuliformis has wide distribution along the coast and on the Adriatic islands, but without the support of morphological and molecular data. Karaman assigned old records of T. brevicauda from Croatia ( Us 1970) to T. cavicola . Using a nuclear marker (16S), Karaman et al. (2011) identified N Dalmatia near Karin as the southern distribution border of T. neglectus . Karaman received Rhaphidophoridae material from Croatia for the study from a well-known Croatian speleologist Roman Ozimec. German lepidopterist and orthopterist Thomas Stalling, specialized in ant-loving crickets ( Myrmecophilidae ), reported Myrmecophilus hirticaudus from Crematogaster scutellaris (Olivier, 1792) nest on Cres Isl. ( Stalling & Birrer 2013). This was the first finding of the species in Croatia. There is a substantial lack of data on this orthopteran group in the country. Together with French orthopterist Michèle Lemonnier-Darcemont, Chobanov described T. balcanica— a new Tettigonia species from the Balkans, morphologically similar to T. cantans and T. silana Capra, 1936 . The authors dealt with specimens from a number of countries, including specimens we collected in Poštak Mt. ( Chobanov et al. 2014). German lepidopterist and orthopterist Wolfgang Wagner reported Rhacocleis buchichii from Pinus nigra forest on Biokovo Mt. ( Wagner 2015). Prior to Wagner’s records, the species was regarded island-endemic. Italian entomologist and evolutionary biologist Giuliana Allegrucci performed comprehensive revisions of Rhaphidophoridae in the Mediterranean (genera Dolichopoda and Troglophilus ) ( Allegrucci et al. 2009, 2011, 2014). Studying phylogeography of Troglophilus in the E Mediterranean, Allegrucci et al. (2017) discovered a new species from the brevicauda group on Mljet Isl. The species is to be described. From 2004 to 2006 the Croatian Natural History Museum in Zagreb was involved in PINMATRA project, that included collecting Orthoptera and assesments for the Habitats Directive ( Saga pedo , Pachytrachis bosniacus , Barbitistes yersini , Prionotropis hystrix , Paracaloptenus caloptenoides , Arcyptera b. brevipennis , Chrysochraon dispar , Acrida ungarica ). Expeditions took place in Gorski kotar, Lika, Biokovo Mt. and Snježnica Mt. The leading orthopterist in the project was Italian expert Filippo Maria Buzzetti (Buzzetti in Tvrtković & Veen 2006). In March 2012, undergraduate biology students Fran Rebrina and Josip Skejo (currently PhD students) of the Faculty of Science (University of Zagreb) founded the Section of Orthopterology (Sekcija za ravnokrilce; currently the Section for grasshoppers and crickets—Sekcija za skakavce i zrikavce) within the Biology Students Association—BIUS, under the mentorship of a Croatian trichopterist Mladen Kučinić, Croatian carabidologist Lucija Šerić Jelaska, and Buzzetti. After the first Section’s field expedition (Dinara Mt.), Rebrina and Skejo started collaboration with a Croatian zoologist Nikola Tvrtković, specialized primarily in Mammalia, a former director of the Croatian Natural History Museum. Striving to compile an annotated list and catalogue of Croatian Orthoptera, Rebrina, Skejo and Tvrtković started a collaboration with Hungarian orthopterists who had data on Croatian fauna—Gergely Szövényi, a senior lecturer at the Eötvös Loránd University and Gellért Puskás, a curator in the Hungarian Natural History Museum in Budapest. Szövényi & Puskás (2012) published the results of Orthoptera fauna survey in Papuk Mt. and surroundings (Slavonija) and recorded several species ( Isophya modestior , Poecilimon affinis , P. fussii , Tetrix bolivari , T. tenuicornis , Pseudopodisma fieberi ) for the first time in the country. From 2013, the authors initiated systematic research of Croatian Orthoptera , performing faunistic surveys and mapping species’ distributions in several parts of the country: the mountains of Hrvatsko Zagorje, Medvednica Mt., the mountains of Slavonija, the sands of N Croatia, Podravina region near Drava River, Mura-Drava Regional Park, the surroundings of Vinkovci (Pannonian region), Žumberak and Samoborsko gorje Mts., Učka Mt. and Ćićarija Mt., Risnjak Mt., Velika and Mala Kapela Mt., Velebit Mt. including Sjeverni Velebit National Park, Paklenica National Park and Velebit Nature Park, Lička Plješivica Mt., Grabovača in Lika region, Dinara Mt. with the surroundings, Troglav Mt., Kamešnica Mt., Moseć Mt., (Dinaric Alps), the surroundings of Rovinj, Pazin and Pula, including Rt Kamenjak (Istria), the surroundings of Senj and Rijeka (Kvarner), Dalmatian coast and Zagora from Zadar to Šibenik, Vransko jezero near Biograd, Krka National Park, Cetinska krajina, Imotsko polje, and isolated mountains Mosor Mt. and Biokovo Mt. (Dalmatia), Krk Isl., Cres Isl., Susak Isl., Unije Isl., Pag Isl., Dugi otok Isl., Čiovo Isl., Šolta Isl., Brač Isl., Hvar Isl., Lokrum Isl. (Adriatic islands). Material collected during the surveys is deposited in the collection of the Natural History Museum in Split (Josip Skejo Orthoptera collection), Fran Rebrina private Orthoptera collection in Zagreb, Gergely Szövényi private collection in Budapest, and the collection of the Hungarian Natural History Museum in Budapest. So far, systematic research resulted in a number of first faunistic records and important findings of rare species: Rammeihippus dinaricus ( Skejo & Rebrina 2013) , Leptophyes discoidalis ( Skejo & Stanković 2013) , Tetrix tuerki , T. undulata (Skejo et al. 2014) , Calliptamus barbarus ( Rebrina et al. 2015) , Acrotylus longipes ( Skejo & Sule 2015) , Gryllotalpa stepposa , G. gryllotalpa ( Bogdanović et al. 2017) , Paramogoplistes novaki deeply inland ( Rebrina & Brigić 2017), Pholidoptera frivaldszkyi ( Szövényi et al. 2018) , Epacromius tergestinus , Leptophyes intermedia , Modicogryllus truncatus , Oecanthus dulcisonans ( Puskás et al. 2018) as well as the description of a new subspecies of Tetrix transsylvanica—T. t. hypsocorypha (Skejo et al. 2014).

Study aims: The main objectives of the study are: (1) To present a list of Ensifera (crickets) and Caelifera (grasshoppers) species inhabiting Croatia, (2) To clarify species’ distributions in the country according to the historical (literature and museum) and new data (fieldwork and online social media), (3) To critically revise historical data, (4) To discuss nomenclatural and taxonomic issues, and (5) To provide a comprehensive bibliography on Croatian Orthoptera . This study is intended to be a base for further research of Orthoptera in Croatia.

Figures

FIGURE 2. History of Orthoptera research in Croatia A. Chart showing the number of first findings of Ensifera (blue) and Caelifera (red) species in Croatia per year. B. Chart showing cumulative number of Ensifera (blue), Caelifera (red) and all Orthoptera (yellow) species recorded in Croatia from the first days of Orthoptera research until today. C. Chart showing the number of first findings of Ensifera (blue) and Caelifera (red) species in Croatia per year and cumulative number of all Orthoptera (black) species recorded in Croatia from the first days of Orthoptera research until today. A and B show only the years in which Orthoptera study was conducted, while C shows full timescale.