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Zootaxa 4533 (1): 001–095 ISSN 1175-5326 (print edition)

http://www.mapress.com/j/zt/
Copyright © 2018 Magnolia Press
Monograph ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4533.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:C0A1AB26-D3D5-4DC3-B894-B18057AED2AC

ZOOTAXA
4533

The first annotated checklist of Croatian crickets and grasshoppers


(Orthoptera: Ensifera, Caelifera)

JOSIP SKEJO1,2, FRAN REBRINA2,3,7, GERGELY SZÖVÉNYI2,4,


GELLÉRT PUSKÁS2,5 & NIKOLA TVRTKOVIĆ6
1
University of Zagreb, Faculty of Science, Department of Biology, Division of Zoology, Evolution Lab, FR: Ecology Lab,
Rooseveltov trg 6 HR-10000 Zagreb, Croatia. E-mail: josip.skejo@biol.pmf.hr
2
IUCN/SSC Grasshopper Specialist Group
3
Bijenička 182, HR-10000 Zagreb, Croatia. E-mail: franrebrina@gmail.com
4
ELTE University, Department of Systematic Zoology & Ecology, Pázmány P. sétány 1/c, H–1117, Budapest, Hungary.
E-mail: szovenyig@gmail.com
5
Hungarian Natural History Museum, Department of Zoology, Baross utca 13, H-1088 Budapest, Hungary.
E-mail: puskas.gellert@nhmus.hu
6
Alagovićeva 21, HR-10000 Zagreb, Croatia. E-mail: nikolatvrtkovic71@gmail.com
7
Corresponding author

Magnolia Press
Auckland, New Zealand

Accepted by D. Rentz: 16 Jul. 201x; published: 21 Dec. 2018


JOSIP SKEJO, FRAN REBRINA, GERGELY SZÖVÉNYI, GELLÉRT PUSKÁS & NIKOLA
TVRTKOVIĆ
The first annotated checklist of Croatian crickets and grasshoppers (Orthoptera: Ensifera, Caelifera)
(Zootaxa 4533)
95 pp.; 30 cm.
21 Dec. 2018
ISBN 978-1-77670-558-0 (paperback)
ISBN 978-1-77670-559-7 (Online edition)

FIRST PUBLISHED IN 2018 BY


Magnolia Press
P.O. Box 41-383
Auckland 1346
New Zealand
e-mail: magnolia@mapress.com
http://www.mapress.com/j/zt

© 2018 Magnolia Press


All rights reserved.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any
means, without prior written permission from the publisher, to whom all requests to reproduce copyright
material should be directed in writing.
This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose
other than private research use.

ISSN 1175-5326 (Print edition)


ISSN 1175-5334 (Online edition)

2 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Sažetak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Orthoptera research in Croatia through history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Suborder Ensifera = zrikavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
infraorder Tettigoniidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
superfamily Tettigonioidea Krauss, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
family Tettigoniidae Krauss, 1902 = konjici . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
subfamily Bradyporinae Burmeister, 1838 = sedlarke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
genus Ephippiger Berthold, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
subfamily Conocephalinae Burmeister, 1838 = konjici čunjoglavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
genus Conocephalus Thunberg, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
genus Ruspolia Schulthess, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
subfamily Meconematinae Burmeister, 1838 = staklasti konjici . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
genus Cyrtaspis Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
genus Meconema Serville, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
subfamily Phaneropterinae Burmeister, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
genus Acrometopa Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
genus Andreiniimon Capra, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
genus Barbitistes Charpentier, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
genus Leptophyes Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
genus Poecilimon Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
genus Polysarcus Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
genus Phaneroptera Serville, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
genus Tylopsis Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
subfamily Saginae Brunner von Wattenwyl, 1878 = konjici vračevi / grabežljivi konjici . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
genus Saga Charpentier, 1825. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
subfamily Tettigoniinae Krauss, 1902 = pravi konjici . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
genus Bicolorana Zeuner, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
genus Decticus Serville, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
genus Eupholidoptera Mařan, 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
genus Gampsocleis Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
genus Metrioptera Wesmaël, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
genus Modestana Beier, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
genus Montana Zeuner, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
genus Pachytrachis Uvarov, 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
genus Pholidoptera Wesmaël, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
genus Platycleis Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
genus Psorodonotus Brunner von Wattenwyl, 1861 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
genus Rhacocleis Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
genus Roeseliana Zeuner, 1941. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
genus Sepiana Zeuner, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
genus Tessellana Zeuner, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
genus Tettigonia Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
genus Yersinella Ramme, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
genus Zeuneriana Ramme, 1951. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
infraorder Gryllidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
superfamily Grylloidea Laicharting, 1781. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
family Mogoplistidae Brunner von Wattenwyl, 1873 = ljuskavi šturci. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
subfamily Mogoplistinae Brunner von Wattenwyl, 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
genus Arachnocephalus Costa, 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
genus Mogoplistes Serville, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
genus Paramogoplistes Gorochov, 1984. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
genus Pseudomogoplistes Gorochov, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
family Trigonidiidae Saussure, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
subfamily Nemobiinae Saussure, 1877 = patuljasti šturci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
genus Pteronemobius Jacobson, 1907. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
genus Stenonemobius Gorochov, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
subfamily Trigonidiinae Saussure, 1874 = trčkoliki šturci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
genus Trigonidium Rambur, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 3
family Gryllidae Laicharting, 1781 = šturci / popci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
subfamily Gryllinae Laicharting, 1781 = pravi šturci / popci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
genus Acheta Fabricius, 1775 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
genus Eumodicogryllus Gorochov, 1986. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
genus Gryllus Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
genus Melanogryllus Chopard, 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
genus Modicogryllus Chopard, 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
subfamily Gryllomorphinae Saussure, 1877 = šturkolike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
genus Gryllomorpha Fieber, 1853. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
subfamily Oecanthinae Blanchard, 1845 = prozirni šturci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
genus Oecanthus Serville, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
superfamily Gryllotalpoidea Leach, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
family Gryllotalpidae Leach, 1815 = rovci / mrmci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
subfamily Gryllotalpinae Leach, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
genus Gryllotalpa Latreille, 1802 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
family Myrmecophilidae Saussure, 1874 = mravoljupci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
subfamily Myrmecophilinae Saussure, 1874. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
genus Myrmecophilus Berthold, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
superfamily Rhaphidophoroidea Walker, 1869 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
family Rhaphidophoridae Walker, 1869 = spiljski konjici / zrikavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
subfamily Dolichopodainae Brunner von Wattenwyl, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
genus Dolichopoda Bolívar, 1880 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
subfamily Troglophilinae Krauss, 1879. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
genus Troglophilus Krauss, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Omitted Ensifera species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
family Tettigoniidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
subfamily Phaneropterinae Burmeister, 1838 = listokrili i ljuskokrili konjici . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
subfamily Tettigoniinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
family Gryllidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
subfamily Nemobiinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
family Myrmecophilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
subfamily Myrmecophilinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
family Rhaphidophoridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
subfamily Dolichopodainae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
subfamily Aemodogryllinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
subfamily Troglophilinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Suborder Caelifera = skakavci. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
infraorder Tridactylidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
superfamily Tridactyloidea Brullé, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
family Tridactylidae Brullé, 1835 = buhoskakavci / ksije . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
subfamily Tridactylinae Brullé, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
genus Xya Latreille, 1809 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
infraorder Acrididea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
superfamily Tetrigoidea Rambur, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
family Tetrigidae Rambur, 1838 = monaški skakavci / trnovratke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
subfamily Tetriginae Rambur, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
genus Paratettix Bolívar, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
genus Tetrix Latreille, 1802. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
superfamily group Acridomorpha MacLeay, 1821 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
superfamily Acridoidea MacLeay, 1821 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
family Acrididae MacLeay, 1821 = pravi skakavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
subfamily Acridinae MacLeay, 1821 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
genus Acrida Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
genus Paracinema Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
subfamily Calliptaminae Tinkham, 1940 = krupnozadi skakavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
genus Calliptamus Serville, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
genus Paracaloptenus Bolívar, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
subfamily Catantopinae Brunner von Wattenwyl, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
genus Pezotettix Burmeister, 184 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
subfamily Cyrtacanthacridinae Kirby, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
tribe Cyrtacanthacridini Kirby, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
genus Anacridium Uvarov, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
genus Schistocerca Stål, 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
subfamily Gomphocerinae Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

4 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


genus Arcyptera Serville, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
genus Chorthippus Fieber, 1851 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
genus Chrysochraon Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
genus Dociostaurus Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
genus Euchorthippus Tarbinsky, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
genus Euthystira Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
genus Gomphocerippus Roberts, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
genus Myrmeleotettix Bolívar, 1914 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
genus Omocestus Bolívar, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
genus Pseudochorthippus Defaut, 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
genus Rammeihippus Woznessenskij, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
genus Stauroderus Bolívar, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
genus Stenobothrus Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
subfamily Melanoplinae Scudder, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
genus Miramella Dovnar-Zapolskij, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
genus Micropodisma Dovnar-Zapolskij, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
genus Odontopodisma Dovnar-Zapolskij, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
genus Podisma Berthold, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
genus Pseudopodisma Mistshenko, 1947 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
subfamily Oedipodinae Walker, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
genus Acrotylus Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
genus Aiolopus Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
genus Epacromius Uvarov, 1942. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
genus Locusta Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
genus Mecostethus Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
genus Oedaleus Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
genus Oedipoda Latreille, 1829 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
genus Psophus Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
genus Sphingonotus Fieber, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
genus Stethophyma Fischer, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
family Pamphagidae Burmeister, 1840 = žaboliki skakavci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
subfamily Thrinchinae Stål, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
genus Prionotropis Fieber, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Omitted Caelifera species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
family Tetrigidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
subfamily Tetriginae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
family Acrididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
subfamily Calliptaminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
subfamily Gomphocerinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
subfamily Melanoplinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
subfamily Oedipodinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Crickets and bush-crickets (Ensifera) overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Overview of Grylloidea families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Overview of Gryllotalpoidea families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Overview of Rhaphidophoroidea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Overview of Tettigonioidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Grasshoppers, groundhoppers and molehoppers (Caelifera) overview. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Overview of Acridoidea families. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Overview of Tetrigoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Overview of Tridactyloidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Acknowledgments and authors' contributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
IUCN Assessments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 5
Abstract

The first annotated checklist of crickets and grasshoppers (Orthoptera) of Croatia is presented. With 184 orthopteran spe-
cies, 103 Ensifera and 81 Caelifera, known to inhabit the country, Croatia is among the richest European countries in terms
of Orthoptera diversity. Altogether 25 species erroneously reported from the country are omitted from the checklist, 16
Ensifera (Isophya speciosa, Poecilimon brunneri, P. jonicus, P. thoracicus, Modestana ebneri, Pachytrachis bosniacus,
Rhacocleis neglecta, Tessellana carinata, T. nigrosignata, Zeuneriana marmorata, Pteronemobius lineolatus, Myrmeco-
philus acervorum, M. ochraceus, Dolichopoda palpata, Diestrammena asynamora, Troglophilus brevicauda) and 9 Cae-
lifera (Tetrix kraussi, Paracaloptenus caloptenoides, Chorthippus albomarginatus, Omocestus viridulus,
Pseudochorthippus montanus, Miramella alpina, Celes variabilis, Oedipoda germanica, O. miniata). First faunistic re-
cords of 10 taxa are reported for Croatia, in total four Ensifera (Leptophyes punctatissima, Metrioptera hoermanni, Ze-
uneriana amplipennis, Gryllotalpa sp.) and six Caelifera (Xya variegata, Chorthippus dichrous, C. loratus, C. mollis
ignifer, Odontopodisma sp., Acrotylus l. longipes). For each listed species, its distribution in Croatia and in Europe is giv-
en, and IUCN European Red List status is shown for species within threatened categories. Numerous distributional, tax-
onomic and nomenclatural problems are discussed. Several taxa with poorly defined diagnostic traits are synonymized,
namely Gampsocleis abbreviata renei syn.nov. (with G. a. abbreviata), Pholidoptera maritima syn.nov. (with P. dalmat-
ica), P. brachynota syn.nov. (with P. dalmatica), Acrida m. mediterranea syn.nov. (with A. u. ungarica), Chrysochraon
dispar intermedius syn.nov. (with C. d. giganteus) and Odontopodisma rammei syn.nov. (with O. fallax).

Key words: taxononomy, distribution, faunistics, Pannonian, Dinaric Alps, Mediteerranean

Sažetak

Predstavljen je prvi popis ravnokrilaca (Orthoptera) Hrvatske. Sa 184 vrste, 103 vrste zrikavaca i 81 vrste skakavaca,
Hrvatska je među najbogatijim europskim zemljama s obzirom na raznolikost ravnokrilaca. Ukupno 25 vrsta koje su
pogrešno zabilježene u Hrvatskoj izostavljeno je s popisa, 16 vrsta zrikavaca (Isophya speciosa, Poecilimon brunneri, P.
jonicus, P. thoracicus, Modestana ebneri, Pachytrachis bosniacus, Rhacocleis neglecta, Tessellana carinata, T. nigrosig-
nata, Zeuneriana marmorata, Pteronemobius lineolatus, Myrmecophilus acervorum, M. ochraceus, Dolichopoda palpa-
ta, Diestrammena asynamora, Troglophilus brevicauda) i 9 vrsta skakavaca (Tetrix kraussi, Paracaloptenus
caloptenoides, Chorthippus albomarginatus, Omocestus viridulus, Pseudochorthippus montanus, Miramella alpina, Ce-
les variabilis, Oedipoda germanica, O. miniata). Objavljeni su prvi faunistički nalazi 10 svojti u Hrvatskoj, ukupno četiri
vrste zrikavaca (Leptophyes punctatissima, Metrioptera hoermanni, Zeuneriana amplipennis, Gryllotalpa sp.) i šest svojti
skakavaca (Xya variegata, Chorthippus dichrous, C. loratus, C. mollis ignifer, Odontopodisma sp., Acrotylus l. longipes).
Za svaku vrstu na popisu navedena je rasprostranjenost u Hrvatskoj i u Europi, a status ugroženosti u Europi prema IUCN-
u prikazan je za vrste koje se nalaze unutar ugroženih kategorija. Raspravljeni su brojni problemi vezani uz rasprostran-
jenost, taksonomiju i nomenklaturu pojedinih vrsta. Sinonimizirano je nekoliko svojti s loše definiranim dijagnostičkim
svojstvima, redom Gampsocleis abbreviata renei syn.nov. (s G. a. abbreviata), Pholidoptera maritima syn.nov. (s P. dal-
matica), P. brachynota syn.nov. (s P. dalmatica), Acrida m. mediterranea syn.nov. (s A. u. ungarica), Chrysochraon dis-
par intermedius syn.nov. (s C. d. giganteus), i Odontopodisma rammei syn.nov. (s O. fallax).

Ključne riječi: taksonomija, rasprostranjenost, faunistika, Panonska nizina, Dinaridi, Mediteran

Introduction

Grasshoppers and crickets (order Orthoptera) are the largest extant polyneopteran order, with about 29 000 described
species and subspecies worldwide (Cigliano et al. 2018). Members of the order are characterized by the presence of
cryptopleuron (pronotum with well-developed paranota), hind legs enlarged for jumping (also used for stridulation in
some groups), tegmina (membranaceous fore wings used in many groups for stridulation), and tympana. The order is
divided into two suborders—(1) Ensifera (crickets, hrv. zrikavci) characterized by long antennae with more than 30
segments, tympana on fore tibiae, tegminal stridulation and long ovipositor, and (2) Caelifera (grasshoppers, hrv.
skakavci) characterized by short antennae with less than 25 segments, tympana on the first abdominal segment,
femoro-tegminal stridulation and short ovipositor. Within Ensifera, the following families are present in Croatia:
Tettigoniidae—bush crickets or katydids (hrv. konjici), Gryllidae—true crickets (hrv. šturci, popci), Mogoplistidae—
scaly (or scale) crickets (hrv. ljuskavi šturci), Myrmecophilidae—ant loving crickets (hrv. mravoljupci),

6 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Gryllotalpidae—mole crickets (hrv. rovci, mrmci), and Rhaphidophoridae—camel or cave crickets (hrv. spiljski
konjici). Within Caelifera, members of the following families inhabit Croatia: Tridactylidae—false (or pygmy) mole
crickets (hrv. ksije), Tetrigidae—pygmy grasshoppers or groundhoppers (hrv. monaški skakavci / trnovratke),
Acrididae—true grasshoppers (hrv. pravi skakavci), and Pamphagidae—toad grasshoppers (hrv. žaboliki skakavci).
Republic of Croatia is a country in the Central and Southeastern Europe. It covers an area of 56 594 km².
Throughout the history, country borders changed substantially. From the early 19th century, it consisted of several
administrative units under French, Italian, Austrian and Hungarian jurisdiction. After the First World War, the
largest part of the current territory was included in the Kingdom of Yugoslavia, while Istria, Zara (= Zadar), islands
Lussin (= Lošinj) and Cherso (= Cres), Lagosta (= Lastovo) Archipelago, Pelagosa (= Palagruža) Archipelago and
Cazza (= Sušac) belonged to the Kingdom of Italy. After the Second World War, Croatia, in its current borders,
became a part of the Federal People's Republic of Yugoslavia, and later a Socialist Republic within the Socialist
Federal Republic of Yugoslavia. In 1991, after the break-up of Yugoslavia, the Republic of Croatia was proclaimed.
Relief of the country is very diverse and, in terms of biodiversity, Dinaric Alps and adjacent Adriatic coast
belong to the world biodiversity hotspots (Myers et al. 2000). Three main relief types can be identified: (1) alluvial
lowlands and isolated low mountains in the SW part of the Pannonian Basin; (2) the Dinaric Alps with karst
plateaus and karst poljes; and (3) Adriatic coast with more than a thousand islands, islets and isolated rocks
(Duplančić Leder et al. 2004). Croatia is officially within three European biogeographic regions—Continental,
Alpine and Mediterranean (European Environmental Agency 2016). Biota typical for the Alpine biogeographic
region do not have continuous distribution in Croatia: in this region only a few isolated subalpine and azonal alpine
and boreal habitats exist, but continental and Mediterranean biota generally dominate. However, in this study we
follow older phytogeographic and biogeographic division(s), dividing the country into two main units—
Mediterranean and Continental [= Eurosiberian—North American] (Horvatić 1967, Horvat et al. 1974), the latter
being divided into Pannonian and Dinaric regions. Thus, we divide Croatia into three regions—the Pannonian
region, the Dinaric Alps region and the Mediterranean region (see Material and methods, Fig. 1).
The Pannonian region: Northern Croatia is situated within the Pannonian region, consisting of two parts: (1)
Pannonian part—alluvial lowland along the rivers Mura, Drava, Danube, and Sava, the hilly area of Bilogora and
Moslavina, isolated Slavonian mountains (Psunj, Ravna gora, Papuk, Krndija) and the westernmost slopes of
Fruška gora Mt. near Ilok in the Croatian part of Srijem (Podunavlje); and (2) Peripannonian part—the area
between the lower reaches of Kupa River and Sava River (Pokuplje and Vukomeričke gorice), shallow karst
plateau of Kordun, Banovina—the area between Una and Kupa rivers, including Petrova gora Mt., Zrinska gora
Mt. and Trgovska gora Mt., and the hills of Hrvatsko Zagorje near the Slovenian border with isolated mountains
(Ravna Gora, Ivanščica, Strahinjčica, Medvednica, Kalnik). The western part of this region is more humid, with the
annual precipitation of about 900 mm, while the eastern part is drier, with the annual precipitation of only 350 mm
near Ilok and in the Croatian part of Baranja. At its eastern border and along the right bank of the Drava River, the
Pannonian region contains some steppic biota, common further northwards and eastwards.
Amphibian fauna typical for the Pannonian region includes Triturus dobrogicus, Bombina bombina, Rana
arvalis, and Pelobates fuscus, while reptiles characteristic for the region are Lacerta agilis argus and Lacerta
viridis. Eastern part of the continental lowlands exhibits some steppic features and is inhabited by Apatura metis
and Colias erate (butterflies), Ablepharus kitaibelii (skink), Cricetus cricetus and Mus spicilegus (rodents). During
the last century, steppe-specialist rodents Spermophilus citellus and Nanospalax leucodon became locally extinct.
The Dinaric Alps region: In comparison to the mountains of the neighboring countries, Croatian Dinaric Alps
are rather low, with peaks under 1800 m, and without the alpine belt. The region contains typical mountainous
continental habitats with a mosaic of Mediterranean habitats penetrating deep in the mainland (e.g. to the Plitvice
Lakes NP) (Horvatić 1967, Trinajstić & Šugar 1968). The highest peaks are characterized by isolated subalpine
habitats with rare alpine and oromediterranean biota (Trinajstić 1988). Relict boreal biota exist also in the
westernmost subalpine belt and cold karst depressions. The western part of the Dinaric Alps has high annual
precipitation (more than 2500 mm), whereas the eastern part is drier. The main border between the continental and
the Mediterranean vegetation is located between the Dinaric Alps and the Mediterranean biogeographic regions.
The border called Adamović's line (defined in Adamović 1909), extends along the southern slopes of the coastal
chain of the Dinaric Alps from the Slovenian border at Ćićarija Mt., Učka Mt., Risnjak Mt., Velika Kapela Mt. and
Velebit Mt. (between 800—900 m a.s.l.), and continues deep in the mainland along Poštak Mt., Dinara Mt.,
Troglav Mt. and Kamešnica Mt. (between 1000—1200 m a.s.l.), on the border with Bosnia and Herzegovina.

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 7
Along the Adamović’s line, cold and strong northern local wind—bora, commonly blows in autumn, winter and
spring months (Bertović 1975). Karst area of the Dinaric limestone abounds in speleological objects. Partially
isolated Istrian mountains Učka Mt. and Ćićarija Mt. are also assigned to this region.
The Dinaric Alps region is home to Salamandra atra prenjensis (salamander) and numerous reptiles—Lacerta
agilis bosnica, Zootoca carniolica, Iberolacerta horvathi, and Vipera ursinii. It is also inhabited by various
mammals—Dryomys nitedula, Chionomys nivalis, Canis lupus, Ursus arctos and Rupicapra rupicapra, and
butterflies, for example Erebia medusa, and E. oeme. One can find boreal biota in the cold mixed and coniferous
forests, including owls Aegolius funereus and Glaucidium passerinum, and a rare bat Eptesicus nilssonii. Isolated
parts of the subalpine belt are inhabited by alpine birds Prunella collaris and Eremophyla alpestris, and numerous
alpine butterflies—Parnassius apollo liburnicus, Erebia pronoe, E. styria, E. gorge, E. triaria, and Colias
caucasica. Butterflies Erebia ottomana and E. melas, and lizard Dinarolacerta mosorensis, are typical
representatives of oromediterranean biota in Croatia.

FIGURE 1. Map of Croatia showing division into regions and subregions applied in this paper. Red and white represent
Continental region, while light and dark blue, yellow, violet and black represent Mediterranean region. Continental region is
divided into Pannonian (red) and the Dinaric Alps (white). Mediterranean region is divided into Istria (light blue), Kvarner
(yellow), Dalmatia (dark blue), Dubrovnik area (violet) and the Adriatic Islands (black).

8 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


The Mediterranean region: Within this region, lowlands of Istrian peninsula (NW part of the region) are a
mosaic of Mediterranean and isolated continental habitats. Kvarner coast (N part of the region) is colder and more
humid (annual precipitation about 1500 mm), while Dalmatian coast (central part of the region) is warmer and drier
(annual precipitation only 700-900 mm). Isolated Dalmatian mountains (Svilaja Mt., Mosor Mt. and Biokovo Mt.)
have a mixed upper belt rich in Mediterranean biota, with rare continental and oromediterranean elements. Finally,
Dubrovnik area (SE part of the region) is warm and humid (annual precipitation about 1500 mm). Islands in the
open sea, e.g. Palagruža Archipelago, Sušac Isl., Vis Archipelago, Lastovo Archipelago and parts of Hvar Isl., are
very sunny and dry (annual precipitation only 270 mm), but with a high dewing rate in summer nights (Bertović
1975).
The Mediterranean region is mostly inhabited by the Mediterranean, Balkan or Eastern Mediterranean species,
such as Melanargia larissa, Papilio alexanor atticus, Charaxes jasius, Gonepteryx cleopatra (butterflies),
Ophisaurus apodus thracicus, Podarcis melisellensis. Algyroides nigropunctatus (lizards), Elaphe quatuorlineata,
Zamenis situlus, Telescopus fallax, Malpolon insignitus, Hieriophis gemonensis (snakes), and shrew Suncus
etruscus. Some species, e.g. scorpion Euscorpius feti, butterflies Kirinia roxelana and Lycaena ottomana, and mole
Talpa caeca (Balkan group), have their western distribution border in the Dubrovnik area. In the Dubrovnik area
and Dalmatia one can find typical butterfly species—Proterebia afra dalmata, Hipparchia syriaca, Gegenes
pumilio, and Tarucus balcanicus, as well as lizard Dalmatolacerta oxycephala, and bird Sitta neumayeri. Western
Mediterranean species, such as butterfly Pyrgus malvoides and Italian frog Rana latastei, are restricted to Istria
peninsula, Western green lizard Lacerta bilineata is found on Cres Isl., while snake Hierophis carbonarius inhabits
Istria, Kvarner and the adjoining part of Dinaric Alps.

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

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 9
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.

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.

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

10 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


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 CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 11
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

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& 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

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 13
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

14 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


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.

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 15
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.

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Material and methods

We (1) examined all literature data on Croatian Orthoptera, (2) visited and revised several museum collections
containing Orthoptera material from Croatia: the collection of the Croatian Natural History Museum (in part),
Košćec Collection at the Varaždin City Museum, Hensch Collection at the Faculty of Forestry in Zagreb (in part),
Orthoptera collection of the Faculty of Science in Novi Sad (Vojvodina, Serbia), the collection of the Museum of
Natural History of Trieste (Italy), the collection of the Hungarian Natural History Museum in Budapest, Fer
Willemse Collection at the Naturalis Biodiversity Center in Leiden (Netherlands) and Filippo Maria Buzzetti
private Orthoptera collection in Arzignano (Italy), (3) performed field studies, and (4) implemented relevant
faunistical data from the photos found in online social media (e.g. Facebook, Flickr, Observado).
Literature examination: We collected all relevant literature on Croatian Orthoptera, the bulk of which is
comprised of faunistic and taxonomic papers published from the middle of the 19th century to the present day (see
Introduction). When assessing a given species’ distribution in Croatia, we considered only true faunistic records, i.e.
records with specify the locality where the species was recorded. All inaccurate records were omitted. Doubtful
records were appropriately discussed and, if considered implausible, omitted from the checklist (see Omitted species).
Sampling methods: In the field, we followed visual and/or auditory cues to detect orthopteran species, and
commonly sampled specimens by hand or using an entomological net. Valuable data for ground-dwelling species
were obtained using pitfall traps, since they were often present as by-catch in field studies investigating other soil
arthropod groups. We performed systematic research aiming to assess species occurrence and diversity of
orthopterans in a number of previously understudied areas throughout the country. Scattered data, collected during
the authors’ holidays, short field excursions etc. were also considered while preparing this checklist.
Identification: Harz (1969) was mostly used for the identification of Ensifera, while Harz (1975) for the
identification of Caelifera. For some orthopteran groups, Harz’ keys were not not satisfactory, so following keys
and papers were used for the identification: Heller et al. (2004) for Isophya, Ingrisch & Pavićević (2010, 2012) for
Leptophyes and P. elegans group, Chobanov & Heller (2010) for Poecilimon ornatus group, Skejo et al. (2015) for
Bicolorana, Chobanov et al. (2014) for Tettigonia, Massa et al. (2013) for Oecanthus, Iorgu et al. (2016, 2017) for
Gryllotalpa, Karaman et al. (2011) for Troglophilus, Devriese (1996) for Tetrigidae, Willemse et al. (2009) for
Chorthippus with angled pronotal keels, and Iorgu & Iorgu (2008) for Chorthippus (dorsatus).
Taxonomy and nomenclature: We followed Orthoptera Species File Online (Cigliano et al. 2018) for valid
nomenclature and current taxonomic placement of the species. All nomenclatural changes are in accordance with
the International Code of the Zoological Nomenclature (ICZN 1999).
Distribution in Croatia and division of the country: For each species, we noted its occurrence in one or
more geographic regions of Croatia: Pannonian, Dinaric Alps and/or Mediterranean, the latter with coastal sub-
regions Istria, Kvarner, Dalmatia and Dubrovnik area. Adriatic islands, because of partial scarcity of data, are
considered a distinct region in this paper. Regions are described in Introduction and schematized in Fig. 1.
Depending on the extent of their occurrence, species were designated as either widespread or localized
(stenoendemic), while depending on the frequency of findings, they were designated as common, not common or
rare. Such designation was possible only if enough faunistic data existed for a given species in Croatia. Otherwise,
its probable distribution in the country was briefly discussed based on the present knowledge and its known
distribution in the Balkans.
Distribution in Europe: Distribution of each species in Europe was described in accordance with the
annotated distributional data published within IUCN Orthoptera Red List for Europe (Hochkirch et al. 2016).
Greater emphasis was put on the distributional data for the Balkans, while peculiarities unrelated to the species'
distribution in Southeast Europe were omitted.
Taxonomic acts: During our research of Croatian Orthoptera, we did not find differences listed in the keys as
relevant for certain species, so we compared Croatian specimens of most of such species (exceptions are endemic
species) with specimens of the same species from outside Croatia, either from online photos, Orthoptera Species
File digitalized material, DORSA project web page with digitalized types, museum material, or fresh material we
collected. Those comparisons made it clear to us that some names should be synonymized and currently create
confusion in identification, especially for the beginners dealing with the group in Croatia. Synonymy has most
often been a result of poor assessment of a species' morphological variability in the past. For all the species and
subspecies we synonymized, we examined the original description, types, photos of types, or drawings of types,
specimens from Croatia and neighboring countries where the species occurs, as well as its intraspecific variability.
We found synonyms of (1) Pholidoptera d. dalmatica (P. dalmatica brachynota syn. nov. and P. d. maritima syn.
nov.), (2) Gampsocleis abbreviata (G. a. renei syn. nov.), (3) Chrysochraon d. giganteus (C. d. intermedius syn.
nov.), (4) Odontopodisma fallax (O. rammei syn. nov.) and (5) Acrida u. ungarica (A. u. mediterranea syn. nov.).

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Also, Chorthippus biguttulus hedickei probably represents a synonym of the nominal subspecies, while C.
bornhalmi could be a synonym of C. maritimus.
IUCN Red List status: Following IUCN Orthoptera Red List for Europe (Hochkirch et al. 2016), we listed
Red List status for the species under threat of extinction (CR, EN, VU, DD) and near the threat of extinction (NT),
which occur in Croatia.

Results

Suborder Ensifera = zrikavci

infraorder Tettigoniidea

superfamily Tettigonioidea Krauss, 1902

family Tettigoniidae Krauss, 1902 = konjici

subfamily Bradyporinae Burmeister, 1838 = sedlarke

genus Ephippiger Berthold, 1827

1. Ephippiger discoidalis Fieber, 1853 = crvenoglava / krška sedlarka (Fig. 3A)


First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Ephippigera limbata Fischer, 1853 (Frauenfeld 1861), Ephippigera limbata variety major
Krauss, 1879, E. l. variety minor Krauss, 1879, E. sphacophila Krauss, 1879 (Krauss 1879)
Distribution in Croatia: Widespread and rather common in the Mediterranean part of the country. In the Dinaric
Alps inhabiting only localities with strong Mediterranean influence.
Distribution in Europe: Common along the eastern Adriatic coast and the neighbouring Dinaric Alps and the
Northern Pindos range, to the north reaching the Eastern Alps in Northeastern Italy (Chobanov et al. 2016).

2. Ephippiger ephippiger (Fiebig, 1784) = crnoglava sedlarka (Fig. 3B)


First faunistic record for Croatia: Graber 1870
Also reported as: Ephippigera vitium Serville, 1831 (Frauenfeld 1861)
Distribution in Croatia: Recorded in numerous localities along the northern part of the Dinaric Alps, as well as in
the Pannonian region. Its presence in some other regions, e.g. Kvarner, although probable, is yet unconfirmed.
Distribution in Europe: Balkan Peninsula from Southern Czech Republic and Northern Slovakia in the north, to
Northern Greece and European Turkey in the south, and Eastern Ukraine in the east (Hochkirch et al. 2016).

3. Ephippiger persicarius (Fruhstorfer, 1921) = istarska (crnoglava) sedlarka


First faunistic record for Croatia: Harz 1966
Also reported as: E. ephippiger vicheti Harz, 1966 (Harz 1966)
Distribution in Croatia: There is a single record for this species in Istria (Harz 1966, 1969) recorded under E ephippiger
vicheti Harz, 1966. Type specimens of E. e. vicheti are found to be conspecific with E. persicarius, described from
Switzerland (Monte Carasso (1600 m a.s.l.) and Alpe di Orino (1200 m a.s.l.)) (Baur & Coray 2004, Barat 2012). Three
male paratypes were collected in Pazin (= Pisino, leg. H. Krauss).
Distribution in Europe: South–western part of the Alps, concentrated in the mountain regions of Italy and Switzerland
between the Lago Maggiore and the region of Trentino, reaching northwards to the southern parts of the Tessin and
Graubünden. Records also exist from the Alpi Marittimi (Italy), Styria (Austria) and Istria (Croatia) (Zuna-Kratky et al.
2016).

subfamily Conocephalinae Burmeister, 1838 = konjici čunjoglavci

genus Conocephalus Thunberg, 1815

4. Conocephalus dorsalis (Latreille, 1804) = kratkokrili (konjic) čunjoglavac

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First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Recorded in all regions except Kvarner, although it was found on adjacent Cres Isl. (Krauss
1879), and Dubrovnik area. Distribution in Croatia scattered.
Distribution in Europe: Southernmost border in Central Portugal, Tuscany (northern Italy) and South Bulgaria,
reaches Scotland and Southern Finland to the north and Eastern European Russia to the east (Hochkirch et al. 2016).

FIGURE 3. Diversity of Croatian Ensifera—Tettigoniidae: Bradyporinae, Conocephalinae, Meconematinae, Phaneropterinae.


A. Ephippiger discoidalis (Lič 9.IX.2014. F. Rebrina), B. Ephippiger ephippiger (Lička Plješivica—Krbavsko polje
2.VIII.2017. N. Tvrtković), C. Ruspolia nitidula (Hrvatsko 30.VIII.2014. F. Rebrina), D. Meconema thalassinum (Kruščica
7.VIII.2014. F. Rebrina), and E. Barbitistes kaltenbachi (Vis Isl. 21.VII.2011. R. Felix).

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5. Conocephalus fuscus (Fabricius, 1793) = dugokrili (konjic) čunjoglavac
First faunistic record for Croatia: Graber 1870
Also recorded as: Conocephalus dilatatus Ramme, 1951 (Ramme 1951).
Distribution in Croatia: Recorded in all regions except Kvarner area, although found on Cres Isl. (Krauss 1879).
Distribution scattered, but generally more common than C. dorsalis.
Distribution in Europe: Southernmost border in Southern Spain and Portugal, Sardinia, Sicily, Crete and Cyprus, to
the north reaching England, the Netherlands and Northern Germany, and Eastern European Russia to the east
(Hochkirch et al. 2016).

genus Ruspolia Schulthess, 1898

6. Ruspolia differens (Serville, 1839) = afrički (konjic) čunjoglavac


First faunistic record for Croatia: Heller 1988 (after Schumacher 1978)
Distribution in Croatia: Rovinj (Istria). Introduced African species, differs from R. nitidula by a smaller number of
stridulatory teeth (64–86 vs. > 120). Another introduced population in Europe recorded in Bodensee (Germany)
(Heller 1988).
Distribution in Europe: Croatia: Rovinj, Germany: Bodensee (see above).

7. Ruspolia nitidula (Scopoli, 1786) = veliki (konjic) čunjoglavac (Fig. 3C)


First faunistic record for Croatia: Graber 1870
Also recorded as: Locusta mandibularis Charpentier, 1825 (Kollar 1846, Krauss 1879), Locusta tuberculata
Rossius, 1790 (Cubich 1875)
Distribution in Croatia: Widely distributed and common in all regions.
Distribution in Europe: From Sicily and Crete in the south to Northern France in the north, and from the Azores in
the west to Southern European Russia in the east (Hochkirch et al. 2016).

subfamily Meconematinae Burmeister, 1838 = staklasti konjici

genus Cyrtaspis Fischer, 1853

8. Cyrtaspis scutata (Charpentier, 1825) = beskrilni staklasti konjic


First faunistic record for Croatia: Krauss 1879
Also reported as: C. variopicta Costa, 1860 (Us 1967)
Distribution in Croatia: Known only from a few localities in the Mediterranean part of the country (Istria, Kvarner,
Dalmatia, Dubrovnik area, Southern Adriatic islands), probably overlooked.
Distribution in Europe: Western Mediterranean, reaching Bretagne (France) in the north, Croatia is the easternmost
border of the distribution. Probably introduced to the Azores (Braud et al. 2016).

genus Meconema Serville, 1831

9. Meconema meridionale Costa, 1860 = ljuskokrili staklasti konjic


First faunistic record for Croatia: Krauss 1879
Also reported as: M. brevipenne Yersin, 1860 (Krauss 1879)
Distribution in Croatia: Widespread but not numerous in records. Present in all regions except Dubrovnik area (not
recorded), but more common in the Mediterranean part. Probably overlooked.
Distribution in Europe: Probably native to Northern Spain, southern France, Italy and Northern Croatian coast, but
range expanded all the way to Ireland and Northern Germany in the north, and Central Slovakia, Southern Poland
and Serbia in the east (Hochkirch et al. 2016).

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10. Meconema thalassinum (De Geer, 1773) = dugokrili staklasti konjic (Fig. 3D)
First faunistic record for Croatia: Us 1938
Also reported as: M. varium (Fabricius, 1775) (Us 1938)
Distribution in Croatia: Widespread but not numerous in records. Present in all regions except Dubrovnik area (not
recorded), but more common in the Pannonian region. Probably overlooked in the past.
Distribution in Europe: From Ireland, Southern Scandinavia and Estonia in the north to Central Portugal, Southern
Italy and Northern Greece in the south, and Eastern Ukraine and Central European Russia in the east (Hochkirch et
al. 2016).

subfamily Phaneropterinae Burmeister, 1838 = listokrili i ljuskokrili konjici

genus Acrometopa Fieber, 1853

11. Acrometopa servillea (Brullé 1832)


11.1. Acrometopa servillea macropoda (Burmeister, 1838) = dugonogi listokrili konjic
First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Common in the Mediterranean part, present also in the Dinaric Alps on localities under
strong influence of Mediterranean climate.
Distribution in Europe: Endemic to the Western Balkans—along the Adriatic Coast from Slovenia to northern
Greece, reaching Albania, FYR Macedonia, and the southeastern tip of Bulgaria (Willemse et al. 2016, Cigliano et
al. 2018).

genus Andreiniimon Capra, 1937

12. Andreiniimon nuptialis (Karny, 1918) = krupnozadi ljuskokrili konjic


First faunistic record for Croatia: Ingrisch 1981
Distribution in Croatia: Only two known localities in Istria (Flengi: Ingrisch 1981; Pazin 21.VII.2015: Plüss—
published online on orthoptera.ch (http://www.orthoptera.ch/images/Home/Galerie/Istrien_2015/
Stefan_Pluess_2015/Andreiniimon_nuptialis_m_Pazin_150721_1728_4.jpg).
Distribution in Europe: Endemic to Balkan Peninsula and Italy, fragmented range with many isolated
subpopulations (Hochkirch et al. 2016).
IUCN Red List status: Vulnerable (Europe and EU28) (Hochkirch et al. 2016).

genus Barbitistes Charpentier, 1825

13. Barbitistes kaltenbachi Harz, 1965 = Kaltenbachov ljuskokrili konjic (Fig. 3E)
First faunistic record for Croatia: Harz 1965
Distribution in Croatia: Stenoendemic, known from central Adriatic islands (Hvar, Vis). Historical records from
Lukovo and Rijeka (Harz 1969) are questionable. Exact position of Lukovo locality is uncertain, since there are
several settlements with that name along the Croatian coast. Presence in Rijeka unconfirmed. Presence on Hvar Isl.
confirmed (Heller pers. comm.), recently recorded on Vis Isl. (R. Felix, Hum 20.VII.2011).
Distribution in Europe: Endemic to Croatia (Chobanov et al. 2016).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

14. Barbitistes ocskayi Charpentier in Ocskay et al., 1850 = Ocskayjev ljuskokrili konjic
First faunistic record for Croatia: Ocskay et al. 1850
Also recorded as: Barbitistes oczkayi (Frauenfeld 1861)
Distribution in Croatia: Along the Adriatic coast—abundant in Istria and Kvarner with adjacent islands, southern
slopes of Velika Kapela Mt. and Velebit Mt. Adamović’s findings (1964) are the only published record from

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Dubrovnik region (there are recent unpublished findings in Dubrovnik area, T. Koren, Konavle VI.2015).
Distribution in Europe: Endemic to the Balkans—widely distributed in the central and western Balkans, its range
extending to the Southeastern Carpathians. Specimens from the western and the eastern population were not
adequately compared so far (Chobanov et al. 2016).

15. Barbitistes serricauda (Fabricius, 1794) = panonski ljuskokrili konjic


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Commonly found in the Pannonian region, but also recorded in the northwesterrn part of
the Dinaric Alps (Velebit Mt.: Senjska Draga: Rebrina, Skejo & Tvrtković 10.VII.2013, Samoborsko gorje:
Plešivica (Vlaškovec): Rebrina 7.VII.2013) and is expected in other areas adjacent to Dinaric region.
Distribution in Europe: Widely distributed, from the Pyrenees, Northern Italy and Central France via Central
Europe to Romania (Hochkirch et al. 2016).

16. Barbitistes yersini Brunner von Wattenwyl, 1878 = Yersinov ljuskokrili konjic
First faunistic record for Croatia: Brunner von Wattenwyl 1878
Also recorded as: Barbitistes dalmatinus Bey-Bienko, 1954 (Bey-Bienko 1954, Us 1967)
Distribution in Croatia: Common in the Mediterranean part of the country, along the coast with adjacent islands,
but also occurs on the southern slopes of the karst mountains deep in the Dinaric Alps region, to Plitvička jezera
(Us 1992).
Distribution in Europe: Distributed in the Western Balkan Peninsula from Slovenia to Northern Greece, present
also in Central Italy (Hochkirch et al. 2016).
genus Isophya Brunner von Wattenwyl, 1878

17. Isophya brevicauda Ramme, 1931 = kratkorepi ljuskokrili konjic


First faunistic record for Croatia: Ramme 1931
Distribution in Croatia: Northwestern Croatia, known from five localities only (Samobor: Ramme 1931, Krapina:
Heller et al. 2004, Mount Medvednica, Sljeme: Puskás et al. 2018, Mts. Strahinjčica and Ivanščica: Gomboc
25.V.2014). Probably overlooked.
Distribution in Europe: Primarily inhabits southeastern Alps and isolated mountains in adjacent Peripannonian area
in Austria, Slovenia and Croatia (Zuna-Kratky et al. 2016).

18. Isophya kraussii Brunner von Wattenwyl, 1878 = Kraussov ljuskokrili konjic
First faunistic record for Croatia: Brunner von Wattenwyl 1882
Also recorded as: Isophya camptoxipha (currently valid name I. camptoxypha (Fieber, 1853)) (Brunner von
Wattenwyl 1882)
Distribution in Croatia: Northwestern Croatia, known from two localities only (Zagreb: Brunner von Wattewyl
1882, Krapina: Heller 1988). Most likely overlooked.
Distribution in Europe: Known range encompasses central and Southern Germany, the Czech Republic, Slovakia,
Southern Poland, Eastern Austria, Eastern Slovenia, Northwestern Croatia, Western and Northern Hungary,
Northwestern and northeastern Romania and Western Ukraine (Chobanov et al. 2016).

19. Isophya modestior Brunner von Wattenwyl, 1882 = dugorepi ljuskokrili konjic
First faunistic record for Croatia: Szövényi & Puskás 2012
Distribution in Croatia: In and close to the mountains of the Pannonian region (Papuk Mt.: Szövényi & Puskás
2012, Psunj Mt., Dilj Mt.: Puskás et al. 2018, Strahinjčica and Ivanščica Mts.: Gomboc 3.VIII.2014). Likely
common in the region, but overlooked.
Distribution in Europe: Northwestern to central part of the Balkan Peninsula, from Northern Italy and Eastern
Austria to Western Bulgaria and Northern FYR Macedonia (Hochkirch et al. 2016).
IUCN Red List status: [Least Concern (Europe)], Vulnerable (EU28) (Hochkirch et al. 2016).

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genus Leptophyes Fieber, 1853

20. Leptophyes albovittata (Kollar, 1833) = bjelorubi ljuskokrili konjic (Fig. 4A)
First faunistic record for Croatia: Graber 1870
Distribution in Croatia: In central (Dinaric) and Pannonian part. Presence in Istria and Kvarner (Us 1967) not
cofirmed.
Distribution in Europe: From Northern Germany and Southern Switzerland in the west to Peloponnesus (Greece) in
the south and Southern European Russia in the east (Hochkirch et al. 2016).

21. Leptophyes boscii Fieber, 1853 = žutoprugi ljuskokrili konjic


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Common in the northern part of the Adriatic coast and the Pannonian region, also in the
western part of the Dinaric Alps (Szövényi et al. 2018).
Distribution in Europe: From the Italian/French border in the west and the Czech Republic in the north to Serbia in
the east and Balkan Peninsula with Greece in the south (Hochkirch et al. 2016).

22. Leptophyes discoidalis (Frivaldszky, 1868) = crnopjegi ljuskokrili konjic


First faunistic record for Croatia: Skejo & Stanković 2013
Distribution in Croatia: Known from a single locality in the eastermost continental part of the country (Privlaka
near Vinkovci, leg. Tvrtković: Skejo & Stanković 2013). Probably overlooked, but most likely not distributed
outside the Pannonian region.
Distribution in Europe: From Southeastern Slovakia and Northern Hungary to Central Balkan Peninsula: Northern
Bulgaria, with isolated subpopulations in Eastern Croatia, Serbia, Montenegro and Bosnia and Herzegovina
(Hochkirch et al. 2016).
IUCN Red List status: Vulnerable (Europe and EU28) (Hochkirch et al. 2016).

23. Leptophyes laticauda (Frivaldszky, 1868) = riđonogi ljuskokrili konjic


First faunistic record for Croatia: Brunner von Wattenwyl 1878
Distribution in Croatia: Common in the entire Adriatic part of the country and in the Dinaric Alps.
Distribution in Europe: From Italy and Southeastern France via Northern Balkan Peninsula to Western Bulgaria
and Southwestern Romania (Hochkirch et al. 2016).

24. Leptophyes intermedia Ingrisch & Pavićević, 2010 = dinarski ljuskokrili konjic (Fig. 4B)
First faunistic record for Croatia: Puskás et al. 2018
Distribution in Croatia: Velebit Mt. (Paklenica: Puskás et al. 2018) and isolated mountains of Dalmatia—Mosor
Mt. (Puskás & Szövényi 25.VII.2016) and Biokovo Mt. (Skejo & Ozimec 27.VII.2016).
Distribution in Europe: Endemic to the Balkans—Dinaric mountains of Bosnia and Herzegovina, Montenegro
(Ingrisch & Pavićević 2010) and Croatia (see above).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

25. Leptophyes punctatissima (Bosc, 1792) = pjegavi ljuskokrili konjic


First faunistic record for Croatia: this study
Distribution in Croatia: Its presence hitherto confirmed only in the Northern Kvarner near Rijeka (Puskás, Skejo &
Szövényi 23.VII.2014) and Istria around and on Učka Mt. in the Dinaric Alps (Roesti & Rutschmann pers. comm.
and orthoptera.ch). Redtenbacher's (1900) records for Dalmatia not confirmed.
Distribution in Europe: From Scotland and Southern Norway in the north to Iberian Peninsula, Sardinia, southern
Italy and Greece in the south, and Bulgaria in the east (Hochkirch et al. 2016).

genus Poecilimon Fischer, 1853

26. Poecilimon affinis (Frivaldszky, 1868) = istočni ljuskokrili konjic


26.1. Poecilimon a. affinis (Frivaldszky, 1868)

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First faunistic record for Croatia: Szövényi & Puskás 2012
Distribution in Croatia: Pannonian region only (Slavonija). Pungur’s (1899) record from Karlovac is dubious, due
to the lack of voucher specimens.
Distribution in Europe: Inhabits Southeastern Europe from Croatia in the north to Ukraine in the east and Greece in
the south (Hochkirch et al. 2016).

FIGURE 4. Diversity of Croatian Ensifera—Tettigoniinae: Phaneropterinae, Saginae. A. Leptophyes albovittata (Malo Polje
4.VIII.2014. F. Rebrina), B. Leptophyes intermedia (Paklenica 28.VII.2014. J. Skejo), C. Poecilimon cf. elegans (Grabovača
6.VIII.2014. F. Rebrina), D. Polysarcus denticauda (Snježnik 25.VIII.2014. F. Rebrina), E. Phaneroptera nana (Proložac
26.VII.2015. J. Skejo), F. Saga pedo (Učka Mt. 24.VII.2014. J. Skejo).

27. Poecilimon ampliatus Brunner von Wattenwyl, 1878 = bradavičasti ljuskokrili konjic
First faunistic record for Croatia: Redtenbacher 1900

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Distribution in Croatia: Not common, present in the Dinaric Alps from Istria (Učka Mt.) to Southeastern Croatia
(Poštak Mt.: Szövényi et al. 2018).
Distribution in Europe: Northwestern Balkan Peninsula from Slovenia to Montenegro and Western Serbia, with
some isolated records in Southeastern Serbia and Southwestern Romania (Chobanov et al. 2016).

28. Poecilimon elegans Fieber in Brunner von Wattenwyl, 1878 = mali ljuskokrili konjic
First faunistic record for Croatia: Brunner von Wattenwyl 1878
Distribution in Croatia: Only specimens from Istria, Učka Mt. and the southern slopes of Velika Kapela and Velebit
Mt. to Karlobag, as well as Cres Island (Mt. Sis, Szövényi 06.VII.2017) belong to P. elegans (Ingrisch & Pavićević
2010).
Distribution in Europe: From Northeastern Italy via Slovenia to Northern Croatian coast (Hochkirch et al. 2016).

29. Poecilimon sp. (sibling species near P. elegans Brunner von Wattenwyl, 1878) [undescribed new species] (Fig.
4C)
First faunistic record for Croatia: Ingrisch & Pavićević 2010
Distribution in Croatia: From Southern Velebit Mt. to Metković (Neretva River) and Korčula Isl.
Distribution in Europe: Endemic to the Balkans—Croatia and probably parts of Bosnia and Herzegovina.
Taxonomic note: According to Ingrisch & Pavićević (2010), some former paralectotypes of P. elegans (specimens
from Obrovac, Metković, Korčula Isl.) deposited in the Natural History Museum in Vienna belong to an
undescribed species. It has the same number of stridulatory teeth as Poecilimon albolineatus Ingrisch & Pavićević,
2010 (95–98), but is morphologically closer to P. elegans. Probably all the populations from the Adriatic coast and
southern slopes of the Dinaric Alps southeastern of Karlobag belong to this new species. There is also a possibility
for the presence of P. albolineatus or another undescribed species (close to P. albolineatus) in Dubrovnik region
and on the western slopes of Orjen Mt. Namely, some specimens from Lovćen Pass near Kotor in Montenegro
examined by Ingrisch & Pavićević (2010) had a higher number of stridulatory teeth (106–114) and their song
differs both from typical P. albolineatus and P. cf. elegans population from Biokovo Mt. in Croatia (K.–G. Heller
pers. comm.).

30. Poecilimon fussii Fieber in Brunner von Wattenwyl, 1878 = Fussov ljuskokrili konjic
First faunistic record for Croatia: Szövényi & Puskás 2012
Distribution in Croatia: Pannonian Mts. (hitherto known only from Papuk Mt.: Szövényi & Puskás 2012 and Dilj
Mt.: Puskás et al. 2018). Us' (1967) record from Istria unconfirmed and highly unlikely.
Distribution in Europe: Northwestern and Western Bulgaria, Northern Serbia, Northeastern Croatia, Western and
Northern Hungary, Southeastern Slovakia and Western Romania (Chobanov et al. 2016).

31. Poecilimon gracilis (Fieber, 1853) = vitki ljuskokrili konjic


First faunistic record for Croatia: Ramme 1931
Distribution in Croatia: In and around the Dinaric Alps (Velika Kapela (Tuk): Rebrina & Tvrtković 23.VII.2014,
Risnjak Mt.: Rebrina, Skejo & Tvrtković 25.VIII.2014, Velebit: Puskás et al. 2018) and mountains of the
Pannonian region (Samobor and Krapina: Ramme 1931, Papuk Mt.: Szövényi & Puskás 2012, Psunj Mt.,
Medvednica Mt.: Puskás et al. 2018, Mts. Strahinjčica and Ivanščica: Gomboc 25.V., 30.VIII.2014).
Distribution in Europe: Along the eastern Adriatic coast from Southern Austria to Northern Greece (Chobanov et
al. 2016).

32. Poecilimon ornatus (Schmidt, 1850) = šareni ljuskokrili konjic


First faunistic record for Croatia: Brunner von Wattenwyl 1878
Also recorded as: Barbitistes Fieberi Ullrich in Fieber, 1853 (Frauenfeld 1861)
Distribution in Croatia: Common in the Dinaric Alps, but recorded also on Medvednica Mt. in Pannonian Croatia
(Tvrtković VII.2013). Present in Istria and Kvarner with adjacent islands. Hitherto no records exist from Dalmatia
and Dubrovnik area.
Distribution in Europe: Western Balkan Peninsula from Northern Italy and Southern Austria to Northern Greece
(Hochkirch et al. 2016).

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33. Poecilimon schmidtii (Fieber, 1853) = Schmidtov ljuskokrili konjic
First faunistic record for Croatia: Ramme 1931
Distribution in Croatia: Common in the Pannonian region, present in the Dinaric Alps, but not as common. Records
from Istria and Dalmatia (Frauenfeld 1861, Redtenbacher 1900, Us 1967, 1992) not confirmed.
Distribution in Europe: Balkan Peninsula from Slovenia in the west and Southern Poland in the north to Northern
Greece, European Turkey and South Caucasus in the south and east (Hochkirch et al. 2016).

genus Polysarcus Fieber, 1853

34. Polysarcus denticauda (Charpentier, 1825) = mesnati bodljorepi konjic (Fig. 4D)
First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Continental belt of the Dinaric Alps in the northern and central Croatia, high density of certain
populations observed (Velebit Mt. (Šegotine krčevine): Rebrina & Tvrtković 18.VII.2013, Velika Kapela Mt.
(Podbitoraj): Mt. (Podbitoraj): Tvrtković 12.VI.2015): Also on isolated Dalmatian mountains (e.g. Biokovo Mt.).
Distribution in Europe: Widely distributed in the Balkan Peninsula, inhabits also Northern and Central Italy, and
Southern France to the Pyrenees, in the north reaching Central Germany and Southern Poland (Hochkirch et al.
2016).

genus Phaneroptera Serville, 1831

35. Phaneroptera falcata (Poda, 1761) = listokrili konjic srponoša


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Northwestern part of the Adriatic coast (Istria, Kvarner) with adjacent islands, in the
Dinaric Alps and the Pannonian region. Historical records from Dalmatia and Dubrovnik area need confirmation,
most likely misidentifications of P. nana as noted in Adamović (1964), and indicated by redetermination of
specimens in MST collection (R. Kleukers, pers. comm.).
Distribution in Europe: Widely distributed, from Northern Iberian Peninsula, Central Italy, Balkan Peninsula with
Greece and European Turkey in the south to England and Southern Sweden in the north, European Russia in the
east (Hochkirch et al. 2016).

36. Phaneroptera nana Fieber, 1853 = obični listokrili konjic (Fig. 4E)
First faunistic record for Croatia: Krauss 1879
Also recorded as: Phaneroptera quadripunctata Brunner von Wattenwyl, 1878 (Krauss 1879) or P. 4-punctata
(Redtenbacher 1900)
Distribution in Croatia: Widely distributed and common in the whole country. Historical findings of P. falcata from
Dalmatia and Dubrovnik area probably belong to this species (see notes under P. falcata).
Distribution in Europe: Widely distributed from the Azores in Southern Europe in the west, and Northern France,
Western Germany and the Czech Republic in the north, to the south of the Iberian Peninsula, Lampedusa (Italy), Crete
and Cyprus in the south. European Russia is the easternmost border of the distribution (Hochkirch et al. 2016).

genus Tylopsis Fieber, 1853

37. Tylopsis lilifolia (Fabricius,1793) = primorski listokrili konjic


First faunistic record for Croatia: Germar 1817
Also recorded as: Locusta gracilis Germar, 1817 (Germar 1817)
Distribution in Croatia: Widely distributed and common along the Adriatic coast and on most islands.
Distribution in Europe: Inhabits Mediterranean region, northernmost border in Western France (area near
Bordeaux) and the Southern Alps in Italy, easternmost in Southern Ukraine. Probably introduced to Normandy
(France) (Hochkirch et al. 2016).

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subfamily Saginae Brunner von Wattenwyl, 1878 = konjici vračevi / grabežljivi konjici

genus Saga Charpentier, 1825

38. Saga natoliae Serville, 1838 = crni konjic vrač / crni grabežljivi konjic
First faunistic record for Croatia: Adamović 1964 [without precise locality]
Distribution in Croatia: Known from Dubrovnik surroundings (Adamović 1964). Its presence is likely, but not yet
confirmed.
Distribution in Europe: Endemic to the Balkans—from Southern Croatia to Southern Bulgaria and Sorthern
Greece, European Turkey (Hochkirch et al. 2016).

39. Saga pedo (Pallas, 1771) = konjic vrac / grabežljivi konjic / jastog Provanse (Fig. 4F)
First faunistic record for Croatia: Krauss 1879
Also reported as: S. serrata (Fabricius, 1793) (Frauenfeld 1861)
Distribution in Croatia: Widely distributed in Istria, Kvarner, on the southern slopes of Dinaric Alps to Kuna
Konavoska in Dubrovnik area (photo Tvrtković 3.VI.2006) and on some Adriatic islands (Cres, Lošinj, Korčula).
Not frequent.
Distribution in Europe: From the Iberian Peninsula in the west and Southern Czech Republic in the north, to Sicily
in the south and the Volga Delta in the east (Hochkirch et al. 2016).

subfamily Tettigoniinae Krauss, 1902 = pravi konjici

genus Bicolorana Zeuner, 1941

40. Bicolorana bicolor (Philippi, 1830) = dvobojni livadni konjic


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Pannonian Croatia, some localities in Kvarner (including a record from Cres Isl.: Schuster
et al. 1998, and Krk Isl., Kampelje: Gomboc 19.VIII.2006), and along the Dinaric Alps (southern border of the
distribution).
Distribution in Europe: From Central France in the west and Southern Sweden in the north to Central Italy in the
south and Eastern European Russia in the east (Hochkirch et al. 2016).

41. Bicolorana kraussi (Padewieth, 1900) = Kraussov livadni konjic (Fig. 5B)
First faunistic record for Croatia: Padewieth 1900
Also reported as: Platycleis padewiethi Krauss in litt. (Padewieth 1900), Metrioptera kuntzeni Ramme, 1931
(Ramme 1931)
Distribution in Croatia: Dinaric mountains in Istria (Ćićarija Mt., Učka Mt.), Gorski kotar (Risnjak Mt.) and the
coastal Dinaric Alps (Velika Kapela Mt., N part of Velebit Mt.) (Skejo et al. 2015).
Distribution in Europe: Endemic to karst area of Northeastern Italy, Southwestern Slovenia and from Slovenian
Slavnik Mt. (Kraški rob area) along the Dinaric arch to North Velebit Mt. in Croatia (Skejo et al. 2015).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

genus Decticus Serville, 1831

42. Decticus albifrons (Fabricius, 1775) = veliki primorski konjic


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Common along the Adriatic coast and on most islands.
Distribution in Europe: Widely distributed throughout the Mediterranean, northernmost points of the distribution
are Northwestern France and Northern Italy, reaching Eastern European Russia in the east (Hochkirch et al. 2016).

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43. Decticus verrucivorus (Linnaeus, 1758) = konjic bradavičar (Fig. 5A)
First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Widely distributed, mostly in the inland part (Pannonian region, Dinaric Alps),
sporadically recorded in the coastal part of Kvarner region.
Distribution in Europe: Widespread throughout the continent, from the Northern Iberian Peninsula, Sardinia and
the Peloponnesus in the south to Central Finland in the north, to the east reaching the Urals in European Russia
(Hochkirch et al. 2016).

FIGURE 5. Diversity of Croatian Ensifera—Tettigoniidae: Tettigoniinae. A. Decticus verrucivorus (Lazac 24.VII.2014. F.


Rebrina), B. Bicolorana kraussi (Francikovac 26.VIII.2014. F. Rebrina), C. Gampsocleis abbreviata (Lisac 27.VI.2015. F.
Rebrina), D. Metrioptera brachyptera (Lazac 24.VII.2014. F. Rebrina), E. Pachytrachis striolatus (Kruščica 4.VIII.2014. F.
Rebrina), F. Pholidoptera dalmatica (Grabovača 7.VI.2014. F. Rebrina).

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genus Eupholidoptera Mařan, 1953

44. Eupholidoptera schmidti (Fieber, 1861) = crno-zeleni kožokrili konjic


First faunistic record for Croatia: Frauenfeld 1861
Also recorded as: Thamnotrizon Chabrieri (Charpentier, 1825) (Frauenfeld 1861)
Distribution in Croatia: Common along the Adriatic coast (including islands) and the Dinaric Alps. Also recorded
in the Pannonian region (Papuk Mt.: Szövényi & Puskás 2012).
Distribution in Europe: Western Balkan Peninsula from northeastern Italy to Western Bulgaria and northern Greece
(Hochkirch et al. 2016).
Taxonomic and distributional note: Us (1967) and Harz (1969) reported only E. chabrieri schmidti from Istria and
along the Adriatic coast, with terra typica in Istria. Since Us (1967), E. chabrieri has been reported from numerous
localities along the Adriatic coast, but Ramme (in Çiplak et al. 2007), Nagy (2006), and Szövényi & Puskás (2012)
extended its known distribution to Pannonian Croatia (Žumberak and Samoborsko gorje Mts., and Papuk Mt.). In
DORSA (2017), specimens from Žumberak and Samoborsko gorje Mts. are reported under E. c. chabrieri (Nagy
2006), but Nadig (1987) considered this subspecies to be a zoogeographical element of the Western Alps.
Adamović (1972) described E. chabrieri usi from Lošinj Isl., which Nadig (1985) later reported also from Krk Isl.
Çiplak et al. (2007) revised the morphology of E. chabrieri complex in the Balkans, concluding that E. c. schmidti
and E. c. usi are conspecific. Massa et al. (2012) listed E. chabrieri and E. schmidti as separate species. The most
reliable discriminating character between the taxa is the shape of the titillators (Harz 1969, Lemonnier-Darcemont
2007, Massa et al. 2012). In a recent phylogenetic study on the genus, Allegrucci et al. (2013) confirmed specific
status of E. schmidti.

genus Gampsocleis Fieber, 1852

45. Gampsocleis abbreviata Brunner von Wattenwyl in Herman, 1874 = bodljoprsi konjic (Fig. 5C)
First faunistic record for Croatia: Brunner von Wattenwyl 1882
Distribution in Croatia: Not abundant, present in Dinaric region and on Pag Isl. (Tvtković 27.VI.2008).
Distribution in Europe: Endemic to the Balkans—western and southern part of the Balkan Peninsula, from Croatia
through Bosnia and Herzegovina, Albania and FYR Macedonia to Greece (Willemse et al. 2016).
Taxonomic note: The great extent of variability in body coloration and morphological traits of G. abbreviata
populations in Croatia (examined specimens from Pag Isl., Lisac Mt., Poštak Mt., Dinara Mt., Troglav Mt.)
indicates that differences used to describe G. abbreviata renei Mikšić, 1973 syn.nov. (described from Livanjsko
polje primarily on the basis of distinct coloration of tegminal veins, being more green than in G. a. abbreviata
(Mikšić 1973)) are not of taxonomic value and fit the variability range of the nominal subspecies. Furthermore, the
population from which G. a. abbreviata was described is probably in contact with the population from Livanjsko
polje, the two localities being less than 30 km apart.

genus Metrioptera Wesmaël, 1838

46. Metrioptera brachyptera (Linnaeus, 1761) = gorski livadni konjic (Fig. 5D)
First faunistic record for Croatia: Nagy 2006
Distribution in Croatia: Restricted to higher altitudes in the Dinaric Alps (from Žumberak–Samoborsko gorje Mts.:
Nagy 2006, Ćićarija Mt.: Skejo & Tvrtković 20.VIII.2013, Kupjak: Puskás et al. 2018, and Risnjak Mt.: Rebrina &
Tvrtković 24.VII.2014, 25.VIII.2014, to Lička Plješivica Mt: Tvrtković 15.VIII.2013).
Distribution in Europe: From the Pyrenees and Southern Scotland in the west, and Northern Scandinavia in the
north, to Southern Serbia in the south and Eastern European Russia in the east (Hochkirch et al. 2016).

47. Metrioptera hoermanni (Werner, 1906) = Hoermannov livadni konjic


First faunistic record for Croatia: this study
Distribution in Croatia: Mountains of Dalmatia. Known only from Biokovo Mt., a single known locality being

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Sveti Jure peak (Skejo & Ozimec VIII.-X.2016).
Distribution in Europe: Stenoendemic to the Balkans—distributed along the Dinaric Alps from Northwestern
Montenegro and Southern Bosnia and Herzegovina (Chobanov et al. 2016) to Croatia (Skejo, this study).
IUCN Red List status: Near Threatened (Europe), assesed as not extant in EU28 (Hochkirch et al. 2016), but our
locality falls within EU28, meaning the species needs to be re-assessed at the EU28 level.

genus Modestana Beier, 1955

48. Modestana modesta (Fieber, 1853) = skromni livadni konjic


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Abundant along the Adriatic coast (including islands Cres: Krauss 1879, Krk: photo
Šegula, http://www.biolib.cz/en/image/id34927/, and Pag, Dinjiška: Skejo 5.IX.2016) and the Dinaric Alps.
Distribution in Europe: Endemic to the Balkans—along the northwestern Adriatic coast from Italy to Northwestern
Albania, but possibly more widely distributed in the West Balkans, as suggested by an isolated record in Western
Serbia (Chobanov et al. 2016).

genus Montana Zeuner, 1941

49. Montana stricta (Zeller, 1849) = uskokrili šikarski konjic


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Present in Istria, Kvarner with adjacent islands (Krk Isl.: Nadig 1987), rather abundant in
the Dinaric Alps. Not common.
Distribution in Europe: Italy and Northwestern Balkan Peninsula: from Northeastern Italy to Montenegro
(Hochkirch et al. 2016).

genus Pachytrachis Uvarov, 1940

50. Pachytrachis frater (Brunner von Wattenwyl, 1882) = južni skroviti konjic
First faunistic record for Croatia: Brunner von Wattenwyl 1882
Distribution in Croatia: Only in the Dalmatian mountains from Mosor Mt. (25.VII.2016. leg. Puskás & Szövényi,
14.VIII.2017. leg. J. Skejo & S. Stermšek) to Dubrovnik area (Adamović 1964) and on Central and Southern
Adriatic islands (e.g. Hvar, Brač Isl., Korčula Isl.). Presence in Istria (Harz 1969) unlikely.
Distribution in Europe: Endemic to the Balkans—from Southern Croatia to Montenegro and Western Serbia.
Records from Slovenia and Northern Croatian coast are doubtful (Hochkirch et al. 2016).
IUCN Red List status: Endangered (Europe and EU28) (Hochkirch et al. 2016).

51. Pachytrachis gracilis (Brunner von Wattenwyl, 1861) = sjeverni skroviti konjic
First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Widely distributed in the Pannonian region, Istria, Kvarner and the Dinaric Alps, not
recorded in Dalmatia and on Adriatic islands. Finding from M. Buzzetti (2006) on Sniježnica Mt. in Dubrovnik
area (under the name P. bosniacus) represents this species.
Distribution in Europe: Balkan Peninsula to Northern Greece in the south, Northeastern Italy, Southern Slovakia
and West Ukraine (Hochkirch et al. 2016).

52. Pachytrachis striolatus (Fieber, 1853) = primorski skroviti konjic (Fig. 3E)
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Thamnotrizon appendiculatus Brunner von Wattenwyl, 1861 (Brunner von Wattenwyl in
Frauenfeld 1861)
Distribution in Croatia: Widespread along the Adriatic coast with the Dinaric Alps, and on some larger Adriatic

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islands (Korčula and Mljet Isl.).
Distribution in Europe: From Southern Switzerland and Northern Italy via the coastal part of the Western Balkan
Peninsula to Montenegro (Hochkirch et al. 2016).

genus Pholidoptera Wesmaël, 1838

53. Pholidoptera aptera (Fabricius, 1793) = gorski kožokrili konjic


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Mountains of Pannonian Croatia and along the Dinaric Alps (absent from mountains with
stronger Mediterranean influence, e.g. Dinara Mt.).
Distribution in Europe: Submontane to alpine regions from Northern Greece, Central Italy and Sardinia in the south
to Southern Poland and Northern Czech Republic in the north, and from Southeastern France in the west to
Romania in the east. Found also in the lowlands of Southern Bulgaria (Hochkirch et al. 2016).
Taxonomic note: There are numerous described subspecies, with either ill–defined diagnostic characters or
descriptions of which were based on small series of specimens, resulting in possible taxonomic irrelevance of the
characters used. Validity and presence in Croatia should be investigated for the following subspecies: P. a. aptera
(Fabricius, 1793) (described from the Alps), P. a. gjorgjevici Karaman, 1960 (described from the Republic of
Macedonia), and P. a. karnyi Ebner, 1908 (described from Bosnia & Herzegovina).

54. Pholidoptera dalmatica (Krauss, 1879) = dinarski kožokrili konjic (Fig. 5F)
First faunistic record for Croatia: Krauss 1879
Also reported as: P. maritima Zeuner, 1931 syn. nov. (Zeuner 1931), P. brachynota Ramme, 1951 syn. nov.
(Ramme 1951), regarded synonymous with P. d. dalmatica (Krauss, 1879)
Distribution in Croatia: Common in the Dinaric Alps and along the coast, on numerous Adriatic islands.
Distribution in Europe: Endemic to the Balkans—Dinaric Alps from Trieste (Italy) via Slovenia and the coastal
part of Croatia to Montenegro (Hochkirch et al. 2016).
Taxonomic note: There are three subspecies of P. dalmatica described from Croatia—(1) P. d. brachynota Ramme,
1951 (described from Mt. Velebit, near Senj), (2) P. d. dalmatica (Krauss, 1879) (described from Grobnik near
Rijeka, and Bakar), (3) P. d. maritima Zeuner, 1931 (described from Hvar Isl.). Harz (1969) (after Ramme 1951)
noted on P. d. brachynota distribution: „In der montanen Region vom Velebit bis in das Innere von BiH, verenzelt
auch in Dalmatien.“ (“In montainous regions from Velebit to the inner BiH, sporadically also in Dalmatia.”).
Difference from P. d. dalmatica in females is the subgenital plate shape (without prominent basal sclerites), and the
great variability of titillators: some specimens have only one or two apical teeth or no apical teeth, like P.
griseoaptera (Harz 1969). There is a continuity in distribution and high morphological variability in shape of cerci,
pronotum, subgenital plate, titillators, length of wings, and coloration of P. dalmatica from Učka Mt. in Istria along
the Dinaric arch to Southern Croatia (e.g. Snježnica Mt. in Dubrovnik area). Morphologically, specimens from
Učka Mt., Ćićarija Mt., Velika Kapela Mt., Velebit Mt. and Plješivica Mt. (Rebrina, Skejo & Tvrtković,
unpublished data) belong to the very same taxon, namely P. d. dalmatica. Since we did not find any relevant
differences on a large amount of material, we synonymize P. d. brachynota syn. nov. with the nominal subspecies.
Some authors consider P. d. maritima a separate species (Ingrisch & Pavićević 2012), but without any evidence
supporting its specific status. However, diagnostic characters of P. (d.) maritima syn. nov. also fit the variability
range of P. d. dalmatica, thus we do not regard it a separate taxon, but a synonym of P. d. dalmatica.

55. Pholidoptera fallax (Fischer, 1853) = bjelorubi kožokrili konjic


First faunistic record for Croatia: Brunner von Wattenwyl 1861
Also reported as: Thamnotrizon austriacus Türk, 1860 (Brunner von Wattenwyl 1861)
Distribution in Croatia: Widely distributed but not common in Pannonian Croatia, present in other regions under
stronger influence of continental climate, including Istria, Kvarner and the larger part of Dinaric Alps.
Distribution in Europe: Large parts of the Balkan Peninsula, also Italy, Sardinia and Sicily, reaching Slovakia in the
north and Southeastern France in the west, easternmost border of its distribution being Moldova (Hochkirch et al.
2016).

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56. Pholidoptera femorata (Fieber, 1853) = mramorni kožokrili konjic
First faunistic record for Croatia: Krauss 1879
Also reported as: Thamnotrizon noctivagus Krauss, 1879 (Krauss 1879)
Distribution in Croatia: Istria, Kvarner (with adjacent islands), Dalmatia and the Dinaric Alps. Not frequent.
Distribution in Europe: Mediterranean part, from Central France to the Peloponnesus (Greece), including islands
such as Corsica, Sardinia and Sicily (Hochkirch et al. 2016).

57. Pholidoptera frivaldszkyi (Herman, 1871) = kožokrili konjic Frivaldszkoga


First faunistic record for Croatia: Szövényi et al. 2018
Distribution in Croatia: Hitherto known from a single locality (Ljubina poljana) on Poštak Mt., Dinaric Alps
(Szövényi et al. 2018). This is the westernmost locality presently known for the species.
Distribution in Europe: From the Croatian/Bosnian border in the west and Slovakia in the north to Southern
Bulgaria and Northern Greece in the south, and eastern Ukraine in the east (Hochkirch et al. 2016).
Nomenclatural note: Herman designated the original description of the species two times, in Hungarian (1871a)
and also in German (1871b). He used in these publications different specific epithetons (Frivaldszkyi, Friwaldszkyi
and Friwaldskyi). However, later a fourth spelling (frivaldskyi) was used most frequently, which originated from
Brunner von Wattenwyl (1882). Puskás (2015) acted as the first reviser and fixed frivaldszkyi as the correct original
spelling. Thus, this latter is the available epitheton according to the article 19.3. of the International Code of the
Zoological Nomenclature (ICZN 1999).

58. Pholidoptera griseoaptera (De Geer, 1773) = riđi kožokrili konjic


First faunistic record for Croatia: Graber 1870
Also reported as: Thamnotrizon cinereus Zetterstedt (Graber 1870) [correct author of Gryllus cinereus is Gmelin,
1790]
Distribution in Croatia: Very common in the inland part (Dinaric Alps and Pannonian region), as well as in Istria
and Kvarner. Not recorded in Dalmatia, Dubrovnik area and the Adriatic islands.
Distribution in Europe: Widely distributed in the most of the continent, from southern Finland in the north and
Ireland in the west to Southern Italy and Northern Greece in the south, reaching Eastern European Russia in the
east (Hochkirch et al. 2016).

59. Pholidoptera littoralis (Fieber, 1853) = zelenotrbi kožokrili konjic


First faunistic record for Croatia: Krauss 1879
Also reported as: Thamnotrizon similis Brunner von Wattenwyl, 1861 (Brunner von Wattenwyl 1861)
Distribution in Croatia: Abundant in Istria and the Dinaric Alps (north of Dinara Mt.), recorded also in Kvarner. Its
presence in Dalmatia (Us 1967) should be confirmed.
Distribution in Europe: Inhabits large, but often discontinuous area in Southern Europe, from the Alpine region at
Italian/French border via Northeastern Balkan Peninsula to the Carpathian arc of Romania (Zuna-Kratky et al.
2016).
Taxonomic note: Terra typica of Pterolepis littoralis in Harz (1969) and on the Orthoptera Species File (Cigliano et
al. 2018) was wrongly designated as „Istrien—Krain“. Namely, in the original description by Fieber (1853), a
single locality was given: Trieste (Italy). Label of the holotype agrees with the description (labeled Tergest[inus] =
Trieste). Furthermore, terra typica of Thamnotrizon similis Brunner von Wattenwyl, 1861 is designated as “Istrien”
in Harz (1969), but the correct terra typica is Mehadia in Romania (Brunner von Wattenyl 1861). The record of P.
l. similis from Josipdol (Brunner von Wattenyl 1861) is questionable, as is the very existence of the subspecies in
Croatia. It could be either a misidentification of P. frivaldszkyi, recently found in Poštak Mt. (see notes under P.
frivaldszkyi) or wrongly identified P. l. littoralis.

genus Platycleis Fieber, 1853

60. Platycleis affinis Fieber, 1853 = jednogrbi šikarski konjic

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First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Along the Adriatic coast, and some Dinaric Mts. (Dinara, Velebit Mt.) influenced by the
Mediterranean climate, numerous Adriatic islands.
Distribution in Europe: Mostly Mediterranean, from Portugal in the west to Southern European Russia in the east,
and from Crete and Cyprus in the south to Eastern Austria, Southern Slovakia, and France in the north (Hochkirch
et al. 2016).

61. Platycleis albopunctata (Goeze, 1778)


61.1. Platycleis albopunctata grisea (Fabricius, 1781) = bezgrbi šikarski konjic
First faunistic record for Croatia: Graber 1870
Also reported as: P. denticulata (Panzer, 1796) (Us 1967)
Distribution in Croatia: Probably widely distributed, found in all regions except coastal Dalmatia and Dubrovnik
area.
Distribution in Europe: P. albopunctata occurs from Southern Spain, Sicily and Crete in the south to Southern
Norway and Estonia in the north, reaching Southern European Russia in the east (Hochkirch et al. 2016).

62. Platycleis intermedia (Serville, 1838) = dvogrbi šikarski konjic


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: In all regions except the Pannonian, numerous findings from Adriatic islands.
Distribution in Europe: Mediterranean region from the Iberian Peninsula to the Romanian Black Sea coast and
Moldova, eastern range border still unclear (Hochkirch et al. 2016).

63. Platycleis romana Ramme, 1927 = istarski šikarski konjic


First faunistic record for Croatia: Kohlich 1975
Distribution in Croatia: Istria and Istrian Dinaric Mts.
Distribution in Europe: Italy, but reaching Slovenia and Northern Croatian coast (Buzzetti et al. 2016).

genus Psorodonotus Brunner von Wattenwyl, 1861

64. Psorodonotus illyricus Ebner, 1923 = ilirski oklopljeni konjic (Fig. 6A)
First faunistic record for Croatia: Pungur 1899
Also reported as: Psorodonotus fieberi (Frivaldszky, in Fieber 1853) (Pungur 1899)
Distribution in Croatia: Along the Dinaric Alps from Ćićarija to Dinara Mt. Records for Istria (Kirby 1906, Ebner
1908) relate to Učka or Ćićarija Mt.
Distribution in Europe: Endemic to the Western Balkans—along the Dinaric Alps from Southern Slovenia and
Northwestern Croatia to the central regions of Montenegro. Contact zone with P. macedonicus is suspected along
the Drina—Tara River Valleys (Chobanov et al. 2016, Kaya et al. 2015).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

genus Rhacocleis Fieber, 1853

65. Rhacocleis buchichii Brunner von Wattenwyl in Herman, 1874 = Bučićev konjic grmušar
First faunistic record for Croatia: Herman 1874
Also reported as: R. bucchichii (Dubrony 1878), R. bucchichi (Bucchich [Bučić] 1886), R. buchicii (Karny 1907a),
R. bucchici (Karny 1907b), R. bucchicii (Heller 1988)
Distribution in Croatia: Described from Hvar Isl., recorded also on Brač and Korčula Isl. (Adriatic islands) with
some new findings from Dinaric mountains of Dalmatia (Biokovo Mt. (Wagner 2015) and on the southern slopes of
Troglav Mt.: Greda—850 m a.s.l. (Skejo 29.VIII.2014).
Distribution in Europe: Endemic to Croatia (Hochkirch et al. 2016).
IUCN Red List status: Endangered (Europe and EU28) (Hochkirch et al. 2016).

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FIGURE 6. Diversity of Croatian Ensifera—Tettigoniidae: Tettigoniinae, Mogoplistidae, Gryllidae, Rhaphidophoridae. A.
Psorodonotus illyricus (Zavižan 30.VIII.2015. F. Rebrina), B. Arachnocephalus vestitus (Lokrum 29.VIII.2017. J. Skejo), C.
Pseudomogoplistes squamiger (Lokrum 29.VIII.2017. J. Skejo), D. Oecanthus pellucens (Đurđevački peski 19.VII.2014. F.
Rebrina), and E. Troglophilus neglectus (Romualdova špilja / cave 19-20.V.2018. M. Malenica).

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66. Rhacocleis germanica (Herrich-Schäffer, 1840) = konjic grmušar
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Thamnotrizon pallidus Brunner von Wattenwyl, 1861 (Brunner von Wattenwyl in Frauenfeld
1861), R. discrepans Fieber, 1853 (Brunner von Wattenwyl 1861)
Distribution in Croatia: Widely distributed in the Mediterranean part and the Dinaric Alps, less common in the
Pannonian region.
Distribution in Europe: From Southeastern France in the west to South Ukraine and Anatolia (Turkey) in the east,
including Mediterranean islands such as Corsica, Crete and Corfu (Hochkirch et al. 2016).

genus Roeseliana Zeuner, 1941

67. Roeseliana roeselii (Hagenbach, 1822) = Roeselov livadni konjic


First faunistic record for Croatia: Graber 1870
Also reported as: Platycleis brevipennis (Charpentier, 1825) (Graber 1870)
Distribution in Croatia: Common in the Pannonian part, Istria, Kvarner, and along the Dinaric Alps, not recorded in
the largest part of the Adriatic coast (Dalmatia, Dubrovnik area) with adjacent islands.
Distribution in Europe: From Northern Spain, Northern Italy and Bulgaria in the south to the north of Sweden and
Finland in the north, and Eastern European Russia in the east (Hochkirch et al. 2016).

genus Sepiana Zeuner, 1941

68. Sepiana sepium (Yersin, 1854) = riđoglavi konjic


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Present in the Mediterranean part and the Dinaric Alps, with most records from the Adriatic
coast and islands.
Distribution in Europe: Widespread in the Mediterranean from Southwestern France and Northeastern Spain to
Southern Balkan Peninsula, all the way to Crimea in the east. Present on large Mediterranean islands such as
Corsica, Sardinia, Sicily and Crete (Hochkirch et al. 2016).

genus Tessellana Zeuner, 1941

69. Tessellana orina (Burr, 1899) = južni šikarski konjic


First faunistic record for Croatia: Brunner von Wattenwyl 1882
Also reported as: Platycleis nigrosignata Costa, 1863 (Brunner von Wattenwyl 1882)
Distribution in Croatia: Rare, in the eastern part of Dinaric Alps from Dinara Mt. (Rebrina et al. 2015) to
Dubrovnik area: Ombla (Ramme 1951).
Distribution in Europe: Endemic to the Balkans—western part of the Balkan Peninsula from Croatia to the
Peloponnesus (Hochkirch et al. 2016).
Distributional note: In Croatia it was recorded under the names (1) T. nigrosignata by Brunner von Wattenwyl
(1882), Redtenbacher (1900), Kuthy (1908), Ramme (1951), and (2) T. orina by Kuthy (1908), Ramme (1951),
Skejo & Rebrina (2013) and Rebrina et al. (2015). Italian endemic T. nigrosignata is not present in the Balkans
(Massa & Fontana 2011), but replaced in the western part of the Balkan Peninsula by T. orina, while in the eastern
by T. carinata (Berland & Chopard, 1922).

70. Tessellana tessellata (Charpentier, 1825) = prugastokrili šikarski konjic


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Along the Adriatic coast and the Dinaric Alps, with no records from Kvarner area and the
Adriatic islands.
Distribution in Europe: Mediterranean part, from the Canary Islands in the west to Dalmatia in the east, reaching

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Northern France in the north. Occurs also from Southern Ukraine to Eastern European Russia (Hochkirch et al.
2016).

genus Tettigonia Linnaeus, 1758

71. Tettigonia balcanica Chobanov & Lemonnier–Darcemont, 2014 = balkanski zeleni konjic
First faunistic record for Croatia: Chobanov et al. 2014
Distribution in Croatia: Dinaric Alps and mountains of Dalmatia, not well documented, being only recently
described: currently known from Poštak Mt. (Chobanov et al. 2014) and Biokovo Mt. (Skejo & Ozimec
27.VII.2016).
Distribution Europe: Endemic to the Balkans—mountains of the Balkan Peninsula, from Southern Croatia, Bosnia
and Herzegovina, and Serbia, to Western and Central Bulgaria, and Northwestern Greece (Chobanov et al. 2016).

72. Tettigonia cantans (Fuessly, 1775) = kratkokrili zeleni konjic


First faunistic record for Croatia: Nagy 2006 [or Padewieth 1900—to be checked for T. balcanica]
Distribution in Croatia: Present in the inland part of the country (Western Dinaric Alps and the Pannonian region),
replaced by T. balcanica in the Dinaric Alps south of Velebit Mt. Kuthy's (1908) record from Krivi put (Velika
Kapela Mt.) should be revised.
Distribution Europe: From Northern Spain in the west and Central Finland in the north, to Sardinia and Southern
Italy in the south and Eastern European Russia in the east (Hochkirch et al. 2016).

73. Tettigonia caudata (Charpentier, 1842) = bodljonogi zeleni konjic


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Not common, present in the Pannonian region, Istria, Kvarner, Dalmatia and the Dinaric
Alps, not recorded in Dubrovnik area and the Adriatic islands
Distribution Europe: From Eastern Switzerland in the west and Northeastern Germany in the north, to the
Peloponnesus (Greece) in the south and Eastern European Russia in the east (Hochkirch et al. 2016).

74. Tettigonia viridissima (Linnaeus, 1758) = veliki zeleni konjic


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Widely distributed, common in all regions.
Distribution Europe: Widespread in the entire continent, from Portugal in the west and Southern Finland in the
north, to Southern Spain and Mediterranean islands (Lampedusa, Crete and Cyprus) in the south and Eastern
European Russia in the east (Hochkirch et al. 2016).

genus Yersinella Ramme, 1933

75. Yersinella raymondi (Yersin, 1860) = krhka jersinela


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Frequently found along the Adriatic coast, the Dinaric Alps, and on most islands (Cres,
Hvar Isl.).
Distribution in Europe: Mediterranean part, from Northeastern Spain to Western Greece, including Corsica,
Sardinia and Sicily (Hochkirch et al. 2016).

genus Zeuneriana Ramme, 1951

76. Zeuneriana amplipennis (Brunner von Wattenwyl, 1882) = (istočni) močvarni konjic
First faunistic record for Croatia: this study
Distribution in Croatia: A large population of the species was recently found in the Pannonian region, Vrbanja

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County (near Strošinci village), close to Jamena, the nearest known locality in Vojvodina, Serbia (Pavićević et al.
2014) (Ivković pers. comm).
Distribution in Europe: Along the lower courses of Sava and Danube rivers, currently known from the Serbian/
Croatian/Bosnian border area (see above), via Serbia (Pavićević et al. 2014) to Romania (Chobanov et al. 2016).
IUCN Red List status: Endangered (Europe), Near Threatened (EU28) (Hochkirch et al. 2016).
Taxonomic note: Taxonomic relationship of Z. amplipennis and Z. marmorata (Fieber, 1853) is unclear:
populations east of Croatia are named Z. amplipennis and west of Croatia Z. marmorata. The latter could simply be
a relict westermost population of the same species, expected to be found on other similar localities along the Sava
River.

infraorder Gryllidea

superfamily Grylloidea Laicharting, 1781

family Mogoplistidae Brunner von Wattenwyl, 1873 = ljuskavi šturci

subfamily Mogoplistinae Brunner von Wattenwyl, 1873

genus Arachnocephalus Costa, 1855

77. Arachnocephalus vestitus Costa, 1855 = ljuskavi šturak grmušar (Fig. 6B)
First faunistic record for Croatia: Brunner von Wattenwyl 1882
Also reported as: A. dalmatinus Saussure, 1877 (Saussure 1877)
Distribution in Croatia: Recorded in the coastal area (including southern slopes of the Dinaric Alps) and on
numerous Adriatic islands. Most likely common but overlooked.
Distribution in Europe: Southern part of the continent, from Southern France, northern Italy and Slovenia in the
north, to Portugal and Mediterranean islands (Lampedusa, Sicily, Crete, Cyprus) in the south, and Southeastern
Ukraine in the east (Hochkirch et al. 2016).

genus Mogoplistes Serville, 1838

78. Mogoplistes brunneus Serville, 1838 = tamni ljuskavi šturak


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Rather widespread and common along the Adriatic coast with islands (Hvar, Mljet, Šipan)
and the southern slopes of the Dinaric Alps.
Distribution in Europe: Widespread in the Mediterranean, from Northeastern Spain in the west and Southern
France and Slovenia in the north, to Sicily in the south and Greece in the east (Hochkirch et al. 2016).

genus Paramogoplistes Gorochov, 1984

79. Paramogoplistes novaki (Krauss, 1888) = Novakov ljuskavi šturak


First faunistic record for Croatia: Krauss 1888
Distribution in Croatia: Until recently known only from Hvar Isl., recorded along the Krčić River (Rebrina &
Brigić 2017), Neretva River mouth (Vujčić–Karlo 12.X.1997) and on Mljet Isl. near temporary pond (Skejo & Zec
23.IX.2013). Probably has much broader distribution both on Adriatic islands and the mainland, but likely
overlooked or misidentified as M. brunneus in the past.
Distribution in Europe: Outside Croatia, historical records exist for Mount Parnassus (Greece), Sardinia and Circeo
in Lazio region (Italy), generally considered doubtful due to possible misidentifications (Hochkirch et al. 2016).
IUCN Red List status: Data Deficient (Europe and EU28) (Hochkirch et al. 2016).

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genus Pseudomogoplistes Gorochov, 1984

80. Pseudomogoplistes squamiger (Fischer, 1853) = obalni ljuskavi šturak (Fig. 6C)
First faunistic record for Croatia: Fischer 1853
Distribution in Croatia: Widely distributed along the Adriatic coast with adjacent islands.
Distribution in Europe: Widespred along the Mediterranean coasts of Spain, France, Italy, the Balkans, and large
Mediterranean islands (Sardinia, Sicily, Corsica) (Hochkirch et al. 2016).

family Trigonidiidae Saussure, 1874

subfamily Nemobiinae Saussure, 1877 = patuljasti šturci

genus Pteronemobius Jacobson, 1907

81. Pteronemobius heydenii (Fischer, 1853) = močvarni patuljasti šturak


First faunistic record for Croatia: Graber 1870
Also reported as: P. lineolatus (Brullé, 1835) (Rebrina et al. 2015)
Distribution in Croatia: Recorded in all regions, not widespread but locally abundant.
Distribution in Europe: From Northern France and the Czech Republic in the north to the Iberian Peninsula and the
Mediterranean islands (Lampedusa, Rhodes, Cyprus) in the south, and Southern European Russia in the east
(Hochkirch et al. 2016).

genus Stenonemobius Gorochov, 1981

82. Stenonemobius bicolor (Saussure, 1877)


82.1. Stenonemobius bicolor ponticus Gorochov, 1984 = dugokrili patuljasti šturak
First faunistic record for Croatia: Chobanov 2003, after Orthoptera Species File and DORSA
Distribution in Croatia: According to data provided by Orthoptera Species File (Cigliano et al. 2018, first time
published in Chobanov 2003), a specimen from Senj (=Zengg) (Kvarner) collected by M. Padewieth (= F. Dobijaš)
is deposited in the collection of Museum für Naturkunde, Berlin (MfN). It was initially identified by Ramme as
Nemobius tartarus and later redetermined by Gorochov as S. b. ponticus. Another specimen from Senj collected
also by Dobijaš and determined by Ebner in 1927 as “Pteronemobius heydenii concolor = Nemobius tartarus
Sauss” is deposited in the Orthoptera collection of Hungarian Natural History Museum in Budapest (Puskás et al.
2018). Kleukers (pers. comm.) found in Trieste Collection a specimen with the following data: Korčula Isl.:
Lumbarda, sex ?, 22.VIII.1925, leg. P. Novak, det. G. Müller, with the name Pteronemobius tartarus written on the
label, which could be either P. heydenii or S. b. ponticus.
Distribution in Europe: Scattered findings in the Crimea, Bulgaria, Hungary, Romania, Croatia and Serbia
(Chobanov et al. 2016).
IUCN Red List status: Data Deficient (Europe and EU28) (Hochkirch et al. 2016).

subfamily Trigonidiinae Saussure, 1874 = trčkoliki šturci

genus Trigonidium Rambur, 1838

83. Trigonidium cicindeloides Rambur, 1838 = trčkoliki šturak


First faunistic record for Croatia: Adamović 1964
Distribution in Croatia: Until recently known only from the surroundings of Dubrovnik, its presence ascertained on
some Adriatic islands (Cres, Krk, Pašman, Dugi Otok, Brač, Molat) (Linić 14.VIII.2014, Rebrina & Brigić
19.IX.2015, Skejo VIII.2016, Rebrina & BIUS Section of Orthopterology 28.–31.IX.2017).

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Distribution in Europe: Found in the Mediterranean region from the Canary Islands, the Iberian Peninsula,
Lampedusa, Crete and Cyprus in the south to Croatian coast and Southern France in the north (Hochkirch et al.
2016).

family Gryllidae Laicharting, 1781 = šturci / popci

subfamily Gryllinae Laicharting, 1781 = pravi šturci / popci

genus Acheta Fabricius, 1775

84. Acheta domesticus (Linnaeus, 1758) = kućni šturak


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Present in all regions, usually associated with human settlements.
Distribution in Europe: Very widespread, present in almost all European countries except Fennoscandian
Peninsula. Natural distribution still unclear, probably all populations north of the Alps introduced by humans
(Hochkirch et al. 2016).

genus Eumodicogryllus Gorochov, 1986

85. Eumodicogryllus bordigalensis (Latreille, 1804) = primorski bjeloprugi šturak


First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Gryllus burdigalensis Latreille, 1804 (Frauenfeld 1861)
Distribution in Croatia: Distributed along the Adriatic coast and on numerous islands.
Distribution in Europe: Southern part of the continent, including many Mediterranean islands, reaching Southern
Germany and the Czech Republic in the north and Eastern European Russia in the east. Range recently expanded to
Eastern Austria, Germany and the Netherlands, but the extent of introduction by humans is unknown. Populations
on the Canary Islands, the Azores and Madeira also likely introduced (Hochkirch et al. 2016).

genus Gryllus Linnaeus, 1758

86. Gryllus bimaculatus De Geer, 1773 = dvopjegi šturak


First faunistic record for Croatia: Graber 1870
Also reported as: Gryllus capensis (Fabricius, 1775) (Graber 1870)
Distribution in Croatia: Recorded in Dalmatia (Karin, Vodice, Šibenik, Primošten, Kaštel Sućurac, Stobreč, Postira
on Brač Isl.: Skejo VI–IX.2016) and Dubrovnik region (Zaton: Skejo 2.VIII.2016) with adjacent islands. Probably
has much broader distribution than previously thought, and was possibly often mistaken for G. campestris. There is
a single historical record of this species in the Pannonian region (Vinkovci; Graber 1870).
Distribution in Europe: Widespread in the Mediterranean, reaching Croatia in the north and Southern Ukraine in
the northeast (Hochkirch et al. 2016).

87. Gryllus campestris Linnaeus, 1758 = livadni šturak


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Widespread and very frequent in all regions.
Distribution in Europe: Common in almost the entire continent, from Southern England, Denmark and Latvia in the
north, to the Iberian Peninsula, the Peloponnesus (Greece) and the Mediterranean islands in the south (Hochkirch et
al. 2016).

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genus Melanogryllus Chopard, 1961

88. Melanogryllus desertus (Pallas, 1771) = tamni šturak


First faunistic record for Croatia: Krauss 1879
Also reported as: Gryllus melas (Charpentier, 1825) (Fieber 1853)
Distribution in Croatia: Widely distributed in all regions.
Distribution in Europe: From Southern Slovakia and Southern Switzerland (where it is extinct) in the north, to the
Iberian Peninsula, Greece and the Mediterranean islands of Sicily and Cyprus in the south (Hochkirch et al. 2016).

genus Modicogryllus Chopard, 1961

89. Modicogryllus frontalis (Fieber, 1844) = crni bjeloprugi šturak


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Known from several localities in the Pannonian region. Not common.
Distribution in Europe: From Northeastern France in the west and Lithuania in the north, via Southern Germany,
Austria and Southern Poland to Northern Bulgaria in the south, and Southern European Russia in the east. Previous
records from FYR Macedonia and Greece probably refer to M. truncatus (Hochkirch et al. 2016).

90. Modicogryllus truncatus (Tarbinsky, 1940) = močvarni bjeloprugi šturak


First faunistic record for Croatia: Puskás et al. 2018
Distribution in Croatia: Recorded for the first time in 2011 on Psunj Mt. and Dilj Mt. in the Pannonian region
(Šumetlica, Donji Slatinik: Puskás et al. 2018). This species is easily mistaken for M. frontalis, reliable diagnostic
characters being male genitalia (Harz 1969) and song (Iorgu & Iorgu 2007).
Distribution in Europe: Southeastern Europe, currently known from Hungary, Croatia, Serbia, FYR Macedonia,
Bulgaria, Romania, Greece and European Turkey. Probably present also in Kosovo and Bosnia and Herzegovina
(Chobanov et al. 2016).

subfamily Gryllomorphinae Saussure, 1877 = šturkolike

genus Gryllomorpha Fieber, 1853

91. Gryllomorpha dalmatina (Ocskay, 1833) = primorska šturkolika


First faunistic record for Croatia: Fischer 1853
Also reported as: Gryllus apterus (Herrich-Schäffer, 1838) (Frauenfeld 1861)
Distribution in Croatia: Common in the Mediterranean region and the Dinaric Alps, inhabiting localities with the
Mediterranean influence.
Distribution in Europe: Widespread in the Mediterranean, from Northeastern Spain in the west and Southern
Switzerland in the north, to Crete and Sicily in the south and Crimea in the east (Hochkirch et al. 2016).

subfamily Oecanthinae Blanchard, 1845 = prozirni šturci

genus Oecanthus Serville, 1831

92. Oecanthus dulcisonans Gorochov, 1993 = veliki prozirni šturak


First faunistic record for Croatia: Puskás et al. 2018
Distribution in Croatia: So far known from two localities—Pristanište, Mljet Isl. (15.X.2001, leg. Gy. Rozner, in
Puskás et al. 2018) and Šibenik (Dalmatia) (Skejo VIII.–X.2016). Probably more widespread along the Dalmatian
coast, but easily misidentified as O. pellucens.
Distribution in Europe: Found on the Canary Islands and in the Mediterranean from the Iberian Peninsula to
Cyprus. Distribution still insufficiently known (Hochkirch et al. 2016).

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93. Oecanthus pellucens (Scopoli, 1763) = vinogradski prozirni šturak (Fig. 6D)
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Acridium italicum Latreille (Kollar 1845) [correct author of Acheta italicus is Fabricius, 1781]
Distribution in Croatia: Widely distributed and rather common, recorded in all regions.
Distribution in Europe: Widespread in the southern part of the continent, to the north presently reaching Southern
England, the Netherlands and Berlin region in Germany (Hochkirch et al. 2016).

superfamily Gryllotalpoidea Leach, 1815

family Gryllotalpidae Leach, 1815 = rovci / mrmci

subfamily Gryllotalpinae Leach, 1815

genus Gryllotalpa Latreille, 1802

94. Gryllotalpa gryllotalpa (Linnaeus, 1758) = sjeverni rovac


First faunistic record for Croatia: Bogdanović et al. 2017
Also reported as: G. vulgaris Latreille, 1804 [partim] (e.g. Pungur 1899, Pokopac 1956)
Distribution in Croatia: Widely distributed according to the present data, recorded in all regions, but existing data
lack information about diagnostic traits (titilator shape, song or molecular characteristics). Therefore, all records
need confirmation (see Potential species).
Distribution in Europe: Widespread, from England, South Sweden and Northwestern European Russia in the north,
to the Iberian Peninsula, Italy and the FYR Macedonia in the south (Hochkirch et al. 2016).

95. Gryllotalpa sp. [undescribed one or two new species, or a sibling species known from Italy and Montenegro]
First faunistic record for Croatia: this study
Also reported as: G. communis (Cubich 1875), G. gryllotalpa [partim] (e.g. Us 1967, Heller et al. 1998, Schuster et
al. 1998) or G. vulgaris Latreille, 1804 [partim] (e.g. Novak 1888, Padewieth 1900)
Distribution in Croatia: First collected and stored specimens from the western part of the Mediterranean region:
Istria, near Šišan: Tvrtković & Malenica 15.V.2013, 29.VI.2013, Krk Island, Ponikve: Skejo & Roesti 8.VI.2014.
All previous records of Gryllotalpa from the Mediterranean region should be revised.
Distribution in Europe: Unknown, possibly endemic to the Balkans, but likely also present in Italy.

96. Gryllotalpa stepposa Zhantiev, 1991 = stepski rovac


First faunistic record for Croatia: Bogdanović et al. 2017
Also reported as: G. gryllotalpa [partim] (e.g. Us 1967) or G. vulgaris Latreille, 1804 [partim] (e.g. Pokopac 1956)
Distribution in Croatia: Heretofore known only from a single locality (Valpovo near Drava River) in the Pannonian
region.
Distribution in Europe: Occurs from Hungary and Eastern Balkans to the southern part of European Russia
(Hochkirch et al. 2016).

family Myrmecophilidae Saussure, 1874 = mravoljupci

subfamily Myrmecophilinae Saussure, 1874

genus Myrmecophilus Berthold, 1827

97. Myrmecophilus hirticaudus Fischer von Waldheim, 1846 = istočni mravoljubac


First faunistic record for Croatia: Stalling & Birrer 2013
Distribution in Croatia: Known only from Cres Isl., probably more widely distributed but overlooked due to the

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lack of research on this family.
Distribution in Europe: Currently known from Crimea, Southern Bulgaria, Southeastern FYR Macedonia,
mainland Greece and Cres Isl. (Croatia). Probably more widely distributed (Chobanov et al. 2016).

98. Myrmecophilus myrmecophilus (Savi, 1819) = mediteranski mravoljubac


First faunistic record for Croatia: Baccetti 1992
Also reported as: Myrmecophila ochracea Fischer, 1853 (Novak 1888), M. acervorum (Panzer, 1799) [partim] (e.g.
Us 1961).
Distribution in Croatia: Known only from Split, probably more widely distributed but overlooked due to the lack of
research on this family.
Distribution in Europe: Widespread in the Northeastern Mediterranean, presently known from France (mainland,
Corsica), Italy (mainland), Croatia (coast) and Greece (Crete, Samos, Thessaloniki region) (Braud et al. 2016).

superfamily Rhaphidophoroidea Walker, 1869

family Rhaphidophoridae Walker, 1869 = spiljski konjici / zrikavci

subfamily Dolichopodainae Brunner von Wattenwyl, 1888

genus Dolichopoda Bolívar, 1880

99. Dolichopoda araneiformis (Germar in Burmeister, 1838) = paukoliki / dugonogi spiljski konjic
First faunistic record for Croatia: Burmeister 1838
Also reported as: D. palpata (Sulzer, 1776) (Fieber 1853, Novak 1888)
Distribution in Croatia: Distributed along the Adriatic coast with adjacent islands from North Dalmatia (Šibenik:
Skejo & Antolos 25.XII.2012) southwards to Dubrovnik area, mostly in warm lowland areas. Found exclusively in
karst caves.
Distribution in Europe: Stenoendemic to the Western Balkans—from Croatia (N Dalmatia) to the south, including
Southwest Bosnia and Herzegovina and the northern coast of Montenegro (Chobanov et al. 2016).
Nomenclatural note: In the original description, Burmeister wrote “Germ.” after the species’ name, implying that
Germar is the author of the description..

subfamily Troglophilinae Krauss, 1879

genus Troglophilus Krauss, 1879

100. Troglophilus cavicola (Kollar, 1833) = obični spiljski konjic


First faunistic record for Croatia: Jurinac 1886
Also reported as: T. brevicauda Chopard, 1934 (Us 1970), T. neglectus Krauss, 1879 [partim] (Bucchich [Bučić]
1886)
Distribution in Croatia: According to Karaman et al. (2011), recorded in all regions of Croatia. In the past
sometimes wrongly identified as Troglophilus brevicauda Chopard, 1934 (records north of Plitvice Lakes: Us
1970).
Distribution in Europe: Central Europe south and east of the Alps and Balkan Peninsula, southern border of its
distribution in Greece (Willemse et al. 2016).

101. Troglophilus neglectus Krauss, 1879 = mramorni spiljski konjic (Fig. 6E)
First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Recorded in the entire western part of Croatia: in Istria and Istrian Dinaric mountains
(Ćićarija Mt. and Učka Mt.), on the islands (Cres and Krk), Gorski kotar (Risnjak Mt. and Kapela Mts.), as well as

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in the karst of Lika and North Dalmatia. There are isolated populations north of Sava River on Ravna Gora Mt. and
Ivanščica Mt., on Papuk Mt. in the Pannonian region (Karaman et al. 2011). However, all older data on the
southern border need redetermination because of the recent records of a sibling species, T. ovuliformis (probably
parapatric).
Distribution in Europe: Two separated areas of distribution—one comprising Republic of Macedonia, Kosovo,
Eastern Serbia, Northeastern Albania and Northwestern Bulgaria, and the other comprising Croatia, Northwestern
Bosnia and Herzegovina, Slovenia, Northeastern Italy and Austria. Introduced to Czechia, Germany and
Switzerland. Origin of the Maltese population uncertain (Chobanov et al. 2016).

102. Troglophilus ovuliformis Karny, 1907 = primorski spiljski konjic


First faunistic record for Croatia: Karaman et al. 2011
Also reported as: T. neglectus Krauss, 1879 [partim] (for details see Karaman et al. 2011)
Distribution in Croatia: Numerous localities, along the Adriatic coast and in the karst mountains of Istria, Dalmatia,
Dubrovnik area and on several islands (Cres, Hvar, Korčula, Mljet). All data need redetermination because of the
recent record of a new species (Allegrucci et al. 2017, see comments under Troglophilus sp. below) and unresolved
morphological differences inside T. neglectus—T. brevicauda—T. ovuliformis groups (Karaman et al. 2011).
Distribution in Europe: Endemic to Western Balkan—along the Adriatic Sea from Istria (Croatia) to Albania
(Chobanov et al. 2016).

103. Troglophilus sp. [undescribed new species]


First faunistic record for Croatia: Allegrucci et al. 2017
Also reported as: T. neglectus Krauss 1879 [partim] (Karny 1907), T. ovuliformis Karny, 1907 [possibly partim]
(Karaman et al. 2011).
Distribution in Croatia: According to the phylogenetic analysis published in Allegrucci et al. 2017, a sample from
Mljet Isl. shows genetic similarity with T. brevicauda group and clear separation from other Balkans Troglophilus
species. Thus, it represents a new, yet undescribed species.
Distribution in Europe: Heretofore known only from Mljet Isl., Croatia.

Omitted Ensifera species

family Tettigoniidae

subfamily Phaneropterinae

1. Isophya speciosa (Frivaldszky, 1868)


Although it is listed for Croatia in IUCN Red List (Hochkirch et al. 2016), no records exist for this species. Thus,
we omit it from this checklist.

2. Poecilimon brunneri Frivaldszky, 1907


A record from Hrvatsko Zagorje (Pannonian Croatia) (Ozimec & Ernoić (2008) after Kodinec (1938)) originating
from the list of orthopteran species presumably found by Franjo Koščec, an entomologist from Varaždin, is
possibly a misidentification. Ozimec & Ernoić (2008) confirmed that no documentation existed about the origin of
the list, or the existence of voucher specimens. There are no other records of this Balkans and Pannonian lowland
species (Hochkirch et al. 2016) in Croatia, and we consider its presence in the country unlikely.

3. Poecilimon jonicus (Fieber, 1853)


Listed for Croatia in Heller & Willemse (2013), probably due to the records from Dalmatia (Redtenbacher 1900),
which most likely refered to historical Austrian Dalmatia, today the coastal part of Montenegro. The closest
faunistic record is from Petrovac (=Castellastua) in Montenegro (Brunner von Wattenwyl 1882), which is currently
known as the westernmost distribution border of P. jonicus on the Balkan Peninsula (Hochkirch et al. 2016).

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4. Poecilimon thoracicus (Fieber, 1853)
Several authors mentioned the presence of this species in Austrian Dalmatia (Redtenbacher 1900, Us 1992), but it
does not necessarily indicate the Croatian part of the Adriatic coast (see comment above). There are no faunistic
records of P. thoracicus from Croatia. Therefore, we omit this species (Hochkirch et al. 2016) from the checklist.

subfamily Tettigoniinae

5. Modestana ebneri (Ramme, 1926)


Harz (1969) cited this species for Istria and Dalmatia, but there are no faunistic records of M. ebneri from Croatia.
It is distributed in the southern part of the Balkan Peninsula (Willemse et al. 2016), and its presence in Croatia is
unlikely.

6. Pachytrachis bosniacus Messina, 1979


Former record of this species from Sniježnica Mt. in the Dinaric Alps (Buzzetti 2006) was based on a
misidentification of P. gracilis (Skejo & Rebrina, new data after insight into Buzzetti’s Collection in Arzignano,
Italy). There are no other faunistic records of this little–known Bosnian species in Croatia.

7. Rhacocleis neglecta (Costa, 1863)


Recorded in Rijeka (= Fiume) by La Greca (1959), probably following an accidental introduction by ship from
Italy, where this species is widely distributed (Massa et al. 2013). There are no other records from Croatia, thus we
omit it from the checklist.

8. Tessellana carinata (Berland & Chopard, 1922)


Although Heller & Willemse (2013) list this species for Croatia, no faunistic records exist. The species is
distributed in the southeastern part of the Balkan Peninsula (Chobanov et al. 2016) and its presence in Croatia is
unlikely.

9. Tessellana nigrosignata (Costa, 1863)


Although old data from Dubrovnik coast exist (Adamović 1964), the presence of this species on the Balkan
Peninsula was disputed by Massa & Fontana (2011). Records from Croatia are probably misidentifications of T.
orina (see comments under T. orina).

10. Zeuneriana marmorata (Fieber, 1853)


Although listed for Croatia by some authors (Kirby 1906, Heller & Willemse 2013), no faunistic records exist.
Presence in Croatia is likely, but its taxonomic relationship with Z. amplipennis, recently recorded for Croatia,
remains unresolved (see comments under Z. amplipennis).

family Gryllidae

subfamily Nemobiinae

11. Pteronemobius lineolatus (Brullé, 1835)


A former record of this species from Sinj (Rebrina et al. 2015) was based on a misidentification of P. heydenii.
Since P. lineolatus is distributed only in the western part of the Mediterranean (Hochkirch et al. 2016), the
historical record from Cres Isl. (Schuster et al. 1998) is probably also a misidentification. Thus, we exclude this
species from the checklist, until firm evidence of its presence in Croatia is provided.

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family Myrmecophilidae

subfamily Myrmecophilinae

12. Myrmecophilus acervorum (Panzer, 1799)


Several authors cited this species for Dalmatia (Saussure 1877, Pungur 1891), but no faunistic records exist.
Considering the known distribution of M. acervorum in Europe (Hochkirch et al. 2016), its presence in Croatia is
likely, especially in the northeastern part of the country (Stalling, pers. comm.). However, we omit it from the
checklist until reliable faunistic records are provided.

13. Myrmecophilus ochraceus Fischer, 1853


The presence of this species in Croatia is improbable considering its known distribution in Europe (Buzzetti et al.
2016). Historical records from Hvar Isl., Split and Zadar (Bucchich 1886, Novak 1888, 1891) are most likely based
on misidentifications (Stalling, pers. comm.).

family Rhaphidophoridae

subfamily Dolichopodainae

14. Dolichopoda palpata (Sulzer, 1776)


Historical records from several localities along the coast and on Adriatic islands (Frauenfeld 1861, Brunner von
Wattenwyl 1882, Redtenbacher 1900, Karny 1908, Werner 1920) proved to be misidentifications of D.
araneiformis, after morphological and molecular data (Allegrucci et al. 2009). An inland record from Raduč, Lika
(Pungur 1899) is doubtful, since no voucher specimen known from that locality, and D. araneiformis has confirmed
data only from North Dalmatia southwards. Therefore, we exclude D. palpata, an Italian endemic species, found in
Sicily and Calabria (Allegrucci et al. 2011, Buzzetti et al. 2016), from the checklist of Croatian Orthoptera.

subfamily Aemodogryllinae Jacobson, 1905

15. Diestrammena asynamora (Adelung, 1902)


A male and a female specimens deposited in the Natural History Museum in Budapest with a label “Dalmácia”
(Puskás et al. 2018) could have been collected in any part of historical Dalmatia (see comment under P. jonicus).
The species was introduced to Europe probably from East Asia and it spreads easily between greenhouses
(Hochkirch et al. 2016), but there are no exact faunistic records from Croatia so far.

subfamily Troglophilinae

16. Troglophilus brevicauda Chopard, 1934


According to Karaman et al. (2011), all material designated as T. brevicauda from Croatia (Us 1970) was most
likely misidentified. Therefore, the former is excluded from the checklist of Croatian Orthoptera.

Suborder Caelifera = skakavci

infraorder Tridactylidea

superfamily Tridactyloidea Brullé, 1835

family Tridactylidae Brullé, 1835 = buhoskakavci / ksije

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subfamily Tridactylinae Brullé, 1835

genus Xya Latreille, 1809

1. Xya pfaendleri (Harz, 1970)


1.1. Xya pfaendleri pfaendleri (Harz, 1970) = crni buhoskakavac / tamna ksija (Fig. 7A)
First faunistic record for Croatia: Szövényi & Puskás 2012
Also reported as: X. variegata Latreille, 1809 [partim] (e.g. Pungur 1891)
Distribution in Croatia: Nominal subspecies only. Pannonian region only. There are suitable habitats in other parts
of Croatia as well, where its presence should be investigated in the future. Since Xya p. pfaedleri was separated
from X. variegata in 1970 (Harz), all older records of the latter species should be revised.
Distribution in Europe: From Cyprus and Rhodes (southern border of the distribution) to Czechia (northern
border), and from Slovenia (western border) to Southern Ukraine (eastern border) (Hochkirch et al. 2016).

2. Xya variegata Latreille, 1809 = šareni buhoskakavac / šarena ksija


First faunistic record for Croatia: this study
Distribution in Croatia: Known with certainty from two localities in Hrvatsko Zagorje, found syntopically with X.
pfaendleri. Reported also from Sisak (Frivaldszky 1868) and Vransko jezero on Cres Isl. (Us 1964), but those
records should be revised.
Distribution in Europe: From Northern Spain (western border of the distribution) to Eastern European Russia
(eastern border) and from the Peloponnesus (southern border) to Czechia (northern border) (Hochkirch et al. 2016).

infraorder Acrididea

superfamily Tetrigoidea Rambur, 1838

family Tetrigidae Rambur, 1838 = monaški skakavci / trnovratke

subfamily Tetriginae Rambur, 1838

genus Paratettix Bolívar, 1887

3. Paratettix meridionalis (Rambur, 1838) = obalni monaški skakavac


First faunistic record for Croatia: Krauss, 1879
Also reported as: Tettix dorhni Fieber, 1853 (Fieber 1853)
Distribution in Croatia: Not common, recorded along the Adriatic coast with adjacent islands. Probably overlooked
in the past.
Distribution in Europe: Mediterranean species, distributed from the Azores and the Canary Islands (eastern
distribution border) to Cyprus and European Turkey (western border) and from Sicily and Crete (southern border)
to the French Atlantic coast and Slovenia (northern border) (Hochkirch et al. 2016).

genus Tetrix Latreille, 1802

4. Tetrix bipunctata (Linnaeus, 1758) = dvopjegi / kratkoticalni monaški skakavac


First faunistic record for Croatia: Graber 1870
Also reported as: T. kraussi Saulcy, 1888 (Us 1967)
Distribution in Croatia: Frequent in Pannonian Croatia and Dinaric Alps north of Velebit Mt., recorded also in
several localities in Istria and Kvarner.
Distribution in Europe: From Northern Greece, the Iberian Peninsula, Sardinia and Central Italy (southern border
of the distribution) to Scandinavia (northern border) and Eastern European Russia (eastern border) (Hochkirch et
al. 2016).

46 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Taxonomic note. Some authors regard Tetrix kraussi Saulcy, 1888 a species distinct from T. bipunctata (e.g.
Lehmann 2004, Defaut 2012b), but morphological and molecular data (Skejo & Hochkirch unpublished data)
suggest it is only a wing morph of T. bipunctata and no synapomorphy (morphological nor genetic) confirms the
specific status of T. kraussi, thus we regard it synonymous with T. bipunctata.

FIGURE 7. Diversity of Croatian Caelifera—Tridactylidae, Tetrigidae, Acrididae: Acridinae, Catantopinae, Gomphocerinae.


A. Xya pfaendleri (Kloštarski peski 9.VIII.2013. J. Skejo), B. Tetrix subulata (Gorski kotar 4.VI.2015. J. Skejo), C. Acrida
ungarica (Murter IX.2012. J. Skejo), D. Pezotettix giornae (Murter IX.2012. J. Skejo), E. Chorthippus apicarius (Paklenica
26.VII.2014. J. Skejo), and F. Chorthippus mollis lesinensis (Biokovo 27.VII.2016. J. Skejo).

5. Tetrix bolivari Saulcy, 1901 = Bolívarov monaški skakavac


First faunistic record for Croatia: Szövényi & Puskás 2012
Also reported as: T. subulata (Linnaeus, 1758) [partim] (e.g. Padewieth 1900)
Distribution in Croatia: Despite being recorded for the first time only recently, the species seems to be widespread

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in Croatia—in both Pannonian and Dalmatian region, but possibly often overlooked (Skejo 2014).
Distribution in Europe: Distribution disjunct, probably a result of the species being overlooked, one part being
from Southern Spain to Northern France and the French Alps; the second from Northeastern Italy across the
Balkans to Southern Greece, Cyprus and Southern European Russia (Hochkirch et al. 2016).

6. Tetrix ceperoi (Bolívar, 1887) = Ceperojev monaški skakavac


First faunistic record for Croatia: Karaman 1960
Also reported as: T. c. balcanicus Karaman 1960 (Karaman 1960)
Distribution in Croatia: Widely distributed in the Mediterranean part of the country. Not recorded, but most likely
present in Kvarner.
Distribution in Europe: Nominal subspecies only. Mediterranean species, distributed from the Iberian Peninsula to
Southern Britain, Northwestern Germany (northern border of the distribution), and Southwestern Ukraine and
Rhodes (eastern border). The distribution not yet fully known as the species has often been confused with other
Tetrix species (Hochkirch et al. 2016).

7. Tetrix depressa Brisout de Barneville, 1848 = udubljeni monaški skakavac


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Common along the Adriatic coast with adjacent islands, found also in the Dinaric Alps. Not
recorded, but most likely present in Istria.
Distribution in Europe: Mediterranean species, present on the Iberian Peninsula, Central and Southern France,
Sardinia, Sicily and Italy, the West and South Balkans, Crete, Rhodes and Cyprus, as well as in the Southern
Crimea (Hochkirch et al. 2016).

8. Tetrix subulata (Linnaeus, 1758) = vitki monaški skakavac (Fig. 7B)


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Common in all regions.
Distribution in Europe: From Southern Spain, Sicily and Northern Greece (S border) to Northern Ireland and the
North of Sweden (northern border) and Eastern European Russia (eastern border) (Hochkirch et al. 2016).

9. Tetrix tenuicornis (Sahlberg, 1891) = dugoticalni monaški skakavac


First faunistic record for Croatia: Szövényi & Puskás 2012
Also reported as: T. bipunctata (Linnaeus, 1758) [partim] (e.g. Graber 1870)
Distribution in Croatia: Common in the Pannonian region, found also in the Dinaric Alps.
Distribution in Europe: From Northern Spain, central Italy and Northern Greece (southern border) to C Finland
(northern border) and Eastern European Russia (eastern border) (Hochkirch et al. 2016).

10. Tetrix transsylvanica (Bazyluk & Kis, 1960)


10.1. Tetrix transsylvanica hypsocorypha Skejo, 2014 = transilvanijski monaški skakavac
First faunistic record for Croatia: Skejo et al. 2014
Distribution in Croatia: Known only from the type locality and old type series (Hrvatsko Zagoje: Gornja Pačetina).
It is either extremely localized or extinct in the country.
Distribution in Europe: The species T. transsylvanica consists of two subspecies: nominal from Southern-
Carpathians of Romania and T. t. hypsocorypha from Hrvatsko Zagorje in Croatia and Boč Mt. in Slovenia (Skejo
et al. 2014, Chobanov et al. 2016). In Croatia only T. transsylvanica hypsocorypha recorded.
IUCN Red List status: Endangered (Europe and EU28) (Hochkirch et al. 2016).

11. Tetrix tuerki (Krauss, 1876) = Türkov monaški skakavac


First faunistic record for Croatia: Skejo et al. 2014
Distribution in Croatia: Known from gravel alluvia of Drava River (Ormoško jezero, Koren 4.IX.2014.). Extremely
localized.
Distribution in Europe: In the mountain systems of the Balkans, Alps and Carpathians (Hochkirch et al. 2016).
IUCN Red List status: Vulnerable (Europe and EU28) (Hochkirch et al. 2016).

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12. Tetrix undulata (Sowerby, 1806) = zapadni / srednjeticalni monaški skakavac
First faunistic record for Croatia: Skejo et al. 2014
Distribution in Croatia: Nominal subspecies only. Reported for Croatia only recently from Pannonian and Dinaric
region. Rare and localized, Croatia being the border of its distribution (Skejo et al. 2014).
Distribution in Europe: From the Iberian Peninsula (southwestern distribution border) to northern Sweden
(northern border) and Northwest European Russia (eastern border) (Hochkirch et al. 2016).

superfamily group Acridomorpha MacLeay, 1821

superfamily Acridoidea MacLeay, 1821

family Acrididae MacLeay, 1821 = pravi skakavci

subfamily Acridinae MacLeay, 1821

genus Acrida Linnaeus, 1758

13. Acrida ungarica (Herbst, 1786)


13.1. Acrida ungarica ungarica (Herbst, 1786) = nosati skakavac (Fig. 7C)
First faunistic record for Croatia: Germar 1817
Also reported as: Truxalis nasuta (Linnaeus, 1758) (Kollar 1846), Truxalis hungaricus Fabricius [correct author of
Truxalis ungaricus is Herbst, 1786] (Germar 1817), Truxalis turrita (Linnaeus, 1758) (Krauss 1879), A. m.
mediterranea (Dirsh 1949) syn. nov. (Dirsh 1949, Adamović 1964), A. meridionalis (Dirsh 1949) (Us 1967) [there
is no species named A. meridionalis and Us (1967) probably confused it for ‘mediterranea’], A. bicolor (Thunberg,
1815) (Cigliano et al. 2018), A. mediterranea lombardica Dirsh, 1949 syn. rev. (Dirsh 1949)
Distribution in Croatia: Nominal subspecies only. According to old division (see Cigliano et al. 2018), A. u.
ungarica is found in sandy areas of Pannonian Croatia, while A. u. mediterranea is found along the Adriatic coast
and on the adjacent islands.
Distribution in Europe: Mediterranean species distributed in the Iberian, Apennine and Balkan Peninsulas, as well
as on Mediterranean islands to Cyprus and reaching European Russia (eastern distribution border). Austria and
Slovakia represent its northern distribution border, while in Czechia the species has been probably introduced
(Hochkirch et al. 2016).
Taxonomic note: We regard A. u. mediterranea synonymous with the nominal subspecies, because no significant
morphological differences were found after comparison of Pannonian (Croatian, Hungarian and Serbian) and
Mediterranean (Italian, Croatian, Montenegrian, Albanian) specimens (unpublished data).

genus Paracinema Fischer, 1853

14. Paracinema tricolor (Thunberg, 1815)


14.1. Paracinema tricolor bisignata (Charpentier, 1825) = zeleni močvarni skakavac
First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Rare, found in a few marsh localities throughout the Mediterranean part of the country.
Distribution in Europe: Europe is inhabited by northern subspecies, P. tricolor bisignata (Charpentier, 1825).
Iberian Peninsula, Southern France (and Corsica), Italy (with Sicily and Sardinia), the Balkans (from Greece to
Slovenia) and Ukraine (Hochkirch et al. 2016).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

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subfamily Calliptaminae Tinkham, 1940 = krupnozadi skakavci

genus Calliptamus Serville, 1831

15. Calliptamus barbarus (Costa, 1836) = stepski krupnozadi skakavac


First faunistic record for Croatia: Rebrina et al. 2015
Also reported as: C. italicus (Linnaeus, 1758) [possibly partim, old records need to be revised]
Distribution in Croatia: Nominal subspecies only. Rare, found in Palagruža Isl. and a few localities in the Dinaric
Alps (Dinara Mt., Dalmatinska zagora around Drniš: Sedramić gornji, Skejo 16.VIII.2016). Further research
required.
Distribution in Europe: From the Canary Islands (Spain) to Cyprus and from France to Southern Russia (Hochkirch
et al. 2016).

16. Calliptamus italicus (Linnaeus, 1758) = talijanski krupnozadi skakavac / skakavac Talijan
First faunistic record for Croatia: Germar 1817
Also reported as: Gryllus germanicus Fabricius, 1775 (Charpentier 1825)
Distribution in Croatia: Nominal subspecies only. Widespread in the entire country.
Distribution in Europe: From the Iberian Peninsula to Greece (including Crete and Rhodes) and from the French
Atlantic coast via Northern Poland to Southern European Russia (Hochkirch et al. 2016).

genus Paracaloptenus Bolívar, 1878

17. Paracaloptenus cristatus Willemse, 1973 = kratkokrili krupnozadi skakavac


First faunistic record for Croatia: Krauss 1879
Also reported as: Caloptenus caloptenoides Brunner von Wattenwyl, 1861 (Krauss 1879), C. brunneri (Stål, 1876)
(Brunner von Wattenwyl 1882)
Distribution in Croatia: Locally abundant, found only in the Dinaric Alps (Učka, Velebit, Mala Kapela, Poštak,
Dinara, Kamešnica). Old records of Paracaloptenus caloptenoides (Brunner von Wattenwyl, 1861) belong to this
species.
Distribution in Europe: Dinaric Alps endemic: along the Adriatic coast and the neighboring Dinaric mountains
from Southwestern Slovenia through Croatia and Western Bosnia and Herzegovina to Northern Montenegro
(Chobanov et al. 2016).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

subfamily Catantopinae Brunner von Wattenwyl, 1893

genus Pezotettix Burmeister, 184

18. Pezotettix giornae (Rossi, 1794) = mali smeđi skakavac (Fig. 7D)
First faunistic record for Croatia: Fischer 1853
Distribution in Croatia: Widespread and very common in the entire country.
Distribution From Southern Iberian Peninsula, Sicily and Crete to Central France, Southern Switzerland, Eastern
Austria and Southern Slovakia (Hochkirch et al. 2016).

subfamily Cyrtacanthacridinae Kirby, 1910

tribe Cyrtacanthacridini Kirby, 1910

genus Anacridium Uvarov, 1923

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19. Anacridium aegyptium (Linnaeus, 1764) = egipatska šaška
First faunistic record for Croatia: Germar 1817
Also reported as: Gryllus lineola Fabricius, 1781 (Germar 1817), Acridium tataricum (Linnaeus, 1758) (Kollar
1846), A. tartaricum [misspelled name] (Pungur 1899),
Distribution in Croatia: Widely distributed and rather common in the Mediterranean part of the country, in the
Dinaric Alps it is found in localities with the Mediterranean influence. There is a doubtful record from Srijem
(=Szerém) (Pungur 1899). Occasionally this species is known to produce swarms in Istria and Dalmatia, and is well
known among local people along the Adriatic coast.
Distribution in Europe: Mediterranean species widely distributed in the Iberian peninsula, Apennine peninsula,
Mediterranean (coastal) part of the Balkans, Mediterranean islands, and Crimea (Hochkirch et al. 2016).

genus Schistocerca Stål, 1873

20. Schistocerca gregaria (Forskål, 1775) = pustinjska šaška


First faunistic record for Croatia: Padewieth 1900
Also reported as: S. peregrina (Olivier, 1804) (Padewieth 1900)
Distribution in Croatia: Doubtful, possibly vagrant. Recorded in the Mediterranean part of the country.
Distribution in Europe: Vagrant species. Swarms have been reported from the southern part of the continent (the
Canary Islands and the Iberian Peninsula), sometimes reaching Southwestern England and Southern Ireland. Single
individuals have also been reported from other countries (including Croatia), but there is no evidence that the
species reproduces in Europe (Hochkirch et al. 2016).
Distributional note: Padewieth (1900) is the only author who reported this species from Croatia (Senj: Nehaj). He
reported the species together with A. aegyptium, meaning he was likely able to distinguish the two. Its presence
was confirmed after the examination of Langhoffer's old material from Hungarian Natural History Museum in
Budapest (Puskás et al. 2018). There are no recent records of the species.
IUCN Red List status: Not Applicable for Europe and EU28 (Hochkirch et al. 2016).

subfamily Gomphocerinae Fieber, 1853

genus Arcyptera Serville, 1838 = žarki skakavci

21. Acryptera brevipennis (Brunner von Wattenwyl, 1861)


21.1. Acryptera brevipennis brevipennis (Brunner von Wattenwyl, 1861) = dinarski žarki skakavac
First faunistic record for Croatia: Brunner von Wattenwyl 1861
Also reported as: Stethophyma variegatum (Panz.) [partim] (Frauenfeld 1861) [correct author of Gryllus (Locusta)
variegatum is Sulzer, 1776], S. variegatum variety brevipennis Brunner von Wattenwyl, 1861 (Brunner von
Wattenwyl 1861)
Distribution in Croatia: Nominal subspecies only. Found in the Mediterranean part of the country (not recorded in
Dubrovnik area) and in the Dinaric Alps, as well as on Cres Island (Mt. Sis, Szövényi 06.VII.2017). Widespread,
but not common.
Distribution in Europe: The species A. brevipennis has disjunct distribution in Southwestern and Southeastern
Europe (two different subspecies). Nominal subspecies (Arcyptera b. brevipennis) inhabits the Balkans from
Croatia (extinct in Slovenia) (northern distribution border) to Montenegro (southern border); A. brevipennis vicheti
Harz, 1975 is found in Northern Spain and in the Mediterranean part of France (Hochkirch et al. 2016).
Taxonomic note: Taxonomic status of the two subspecies needs to be studied as these may represent unique
species.
IUCN Red List status: Vulnerable (Europe and EU28) (Hochkirch et al. 2016).

22. Arcyptera fusca (Pallas, 1773) = dugokrili žarki skakavac


First faunistic record for Croatia: Brunner von Wattenwyl 1861

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Also reported as: Stethophyma variegatum (Sulzer, 1776) [partim] (see Brunner von Wattenwyl 1861), S.
variegatum variety genubus posticis pallidis Brunner von Wattenwyl, 1861 (Brunner von Wattenwyl 1861),
Arcyptera stollii (Fieber, 1853) (Karny 1907)
Distribution in Croatia: Restricted to the Dinaric Alps, not common.
Distribution in Europe: Mainly in montane areas, including the Alps, Pyrenees, Carpathians and mountain systems
of the Balkans (Hochkirch et al. 2016).

23. Arcyptera microptera (Fischer von Waldheim, 1833) = istarski žarki skakavac
First faunistic record for Croatia: Bellmann 1993 [Pungur’s (1899) records of Stethophyma flavicosta Fisch. are to
be checked]
Also reported as: Stethophyma flavicosta (Fischer, 1853) (Pungur 1899)
Distribution in Croatia: Recorded only on a few localities in Istria. There are literature records for this species from
Croatia, but all revised museum material from Croatia identified as A. microptera was found to belong to A.
brevipennis. First exact records of the species are photos of a male and a female from Marčana (Bellmann 1993,
2006). Its known distribution in Istria was expanded recently after the research of Roesti and Rutschmann (http://
www.orthoptera.ch/orthoptera-galerie/item/istrien-2014).
Distribution in Europe: Mosaic distribution (without continuity in distribution area), a lot of severely fragmented
subpopulations exist in Southern Europe (including the Iberian peninsula, Southern France, Southern Italy and the
Balkans), Pannonian Basin, and Ukraine (Hochkirch et al. 2016).
IUCN Red List status: [Least Concern (Europe)], Vulnerable (EU28) (Hochkirch et al. 2016).

genus Chorthippus Fieber, 1851 = livadni skakavci

Taxonomy of the genus is unclear, so here we list the species alphabetically, not assigned to subgenera, as proposed
in Cigliano et al. (2017). The genus can be divided into species groups, of which the following groups and species
are present in Croatia—(1) C. albomarginatus group (including only C. oschei pusztaensis), (2) C. apricarius
group (including only C. apricarius), (3) C. biguttulus group (including six species, and two additional
subspecies—C. b. biguttulus, C. bornhalmi, C. brunneus, C. eisentrauti, C. mollis with subspecies C. m. ignifer, C.
m lesinensis, C. m. mollis, and C. v. vagans) and (4) C. dorsatus group (including three species—C. dichrous, C.
dorsatus, C. loratus). The group of Chorthippus parallelus (in Europe includes C. montanus, C. parallelus, C.
smardai, C. tatrae, in Croatia only C. parallelus present) was assigned to a separate genus by Defaut (2012a),
namely Pseudochorthippus, and C. parallelus is here accordingly treated as a species of that genus.

24. Chorthippus apricarius (Linnaeus, 1758) = planinski livadni skakavac (Fig. 7E)
First faunistic record for Croatia: Padewieth 1900
Distribution in Croatia: Rare, recorded only from a few localities in the Dinaric Alps.
Distribution in Europe: Found in most parts of continental Europe, occuring from Northeastern Spain, Central Italy
and Northern Greece (southern border) to Southern Sweden and Latvia (northern border). In the south of its range,
it is restricted to montane regions (Hochkirch et al. 2016).

25. Chorthippus biguttulus (Linnaeus, 1758)


25.1. Chorthippus biguttulus biguttulus (Linnaeus, 1758) = širokokrili livadni skakavac
First faunistic record for Croatia: Graber 1870
Also reported as: Chorthippus variabilis Fieber, 1852 (Frauenfeld 1861), C. biguttulus hedickei (Ramme, 1942)
(see Taxonomic note)
Distribution in Croatia: Nominal subspecies only. Widespread and common in all regions.
Distribution in Europe: Widespread, from the Pyrennees to the Urals and from Greece to Southern Scandinavia. In
Italy occurs only north of the Po Valley (Hochkirch et al. 2016).
Taxonomic note: Populations from the Pannonian lowland and the Balkans were assigned to C. b. hedickei (e.g.
Mikšić 1970, Helversen 1989). However, no morphological or bioacoustic differences that would evidence
separation of more than one group within this species were found among Croatian populations (unpublished data).

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Thus, we consider further investigations necessary to clear out the infraspecific taxonomy of this species.

26. Chorthippus bornhalmi Harz, 1971 = primorski (dugokrili) livadni skakavac


First faunistic record for Croatia: Harz 1971
Also reported as: Stenobothrus bicolor (Charpentier, 1825) [partim] (Bucchich [Bučić] 1886), C. brunneus
(Thunberg, 1815) (all C. brunneus records from the Mediterranean region and the Dinaric Alps prior to Harz’s
(1971) description of C. bornhalmi).
Distribution in Croatia: Widespread and common in the Mediterranean region (Istria, Kvarner, Dalmatia,
Dubrovnik, Adriatic Islands) and in the Dinaric Alps. Not reported from the Pannonian region, where it is replaced
by C. brunneus.
Distribution in Europe: Southeastern Europe along the Adriatic coast from Slovenia to Greece as well as on the
Greek islands, Southern Bulgaria and the European Turkey. At the northern range border it hybridizes with
Chorthippus brunneus (Skejo & Ivković 2015).
Taxonomic note: Likely synonymous with C. maritimus Mistshenko, 1951 (Vedenina pers. comm.).

27. Chorthippus brunneus (Thunberg, 1815) = dugokrili livadni skakavac


First faunistic record for Croatia: Pongrácz 1944 [we consider this, the first Pannonian record of the species, the
first exact record, because records prior to 1944 referred to the coastal area, where C. bornhalmi generally replaces
C. brunneus]
Also reported as: Gryllus bicolor Charpentier, 1825 (Pungur 1899) (see Taxonomic note)
Distribution in Croatia: Widespread and common in the Pannonian region and Northern Dinaric Alps. In the
Mediterranean area locally common, but generally replaced by C. bornhalmi, with which it hybridizes in numerous
localities from Istria to Dinara Mt. (central part of the Dinaric Alps) (Skejo & Ivković 2015). All records from the
coastal area before the description of C. bornhalmi are regarded doubtful (especially from the Southern Adriatic
Islands and Dubrovnik area).
Distribution in Europe: Widespread, from the Northern Iberian Peninsula to the Balkans and from Ireland to
Northern Finland (Hochkirch et al. 2016).
Taxonomic note: Gryllus bicolor Charpentier, 1825 is a junior subjective synonym of C. brunneus. However, the
name has been used in the past for numerous Chorthippus spp. (e.g. C. biguttulus, C. mollis…), thus one should be
careful when referring to historical records under this name.

28. Chorthippus dichrous (Eversmann, 1859) = dvobojni livadni skakavac


First faunistic record for Croatia: this study
Distribution in Croatia: Only in the eastern part of Pannonian Croatia. Specimens from Dalmatia are atypical,
probably hybrids with C. dorsatus, similar to those observed by Chobanov & Mihajlova (2010).
Distribution in Europe: From Eastern Austria, Czechia and Central Italy to Southern Greece and Southern Russia.
Probably extinct in Slovenia (Hochkirch et al. 2016).

29. Chorthippus dorsatus (Zetterstedt, 1821)


29.1. Chorthippus dorsatus dorsatus (Zetterstedt, 1821) = zeleni livadni skakavac
First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Nominal subspecies only. Common and very abundant in the Pannonian region and the
Dinaric Alps. Present in Istria and Dalmatia, but not common in the Mediterranean region. In numerous localities
(e.g. Varaždin, Vinkovci) specimens have intermediate characters between C. dorsatus and C. dichrous—this
should be studied in the future.
Distribution in Europe: From Spain to Eeastern European Russia and from Southern Italy (inhabited by endemic
brachypterous subspecies) to Southern Sweden (Hochkirch et al. 2016).

30. Chorthippus eisentrauti (Ramme, 1931) = alpski livadni skakavac


First faunistic record for Croatia: Ebner 1951
Distribution in Croatia: Reported from NW Dinaric Alps (e.g. Ebner 1951, Nagy 2006). Male calling song from
Učka Mt., Kapela Mt. and Velebit Mt. close to Senj is similar to that of typical C. eisentrauti (Skejo, unpublished

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data). Thorough taxonomic and biogeographic study has not been carried out yet, but is necessary to determine
whether this species is present in Croatia, or these are hybrid populations between C. eisentrauti and C. biguttulus.
Distribution in Europe: Endemic to the southern and eastern parts of the Alps. It ranges from the Haute-Alpes, just
at the border to Italy in the west, over Val d´Aosta, Tessin and Graubünden in Switzerland along the southern edge
of the Italian Alps eastwards to Southern Austria Northern Slovenia. It is very similar to the closely related
Chorthippus biguttulus, so the precise distribution is often unclear and misidentifications (and even hybrids) are
not rare (Zuna-Kratky et al. 2016).

31. Chorthippus loratus (Fischer von Waldheim, 1846) = stepski livadni skakavac
First faunistic record for Croatia: this study
Distribution in Croatia: There is a single confirmed old record of this species from the northernmost part of the
country (Varaždin, Košćec Collection). The species is probably extinct in Croatia (Hochkirch et al. 2016) and
detailed research is necessary to confirm its potential presence in the country. Thorough morphological and
bioacoustic comparison with other species from C. dorsatus aggregate—C. dorsatus and C. dichrous, is neccessary
to assess true taxonomic state.
Distribution in Europe: Balkan Peninsula from Slovenia and Croatia (where it is probably extinct) (northern
distrubution border) to Northern Greece (southern border) and Southern European Russia (eastern border)
(Hochkirch et al. 2016).

32. Chorthippus mollis (Charpentier, 1825) = mali livadni skakavac


First faunistic record in Croatia: Krauss 1888 [description of Stenobothrus lesinensis, currently C. m. lesinensis]
Distribution in Croatia: see below for each subspecies
Distribution in Europe: From the Pyrenees to Eastern European Russia and from Greece and Northern Italy to
Estonia (Hochkirch et al. 2016).

32.1. Chorthippus mollis ignifer Ramme, 1923 = istarski (mali) livadni skakavac
First faunistic record for Croatia: this study [mentioned without precise locality in Fontana et al. (2002)]
Also reported as: C. mollis (Charpentier, 1825) (Us 1967)
Distribution in Croatia: Istria with Učka Mt. (Dinaric Alps). Further research is required to determine complete
distribution.

32.2. Chorthippus mollis lesinensis (Krauss, 1888) = glavati (mali) livadni skakavac (Fig. 7F)
First faunistic record for Croatia: Krauss 1888
Also reported as: C. mollis (Charpentier, 1825) (Adamović 1964), C. lesinensis lastovensis Mařan, 1965, C.
lesinensis svarci Mařan, 1965 (Mařan 1965)
Distribution in Croatia: Dalmatia with Southern Adriatic islands, the Dinaric Alps from Troglav Mt. to south, and
mountains of Dalmatia—from Biokovo Mt. to the southernmost Croatia (Sniježnica), and Dubrovnik region.
Further research is required to determine its complete distribution.

32.3. Chorthippus mollis mollis (Charpentier, 1825) = mali livadni skakavac


First faunistic record for Croatia: Szövenyi & Puskás 2012
Distribution in Croatia: Panonian region, the Dinaric Alps from Risnjak Mt. in the north to Dinara Mt. in the south,
Pag Isl. Further research is required to determine complete distribution.
Taxonomic note: There are populations intermediate in morphology and bioacoustics between typical forms of
these subspecies.

33. Chorthippus oschei Helversen, 1986


33.1. Chorthippus oschei pusztaensis Vedenina & Helversen, 2009 = panonski livadni skakavac
First faunistic record for Croatia: Vedenina & Helversen 2009 [possibly Krauss 1879 as Stenobothrus elegans
Charpentier]
Also reported as: Stenobothrus elegans (Charpentier, 1825) (Krauss 1879), Chorthippus albomarginatus (De Geer,
1773) (Padewieth 1900)

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Distribution in Croatia: Subspecies C. o. pusztaensis Vedenina & Helversen, 2009 is present in the whole country,
except on the Adriatic Islands, in Dalmatia and Dubrovnik area. Common in the Pannonian region and the Dinaric
Alps.
Distribution in Europe: Species endemic to the Balkans and surroundings, from Eastern Austria and Slovakia to
Northern Greece (southern distribution border) and Ukraine (southern border). On northern range border, the
species can hybridize with C. albomarginatus, which occurs from Spain and the United Kingdom to Finland,
Eastern European Russia and Northern Romania. C. oschei is divided into two subspecies—nominal restricted to
Greece, and C. o. pusztaensis (after Puszta, Hungarian word for waste plain steppic grassland), inhabiting other
parts of the species' distribution—in and around the Pannonian Basin (Hochkirch et al. 2016).

34. Chorthippus vagans (Eversmann, 1848) = velikouhi livadni skakavac


34.1. Chorthippus vagans vagans (Eversmann, 1848)
First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Nominal subspecies only. Rare, known from a few records from most of the regions, but
the majority of the records have not been confirmed for more than 50 years. Pannonian region (mostly old
records—Graber 1870 and Pungur 1899; new record Mura-Drava Regional Park: Rebrina and BIUS, Section of
Orthopterology 1.-5.VIII.2016), the Dinaric Alps (Dinara—Rebrina et al. 2015, new data), Istria (old record
only—Redtenbacher 1900), Kvarner (old record only—Krauss 1879), Adriatic Islands (old record only—Us 1964),
and Dubrovnik region (old record only—Frauenfeld 1861). Not recorded from Dalmatia.
Distribution in Europe: Widespread from the Iberian Peninsula, Southern Italy and Cyprus (southern distribution
border) to South England, Denmark and Northern Poland (northern border) and Siberia (eastern border)
(Hochkirch et al. 2016).

genus Chrysochraon Fischer, 1853

35. Chrysochraon dispar (Germar, 1834) = veliki zlatni skakavac (Fig. 8A)
First faunistic record for Croatia: Us 1938 [without subspecies designation]
Distribution in Croatia [of both 35.1 and 35.2]: Present in the Pannonian region, Dinaric Alps and Istria, localized.
Subspecies hitherto known from Croatia: C. d. dispar, C. d. giganteus Harz, 1975. The latter is known from a single
record in Istria (Ingrisch 1981), but in other regions of the country the authors did not specify the subspecies.
Hence, it is necessary to test the presence of C. d. giganteus in the rest of the country. Presence of C. d. dispar is
hitherto confirmed in Žumberak (Nagy 2006) and Papuk Mt. (Szövenyi & Puskás 2012). After the comparison of
the material from Serbia, Croatia, Italy, Slovenia and Macedonia, we found that the subspecies described from
Bosnia by Mikšić—C. d. intermedius Mikšić, 1978 syn. nov. is synonymous with C. d. giganteus, because it fits
the variability range of the latter. Statuses of the nominal subspecies and of C. d. giganteus are unclear and need to
be resolved.
Distribution in Europe: Widely distributed from the Spanish Pyrenees, Northern Italy and Northeastern Greece
(southern distribution border) to Northern Sweden (northern border) and Eastern European Russia (eastern border)
(Hochkirch et al. 2016).

35.1. Chrysochraon dispar dispar (Germar, 1834)


First exact record in Croatia: Nagy 2006

35.2. Chrysochraon dispar giganteus Harz, 1975


First exact record in Croatia: Ingrisch, 1981

genus Dociostaurus Fieber, 1853

36. Dociostaurus brevicollis (Eversmann, 1848) = kratkovrati X-skakavac


First faunistic record for Croatia: Pongrácz 1944

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Distribution in Croatia: Known only from the easternmost part of the Pannonian Basin in Croatia (Kopački rit).
Pungur’s (1899) record from Srijem is not precise, since it can be either in Croatia or Serbia.
Distribution in Europe: From Austria and Czechia (western distribution border) to Greece and the European part of
Turkey (southern border) and Eastern European Russia (Eastern border) (Hochkirch et al. 2016).

FIGURE 8. Diversity of Croatian Caelifera—Acrididae: Gomphocerinae, Melanoplinae. A. Chrysochraon dispar (Plešivica


25.VI.2018. I. Grgić), B. Omocestus rufipes (Biokovo 27.VII.2016. J. Skejo), C. Rammeihippus dinaricus (Vrdovo, Troglav
30.V.2014. J. Skejo), D. Stenobothrus croaticus (Velebit 25.VII.2014. J. Skejo), and E. Miramella irena (Buzim 9.VII.2006. N.
Tvrtković).

37. Dociostaurus genei (Ocskay, 1833) = otočni X-skakavac


First faunistic record for Croatia: Bucchich [Bučić] 1886
Distribution in Croatia: Known only from Hvar Isl. (Bucchich 1886, Novak 1888).
Distribution in Europe: Southern Europe from the Iberian Peninsula, Southern France and Italy to the Balkans
(Hochkirch et al. 2016).

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38. Dociostaurus maroccanus (Thunberg, 1815) = crvenonogi X-skakavac
First faunistic record for Croatia: Novak 1928
Distribution in Croatia: Known from the Mediterranean part (Istria, Kvarner, Dalmatia, Dubrovnik region, Adriatic
Islands), and from some parts of the Dinaric Alps characterized by the Mediterranean climate.
Distribution in Europe: Widespread in the southern part of Europe (Iberian Peninsula, Apennine Peninsula, the
Balkans and surroundings, Crimea) (Hochkirch et al. 2016).

genus Euchorthippus Tarbinsky, 1926

39. Euchorthippus declivus (Brisout de Barneville, 1848) = obični žitni skakavac


First faunistic record for Croatia: Graber 1870
Also reported as: Stenobothrus pulvinatus (Fischer von Waldheim) [partim] (Padewieth 1900)
Distribution in Croatia: Nominal subspecies only. Widespread in the whole country.
Distribution in Europe: Widely distributed in Southern Europe from Spain to the Ukraine. Northern France
represents its northern distribution border. The species is expanding its range northwards (Austria, Czechia)
(Hochkirch et al. 2016).

40. Euchorthippus pulvinatus (Fischer von Waldheim, 1846) = planinski žitni skakavac
First faunistic record for Croatia: Us 1938
Distribution in Croatia: Nominal subspecies only. Present in the Dinaric Alps and Dalmatia, rare and localized.
Distribution in Europe: From Hungary and Slovakia (northwestern distribution border) to Greece, European
Turkey (southern border) and Eastern European Russia (Hochkirch et al. 2016).
IUCN Red List status: [Least Concern (Europe)], Vulnerable (EU28) (Hochkirch et al. 2016).

genus Euthystira Fieber, 1852

41. Euthystira brachyptera Ocskay, 1826 = mali zlatni skakavac


First faunistic record for Croatia: Us 1938
Distribution in Croatia: Widespread in the Pannonian region, Dinaric Alps, Istria and Kvarner. Not recorded in
Dalmatia, Dubrovnik region and on the Adriatic islands.
Distribution in Europe: Widely distributed from Italy and Greece to Estonia and Eastern European Russia
(Hochkirch et al. 2016).

genus Gomphocerippus Roberts, 1941

Taxonomic note: Species of this genus (G. rufus (Linnaeus, 1758) and recently described G. longipennis Li & Ren,
2016) belong to Chorthippus biguttulus group according to morphological and bioacoustic characters (unpublished
data). We do not perform any taxonomic act because Gomphocerippus is a widely accepted name in orthopterists'
community, experts being aware of this problem. The Chorthippus genus group is in need of comprehensive
revision.

42. Gomphocerippus rufus (Linnaeus, 1758) = skakavac kijačoticalac


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Not common, reported from almost all regions (except Dalmatia, Dubrovnik area and
Adriatic islands).
Distribution in Europe: From Southern Italy and Northern Greece (southern distribution border) to Northern
Sweden (northern border) and Eastern European Russia (eastern border) (Hochkirch et al. 2016).

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genus Myrmeleotettix Bolívar, 1914

43. Myrmeleotettix maculatus (Thunberg, 1815) = patuljasti skakavac kijačoticalac


First faunistic record for Croatia: Krauss 1879
Also repored as: Gomphocerus biguttatus (Charpentier, 1825) (Krauss 1879)
Distribution in Croatia: Not common, known only from a few localities in Kvarner, Dinaric Alps and Pannonian
region.
Distribution in Europe: Found all across the continent. Absent from most of the Mediterranean islands, the
southernmost part of the Mediterranean and the northernmost region of Fennoscandia (Hochkirch et al. 2016).

genus Omocestus Bolívar, 1878

44. Omocestus haemorrhoidalis (Charpentier, 1825) = brdski travnjački skakavac


First faunistic record for Croatia: Graber 1870
Distribution in Croatia: Commonly found in the Dinaric Alps. Present but rare in the Pannonian region, Adriatic
islands, and Dubrovnik area. Probably also present in other regions, but not yet recorded.
Distribution in Europe: From Italy and Greece to the Swedish islands Öland and Gotland and Northwestern
European Russia (Hochkirch et al. 2016).

45. Omocestus minutus (Brullé, 1832) = vitki travnjački skakavac


First faunistic record for Croatia: Baccetti 1992
Also reported as: O. petraeus (Brisout de Barneville, 1856) [partim] (e.g. Padewieth 1900)
Distribution in Croatia: No records since the first report from the Adriatic islands (Biševo: Baccetti 1992) until
recently, when the species was found in Dalmatia (Biokovo Mt., around Šibenik, Kaštela, and Trogir), Dubrovnik
area (Pelješac) and on other Adriatic islands (Brač, Šolta) (Skejo & Sule 2015).
Distribution in Europe: Pannonian Plain and the Balkans from Hungary, Croatia and Greece, to the southern part of
European Russia and Anatolia (Hochkirch et al. 2016).

46. Omocestus petraeus (Brisout de Barneville, 1856) = glavati travnjački skakavac


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Widespread and common in the Dinaric Alps and the Mediterranean region, also found in
the Pannonian region.
Distribution in Europe: From Italy and Greece (southern distribution border) to Czechia (northern border) and
Eastern European Russia (eastern border) (Hochkirch et al. 2016).

47. Omocestus rufipes (Zetterstedt, 1821) = crvenozadi travnjački skakavac (Fig. 8B)
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: O. ventralis (Zetterstedt, 1821) (Adamović 1964), O. viridulus (Linnaeus, 1758) (Pongrácz 1944)
Distribution in Croatia: Widespread and common in the whole country.
Distribution in Europe: From the Iberian Peninsula, Sicily and the Greek mainland to Southern England, Southern
Sweden and the Baltic states (northern distribution border), to Eastern European Russia (eastern border)
(Hochkirch et al. 2016).

genus Pseudochorthippus Defaut, 2012

48. Pseudochorthippus parallelus (Zetterstedt, 1821) = obični livadni skakavac


First faunistic record for Croatia: Graber 1870
Also reported as: Stenobothrus pratorum (Fieber, 1852) (Graber 1870), Chorthippus longicornis Latreille [correct
author of Stenobothrus longicornis is Jacobson, 1905] (Adamović 1964), P. montanus (Charpentier, 1825) (Rebrina
et al. 2015)

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Distribution in Croatia: Population not yet assigned to subspecies. Common and widespread in the whole country.
Some individuals from the populations in the Dinaric Alps resemble P. montanus in morphology, but the latter is
not present in Croatia according to the current knowledge. Perceived similarity is a consequence of high
morphological variability of P. parallelus in the West Balkans (Rebrina et al. 2015).
Distribution in Europe: Widespread, from the southern part of the Iberian Peninsula and the Peloponnesus
(southern distribution border) to Northern Finland (northern border) and from the Portuguese Atlantic coast
(western border) to Eastern European Russia (Eastern border) (Hochkirch et al. 2016).

genus Rammeihippus Woznessenskij, 1996

49. Rammeihippus dinaricus Götz, 1970 = mali dinarski skakavac (Fig. 8C)
First faunistic record for Croatia: Skejo & Rebrina 2013
Distribution in Croatia: Known from three localities in the Dinaric Alps—Paklenica NP on S Velebit Mt., Dinara
Mt. and Troglav Mt.
Distribution in Europe: Endemic to the Croatian and Bosnian and Herzegovinian Dinaric Alps, where it occurs on
Southern Velebit Mt., Dinara Mt., Troglav Mt., Kamešnica Mt. and Čvrsnica Mt. (Hochkirch et al. 2016), and on
Vrant Mt. (Puskás and Szövényi unpublished).
IUCN Red List status: Near Threatened (Europe and EU28) (Hochkirch et al. 2016).

genus Stauroderus Bolívar, 1897

50. Stauroderus scalaris (Fischer von Waldheim, 1846) = rokćući skakavac


First faunistic record for Croatia: Padewieth 1900
Also reported as: Stenobothrus morio Fabricius [missaplied name, nec Acheta morio Fabricius, 1871] (Padewieth
1900)
Distribution in Croatia: Common in the Dinaric Alps, no exact records or confirmed localities in other regions.
Distribution in Europe: In the Alps, the Carpathians, mountain ranges of the Iberian Peninsula, Italy and the
Balkans to Greece. At northern latitudes also found at lower elevations (Hochkirch et al. 2016).

genus Stenobothrus Fischer, 1853

51. Stenobothrus crassipes (Ocskay in Charpentier, 1825) = kratkokrili panonski skakavac


First faunistic record for Croatia: Šerić Jelaska & Skejo 2014
Distribution in Croatia: Known from a single historical record in N Croatia, Pannonian region (Bjelovar, Šerić
Jelaska & Skejo 2014, after Pokopac' material from the 1940s (Pokopac 1956)).
Distribution in Europe: Central and Southeast Europe—Germany, Austria, Czechia, Slovakia, Southwestern
Ukraine, Hungary, Eastern Slovenia and Croatia, Bosnia and Herzegovina, Serbia, Western Romania and Western
Bulgaria (Chobanov et al. 2016).

52. Stenobothrus croaticus (Ramme, 1933) = hrvatski tamnokrili skakavac (Fig. 8D)
First faunistic record for Croatia: Ramme 1933
Also reported as: S. nigromaculatus variety istrianus (Krauss, 1879) [partim] (Padewieth 1900)
Distribution in Croatia: Stenoendemic, known only from the southern slopes of Velebit Mt. The species' presence is
confirmed at a single locality (above Jurjevo on Velebit Mt., close to Gornji Lopci), while the other locality (on a
hill above Senj, not precisely reported, cited by Ramme in the original description from 1933) was never identified.
Distribution in Europe: Endemic to Croatia. Known from two localities in the northern part of Velebit Mountain
(Chobanov et al. 2016).
IUCN Red List status: Critically Endangered (Europe and EU28) (Hochkirch et al. 2016).

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53. Stenobothrus fischeri (Eversmann, 1848) = bjeloglavi tamnokrili skakavac
First faunistic record for Croatia: Krauss 1879
Also reported as: S. nigrogeniculatus (Krauss, 1879) (Krauss 1879), S. geniculatus Eversmann [nec Oedipoda
geniculata Brullé, 1832 nor Oedipoda geniculata Fischer von Waldheim, 1846; Eversmann did not describe any
taxon with geniculatus as specific epitheton] (Pungur 1899)
Distribution in Croatia: Not common, present in the Mediterranean region (Istria, Kvarner, Dalmatia, Dubrovnik
area, Adriatic Islands) and the Dinaric Alps.
Distribution in Europe: From Southern Italy and Greece (Peloponnesus) to Southern France and Eastern Austria
(northern distribution border) and Eastern European Russia (eastern border) (Hochkirch et al. 2016).

54. Stenobothrus lineatus (Panzer, 1796) = bjeloprugi tamnokrili skakavac


First faunistic record for Croatia: Frauenfeld 1861
Distribution in Croatia: Widespread and very common in the whole country.
Distribution in Europe: From Central Spain, Sicily and Northern Greece (southern distribution border) to Southern
England, the Netherlands and Latvia (northern border) and Eastern European Russia (eastern border) (Hochkirch et
al. 2016).

55. Stenobothrus nigromaculatus (Herrich-Schäffer, 1840) = pjegavi tamnokrili skakavac


First faunistic record for Croatia: Krauss 1879
Also reported as: Stenobothrus nigromaculatus istrianus Krauss, 1879 [partim] (Krauss 1879)
Distribution in Croatia: Common in Istria and the Dinaric Alps, present on some Adriatic islands (Hvar, Brač).
Records from Srijem (Graber 1870) are not confirmed and there are no new data on occurrence of this species in
the Pannonian region.
Distribution in Europe: From Southern Italy and Northern Greece (southern distribution border) to Central
Germany (northern border) and Eastern European Russia (eastern border) (Hochkirch et al. 2016).

56. Stenobothrus rubicundulus Kruseman & Jeekel, 1967 = tamnokrili skakavac čegrtaš
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: S. miniatus (Charpentier, 1825) (Frauenfeld 1861), S. rubicundus [missaplied name, nec Gryllus
(Tettigonia) rubicundus Goeze, 1778] (Germar 1817)
Distribution in Croatia: Common and widespread in the Dinaric Alps, found also in Kvarner. Records from
Dalmatia (Redtenbacher 1900) and Zadar (Frauenfeld 1861), respectively, refer to Dalmatian mountains. There are
no new records from Dalmatia.
Distribution in Europe: Endemic to Europe—found in the Balkans from the Peloponnesus to Austria and the
Southwestern Carpathians, in the northern half of Italy and the Alps to Southeastern France (Hochkirch et al.
2016).

57. Stenobothrus stigmaticus (Rambur, 1838) = mali tamnokrili skakavac


57.1. Stebobothrus stigmaticus faberi Harz, 1975
First faunistic record for Croatia: Padewieth 1900
Distribution in Croatia: Rare, present in the Dinaric Alps (literature and new records) and in Northern Croatia,
Pannonian region (grasslands near Bjelovar, Šerić Jelaska & Skejo 2014 after Pokopac 1956, Papuk (Szövényi &
Puskás 2012)).
Distribution in Europe: From the Iberian Peninsula (nominal subspecies) and Northern Greece to Latvia and
Eastern European Russia (S. stigmaticus faberi Harz, 1975), nearly extinct in Italy (Hochkirch et al. 2016).

subfamily Melanoplinae Scudder, 1897

genus Miramella Dovnar-Zapolskij, 1932

58. Miramella irena (Fruhstorfer, 1921) = lijepa irena (Fig. 8E)

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First faunistic record for Croatia: Padewieth 1900 [as M. alpina; first reported as M. irena in Galvagni 1986, 1987]
Also reported as: Pezotettix alpina Kollar, 1833 (Padewieth 1900)
Distribution in Croatia: Not common, present in the Pannonian region and the northern slopes of the Dinaric Alps
(north of Poštak Mt.). Recorded also on Rab Island (Puskás et al. 2018), but its presence there should be confirmed.
Distribution in Europe: From the southern edge of the Eastern Alps in Italy, Austria and Slovenia (Western
distribution border) to the Slavonian Mountains of Croatia, and through Bosnia and Herzegovina to Kosovo
(southern border), and to Southern Romania (eastern border) (Zuna-Kratky et al. 2016).

genus Micropodisma Dovnar-Zapolskij, 1932

59. Micropodisma salamandra (Fischer, 1853) = beskrilni skakavac listoplaz


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Not common, but locally abundant, present mostly in mountainous areas—the Dinaric Alps
(Učka Mt., Žumberak—Samoborsko gorje Mts., Velika Kapela Mt., Velebit Mt.) and Pannonian mountains (Papuk
Mt.).
Distribution in Europe: Fragmented, present in Northeastern Italy, Southern Austria, Slovenia, Croatia, Serbia,
Bosnia and Herzegovina and Northern Albania (Buzzetti et al. 2016).

genus Odontopodisma Dovnar-Zapolskij, 1932

60. Odontopodisma decipiens Ramme, 1951


60.1. Odontopodisma decipiens decipiens Ramme, 1951 = crvenokrili skakavac listoplaz (Fig. 9A)
First faunistic record for Croatia: Ramme 1951 [described from Croatia: Pannonian region: Krapina]
Also reported as: Pezotettix mendax Fischer, 1853 [partim] (Padewieth 1900)
Distribution in Croatia: Nominal subspecies only. From the Pannonian region via the Dinaric Alps and Kvarner to
northern Dalmatia.
Distribution in Europe: From Southern Switzerland, Northern Italy, Austria and Slovakia (northern distribution
border) via Balkans to Greece, European Turkey (southern border) and Ukraine (eastern border) (Hochkirch et al.
2016).

61. Odontopodisma fallax Ramme, 1951 = istarski (crvenokrili) listoplaz


First faunistic record for Croatia: Ramme 1951 [described from Croatia: the Dinaric Alps: Učka Mt.]
Also reported as: Pezotettix mendax Fischer, 1853 [partim] (Padewieth 1900), O. schmidti (Fieber, 1853) [partim]
(see O. schmidti), O. rammei Harz, 1973 syn. nov. (Harz 1973)
Distribution in Croatia: Present in Istria, Kvarner, Northern Adriatic islands (Krk), and the Dinaric Alps north of
Dinara Mt. (probably including the latter).
Distribution in Europe: Endemic to Western Dinaric Alps from Dinara to Učka and Ćicarija in Croatia (including
Krk Island), and karstic regions with Mediterranean influence in Slovenia and Northeastern taly (Chobanov et al.
2016).
IUCN Red List status: Near Threatened (Europe and EU28). Its new synonym, O. rammei syn. nov., was assessed
in IUCN as Data Deficient (Hochkirch et al. 2016).
Taxonomic note: O. rammei Harz, 1973 syn. nov. was described from Rijeka or hills above Rijeka (Kvarner or the
Dinaric Alps) by Harz (1973). Since then, the species has not been recorded and no studies aiming to assess its
identity and/or distribution have been conducted. According to our morphological studies on the populations of O.
fallax/O. rammei syn. nov. from seven different localities in Croatia and after examination of the type material of
both species, we found: 1) no differences in external morphology exist between the species, while the shape of
genitalia varies equally in all examined populations, differences reported by Harz (1973) being of no diagnostic
value; 2) no differences exist in morphometric traits between the species, especially in vertex width/eye width
ratio, used by Harz (1973) as a diagnostic trait—measurements taken from altogether 48 specimens (including
holotypes of both species, two paratypes of O. rammei syn. nov. and an allotype of O. fallax) indicated only sexual

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dimorphism-related size differences in the sample (females being on average larger); 3) preliminary molecular
analysis on freshly collected specimens resulted in a single monophyletic cluster without separation of subordinate
evolutionary units (Skejo, Rebrina, Tvrtković—fieldwork and morphological study, L. Lovrenčić—molecular
study, unpublished data). No geographical, morphological or molecular evidence imply separation of O. rammei
syn. nov. from O. fallax, hence we regard it a junior synonym of O. fallax. Within-species variability of genitalia
shape, along with the small size of existing series, were likely responsible for its separation from the latter species.

FIGURE 9. Diversity of Croatian Caelifera—Acrididae: Melanoplinae, Oedipodinae, Pamphaginae. A. Odontopodisma


decipiens (Kruščica 4.VIII.2014. F. Rebrina), B. Podisma pedestris (Dinara N. Tvrtković), C. Pseudopodisma fieberi (Lepenice
30.VIII.2014. F. Rebrina), D. Acrotylus patruelis (Čiovo 17.VI.2017. F. Rebrina), E. Psophus stridulus (Učka 24.VII.2014. J.
Skejo), and Prionotropis hystrix (Lisac 29.VII.2015. F. Rebrina).

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62. Odontopodisma schmidtii (Fieber, 1853) = crnozadi (crvenokrili) listoplaz
First faunistic record for Croatia: Ramme 1951 [we regard this to be the first exact record because older records
could also belong to O. decipiens or O. fallax, both later described from Croatia]
Also reported as: Pezotettix mendax Fischer, 1853 [partim] (Padewieth 1900)
Distribution in Croatia: Present in Istria, Kvarner and the Dinaric Alps north of Velebit Mt., common in the Pannonian
region.
Distribution in Europe: Northeastern Italy, Southern Austria, Southwestern Hungary, Slovenia, Pannonian and
partly Dinaric Croatia, Bosnia and Herzegovina and northern Serbia (Chobanov et al. 2016).

63. Odontopodisma sp. [undescribed species]


First faunistic record for Croatia: this study
Also reported as: Pezotettix salamandra Fischer, 1853 [partim] (Padewieth 1900)
Distribution in Croatia: Only two female specimens known from a single locality in the Dinaric Alps—Krivi Put
(Velika Kapela Mt.), collected at the end of the XIX. century by Padewieth (= F. Dobijaš).
Taxonomic note: Two females deposited in the Croatian Natural History Museum in Zagreb have their tegmina cut
off, which was ascertained after careful examination of the specimens under stereomicroscope. Remains of the
basal portions of reddish tegmina are clearly visible. The specimens are labeled ‘Pezotettix salamandra’ and were
later published under that name (Padewieth 1900). We do not have any information about other Padewieth’s ‘P.
salamadra’ material. The species is characterized by bilobate subgenital plate with a deep apical excision, similar
to that of O. montana Kis, 1962, but without black markings.

genus Podisma Berthold, 1827

64. Podisma pedestris (Linnaeus, 1758) = gorski skakavac stjenoplaz (Fig. 9B)
First faunistic record for Croatia: Fischer 1853
Distribution in Croatia: Nominal subspecies only. Not common, found in isolated localities in the Dinaric Alps and
possibly Pannonian mountains (a record by Jurinac 1887a needs to be checked). A specimen labeled 'Dalmatia:
Split' from Hungarian Natural History Museum in Budapest (Puskás et al. 2018) was probably caught in the
surrounding Dinaric Alps.
Distribution in Europe: Disjunct distribution with the southern range extending from the Pyrenean mountains
across the Alps and the lower mountain areas of Central Europe into the Balkans, reaching the Black Sea. The
northern range covers Scandinavia, Finland and Northwestern Russia. Two range-restricted subspecies inhabit the
Italian Alps (P. p. caprai in the Alpi Finestre in Piemonte and P. p. nadigi in the Northeastern Trentino). Apart from
the Alps, occurrences are often isolated and separated by vast uninhabited areas (Zuna-Kratky et al. 2016).

genus Pseudopodisma Mistshenko, 1947

65. Pseudopodisma fieberi (Scudder, 1897) = crvenonogi skakavac listoplaz (Fig. 9C)
First faunistic record for Croatia: Szövényi & Puskás 2012
Distribution in Croatia: Common in mountains and lowlands of the Pannonian region, present also in the Dinaric
Alps NW of Poštak Mt. (including the latter).
Distribution in Europe: Endemic to the Southeastern ridges of the Alps (Italy, Austria, Slovenia) and to the
Northern Balkans (Croatia, Bosnia and Herzegovina, Serbia, Kosovo, Bulgaria, Romania) (Zuna-Kratky et al.
2016).

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subfamily Oedipodinae Walker, 1871

genus Acrotylus Fieber, 1853

66. Acrotylus insubricus (Scopoli, 1786) = debelonogi pješčarski skakavac


First faunistic record for Croatia: Pungur 1899
Distribution in Croatia: Nominal subspecies only. Confirmed in a few localities in the Pannonian region (Đurđevački
peski). Older records from the coastal area (e.g. Baccetti & Capra 1988, Sombke & Schlegel 2007) should be revised
in the future.
Also reported as: A. patruelis (see comments under A. patruelis (Herrich-Schäffer, 1838)), Acrotylus versicolor Burr,
1898 [partim?] (Werner 1920)
Distribution in Europe: in the southern and central part of Europe and on the Canary Islands (Hochkirch et al. 2016).
Identification and distribution note: Novak (1888) and Karny (1907) denied the presence of A. insubricus on the
coast, regarding only A. patruelis to be present in the coastal area. The species is easily misidentified with A.
patruelis if the identification is based only on alae coloration (used in most keys). Black band on the hind wings is
highly variable and there are overlaps in coloration between the two species. On the other hand, arolium
morphology is a reliable diagnostic character—it is slender and 1/3 as long as tarsal claws in A. insubricus, while
robust and 1/2 as long as tarsal claws in A. patruelis. Furthermore, A. patruelis has slender hind femora, while they
are robust in A. insubricus (a character helpful for discrimination, but not used in available keys). We confirmed the
presence of both species in Croatia.

67. Acrotylus longipes (Charpentier, 1845) = žutokrili pješčarski skakavac


67.1. Acrotylus longipes longipes (Charpentier, 1845)
First faunistic record for Croatia: this study (in Skejo & Sule 2015 noted as Acrotylus cf. longipes)
Distribution in Croatia: Nominal subspecies only. Known exclusively from Dubrovnik area (Zaton by Dubrovnik:
František Chladek, photo in www.biolib.cz) and the Adriatic Islands (Šolta Isl.: Skejo & Sule 2015). The species
probably has wider distribution in Southern Croatia.
Distribution in Europe: Mediterranean species, from the Canary Islands, Southern Italy and the Balkans (southern
distribution border), to Hungary (northern border) and Ukraine (eastern border). Presence in Slovakia not
confirmed (Hochkirch et al. 2016).
IUCN Red List status: Near Threatened (Europe), Vulnerable (EU28) (Hochkirch et al. 2016).

68. Acrotylus patruelis (Herrich-Schäffer, 1838)


68.1. Acrotylus patruelis patruelis (Herrich-Schäffer, 1838) = vitki pješčarski skakavac (Fig. 9D)
First faunistic record for Croatia: Frauenfeld 1861
Also reported as: Also reported as: A. insubricus (see comments under A. insubricus (Scopoli, 1786)), Acrotylus
versicolor Burr, 1898 [partim?] (Werner 1920)
Distribution in Croatia: Nominal subspecies only. Widespread in the Mediterranean part of the country—Istria,
Kvarner, Dalmatia, Dubrovnik area and the Adriatic islands (e.g. Mljet, Korčula, Hvar, Brač, Šolta, Zlarin, Prvić).
A record from the Pannonian region (Pungur 1899) should be revised.
Distribution in Europe: Widespread Mediterranean species, found in the peninsulas of Southern Europe (Pyrenean,
Apennine, Balkans) and adjacent islands (Hochkirch et al. 2016).

genus Aiolopus Fieber, 1852

69. Aiolopus strepens (Latreille, 1804) = debelonogi pjegavi skakavac


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Common and widespread in all regions, more common in the Mediterranean part of the
country.
Distribution in Europe: Widespread Mediterranean species, with the northern distribution border in Central France
and Slovenia—Austria—Hungary border (Hochkirch et al. 2016).

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70. Aiolopus thalassinus (Fabricius, 1781) = močvarni pjegavi skakavac
70.1. Aiolopus thalassinus thalassinus (Fabricius, 1781)
First faunistic record for Croatia: Krauss 1879 [Frauenfeld's (1861) records are older, but could represent A.
strepens, since the author did not report this species in the coastal area]
Distribution in Croatia: Nominal subspecies only. Recorded in all regions, but older records need to be revised.
Distribution in Europe: Widespread, from the Canary Islands (southwestern distribution border) to Western
Germany (northern border) and Russia (eastern border) (Hochkirch et al. 2016)

genus Epacromius Uvarov, 1942

71. Epacromius coerulipes (Ivanov, 1888) = tamnonogi pjegavi skakavac


First faunistic record for Croatia: Ingrisch 1981
Distribution in Croatia: Very rare, known from two localities in Istria—Poreč: Plava laguna (Ingrisch 1981), and
Valbandon, 6 km north of Pula (Kleukers new data, pers. comm.). In the past, a few authors cited this species for
Croatia (Istria—Ebner 1951, Us 1967, Kaltenbach 1970), but without precise localities.
Distribution in Europe: From Northeastern Italy and Eastern Austria through Croatia, Hungary, Romania and
Ukraine to Southern European Russia (Hochkirch et al. 2016).
IUCN Red List status: Near Threatened (Europe), Endangered (EU28) (Hochkirch et al. 2016).
Taxonomic and distribution note: E. coerulipes has been synonymized with E. pulverulentus (Fischer von
Waldheim, 1846) by Mistshenko (1968), but it is unclear which name should be used, since the synonymy is not
widely accepted (Cigliano et al. 2018). Taxonomic revision of these two species is required. Furthermore, E.
coerulipes is usually divided into two subspecies in Europe—nominal subspecies and E. c. panonnicus (Karny,
1907). However, differences between them are unclear, thus we do not follow this division.

72. Epacromius tergestinus (Megerle von Mühlfeld in Charpentier, 1825)


72.1. Epacromius tergestinus tergestinus (Megerle von Mühlfeld in Charpentier, 1825) = otočni pjegavi
skakavac
First faunistic record for Croatia: Puskás et al. 2018
Distribution in Croatia: Nominal subspecies only. Very rare, known from a single locality (alkaline seashore
grassland near Osor, Cres Island: Szövenyi 10.VIII.2000, 07.VII.2017 (Puskás et al. 2018)). Historical records of
this species in Istria (Fieber 1853, Fischer 1853, Redtenbacher 1900) are not confirmed.
Distribution in Europe: Rare species with heavily fragmented distribution. Two subspecies inhabit Europe—(1) E.
t. tergestinus along the coastlines of the Altlantic ocean in France, the Adriatic Sea in Italy and Croatia, and the
coast of the Black Sea in Romania (all occurrences isolated and small), (2) E. t. ponticus inhabits several alpine
rivers in the French Alps (Zuna-Kratky et al. 2016).
IUCN Red List status: [Least Concern (Europe)], Endangered (EU28) (Hochkirch et al. 2016).

genus Locusta Linnaeus, 1758

73. Locusta migratoria (Linnaeus, 1758) = skakavac selac / migratorna šaška


First faunistic record for Croatia: Cubich 1875
Also reported as: Locusta cinerascens Fabricius, 1781 (Kollar 1846), Pachytylus danicus (Linnaeus, 1767) (Karny
1908)
Distribution in Croatia: From the Pannonian region there are historical records only (Graber 1870, Pungur 1899).
Generally not common, present in the Dinaric Alps, in the Mediterranean region more common towards the South
(Adriatic islands, Istria, Kvarner, Dalmatia, Dubrovnik area). In the last few years recorded at some localities in
higher numbers.
Distribution in Europe: Found in the Mediterranean, but its swarms reached Northern European countries in the
past and reproduced there. Recently, the species has been frequently found in northern areas, as it often escapes
from captivity (pet food) (Hochkirch et al. 2016).

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Taxonomic note: Two subspecies are currently recognized—nominal subspecies and L. m. migratorioides (Reiche
& Fairmaire, 1849) (Ma & Kang 2013). According to Ma & Kang (2013), it is clear that molecular studies should
be performed in Croatia, since it could be a contact zone between the two subspecies.

genus Mecostethus Fieber, 1852

74. Mecostethus parapleurus (Hagenbach, 1822) = prugasti močvarni skakavac


First faunistic record for Croatia: Pongrácz 1944
Also reported as: Parapleurus alliaceus Germar, 1825 (Pongrácz 1944)
Distribution in Croatia: Not common, found in wet lowland habitats in the Pannonian region and known from a
single locality in the Dinaric Alps (Poštak Mt., Szövenyi et al. 2018).
Distribution in Europe: The very north of Spain, most of France, the South of Germany, in Switzerland and Austria,
Northern Italy and the northern part of the Balkan Peninsula, to Northern Greece in the south, and to Eastern
European Russia in the east (Hochkirch et al. 2016).

genus Oedaleus Fieber, 1853

75. Oedaleus decorus (Germar, 1825) = šareni X-skakavac


First faunistic record for Croatia: Krauss 1879
Also reported as: Pachytylus nigrofasciatus Latreille [correct author of Acrydium nigrofasciatum is De Geer, 1773]
(Frauenfeld 1861), O. flavus (Linnaeus, 1758) (Kuthy 1908)
Distribution in Croatia: Locally common, but records generally not numerous. Present in the Pannonian region and
the Dinaric Alps, but most of the records are from the Mediterranean part of the country—Istria, Kvarner,
Dalmatia, Dubrovnik area and the Adriatic islands.
Distribution in Europe: From the Canary Islands and Madeira to Southern Russia and from Sicily to Central
Germany and Southern Slovakia. In Austria and the Czech Republic, the species is regionally extinct (Hochirch et
al. 2016).

genus Oedipoda Latreille, 1829

76. Oedipoda caerulescens (Linnaeus, 1758) = plavokrili kamenjarski skakavac


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Common and widespread in all regions.
Distribution in Europe: From Sicily, Crete and Cyprus (southern distribution border) to Latvia (northern border)
and from Portugal (western border) to Eastern European Russia (eastern border). Presence in Sweden uncertain
(Hochkirch et al. 2016).

77. Oedipoda meridionalis Ramme, 1913 = crvenokrili kamenjarski skakavac


Nomenclatural note: Hitherto reported either as O. germanica (Latreille, 1804), and O. miniata (Pallas, 1771).
According to Hochkirch et al. (2016), the Balkans lineage represents a distinct species. The linage is not officially
regarded as O. germanica any more (see IUCN assessment and map of O. germanica, Hochkirch et al. 2016).
Available name for this lineage is Oedipoda miniata meridionalis Ramme, 1913, described from Istria and
Dalmatia. Thus, here we use that name for the species, as a binomen.
First faunistic record for Croatia: Krauss 1879
Also reported as: O. miniata (Pallas, 1771) (Bucchich [Bučić] 1886), O. gratiosa Serville, 1838 (Frauenfeld 1861),
O. germanica (Latreille, 1804) (Harz 1975)
Distribution in Croatia: Common in the Mediterranean part of the country (Istria, Kvarner, Dalmatia, Dubrovnik
area, Adriatic islands), found in the Dinaric Alps.
Distribution in Europe: O. germanica species group—(1) Oedipoda germanica (Latreille, 1804) from Northern

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Spain and Northern Italy across France, Switzerland and Austria to Germany (northern distribution border) and
Czechia (eastern border), (2) the Balkans is inhabited by O. meridionalis Ramme, 1913, while (3) in Italy south of
the Po valley there is a third distinct species (Hochkirch et al. 2016).
IUCN Red List status: Not assessed.

genus Psophus Fieber, 1852

78. Psophus stridulus (Linnaeus, 1758) = crvenokrili (skakavac) čegrtaš (Fig. 9E)
First faunistic record for Croatia: Krauss 1879 [possibly Cubich 1875 on Krk Isl.]
Distribution in Croatia: Mountains of the Pannonian region (Papuk Mt., Strahinjščica Mt.), Ćićarija Mt., Kvarner,
the Dinaric Alps to Poštak Mt., and on the Adriatic islands of Krk and Pag.
Distribution in Europe: From Northern Iberian Peninsula, Central Italy and Northern Greece (southern distribution
border) to Central Sweden and Central Finland (northern border) and Northern European Russia (eastern border)
(Hochkirch et al. 2016).

genus Sphingonotus Fieber, 1852

79. Sphingonotus caerulans (Linnaeus, 1767) = plavokrili pješčarski skakavac


First faunistic record for Croatia: Krauss 1879
Distribution in Croatia: Not common, present in the Pannonian region (Bjelovar, Kloštarski and Đurđevački peski),
the Dinaric Alps (Velika Kapela Mt. and Velebit Mt.), Kvarner (Grobnik near Rijeka) and northern Adriatic islands
(Rab, Pag).
Distribution in Europe: From the Southern Iberian Peninsula, Italy and Greece to Southern Finland and Estern
Russia. Recent molecular studies show there are more lineages that could represent distinct species (especially
Iberian population) (Hochkirch et al. 2016).

genus Stethophyma Fischer, 1853

80. Stethophyma grossum (Linnaeus, 1758) = veliki močvarni skakavac


First faunistic record for Croatia: Padewieth 1900
Distribution in Croatia: Not common, present in the Pannonian region, Istria, and the Dinaric Alps (Velebit Mt.,
Lička Plješivica Mt.).
Distribution in Europe: From Northern Iberian Peninsula, and Northern Italy and Bulgaria (southern distribution
border) to Northern Sweden and Finland (northern border), present also in England and Ireland (Hochkirch et al.
2016).

family Pamphagidae Burmeister, 1840 = žaboliki skakavci

subfamily Thrinchinae Stål, 1876

genus Prionotropis Fieber, 1853

81. Prionotropis hystrix (Germar 1817) = krški žaboliki skakavac (Fig. 9F)
First faunistic record for Croatia: Germar 1817 [described from Krk Island]
Also reported as: Pamphagus marmoratus Burmeister, 1838 (Kollar 1840)
Distribution in Croatia: Not common, locally abundant, present along the coast (Istria, Kvarner, Dalmatia and
Dubrovnik area), but more common on the southern slopes of the Dinaric Alps and the Adriatic islands (Cres, Krk,
Pag) in stony areas.

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 67
Distribution in Europe: From the Adriatic coast of Northeastern Italy to South Bosnia and Herzegovina (Hochkirch
et al. 2016).
IUCN Red List status: Vulnerable (Europe and EU28) (Hochkirch et al. 2016).

Omitted Caelifera species

family Tetrigidae

subfamily Tetriginae

1. Tetrix kraussi Saulcy, 1888


Not regarded a valid species, but a synonym of T. bipunctata (SIGTET 2017). Thus, all former records of T. kraussi
(Padewieth 1900, Kuthy 1908, Skejo 2014, Skejo et al. 2014) belong to the latter species.

family Acrididae

subfamily Calliptaminae

2. Paracaloptenus caloptenoides (Brunner von Wattenwyl, 1861)


The species is replaced by its sibling species, P. cristatus, in this part of Europe (Chobanov et al. 2016). All older
records of this species (Krauss 1879, Pungur 1899, Padewieth 1900, Redtenbacher 1900, Karny 1907a, Kuthy
1908, Us 1967) belong to P. cristatus (Sardet et al. 2014). P. cristatus had not yet been described when these
authors reported P. caloptenoides for Croatia. The only mountain hitherto known to be inhabited by both species is
Durmitor Mt. in Montenegro (Ingrisch & Pavićević 2012).

subfamily Gomphocerinae

3. Chorthippus albomarginatus (De Geer, 1773)


This species is replaced by its sibling species, C. oschei (subspecies C. o. pusztaensis), in this part of Europe. All
former records of C. albomarginatus (Krauss 1879, Pungur 1899, Redtenbacher 1900, Padewieth 1900, Us 1938,
Pongrácz 1944, Pokopac 1956, Nagy 2006) belong to C. o. pusztaensis (Vedenina & Helversen 2009, Szövényi &
Puskás 2012). It is thus omitted from the checklist of Croatian Orthoptera.

4. Omocestus viridulus (Linnaeus, 1758)


A record of the species (Pongrácz 1944) from the surroundings of Kopački Rit is questionable, but impossible to
review since the collection in Hungarian Natural History Museum was destroyed in 1956. We did not find the
species as there are no suitable habitats in Croatia (Alpine meadows). Revised material (Croatian Natural History
Museum, Faculty of Science in Zagreb) previously identified as O. viridulus turned out to be O. rufipes (in some
cases green Chorthippus sp. or Stenobothrus sp.). Since no evidence exists confirming its presence in the country,
we omit the species from the checklist.

5. Pseudochorthippus montanus (Charpentier, 1825)


A single published record of this species was a misidentified P. parallelus (Rebrina et al. 2015).

subfamily Melanoplinae

6. Miramella alpina Kollar, 1833


After the examination of specimens collected by Padewieth in the Dinaric Alps near Senj (deposited in Croatian

68 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Natural History Museum) and published as M. alpina (Padewieth 1900, cited in Us 1967), we confirm they belong
to M. irena, described in 1921. It is highly unlikely that M. alpina, a true alpine species, is present in Croatia, the
closest populations inhabiting Westernmost Hungary and Austrian Alps.

subfamily Oedipodinae

7. Celes variabilis (Pallas, 1771)


A record from Hrvatsko Zagorje (Pannonian Croatia) (Kodinec 1938) originating from the list of orthopteran
species presumably found by Franjo Koščec, an entomologist from Varaždin, is possibly a misidentification of
some more common species (either Psophus stridulus or Oedipoda sp.). Ozimec & Ernoić (2008) confirmed that
no documentation existed about the origin of the list, or the existence of voucher specimens. There are no other
records of this species in Croatia, so we omit it from the checklist.

TABLE 1. List of Orthopteran species inhabiting Croatia, sorted and enumerated within each suborder independently
(Ensifera 1–103, Caelifera 1–81), assigned to infraorder, superfamily, family, and subfamily. Within each subfamily,
species are ordered alphabetically. Presence of a species within a geographical region is marked with (+). Abbreviations
for regions: PA—Pannonian region, DI—Dinaric Alps region, Mediterranean subregions IS—Istria, KV—Kvarner,
DA—Dalmatia and DU—Dubrovnik area, AI—Adriatic Islands.
N TAXON PA DI IS KV DA DU AI
SUBORDER ENSIFERA Chopard, 1921
infraorder Tettigoniidea Krauss, 1902
superfamily Tettigonioidea Krauss, 1902
family Tettigoniidae Krauss, 1902
subfamily Bradyporinae Burmeister, 1838
1 Ephippiger discoidalis Fieber, 1853 + + + + + +
2 Ephippiger ephippiger (Fiebig, 1784) + +
3 Ephippiger persicarius Fruhstorfer, 1921 +
subfamily Conocephalinae Burmeister, 1838
4 Conocephalus dorsalis (Latreille, 1804) + + + + +
5 Conocephalus fuscus (Fabricius, 1793) + + + + + +
6 Ruspolia differens (Serville, 1839) +
7 Ruspolia nitidula (Scopoli, 1786) + + + + + + +
subfamily Meconematinae Burmeister, 1838
8 Cyrtaspis scutata (Charpentier, 1825) + + + + +
9 Meconema meridionale Costa, 1860 + + + + + +
10 Meconema thalassinum (De Geer, 1773) + + + + + +
subfamily Phaneropterinae Burmeister, 1838
11 Acrometopa s. macropoda (Burmeister, 1838) + + + + + +
12 Andreiniimon nuptialis (Karny, 1918) +
13 Barbitistes kaltenbachi Harz, 1965 +
14 Barbitistes ocskayi Charpentier in Ockay, 1850 + + + +
15 Barbitistes serricauda (Fabricius, 1794) + +
16 Barbitistes yersini Brunner von Wattenwyl, 1878 + + + + + +
17 Isophya brevicauda Ramme, 1931 +
18 Isophya kraussii Brunner von Wattenwyl, 1878 +
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TABLE 1. (Continued)
N TAXON PA DI IS KV DA DU AI
19 Isophya modestior Brunner von Wattenwyl, 1882 +
20 Leptophyes albovittata (Kollar, 1833) + +
21 Leptophyes boscii Fieber, 1853 + + + +
22 Leptophyes discoidalis (Frivaldszky, 1868) +
23 Leptophyes laticauda (Frivaldszky, 1868) + + + + + +
24 Leptophyes intermedia Ingrisch & Pavićević, 2010 + +
25 Leptophyes punctatissima (Bosc, 1792) + + +
26 Poecilimon a. affinis (Frivaldszky, 1868) +
27 Poecilimon ampliatus Brunner von Wattenwyl, 1878 +
28 Poecilimon elegans (Brunner von Wattenwyl, 1878) + + + +
29 Poecilimon sp. + + +
30 Poecilimon fussii Fieber, 1878 +
31 Poecilimon gracilis (Fieber, 1853) + +
32 Poecilimon ornatus (Schmidt, 1850) + + + + +
33 Poecilimon schmidtii (Fieber, 1853) + +
34 Polysarcus denticauda (Charpentier, 1825) + +
35 Phaneroptera falcata (Poda, 1761) + + + + +
36 Phaneroptera nana Fieber, 1853 + + + + + + +
37 Tylopsis lilifolia (Fabricius , 1793) + + + + +
subfamily Saginae Brunner von Wattenwyl, 1878
38 Saga natoliae Serville, 1839 +
39 Saga pedo (Pallas, 1771) + + + + +
subfamily Tettigoniinae Krauss, 1902
40 Bicolorana bicolor (Philippi, 1830) + + + +
41 Bicolorana kraussi (Padewieth, 1900) + +
42 Decticus albifrons (Fabricius, 1775) + + + + +
43 Decticus verrucivorus (Linnaeus, 1758) + + +
44 Eupholidoptera schmidti (Fieber 1861) + + + + + + +
45 Gampsocleis abbreviata Herman, 1874 + +
46 Metrioptera brachyptera (Linnaeus, 1761) +
47 Metrioptera hoermanni (Werner, 1906) +
48 Modestana modesta (Fieber, 1853) + + + + + +
49 Montana stricta (Zeller, 1849) + + + +
50 Pachytrachis frater (Brunner von Wattenwyl, 1882) + + +
51 Pachytrachis gracilis (Brunner von Wattenwyl, 1861) + + + + +
52 Pachytrachis striolatus (Fieber, 1853) + + + + + +
53 Pholidoptera aptera (Fabricius, 1793) + +
54 Pholidoptera dalmatica (Krauss, 1879) + + + + + +
55 Pholidoptera fallax (Fischer, 1853) + + + +
56 Pholidoptera femorata (Fieber, 1853) + + + + +
57 Pholidoptera frivaldszkyi (Herman, 1871) +
58 Pholidoptera griseoaptera (De Geer, 1773) + + + +
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TABLE 1. (Continued)
N TAXON PA DI IS KV DA DU AI
59 Pholidoptera littoralis (Fieber, 1853) + + +
60 Platycleis affinis Fieber, 1853 + + + + + +
61 Platycleis a. grisea (Fabricius, 1781) + + + + +
62 Platycleis intermedia (Serville, 1838) + + + + + +
63 Platycleis romana Ramme, 1927 + +
64 Psorodonotus illyricus Ebner, 1923 +
65 Rhacocleis buchichii Herman, 1874 + +
66 Rhacocleis germanica (Herrich-Schäffer, 1840) + + + + + + +
67 Roeseliana roeselii (Hagenbach, 1822) + + + +
68 Sepiana sepium (Yersin, 1854) + + + + + +
69 Tessellana orina (Burr, 1899) + +
70 Tessellana tessellata (Charpentier, 1825) + + + +
71 Tettigonia balcanica Chobanov & Lemonnier-Darcemont, 2014 + +
72 Tettigonia cantans (Fuessly, 1775) + +
73 Tettigonia caudata (Charpentier, 1842) + + + + +
74 Tettigonia viridissima (Linnaeus, 1758) + + + + + + +
75 Yersinella raymondi (Yersin, 1860) + + + + + +
76 Zeuneriana amplipennis (Brunner von Wattenwyl, 1882) +
superfamily Grylloidea Laicharting, 1781
family Mogoplistidae Brunner von Wattenwyl, 1873
subfamily Mogoplistinae Brunner von Wattenwyl, 1873
77 Arachnocephalus vestitus Costa, 1855 + + + + +
78 Mogoplistes brunneus Serville, 1838 + + + + + +
79 Paramogoplistes novaki (Krauss, 1888) + + +
80 Pseudomogoplistes squamiger (Fischer, 1853) + + + + +
family Trigonidiidae Saussure, 1874
subfamily Nemobiinae Saussure, 1877
81 Pteronemobius heydenii (Fischer, 1853) + + + + + + +
82 Stenonemobius b. ponticus Gorochov, 1984 +
subfamily Trigonidiinae Saussure, 1874
83 Trigonidium cicindeloides Rambur, 1838 + +
family Gryllidae Laicharting, 1781
subfamily Gryllinae Laicharting, 1781
84 Acheta domesticus (Linnaeus, 1758) + + + + + + +
85 Eumodicogryllus bordigalensis (Latreille, 1804) + + + + +
86 Gryllus bimaculatus De Geer, 1773 + + + +
87 Gryllus campestris Linnaeus, 1758 + + + + + + +
88 Melanogryllus desertus (Pallas, 1771) + + + + + + +
89 Modicogryllus frontalis (Fieber, 1844) +
90 Modicogryllus truncatus (Tarbinsky, 1940) +
subfamily Gryllomorphinae Saussure, 1877
91 Gryllomorpha dalmatina (Ocskay, 1832) + + + + + +
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TABLE 1. (Continued)
N TAXON PA DI IS KV DA DU AI
subfamily Oecanthinae Blanchard, 1845
92 Oecanthus dulcisonans Gorochov, 1993 + +
93 Oecanthus pellucens (Scopoli, 1763) + + + + + + +
superfamily Gryllotalpoidea Leach, 1815
family Gryllotalpidae Leach, 1815
subfamily Gryllotalpinae Leach, 1815
94 Gryllotalpa gryllotalpa (Linnaeus, 1758) + +
95 Gryllotalpa sp. + +
96 Gryllotalpa stepposa Zhantiev, 1991 +
family Myrmecophilidae Saussure, 1874
subfamily Myrmecophilinae Saussure, 1874
97 Myrmecophilus hirticaudus Fischer von Waldheim, 1846 +
98 Myrmecophilus myrmecophilus (Savi, 1819) +
superfamily Rhaphidophoroidea Walker, 1869
family Rhaphidophoridae Walker, 1869
subfamily Dolichopodainae Brunner von Wattenwyl, 1888
99 Dolichopoda araneiformis (Burmeister, 1838) + + +
subfamily Troglophilinae Krauss, 1879
100 Troglophilus cavicola (Kollar, 1833) + + + + + + +
101 Troglophilus neglectus Krauss, 1878 + + + + + +
102 Troglophilus ovuliformis Karny, 1907 + + + +
103 Troglophilus sp. +
SUBORDER CAELIFERA Ander, 1936
infraorder Tridactylidea Brullé, 1835
superfamily Tridactyloidea Brullé, 1835
family Tridactylidae Brullé, 1835
subfamily Tridactylinae
1 Xya p. pfaendleri Harz, 1970 +
2 Xya variegata (Latreille, 1809) +
infraorder Acrididea MacLeay, 1821
superfamily Tetrigoidea Rambur, 1838
family Tetrigidae Rambur, 1838
subfamily Tetriginae Rambur, 1838
3 Paratettix meridionalis (Rambur, 1838) + + + + +
4 Tetrix bipunctata (Linnaeus, 1758) + + + +
5 Tetrix bolivari Saulcy, 1901 + +
6 Tetrix ceperoi (Bolívar, 1887) + + + +
7 Tetrix depressa Brisout de Barneville, 1848 + + + + +
8 Tetrix subulata (Linnaeus, 1758) + + + + + + +
9 Tetrix tenuicornis (Sahlberg, 1891) + +
10 Tetrix transsylvanica hypsocorypha Skejo, 2014 +
11 Tetrix tuerki (Krauss, 1876) +
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TABLE 1. (Continued)
N TAXON PA DI IS KV DA DU AI
12 Tetrix undulata (Sowerby, 1806) + +
superfamily group Acridomorpha MacLeay, 1821
superfamily Acridoidea MacLeay, 1821
family Acrididae MacLeay, 1821
subfamily Acridinae
13 Acrida u. ungarica (Herbst, 1786) + + + + + +
14 Paracinema tricolor bisignata (Charpentier, 1825) + + + + +
subfamily Calliptaminae Tinkham, 1940
15 Calliptamus barbarus (Costa, 1836) + +
16 Calliptamus italicus (Linnaeus, 1758) + + + + + + +
17 Paracaloptenus cristatus Willemse, 1973 +
subfamily Catantopinae Brunner von Wattenwyl, 1893
18 Pezotettix giornae (Rossi, 1794) + + + + + + +
subfamily Cyrtacanthacridinae Kirby, 1910
tribe Cyrtacanthacridini Kirby, 1910
19 Anacridium aegyptium (Linnaeus, 1764) + + + + + +
20 Schistocerca gregaria (Forskål, 1775) + + + + +
subfamily Gomphocerinae Fieber, 1853
21 Arcyptera b. brevipennis (Brunner von Wattenwyl, 1861) + + + + + +
22 Arcyptera fusca (Pallas, 1773) +
23 Arcyptera microptera (Fischer von Waldheim, 1833) +
24 Chorthippus apricarius (Linnaeus, 1758) +
25 Chorthippus b. biguttulus (Linnaeus, 1758) + + + + + + +
26 Chorthippus bornhalmi Harz, 1971 + + + + + +
27 Chorthippus brunneus (Thunberg, 1815) + + + + +
28 Chorthippus dichrous (Eversmann, 1859) +
29 Chorthippus d. dorsatus (Zetterstedt, 1821) + + + +
30 Chorthippus eisentrauti (Ramme, 1931) +
31 Chorthippus loratus (Fischer von Waldheim, 1846) +
32.1 Chorthippus mollis ignifer Ramme, 1923 + +
32.1 Chorthippus mollis lesinensis (Krauss, 1888) + + + +
32.3 Chorthippus m. mollis (Charpentier, 1825) + + +
33 Chorthippus oschei pusztaensis Vedenina & Helversen, 2009 + + + +
34 Chorthippus vagans (Eversmann, 1848) + + + + + +
35.1 Chrysochraon d. dispar (Germar, 1834) + +
35.2 Chrysochraon d. giganteus Harz, 1975 +
36 Dociostaurus brevicollis (Eversmann, 1848) +
37 Dociostaurus genei (Ocskay, 1832) +
38 Dociostaurus maroccanus (Thunberg, 1815) + + + + + +
39 Euchorthippus declivus (Brisout de Barneville, 1848) + + + + + + +
40 Euchorthippus pulvinatus (Fischer von Waldheim, 1846) + +
41 Euthystira brachyptera (Ocskay, 1826) + + + +
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TABLE 1. (Continued)
N TAXON PA DI IS KV DA DU AI
42 Gomphocerippus rufus (Linnaeus, 1758) + + + +
43 Myrmeleotettix maculatus (Thunberg, 1815) + + +
44 Omocestus haemorrhoidalis (Charpentier, 1825) + + + +
45 Omocestus minutus (Brullé, 1832) + + +
46 Omocestus petraeus (Brisout de Barneville, 1856) + + + + + + +
47 Omocestus rufipes (Zetterstedt, 1821) + + + + + + +
48 Pseudochorthippus parallelus (Zetterstedt, 1821) + + + + + + +
49 Rammeihippus dinaricus (Götz, 1970) +
50 Stauroderus scalaris (Fischer von Waldheim, 1846) +
51 Stenobothrus crassipes (Charpentier, 1825) +
52 Stenobothrus croaticus Ramme, 1933 + +
53 Stenobothrus fischeri (Eversmann, 1848) + + + + + +
54 Stenobothrus lineatus (Panzer, 1796) + + + + + + +
55 Stenobothrus nigromaculatus (Herrich-Schäffer, 1840) + + +
56 Stenobothrus rubicundulus Kruseman & Jeekel, 1967 + + +
57 Stenobothrus stigmaticus faberi Harz, 1975 + +
subfamily Melanoplinae Scudder, 1897
58 Miramella irena (Fruhstorfer, 1921) + +
59 Micropodisma salamandra (Fischer, 1853) + +
60 Odontopodisma d. decipiens Ramme, 1951 + + + +
61 Odontopodisma fallax Ramme, 1951 + + + +
62 Odontopodisma schmidti (Fieber, 1853) + + + +
63 Odontopodisma sp. [undescribed sp. or ssp.] +
64 Podisma pedestris (Linnaeus, 1758) +
65 Pseudopodisma fieberi (Scudder, 1897) + +
subfamily Oedipodinae Walker, 1871
66 Acrotylus insubricus (Scopoli, 1786) +
67 Acrotylus l. longipes (Charpentier, 1845) + +
68 Acrotylus p. patruelis (Herrich-Schäffer, 1838) + + + + +
69 Aiolopus strepens (Latreille, 1804) + + + + + + +
70 Aiolopus t. thalassinus (Fabricius, 1781) + + + + + + +
71 Epacromius coerulipes (Ivanov, 1888) +
72 Epacromius t. tergestinus (Charpentier, 1825) +
73 Locusta migratoria (Linnaeus, 1758) + + + + + + +
74 Mecostethus parapleurus (Hagenbach, 1822) + +
75 Oedaleus decorus (Germar, 1825) + + + + + + +
76 Oedipoda caerulescens (Linnaeus, 1758) + + + + + + +
77 Oedipoda meridionalis (Ramme, 1913) + + + + + +
78 Psophus stridulus (Linnaeus, 1758) + + + +
79 Sphingonotus caerulans (Linnaeus, 1767) + + + +
80 Stethophyma grossum (Linnaeus, 1758) + + +
subfamily Pamphagidae Burmeister, 1840
subfamily Thrinchinae Stål, 1876
81 Prionotropis hystrix (Germar, 1817) + + + + + +

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8. Oedipoda germanica (Latreille, 1804)
According to Hochkirch et al. (2016), O. germanica is replaced by O. meridionalis in the Balkan Peninsula (see
comments under Oedipoda meridionalis). Thus, all former records of O. germanica from Croatia belong to the
latter species.

9. Oedipoda miniata (Pallas, 1771)


In the past, authors reported all red-winged Oedipoda specimens as O. miniata (Frauenfeld 1861, Krauss 1879,
Bucchich 1886, Novak 1888, Pungur 1899, Padewieth 1900, Burr 1906, Karny 1907, 1908), probably not being
familiar with differences from O. germanica (s.l.). Furthermore, Ramme (1913) assigned O. meridionalis a subspe-
cies status within O. miniata. Identity and distribution of O. miniata are still under discussion (Hochkirch pers.
comm.).

Discussion

By the middle of the 19th century, about 30 species have been reported from the country, together with 11 species
new to science, which are currently valid—Acrometopa servillea macropoda, Barbitistes ocskayi, Dolichopoda
araneiformis, Gryllomorpha dalmatina, and Pholidoptera femorata from Dalmatia (without precise locality),
Prionotropis hystrix from the hills on the island of Krk, Ephippiger discoidalis from Istria, Leptophyes boscii and
Modestana modesta from the surroundings of Rijeka (Učka Mt.), Pholidoptera l. littoralis from Istria and Kranj in
Slovenia (Istria is without precise locality, so it could belong to Slovenian or Croatian part), Pseudomogoplistes
squamiger from Dubrovnik area (Germar 1817, Charpentier 1825, Fischer 1853, Krauss 1879, Nonveiller 1999).
In the second half of the 19th century, about 130 species were known to inhabit Croatia. Also, 11 currently valid
species with a type locality in Croatia were described during this period—Arcyptera b. brevipennis from the
surroundings of Senj and Rijeka (Velebit Mt., Kapela Mt., Učka Mt.), Barbitistes yersini from Obrovac, Bicolorana
kraussi from Senjska Draga, Eupholidoptera schmidti from Istria (without precise locality, could be Slovenian or
Croatian part), Gampsocleis abbreviata from the surroundings of Sinj, Pachytrachis frater from Dubrovnik area,
Pholidoptera dalmatica from the surroundings of Rijeka, Poecilimon elegans from Učka Mt., Rhacocleis
buchichii, Chorthippus mollis lesinensis and Paramogoplistes novaki from the island of Hvar, Troglophilus
neglectus from a cave near Rijeka, probably the entrance of Klana pit (Krauss 1879, 1888, Brunner von Wattenwyl
1861, 1882, Padewieth 1900).
Since two world wars took place in the first half of the 20th century, the research of Orthoptera stagnated. Only
few papers were published during this period, adding not more than ten species to the national fauna, including the
description of Isophya brevicauda from Zagreb and Krapina surroundings, Oedipoda meridionalis from Istria and
Kvarner, Psorodonotus illyricus from Učka Mt. (among other localities in the Balkan peninsula), and Stenobothrus
croaticus from Velebit Mt. or Velika Kapela Mt. above Senj (Ebner 1923, Ramme 1913, 1931, 1933, Nonveiller
1999).
The second half of the 20th century, during Tito's Yugoslavia, was more fruitful in terms of Orthoptera research
than the first half. Five species were described from Croatia during this period—Barbitistes kaltenbachi from Hvar
Island, Chorthippus bornhalmi from Dubrovnik, Odontopodisma decipiens from Krapina, Odontopodisma fallax
and Paracaloptenus cristatus from Učka Mt. (Harz 1965, 1971, 1973, Ramme 1951, Willemse 1973). Us (1967)
gathered all data on orthopterans from Croatia up to that point and pubished it within the catalog of Yugoslavian
Orthopteroidea (Polyneoptera without Plecoptera). For Croatia, Us (1967) reported altogether 161 species, of
which 145 are currently valid.
Since Us' (1967) checklist was the first and the last for Croatia, it was the basic literature for most of foreign
researchers studying grasshoppers and crickets of this area in the 20th and 21st century. Thus, we decided to compile
all data on Orthoptera of Croatia and make a new and annotated checklist. We present a lot of changes in
comparison with Us' (1967) checklist, taxonomic and distributional.
Altogether 184 species (103 Ensifera, 81 Caelifera) are currently known to inhabit Croatia (Fig. 2A, B, C),
while 25 previously reported species are omitted (16 Ensifera, 9 Caelifera), either because of misidentification (e.g.
Pteronemobius lineolatus, Pseudochorthippus parallelus) or the change in the historical usage of names (e.g.
Chorthippus albomarginatus, Oedipoda germanica). In comparison to Us' (1967) checklist, 39 species are added to

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 75
the national list. The following species present in the neighboring countries could be expected in Croatia, since
there are suitable habitats in the country—Roeseliana brunneri Ramme, 1951 and Zeuneriana marmorata (Fieber,
1853) in the northern part of Istria, Isophya camptoxypha (Fieber, 1853), Nemobius sylvestris (Bosc, 1792), and
Myrmecophilus acervorum (Panzer, 1799) in the Pannonian region, Stenonemobius gracilis (Jakovlev, 1871) and
Natula averni (Costa, 1855) in the Adriatic part of the country and Adriatic islands, whereas Poecilimon
pseudornatus Ingrisch et Pavićević, 2010, P. thoracicus (Fieber, 1853), Metrioptera prenjica (Burr, 1899) and
Pachytrachis tumidus Ingrisch et Pavićević, 2010 in the mountains of southern Croatia (Dubrovnik area), and
finally Tropidopola graeca Uvarov, 1926 in the southernmost part of the Dubrovnik area. Thus, the real number of
Orthoptera species inhabiting Croatia might be around 200, if some of the aforementioned species will be found
and/or if future phylogeographic studies reveal old and isolated lineages within currently known species.
Inhabited by 184 species, Croatia is one of the richest countries in Europe concerning Orthoptera diversity.
Central European countries have significantly lower number of species—Czechia 96 (Holuša et al. 2013), Austria
139 (Berg et al. 2005), Germany and Switzerland 119 together (Maas et al. 2002, Baur et al. 2006), Hungary 128
(Szövényi et al. 2016), even if they cover larger area. Countries with similar number of species are SE European
countries, such as Romania 182 (Iorgu et al. 2008), Serbia 179 (Pavićević & Ivković 2015), Macedonia 175
(Lemmonier-Darcemont et al. 2014), Slovenia 157 (Gomboc & Šegula 2014), Albania 187 (Puskás 2016), Bulgaria
218 (Chobanov 2009). Italy and Greece have significantly higher diversity, due to their refugial geographic
position, size and accompanying large and remote mountains, islands and islets (where speciation of numerous
local endemics took place). Italy is inhabited by about 350 species (Massa et al. 2012), and Greece by about 400
(Willemse & Willemse 2008).
This checklist possibly includes complete bibliography on Croatian Orthoptera, as well as all the publications
that are relevant for future Orthoptera studies in Croatia. It is thus practical for every reader and future student of
Croatian orthopterology interested in past research. All the bibliography is listed under References (Literature), and
besides references cited in other parts of this papers, those papers can be found in the list as well, because it
contains data helpful for Croatian Orthoptera research: Novak (1890), Babić & Rössler (1912), Beier (1954a,
1954b), Gorochov (1984a, 1984b), Harz & Kaltenbach (1976), Jacobson & Bianchi (1905), Kolosváry (1938),
Kovačević (1952, 1956), Mařan (1952), Mikšić (1965), Pavićević & Karaman (2001), Puschnig (1896), Serville
(1839), Weidner (1959), Werner (1905), Willemse (1982, 1985), Zacher (1910), and Zeuner (1941).

Crickets and bush-crickets (Ensifera) overview

Overview of Grylloidea families

True crickets (Gryllidae) are among the largest and most widespread orthopteran families, consisting
predominantly of ground-dwelling, omnivorous species. Whereas acoustic communication and behavior of some
species have been studied in detail, their nocturnal habits and tricky identification led to a substantial lack of
research on this group in Europe and, accordingly, in Croatia. This resulted in numerous taxonomic problems and
fragmentary knowledge of the distribution of many species. The entire family is in need of a comprehensive
taxonomic revision at the European level (Hochkirch et al. 2016). Altogether 12 true cricket species are currently
known from Croatia. Some of them, such as Modicogryllus truncatus and Oecanthus dulcisonans, were recently
recorded for the first time in Croatia and are known from only a few localities so far. Morphological similarity to
related species, namely M. frontalis and O. pellucens, rather than rarity, was most likely responsible for those
species being overlooked, and we believe future studies will reveal much wider distribution in the country.
Scaly crickets (Mogoplistidae) are a globally understudied family of small, predominantly ground-dwelling
orthopterans, which occur in Europe with four genera—Arachnocephalus, Mogoplistes, Paramogoplistes and
Pseudomogoplistes. Each of those occurs in Croatia with a single species. Until recently, very few data existed for
P. novaki, a species described from Hvar Isl. (Krauss 1888). Reliable records exist from only four localities in
Croatia, and future research should aim to uncover its full distribution, as well as taxonomic status. There are
morphological and anatomical cues suggesting that the species might be an apterous member of Mogoplistes
(Rebrina & Brigić 2017). In that case, Paramogoplistes could be a polyphyletic genus (homoplasy being the lack of
tegmina) within paraphyletic Mogoplistes. In recent years, a few new species of Pseudomogoplistes have been

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described, but since the type species, P. squamiger (described from Dubrovnik area), is typeless, the necessity of
designating a neotype is evident.
Trigs' family (Trigonidiidae) are represented by three species in Croatia—Stenonemobius bicolor, locally
common Pteronemobius heydenii and common trig (Trigonidium cicindeloides), small and elusive species, whose
known distribution in Croatia has been substantially expanded with recent data. It is a cosmopolitan species.
Common trig was hitherto recorded from a few coastal localities (in some localities short winged, in some long-
winged form) and is probably widespread in the marshy habitats of the coastal area. Reed cricket (Natula averni) is
expected to be found in marshy habitats in the southern part of the country (Dubrovnik area). Future studies should
aim to confirm the presence of S. b. ponticus in the country and assess the extent of its occurrence, which is highly
important for the conservation of this enigmatic subspecies (Hochkirch et al. 2016).

Overview of Gryllotalpoidea families

Mole crickets (Gryllotalpidae) are a family of burrowing orthopterans, comprised mostly of large species. Of 13
European species, six were originally described from Italy and adjacent islands, primarily based on the karyotype
differences (number and appearance of the chromosomes). Due to cytotaxonomic nature of past revisions,
morphological and bioacoustic data on the genus in Europe remain scarce, making further revisions especially
difficult, but necessary. For now, two species are recorded in Croatia—Gryllotalpa gryllotalpa and G. stepposa.
However, morphologically distinct Adriatic populations are still not assigned to any known species. Those
populations could belong to one or more new species, or even to an already described Italian or southern Balkan
species.
Ant-loving crickets (Myrmecophilidae) are a family of small orthopterans (most species less than 5 mm in
length) inhabiting ants' nests. They are morphologically homogenous, which led to their obscurity on the
worldwide level. Reliable data for Croatia did not exist before Stalling & Birrer (2013). Only two species are
undoubtedly present in the country, namely Myrmecophilus hirticaudus and M. myrmecophilus. Several species
have been described from Southestern Europe in recent years, the occurrence of which should be investigated in
Croatia. Potential species include: M. acervorum (Panzer, 1799) in the Pannonian region, M. balcanicus Stalling,
2013 and M. nonveilleri Ingrisch & Pavićević, 2008 in the Adriatic, Dinaric and/or Pannonian regions.

Overview of Rhaphidophoroidea

Cave or camel crickets (Rhaphidophoridae) are represented by two genera in Croatia—troglobiotic Dolichopoda
(1 species) easily recognizable by long appendages and small eyes, and troglophilous Troglophilus (4 species). For
the first, a comprehensive phylogeographical study needs to be done, since the presence of yet unrecognized old
lineages is possible, of which some could be morphologically hardly distinguishable (as in Proteus: Gorički &
Trontelj (2006) or Dalmatolacerta: Podnar et al. (2013)). Recently, Karaman et al. (2011) performed preliminary
molecular and morphological revision of the genus Troglophilus in the Western Balkans. However, we still do not
have satisfactory knowledge on differential characters, so we are still unable to identify species with certainty.
Namely, T. cavicola is easily distinguished from other species, but exact differences between T. neglectus and T.
ovuliformis are unclear, the main difference allegedly being coloration of live specimens (Karaman et al. 2011).
Also, type series of T. neglectus and D. araneiformis are lost and a neotype for each needs to be designated in order
to have a referent specimen for further analysis. Furthermore, Allegrucci et al. (2017) found an unknown old
isolated lineage of T. brevicauda species group on the island of Mljet. After preliminary morphological analyses,
we confirm that the species is close to T. brevicauda species group (including T. brevicauda and T. zorae), but does
not belong to any of the described taxa. Description of the new species is in preparation.

Overview of Tettigonioidea

Bush crickets or katydids (Tettigoniidae) are the largest orthopteran family, including mainly plant-dwelling

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species, well-known for their cryptic coloration, characteristic songs and, often, large body size. The majority of
bush crickets are omnivorous (e.g. most representatives of the largest subfamily, Tettigoniinae), but there are also
predatory (e.g. Saginae) and herbivorous groups (e.g. Phaneropterinae), although they are rarely specialized for a
single food source. With altogether 76 species currently known from Croatia, bush cricket family is the largest
orthopteran group inhabiting the country. Diverse bush cricket fauna of Croatia includes two species endemic to the
country—B. kaltenbachi, known from only two Croatian islands, and R. buchichii, recorded on some islands and
two localities in the Dinaric mountains. However, these species were poorly studied in the past and the full extent
of their distribution is yet to be uncovered. This is prerequisite for reliable extinction risk assessment, as well as
successful monitoring and conservation of their populations. A substantial portion of Croatian bush cricket fauna
(15 species, about 20%) is comprised of Balkan endemics or stenoendemics, e.g. Leptophyes intermedia, P.
dalmatica, Metrioptera hoermanni, Modestana modesta, Tessellana orina etc., and other regional endemics, such
as B. kraussi. Relatively high level of bush cricket endemism in the Balkan Peninsula (Kenyeres et al. 2009) is
mainly associated with heterogeneous landscape abounding in physical barriers (e.g. the Dinaric Alps), varied
habitats and the existence of numerous glacial microrefugia in the past. Unsurprisingly, most endemics are
flightless, brachypterous or squamipterous species with specific habitat requirements. In spite of its perceived
diversity, bush cricket fauna of Croatia is far from being thoroughly studied, with only a few historical records of
some species, either yet unconfirmed (e.g. Isophya kraussii, Saga natoliae, Ephippiger persicarius) or only
recently confirmed (e.g. Andreiniimon nuptialis, Leptophyes punctatissima). Also, some species, such as
Leptophyes discoidalis, L. intermedia, Pholidoptera frivaldszkyi, M. hoermanni and Zeuneriana amplipennis, have
been recorded for the first time in Croatia fairly recently, and are known from a single or only a few localities so
far. Thus, future studies should aim to confirm historical records and evaluate the true extent of occurrence of those
species in the country. Furthermore, quite a few taxonomic problems remain to be resolved, notably the validity
and distribution of the subspecies within P. aptera (P. a. aptera, P. a. gjorgjevici and P. a. karnyi) and P. littoralis
(P. l. similis). Finally, as suggested by Ingrisch & Pavićević (2010), the exact number (one or two) and distribution
of the yet undescribed Poecilimon species (elegans group) from Croatia should be ascertained, and detailed
morphological and bioacoustic description should be done.

Grasshoppers, groundhoppers and molehoppers (Caelifera) overview

Overview of Acridoidea families

True grasshoppers (Acrididae) are the most specialized and diverse family of caeliferans, after Tettigoniidae the
second most diverse within Orthoptera. The family bloomed recently, its main lineages radiated in Caenozoic
(Song et al. 2015). Today there are more than 6 800 known species (Cigliano et al. 2018). Croatia is inhabited by
almost 80 species, and numerous are likely yet undiscovered. All species are herbivorous, although in some species
carnivorous behavior has also been observed. Some of the best known representatives of the family are swarming
locusts, due to their economic importance all over the world. Croatia is inhabited by a few locust species, of which
only some make occasional swarms in the Adriatic coast (e.g. Anacridium aegyptium, Calliptamus italicus,
Dociostaurus maroccanus, but not Schistocerca gregaria and Locusta migratoria). True grasshoppers are known
for their songs, and for being good jumpers and flyers (e.g. Massa et al. 2012). This is why endemism is not as high
as in Tettigoniidae (Kenyeres et al. 2009). Calling and courtship songs are usually species-specific and knowledge
from bioacoustics should be implemented into taxonomy, ecology, and evolution research of the group (e.g.
Vedenina & Helversen 2009; Willemse et al. 2009). Concerning bioacoustics, members of the subfamily
Gomphocerinae are diverse and interesting, but rather complex and thus still not understood very well. Arcyptera
brevipennis brevipennis is a taxon endemic to Dinaric mountains, while relic A. microptera was recently
rediscovered in the country. The genus Chorthippus is especially diverse in the Balkans. Recent revision of its
members inhabiting Greece discovered a few new lineages (Willemse et al. 2009), and the authors defined them as
new species and subspecies. Detailed bioacoustic analysis should be done in Croatia, since most of the records
were based on morphology. Particularly interesting is the situation with C. mollis—typical morphology is not
common in the Balkans, and the species can be misidentified for C. vagans or even C. biguttulus. Northern (central
European, C. m. mollis) form, rare in the Balkans, probably had higher selective advantage in cold north than in the

78 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


warm south. C. m. lesinensis is characteristic for southern Adriatic area and its mountains, while Alpine C. m.
ignifer inhabits Istria. We propose a new synonym based on the interpretation of bioacoustics, morphology, and
phylogeny (Helversen 1989, Ingrisch 1995, Willemse et al. 2009) within Chorthippus (biguttulus) species group
(C. b. hedickei = C. b. biguttulus). Steppic C. dichrous and C. loratus are very rare, almost extinct in Croatia.
Members of the tribe Stenobothrini (Stenobothrus, Myrmeleotettix, Rammeihippus, Omocestus) are in need of
revision. There are some endemic species within this lineage, but their relationship to the widely distributed species
remains unclear. Dinaric grasshopper (Rammeihippus dinaricus), recently reported from the country, is an
especially interesting stenobothrin, with unusual and complex calling and courtship song (Vedenina & Skejo,
unpublished). The species is not a good flier (especially squamipterous females) and is restricted to karst Dinaric
Alps. Another species recorded at an even smaller number of localities in the Dinaric Alps, Stenobothrus croaticus,
is restricted (as far as we know) to a single hill on the Velebit Mt. (Berger 2008). The species can fly (both males
and females) and the reason for its small area of occurrence is not yet understood. Interesting are members of the
subfamily Melanoplinae, probably a part of the relic group that was dominant in Europe a long time ago
(Chintauan-Marquier et al. 2011, 2014). Most of the species are brachypterous, squamipterous or apterous, and
because of their restricted dispersal ability, they usually inhabit small geographical areas (e.g. Odonopodisma
fallax, Micropodisma salamandra). Within the band winged grasshoppers, relic populations of Epacromius
tergenstinus and E. coerulipes, very rare in Europe (Hochkirch et al. 2016), inhabit parts of Istria and Cres Isl., and
are likely remains of the ancient Po river valley fauna.
Toad grasshoppers (Pamphagidae) are large, armored herbivorous orthopterans. The family is especially
diverse in the southernmost parts of the continent (there are numerous genera and species in the Iberian peninsula,
Italy, and Greece), whereas only eastern stone grasshopper (P. hystrix) inhabits Adriatic karst. Until recently, P.
hystrix was divided into three subspecies, all subsequently elevated to the rank of species, nominotypical inhabiting
Dinaric Alps (Massa et al. 2015). Eastern stone grasshopper is well known among natives in Croatia—sometimes it
swarms in the mountains, and there are stories of local children playing with the specimens instead of toy cars
(story of Ante Vukušić, famous meteorologist from Zavižan Meterological station on Velebit Mt.).

Overview of Tetrigoidea

Groundhoppers or pygmy grasshoppers (Tetrigidae) are a family of small grasshoppers, generally less than 2 cm in
length, which mostly feed on moss, algae and detritus. They inhabit humid microhabitats (e.g. around ponds, lakes,
rivers etc.). Southeastern Europe, including Croatia, harbors the majority of European species. Of 12 European
species, altogether 10 occur in Croatia (Skejo 2014). Noteworthy are T. tuerki, a relict species inhabiting alluvia of
Drava River, and T. transsylvanica hypsocorypha, a subspecies stenoendemic to the area on the border between
Croatia and Slovenia. Exact distribution areas of Croatian species are still not known, both because of the lack of
research and numerous misidentifications. Members of this family are not easily identifiable by non-experts due to
subtle morphological differences between the species (Devriese 1996, Skejo et al. 2014).

Overview of Tridactyloidea

Molehoppers (Tridactylidae), also known as pygmy or false mole crickets, are a tropical family with only a few
representatives in Holarctic. They are small (up to 3 mm in length), sand-digging grasshoppers which inhabit river
banks. Three species of the genus Xya occur in Europe. One of them is an Iberian endemic (X. iberica) with unclear
taxonomic status, while the remaining two species are widespread and both present in Croatia. They are easily
separated already by coloration. Distribution of these species in Croatia is not fully understood, because X.
pfaendleri was described relatively recently (Harz 1970). Prior to its description, only the records of X. variegata
existed for Croatia (Us 1967). Since the first record of X. pfaendleri (Szövényi & Puskás 2012), it became evident
that this species is more common than X. variegata, hitherto known from only a few localities, where it occurs
together with X. pfaendleri.

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 79
Conclusive thoughts

This checklist not only lists all the known crickets and grasshoppers inhabiting Croatia, but is intended to serve any
Orthoptera-lover as a basis for the research on this group of insects in Croatia. Numerous taxonomic, geographic
and morphological peculiarities are mentioned, and a detailed history of Orthoptera research in Croatia is given.
For every species, we checked and discussed its distribution, comparing it against the species' distribution in
Europe and in the neighboring countries. We attempted to bring up all the issues and criticisms concerning our
predecessors’ work, as well as our own work.
We are aware that this list is only a basis for further research and it will change in the future, hopefully as a
result of work of future authors who will invest their time and passion into uncovering new and interesting facts
about the grasshoppers and crickets of Croatia.

Acknowledgments and authors' contributions

The idea to make the first annotated checklist of Croatian Orthoptera was conceived by JS, FR and NT in 2011 and
2012, when the first two authors founded the Section of Orthopterology within the Biology students' association—
BIUS, under the mentorship of GP, GS and NT, and at the same time independently by GP and GS, who started to
study Orthoptera of Croatia at that time. Parts of this study are the result of numerous discussions and situations in
the last 6 years that included numerous people, namely European orthopterists (Dirk Berger, Filippo M. Buzzetti,
Dragan P. Chobanov, Battal Çiplak, Pedro J. Cordero Tapia, Bernard Defaut, Hendrik Devriese, Rob Felix, Paolo
Fontana, Stanislav Gomboc, Andrey V. Gorokhov, Klaus-Gerhard Heller, Axel Hochkirch, Martin Husemann,
Sigfrid Ingrisch, Elena I. Iorgu, Ionuț Ș. Iorgu, Slobodan Ivković, Ivo M. Karaman, Roy Kleukers, Arne Lehmann,
Gerlind Lehmann, Michele Lemonnier-Darcemont, Paulo Lemos, Vicenta Llorente, Antal Nagy, Barnabás Nagy,
Kirill Márk Orci, Dragan Pavićević, Taras I. Pushkar, István Rácz, Klaus Riede, Christian Roesti, Florin
Rutschmann, Hasan Sevgili, Sergey Yu. Storozhenko, Thomas Stalling, Deniz Şirin, Varvara Vedenina, Luc
Willemse), colleagues from the Faculty of Science in Zagreb (especially Lucija Šerić Jelaska), Natural History
Museum in Zagreb (especially Darija Ćaleta), Natural History Museum in Split, students from the Biology
students’ association—BIUS, and our friends. We sincerely thank all of them for giving us time, necessary
facilities, help and knowledge. Finally, we thank the anonymous reviewers for improving the quality of this
manuscript. Special thanks to Ivan Šapina for reading the proof and helping with corrigenda. JS created the
structure of the manuscript, JS and FR wrote the manuscript (FR in charge of Ensifera part, JS of Caelifera), GS
and GP contributed to all parts of the study, NT wrote historical introduction and division of the country into
regions, and contributed with organizing and funding the initial stages of the study. All the authors analyzed the
data and agree with all parts of the study. JS, FR and NT are of equal contribution.

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IUCN Assessments

Braud, Y., Hochkirch, A., Presa, J.J., Fontana, P., Roesti, C., Rutschmann, F., Monnerat, C., Zuna-Kratky, T. & Dusoulier, F.
(2016) Cyrtaspis scutata. The IUCN Red List of Threatened Species, 2016, e.T68286655A70649113. (accessed 29 March
2017).
Braud, Y., Hochkirch, A., Stalling, T., Fontana, P., Roesti, C., Rutschmann, F., Monnerat, C., Presa, J.J., Zuna-Kratky, T. &
Dusoulier, F. (2016) Myrmecophilus myrmecophilus. The IUCN Red List of Threatened Species, 2016,
e.T68441393A70648409. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68441393A70648409.en (accessed

88 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


30 March 2017).
Buzzetti, F.M., Hochkirch, A., Fontana, P., Kleukers, R., Massa, B. & Odé, B. (2016) Dolichopoda palpata
(e.T47709592A95721485. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T47709592A95721485.en.),
Platycleis romana (e.T47712879A47716262. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T47712879A47716262.en). The IUCN Red List of Threatened Species, 2016. (accessed 28 March 2017).
Buzzetti, F.M., Hochkirch, A., Fontana, P., Massa, B., Kleukers, R. & Odé, B. (2016) Micropodisma salamandra. The IUCN
Red List of Threatened Species, 2016, e.T15039570A70737661. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T15039570A70737661.en. (accessed 12 May 2017).
Buzzetti, F.M., Hochkirch, A., Stalling, T., Massa, B., Fontana, P., Odé, B. & Kleukers, R. (2016) Myrmecophilus ochraceus.
The IUCN Red List of Threatened Species, 2016, e.T47696335A47696362. (accessed 20 May 2017).
Chobanov, D.P., Hochkirch, A., Iorgu, I.Ș., Ivković, S., Krištín, A., Lemonnier-Darcemont, M., Pushkar, T., Şirin, D., Skejo, J.,
Szövényi, G., Vedenina, V. & Willemse, L.P.M. (2016) Odontopodisma fallax (e.T16084359A70204980. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16084359A70204980.en), Odontopodisma schmidti
(e.T16084382A70205163. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16084382A70205163.en),
Paracaloptenus cristatus (e.T15038919A70205175. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T15038919A70205175.en), Stenobothrus crassipes (e.T16084425A70215461. DOI: http://dx.doi.org/10.2305/
IUCN.UK.2016-3.RLTS.T16084425A70215461.en.), Stenobothrus croaticus (2016: e.T16084586A70214986. DOI: http:/
/dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16084586A70214986.en.) Tetrix transsylvanica (e.T2805A69677917.
DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T2805A69677917.en). The IUCN Red List of Threatened Species,
2016. (accessed 08 April 2017).
Chobanov, D.P., Hochkirch, A., Iorgu, I.Ș., Ivković, S., Krištín, A., Lemonnier–Darcemont, M., Pushkar, T., Şirin, D., Skejo, J.,
Stalling, T., Szövényi, G., Vedenina, V. & Willemse, L.P.M. (2016) Myrmecophilus hirticaudus. The IUCN Red List of
Threatened Species 2016: e.T68441316A70217343. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68441316A70217343.en. (accessed 30 March 2017).
Chobanov, D.P., Hochkirch, A., Iorgu, I.Ș., Ivković, S., Krištín, A., Lemonnier–Darcemont, M., Pushkar, T., Şirin, D., Skejo, J.,
Szövényi, G., Vedenina, V. & Willemse, L.P.M. (2016) Barbitistes kaltenbachi (e.T64549053A70217489. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T64549053A70217489.en), Barbitistes ocskayi (e.T64549495A70217509.
DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T64549495A70217509.en.), Dolichopoda araneiformis
(e.T68313567A70271472. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68313567A70271472.en),
Ephippiger discoidalis (e.T68315330A70271683. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68315330A70271683.en), Isophya kraussii (e.T68399046A70268900. DOI: http://dx.doi.org/10.2305/
IUCN.UK.2016-3.RLTS.T68399046A70268900.en), Metrioptera hoermanni (e.T68427655A70271693. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68427655A70271693.en), Modestana modesta (e.T68439525A70271763.
DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68439525A70271763.en), Modicogryllus truncatus
(e.T68439685A70627883), Poecilimon ampliatus (e.T45463520A70270406. DOI: http://dx.doi.org/10.2305/
IUCN.UK.2016-3.RLTS.T45463520A70270406.en), Poecilimon fussii (e.T68454987A70270595. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T68454987A70270595.en), Poecilimon gracilis (e.T45466571A70270600. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T45466571A70270600.en), Psorodonotus illyricus
(e.T68462447A85851396, DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68462447A85851396.en),
Stenonemobius bicolor (e.T68469061A70216325), Tessellana carinata (e.T68469288A70272066. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T68469288A70272066.en), Tettigonia balcanica (e.T70613909A70613914. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T70613909A70613914.en), Troglophilus neglectus
(e.T68485569A70271527. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68485569A70271527.en),
Troglophilus ovuliformis (e.T68485573A70271549. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68485573A70271549.en), Zeuneriana amplipennis (e.T68486506A70272132. DOI: http://dx.doi.org/10.2305/
IUCN.UK.2016-3.RLTS.T68486506A70272132.en). The IUCN Red List of Threatened Species, 2016. (accessed 15
January 2018).
Hochkirch, A., Chobanov, D.P., Heller, K.–G., Iorgu, I.Ș., Willemse, L.P.M., Szövényi, G., Şirin, D., Krištín, A., Ivković, S.,
Lemonnier–Darcemont, M., Pushkar, T., Skejo, J. & Vedenina, V. (2016) Poecilimon affinis. The IUCN Red List of
Threatened Species, 2016, e.T45466482A74527328. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T45466482A74527328.en (accessed 27 March 2017).
Hochkirch, A., Chobanov, D.P., Ivković, S., Lemonnier-Darcemont, M., Iorgu, I.Ș., Szövényi, G., Şirin, D., Willemse, L.P.M.,
Skejo, J., Pushkar, T., Vedenina, V. & Krištín, A. (2016) Isophya speciosa. The IUCN Red List of Threatened Species,
2016, e.T64869542A74525748. http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T64869542A74525748.en (accessed
30 October 2017).
Hochkirch, A., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J., Rutschmann, F., Szövényi, G. & Willemse, L.P.M. (2016)
Dociostaurus maroccanus (e.T16084458A74491017), Chorthippus dorsatus (e.T16084487A74245450). The IUCN Red
List of Threatened Species, 2016. (accessed 11 April 2017).
Hochkirch, A., Chobanov, D.P., Kleukers, R., Rutschmann, F., Krištín, A., Szövényi, G., Presa, J.J. & Willemse, L.P.M. (2016)
Acrida ungarica. The IUCN Red List of Threatened Species, 2016, e.T15037223A70632622. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T15037223A70632622.en (accessed10 April 2017).

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 89
Hochkirch, A., Chobanov, D.P., Krištín, A., Kleukers, R., Szövényi, G., Rutschmann, F., Willemse, L.P.M. & Presa, J.J. (2016)
Oedaleus decorus. The IUCN Red List of Threatened Species, 2016, e.T16084481A74495965. (accessed 08 May 2017).
Hochkirch, A., Chobanov, D.P., Krištín, A., Szövényi, G., Rutschmann, F., Willemse, L.P.M., Kleukers, R. & Presa, J.J. (2016)
Pseudochorthippus parallelus. The IUCN Red List of Threatened Species, 2016, e.T16084537A74250183. (accessed 05
May 2017).
Hochkirch, A., Chobanov, D.P., Krištín, A., Zuna-Kratky, T., Puskás, G., Lemonnier-Darcemont, M., Iorgu, I.Ș., Buzzetti, F.M.,
Willemse, L.P.M., Şirin, D., Skejo, J., Ivković, S., Szövényi, G. & Vedenina, V. (2016) Odontopodisma decipiens. The
IUCN Red List of Threatened Species, 2016, e.T16084581A74495954. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T16084581A74495954.en. (accessed 12 May 2017).
Hochkirch, A., Chobanov, D.P., Massa, B., Skejo, J., Ivković, S., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A.,
Presa, J.J. & Szövényi, G. (2016) Eupholidoptera schmidti. The IUCN Red List of Threatened Species, 2016,
e.T68379164A74538718. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68379164A74538718.en (accessed
28 March 2017).
Hochkirch, A., Chobanov, D.P., Pushkar, T., Korsunovskaya, O., Vedenina, V., Willemse, L.P.M., Szövényi, G., Şirin, D., Nieto,
A., Krištín, A., Ivković, S., Iorgu, I.Ș., Kranz, M., Lemonnier–Darcemont, M. & Skejo, J. (2016) Gryllotalpa stepposa.
The IUCN Red List of Threatened Species, 2016, e.T68382002A74521314. (accessed 13 July 2017).
Hochkirch, A., Chobanov, D.P., Şirin, D., Savitsky, V., Lemonnier-Darcemont, M., Szövényi, G., Iorgu, I.Ș., Willemse, L.P.M.,
Skejo, J., Krištín, A., Ivković, S., Pushkar, T. & Vedenina, V. (2016) Omocestus minutus. The IUCN Red List of Threatened
Species, 2016, e.T16084615A74507804. (accessed 05 May 2017).
Hochkirch, A., Chobanov, D.P., Szövényi, G., Krištín, A., Kleukers, R., Presa, J.J., Rutschmann, F. & Willemse, L.P.M. (2016)
Aiolopus strepens. The IUCN Red List of Threatened Species, 2016, e.T15431303A72593093. (accessed 08 May 2017).
Hochkirch, A., Chobanov, D.P., Szövényi, G., Krištín, A., Rutschmann, F., Willemse, L.P.M., Kleukers, R. & Presa, J.J. (2016)
Chorthippus mollis. The IUCN Red List of Threatened Species, 2016, e.T16084500A74247874. (accessed 05 May 2017).
Hochkirch, A., Chobanov, D.P., Szövényi, G., Krištín, A., Willemse, L.P.M., Presa, J.J., Kleukers, R. & Rutschmann, F. (2016)
Calliptamus barbarus. The IUCN Red List of Threatened Species, 2016, e.T15037702A72598041. Downloaded on 10
April 2017.
Hochkirch, A., Chobanov, D.P., Szövényi, G., Rutschmann, F., Krištín, A., Kleukers, R., Willemse, L.P.M. & Presa, J.J. (2016)
Acrotylus patruelis. The IUCN Red List of Threatened Species, 2016, e.T15431178A71713869. (accessed 08 May 2017).
Hochkirch, A., Chobanov, D.P., Vedenina, V., Iorgu, I.Ș., Zuna-Kratky, T., Szövényi, G., Fontana, P., Gomboc, S., Willemse,
L.P.M., Krištín, A., Ivković, S. & Pushkar, T. (2016) Chorthippus dichrous. The IUCN Red List of Threatened Species,
2016, e.T16084550A74244816. (accessed 05 May 2017).
Hochkirch, A., Chobanov, D.P., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016)
Poecilimon jonicus. The IUCN Red List of Threatened Species, 2016, e.T47709322A74528383. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016–3.RLTS.T47709322A74528383.en (accessed 16 May 2017).
Hochkirch, A., Chobanov, D.P., Zuna-Kratky, T., Gomboc, S., Puskás, G., Ivković, S., Rutschmann, F., Şirin, D., Defaut, B.,
Korsunovskaya, O., Krištín, A., Willemse, L.P.M., Presa, J.J., Kleukers, R. & Szövényi, G. (2016) Melanogryllus desertus.
The IUCN Red List of Threatened Species, 2016, e.T68427425A74519629. (accessed 30 March 2017).
Hochkirch, A., Devriese, H., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Paratettix meridionalis. The IUCN Red List of Threatened Species, 2016, e.T68449531A74530062.
(accessed 08 April 2017).
Hochkirch, A., Ferreira, S., Monnerat, C., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa,
J.J. & Szövényi, G. (2016) Pteronemobius lineolatus. The IUCN Red List of Threatened Species, 2016,
e.T68463350A74520487. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016–3.RLTS.T68463350A74520487.en. (accessed
18 May 2017).
Hochkirch, A., Fontana, P., Rutschmann, F., Presa, J.J., Szövényi, G., Chobanov, D.P., Kleukers, R., Willemse, L.P.M. &
Krištín, A. (2016) Calliptamus italicus. The IUCN Red List of Threatened Species, 2016, e.T15038693A72598188.
(accessed 10 April 2017).
Hochkirch, A., Gomboc, S., Lemonnier-Darcemont, M., Şirin, D., Chobanov, D.P., Skejo, J., Fontana, P., Willemse, L.P.M.,
Ivković, S., Szövényi, G., Iorgu, I.Ș., Pushkar, T., Vedenina, V. & Krištín, A. (2016) Chorthippus bornhalmi. The IUCN
Red List of Threatened Species, 2016, e.T16084529A74241469. (accessed 06 May 2017).
Hochkirch, A., Gomboc, S., Skejo, J., Szövényi, G., Ivković, S., Krištín, A., Lemonnier-Darcemont, M., Chobanov, D.P., Iorgu,
I.Ș., Willemse, L.P.M., Vedenina, V., Pushkar, T. & Şirin, D. (2016) Prionotropis hystrix. The IUCN Red List of Threatened
Species, 2016, e.T15038381A74523729. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T15038381A74523729.en. (accessed 12 May 2017).
Hochkirch, A., Holuša, J., Monnerat, C., Iorgu, I.Ș., Braud, Y., Skejo, J., Krištín, A., Chobanov, D.P., Liana, A., Zuna-Kratky,
T., Roesti, C., Fontana, P., Dusoulier, F., Presa, J.J. & Rutschmann, F. (2016) Tetrix tuerki. The IUCN Red List of
Threatened Species, 2016, e.T68469431A74530776. (accessed 08 April 2017).
Hochkirch, A., Iorgu, I.Ș., Chobanov, D.P., Szövényi, G., Krištín, A., Rutschmann, F., Kleukers, R., Willemse, L.P.M. & Presa,
J.J. (2016) Locusta migratoria. The IUCN Red List of Threatened Species, 2016, e.T16084575A74494334. (accessed 08
May 2017).
Hochkirch, A., Iorgu, I.Ș., Ivković, S., Chobanov, D.P., Szövényi, G., Krištín, A., Willemse, L.P.M., Skejo, J., Pushkar, T.,

90 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Lemonnier–Darcemont, M., Şirin, D. & Vedenina, V. (2016) Leptophyes discoidalis. The IUCN Red List of Threatened
Species, 2016, e.T44709134A55298760. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T44709134A55298760.en (accessed 27 March 2017).
Hochkirch, A., Iorgu, I.Ș., Lemonnier–Darcemont, M., Szövényi, G., Krištín, A., Ivković, S., Chobanov, D.P., Willemse,
L.P.M., Skejo, J., Pushkar, T., Puskás, G., Şirin, D. & Vedenina, V. (2016) Pholidoptera frivaldszkyi. The IUCN Red List of
Threatened Species, 2016, e.T62148031A74621932. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T62148031A74621932.en (accessed 28 March 2017).
Hochkirch, A., Iorgu, I.Ș., Puskás, G., Zuna-Kratky, T., Chobanov, D.P., Presa, J.J., Skejo, J., Willemse, L.P.M., Rutschmann,
F., Kleukers, R., Krištín, A. & Szövényi, G. (2016) Tetrix bolivari. The IUCN Red List of Threatened Species, 2016,
e.T68469357A74530185. (accessed 08 April 2017).
Hochkirch, A., Iorgu, I.Ș., Szövényi, G., Chobanov, D.P., Şirin, D., Lemonnier–Darcemont, M., Ivković, S., Willemse, L.P.M.,
Skejo, J., Krištín, A., Pushkar, T. & Vedenina, V. (2016) Poecilimon brunneri. The IUCN Red List of Threatened Species,
2016, e.T68454861A74528033. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016–3.RLTS.T68454861A74528033.en.
(accessed 16 May 2017).
Hochkirch, A., Iorgu, I.Ș., Vedenina, V., Chobanov, D.P., Willemse, L.P.M., Skejo, J., Pushkar, T., Ivković, S., Lemonnier-
Darcemont, M., Şirin, D., Szövényi, G. & Krištín, A. (2016) Chorthippus loratus. The IUCN Red List of Threatened
Species, 2016, e.T16084376A74246343. (accessed 05 May 2017).
Hochkirch, A., Iorgu, I.Ș., Zuna-Kratky, T., Szövényi, G., Ivković, S., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P.,
Kleukers, R., Krištín, A. & Presa, J.J. (2016) Leptophyes boscii. The IUCN Red List of Threatened Species, 2016,
e.T44709104A74526085. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T44709104A74526085.en (accessed
27 March 2017).
Hochkirch, A., Ivković, S., Gomboc, S., Chobanov, D.P., Presa, J.J., Monnerat, C., Lemonnier-Darcemont, M., Puskás, G.,
Korsunovskaya, O., Ferreira, S., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi, G. (2016)
Paracinema tricolor. The IUCN Red List of Threatened Species, 2016, e.T16084490A74509446. (accessed 10 April 2017).
Hochkirch, A., Ivković, S., Puskás, G., Lemonnier–Darcemont, M., Chobanov, D.P., Willemse, L.P.M., Rutschmann, F.,
Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016) Andreiniimon nuptialis. The IUCN Red List of Threatened
Species, 2016, e.T44710118A74525015. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T44710118A74525015.en (accessed 27 March 2017).
Hochkirch, A., Ivković, S., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Trigonidium cicindeloides. The IUCN Red List of Threatened Species, 2016,
e.T68485103A74520982. (accessed 30 March 2017).
Hochkirch, A., Kleukers, R., Rutschmann, F., Presa, J.J., Willemse, L.P.M., Krištín, A., Szövényi, G. & Chobanov, D.P. (2016)
Oedipoda germanica. The IUCN Red List of Threatened Species, 2016, e.T16084601A74507300. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T16084601A74507300.en. (accessed 08 May 2017).
Hochkirch, A., Korsunovskaya, O., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Tessellana tessellata. The IUCN Red List of Threatened Species, 2016, e.T68469317A74624644.
(accessed 28 March 2017).
Hochkirch, A., Krištín, A., Chobanov, D.P., Ivković, S., Skejo, J., Zuna-Kratky, T., Willemse, L.P.M., Rutschmann, F.,
Kleukers, R., Presa, J.J. & Szövényi, G. (2016) Ephippiger ephippiger. The IUCN Red List of Threatened Species, 2016,
e.T68315499A74537999. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016–3.RLTS.T68315499A74537999.en. (accessed
29 March 2017).
Hochkirch, A., Krištín, A., Chobanov, D.P., Skejo, J., Ivković, S., Gomboc, S., Willemse, L.P.M., Szövényi, G., Şirin, D.,
Heller, K.–G., Iorgu, I.Ș., Lemonnier–Darcemont, M., Pushkar, T. & Vedenina, V. (2016) Poecilimon schmidtii. The IUCN
Red List of Threatened Species, 2016, e.T68459063A74528709. (accessed 27 March 2017).
Hochkirch, A., Krištín, A., Kleukers, R., Szövényi, G., Chobanov, D.P., Rutschmann, F., Presa, J.J. & Willemse, L.P.M. (2016)
Sphingonotus caerulans. The IUCN Red List of Threatened Species, 2016, e.T16084628A74511116. (accessed 08 May
2017).
Hochkirch, A., Krištín, A., Szövényi, G., Chobanov, D.P., Rutschmann, F., Kleukers, R., Presa, J.J. & Willemse, L.P.M. (2016)
Aiolopus thalassinus. The IUCN Red List of Threatened Species, 2016, e.T15431312A72593174. (accessed 08 May 2017).
Hochkirch, A., Krištín, A., Szövényi, G., Chobanov, D.P., Rutschmann, F., Willemse, L.P.M., Kleukers, R. & Presa, J.J. (2016)
Oedipoda caerulescens. The IUCN Red List of Threatened Species, 2016, e.T16084576A74496127. (accessed 08 May
2017).
Hochkirch, A., Krištín, A., Szövényi, G., Presa, J.J., Rutschmann, F., Chobanov, D.P., Kleukers, R. & Willemse, L.P.M. (2016)
Anacridium aegyptium. The IUCN Red List of Threatened Species, 2016, e.T16084533A72594185. (accessed 10 April
2017).
Hochkirch, A., Krištín, A., Zuna-Kratky, T., Ivković, S., Puskás, G., Chobanov, D.P., Şirin, D., Iorgu, I.Ș., Willemse, L.P.M.,
Rutschmann, F., Kleukers, R., Presa, J.J. & Szövényi, G. (2016) Pachytrachis gracilis. The IUCN Red List of Threatened
Species, 2016, e.T68446261A74542870. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68446261A74542870.en (accessed 28 March 2017).
Hochkirch, A., Lemonnier-Darcemont, M., Chobanov, D.P., Puskás, G., Presa, J.J., Skejo, J., Willemse, L.P.M., Rutschmann, F.,
Kleukers, R., Krištín, A. & Szövényi, G. (2016) Arcyptera microptera. The IUCN Red List of Threatened Species, 2016,

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 91
e.T16084361A72594357. (accessed 10 April 2017).
Hochkirch, A., Lemonnier–Darcemont, M., Iorgu, I.Ș., Chobanov, D.P., Ivković, S., Willemse, L.P.M., Szövényi, G., Şirin, D.,
Nieto, A., Krištín, A., Heller, K.–G., Kranz, M., Pushkar, T., Skejo, J. & Vedenina, V. (2016) Poecilimon thoracicus. The
IUCN Red List of Threatened Species, 2016, e.T68459410A74529086. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016–
3.RLTS.T68459410A74529086.en. (accessed 16 May 2017).
Hochkirch, A., Lemonnier–Darcemont, M., Skejo, J., Chobanov, D.P., Willemse, L.P.M., Szövényi, G., Şirin, D., Pushkar, T.,
Iorgu, I.Ș., Ivković, S., Krištín, A. & Vedenina, V. (2016) Tessellana orina. The IUCN Red List of Threatened Species,
2016, e.T68469308A74624624. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68469308A74624624.en
(accessed 28 March 2017).
Hochkirch, A., Lemonnier–Darcemont, M., Skejo, J., Ivković, S., Gomboc, S., Willemse, L.P.M., Rutschmann, F., Chobanov,
D.P., Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016) Barbitistes yersini. The IUCN Red List of Threatened
Species, 2016, e.T64549979A74525174. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T64549979A74525174.en (accessed 27 March 2017).
Hochkirch, A., Lemonnier–Darcemont, M., Skejo, J., Szövényi, G., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P.,
Kleukers, R., Krištín, A. & Presa, J.J. (2016) Rhacocleis germanica. The IUCN Red List of Threatened Species, 2016,
e.T44698061A74623844. (accessed 28 March 2017).
Hochkirch, A., Massa, B., Fontana, P., Presa, J.J., Iorgu, I.Ș., Şirin, D., Chobanov, D.P., Skejo, J., Willemse, L.P.M.,
Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi, G. (2016) Platycleis intermedia. The IUCN Red List of Threatened
Species, 2016, e.T68452838A74622481. (accessed 28 March 2017).
Hochkirch, A., Massa, B., Iorgu, I.Ș., Korsunovskaya, O., Presa, J.J., Chobanov, D.P., Ivković, S., Skejo, J., Morin, D.,
Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi, G. (2016) Decticus albifrons. The IUCN Red List
of Threatened Species, 2016, e.T68288297A74536527. (accessed 28 March 2017).
Hochkirch, A., Massa, B., Monnerat, C., Skejo, J., Gomboc, S., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers,
R., Krištín, A., Presa, J.J. & Szövényi, G. (2016) Yersinella raymondii. The IUCN Red List of Threatened Species, 2016,
e.T68486449A74625126. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68486449A74625126.en (accessed
28 March 2017).
Hochkirch, A., Massa, B., Skejo, J., Presa, J.J., Zuna-Kratky, T., Krištín, A., Ivković, S., Korsunovskaya, O., Monnerat, C.,
Puskás, G., Chobanov, D.P., Szövényi, G., Kleukers, R. & Rutschmann, F. (2016) Saga pedo. The IUCN Red List of
Threatened Species, 2016, e.T19811A74624296. (accessed 29 March 2017).
Hochkirch, A., Massa, B., Zuna-Kratky, T., Krištín, A., Puskás, G., Ivković, S., Presa, J.J., Skejo, J., Monnerat, C., Szövényi,
G., Kleukers, R., Chobanov, D.P. & Rutschmann, F. (2016) Meconema meridionale, The IUCN Red List of Threatened
Species, 2016, e.T68427413A74540416. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68427413A74540416.en (accessed 29 March 2017).
Hochkirch, A., Monnerat, C., Szövényi, G., Skejo, J., Gomboc, S., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P.,
Kleukers, R., Krištín, A. & Presa, J.J. (2016) Pachytrachis striolatus. The IUCN Red List of Threatened Species, 2016,
e.T68446326A74542911. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68446326A74542911.en (accessed
28 March 2017).
Hochkirch, A., Odé, B., Lemos, P., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Oecanthus dulcisonans. The IUCN Red List of Threatened Species, 2016, e.T68444276A74520421.
(accessed 30 March 2017).
Hochkirch, A., Presa, J.J., Chobanov, D.P., Krištín, A., Szövényi, G., Kleukers, R., Rutschmann, F. & Willemse, L.P.M. (2016)
Schistocerca gregaria. The IUCN Red List of Threatened Species, 2016, e.T16084387A74510644. (accessed 10 April
2017).
Hochkirch, A., Presa, J.J., Rutschmann, F., Szövényi, G., Kleukers, R., Chobanov, D.P., Willemse, L.P.M. & Krištín, A. (2016)
Arcyptera brevipennis. The IUCN Red List of Threatened Species, 2016, e.T16084569A72594260. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T16084569A72594260.en. (accessed 10 April 2017).
Hochkirch, A., Presa, J.J., Skejo, J., Chobanov, D.P., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi,
G. (2016) Tetrix depressa. The IUCN Red List of Threatened Species, 2016, e.T69674446A69674726. (accessed 08 April
2017).
Hochkirch, A., Presa, J.J., Skejo, J., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A. & Szövényi,
G. (2016) Pseudomogoplistes squamiger. The IUCN Red List of Threatened Species, 2016, e.T68460190A74521817.
(accessed 30 March 2017).
Hochkirch, A., Puskás, G., Chobanov, D.P., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Arachnocephalus vestitus. The IUCN Red List of Threatened Species, 2016, e.T64547639A74521330.
(accessed 30 March 2017).
Hochkirch, A., Puskás, G., Şirin, D., Ivković, S., Szövényi, G., Chobanov, D.P., Lemonnier-Darcemont, M., Skejo, J.,
Rutschmann, F., Presa, J.J., Krištín, A., Willemse, L.P.M. & Kleukers, R. (2016) Acrotylus longipes. The IUCN Red List of
Threatened Species, 2016, e.T15431174A70826598. (accessed 08 May 2017).
Hochkirch, A., Puskás, G., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Poecilimon elegans. The IUCN Red List of Threatened Species, 2016, e.T68454968A74528219. DOI:
http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68454968A74528219.en (accessed 27 March 2017).

92 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Hochkirch, A., Puskás, G., Zuna-Kratky, T., Chobanov, D.P., Ivković, S., Monnerat, C., Willemse, L.P.M., Rutschmann, F.,
Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016) Pteronemobius heydenii. The IUCN Red List of Threatened
Species, 2016, e.T68463301A74520473. (accessed 30 March 2017).
Hochkirch, A., Rutschmann, F., Chobanov, D.P., Presa, J.J., Kleukers, R., Willemse, L.P.M., Szövényi, G. & Krištín, A. (2016)
Mecostethus parapleurus. The IUCN Red List of Threatened Species, 2016, e.T16084608A74495170. (accessed 08 May
2017).
Hochkirch, A., Rutschmann, F., Willemse, L.P.M., Krištín, A., Chobanov, D.P., Szövényi, G., Presa, J.J. & Kleukers, R. (2016)
Chorthippus biguttulus. The IUCN Red List of Threatened Species, 2016, e.T16084474A74239267. (accessed 05 May
2017).
Hochkirch, A., Sergeev, M.G., Chobanov, D.P., Krištín, A., Szövényi, G., Rutschmann, F., Kleukers, R., Willemse, L.P.M. &
Presa, J.J. (2016) Chorthippus brunneus. The IUCN Red List of Threatened Species, 2016, e.T16084483A74243716.
(accessed 05 May 2017).
Hochkirch, A., Şirin, D., Presa, J.J., Chobanov, D.P., Puskás, G., Dusoulier, F., Lemonnier-Darcemont, M., Iorgu, I.Ș.,
Monnerat, C., Ivković, S., Korsunovskaya, O., Willemse, L.P.M., Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi, G.
(2016) Xya variegata. The IUCN Red List of Threatened Species, 2016, e.T68486396A74625447. (accessed 08 April
2017).
Hochkirch, A., Skejo, J. & Szövényi, G. (2016) Rammeihippus dinaricus. The IUCN Red List of Threatened Species, 2016,
e.T16084399A74510426. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16084399A74510426.en. (accessed
05 May 2017).
Hochkirch, A., Skejo, J., Buzzetti, F.M., Zuna-Kratky, T., Gomboc, S., Willemse, L.P.M., Szövényi, G., Şirin, D., Krištín, A.,
Ivković, S., Chobanov, D.P., Iorgu, I.Ș., Lemonnier–Darcemont, M., Pushkar, T., Vedenina, V. & Heller, K.–G. (2016)
Poecilimon ornatus. The IUCN Red List of Threatened Species, 2016, e.T68458496A74528658. (accessed 27 March
2017).
Hochkirch, A., Skejo, J., Fontana, P., Szövényi, G., Lemonnier–Darcemont, M., Chobanov, D.P., Şirin, D., Krištín, A., Pushkar,
T., Ivković, S., Vedenina, V., Willemse, L.P.M. & Iorgu, I.Ș. (2016) Rhacocleis buchichii. The IUCN Red List of
Threatened Species, 2016, e.T44698046A74622946. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T44698046A74622946.en (accessed 28 March 2017).
Hochkirch, A., Skejo, J., Gomboc, S., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J.
& Szövényi, G. (2016) Montana stricta. The IUCN Red List of Threatened Species, 2016, e.T68440172A74542394. DOI:
http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68440172A74542394.en (accessed 28 March 2017).
Hochkirch, A., Skejo, J., Gomboc, S., Willemse, L.P.M., Szövényi, G., Şirin, D., Krištín, A., Ivković, S., Chobanov, D.P., Iorgu,
I.Ș., Lemonnier–Darcemont, M., Pushkar, T. & Vedenina, V. (2016) Pholidoptera dalmatica. The IUCN Red List of
Threatened Species, 2016, e.T44709923A74621900. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T44709923A74621900.en (accessed 28 March 2017).
Hochkirch, A., Skejo, J., Lemonnier–Darcemont, M., Ivković, S., Chobanov, D.P., Şirin, D., Willemse, L.P.M., Szövényi, G.,
Krištín, A., Iorgu, I.Ș., Pushkar, T. & Vedenina, V. (2016) Saga natoliae. The IUCN Red List of Threatened Species, 2016,
e.T68468842A74624210. (accessed 29 March 2017).
Hochkirch, A., Skejo, J., Massa, B., Presa, J.J., Puskás, G., Şirin, D., Chobanov, D.P., Gomboc, S., Willemse, L.P.M.,
Rutschmann, F., Kleukers, R., Krištín, A. & Szövényi, G. (2016) Sepiana sepium. The IUCN Red List of Threatened
Species, 2016, e.T68468982A74624610. (accessed 28 March 2017).
Hochkirch, A., Skejo, J., Szövényi, G., Lemonnier–Darcemont, M., Ivković, S., Willemse, L.P.M., Şirin, D., Krištín, A.,
Chobanov, D.P., Iorgu, I.Ș., Pushkar, T. & Vedenina, V. (2016) Pachytrachis frater. The IUCN Red List of Threatened
Species, 2016, e.T68446184A74542746. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T68446184A74542746.en (accessed 28 March 2017).
Hochkirch, A., Skejo, J., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. & Szövényi,
G. (2016) Paramogoplistes novaki. The IUCN Red List of Threatened Species, 2016, e.T68446520A74521769. DOI: http:/
/dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68446520A74521769.en (accessed 30 March 2017).
Hochkirch, A., Stalling, T., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Myrmecophilus acervorum. The IUCN Red List of Threatened Species, 2016,
e.T68441140A74521979. (accessed 20 May 2017).
Hochkirch, A., Szövényi, G., Buzzetti, F.M., Lemonnier–Darcemont, M., Skejo, J., Gomboc, S., Willemse, L.P.M., Presa, J.J.,
Kleukers, R., Chobanov, D.P., Krištín, A. & Rutschmann, F. (2016) Pholidoptera femorata. The IUCN Red List of
Threatened Species, 2016, e.T68451595A74621927. (accessed 28 March 2017).
Hochkirch, A., Szövényi, G., Chobanov, D.P., Fontana, P., Buzzetti, F.M., Gomboc, S., Ivković, S., Iorgu, I.Ș., Lemonnier–
Darcemont, M., Puskás, G., Zuna-Kratky, T., Willemse, L.P.M., Skejo, J., Pushkar, T., Krištín, A., Şirin, D. & Vedenina, V.
(2016) Isophya modestior. The IUCN Red List of Threatened Species, 2016, e.T68399393A74525391. DOI: http://
dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68399393A74525391.en (accessed 27 March 2017).
Hochkirch, A., Szövényi, G., Holuša, J., Zuna-Kratky, T., Krištín, A., Şirin, D., Savitsky, V., Lemonnier-Darcemont, M.,
Chobanov, D.P., Willemse, L.P.M., Skejo, J., Ivković, S., Iorgu, I.Ș., Pushkar, T. & Vedenina, V. (2016) Dociostaurus
brevicollis. The IUCN Red List of Threatened Species, 2016, e.T16084356A74253475. (accessed 11 April 2017).
Hochkirch, A., Szövényi, G., Ivković, S., Puskás, G., Skejo, J., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers,

FIRST CROATIAN CRICKETS AND GRASSHOPPERS Zootaxa 4533 (1) © 2018 Magnolia Press · 93
R., Krištín, A. & Presa, J.J. (2016) Leptophyes laticauda. The IUCN Red List of Threatened Species, 2016,
e.T44709490A74526282. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T44709490A74526282.en (accessed
27 March 2017).
Hochkirch, A., Szövényi, G., Kleukers, R., Krištín, A., Chobanov, D.P., Rutschmann, F., Willemse, L.P.M. & Presa, J.J. (2016)
Chorthippus apricarius. The IUCN Red List of Threatened Species, 2016, e.T16084489A74239260. (accessed 05 May
2017).
Hochkirch, A., Szövényi, G., Krištín, A., Chobanov, D.P., Rutschmann, F., Willemse, L.P.M., Kleukers, R. & Presa, J.J. (2016)
Acrotylus insubricus. The IUCN Red List of Threatened Species, 2016, e.T15431156A70802618. (accessed 08 May 2017).
Hochkirch, A., Szövényi, G., Zuna-Kratky, T., Gomboc, S., Korsunovskaya, O., Chobanov, D.P., Iorgu, I.Ș., Defaut, B.,
Willemse, L.P.M., Presa, J.J., Kleukers, R., Krištín, A. & Rutschmann, F. (2016) Omocestus petraeus. The IUCN Red List
of Threatened Species, 2016, e.T16084565A74507810. (accessed 05 May 2017).
Hochkirch, A., Szövényi, G., Zuna-Kratky, T., Korsunovskaya, O., Gomboc, S., Presa, J.J., Monnerat, C., Chobanov, D.P.,
Iorgu, I.Ș., Defaut, B., Willemse, L.P.M., Krištín, A., Kleukers, R. & Rutschmann, F. (2016) Omocestus haemorrhoidalis.
The IUCN Red List of Threatened Species, 2016, e.T16084427A74507786. (accessed 05 May 2017).
Hochkirch, A., Szövényi, G., Zuna-Kratky, T., Korsunovskaya, O., Ivković, S., Iorgu, I.Ș., Lemonnier-Darcemont, M., Defaut,
B., Presa, J.J., Krištín, A., Willemse, L.P.M., Kleukers, R., Chobanov, D.P. & Rutschmann, F. (2016) Stenobothrus fischeri.
The IUCN Red List of Threatened Species, 2016, e.T16084438A74514425. (accessed 05 May 2017).
Hochkirch, A., Szövényi, G., Zuna-Kratky, T., Krištín, A., Iorgu, I.Ș., Skejo, J., Korsunovskaya, O., Willemse, L.P.M., Presa,
J.J., Chobanov, D.P., Kleukers, R. & Rutschmann, F. (2016) Epacromius coerulipes. The IUCN Red List of Threatened
Species, 2016, e.T16084393A74491103. (accessed 08 May 2017).
Hochkirch, A., Szövényi, G., Zuna-Kratky, T., Vedenina, V., Gomboc, S., Krištín, A., Willemse, L.P.M., Iorgu, I.Ș., Chobanov,
D.P., Lemonnier-Darcemont, M., Ivković, S., Pushkar, T., Skejo, J. & Şirin, D. (2016) Chorthippus oschei. The IUCN Red
List of Threatened Species, 2016, e.T16084560A74249405. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-
3.RLTS.T16084560A74249405.en. (accessed 05 May 2017).
Hochkirch, A., Vedenina, V., Szövényi, G., Chobanov, D.P., Şirin, D., Gomboc, S., Krištín, A., Savitsky, V., Iorgu, I.Ș.,
Willemse, L.P.M., Lemonnier-Darcemont, M., Ivković, S. & Pushkar, T. (2016) Euchorthippus pulvinatus. The IUCN Red
List of Threatened Species, 2016, e.T16084621A74492473. (accessed 11 April 2017).
Hochkirch, A., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016)
Acheta domesticus (e.T64336581A74517796), Barbitistes serricauda (e.T64549568A74525163. DOI: http://dx.doi.org/
10.2305/IUCN.UK.2016-3.RLTS.T64549568A74525163.en), Diestrammena asynamora (e.T69673959A69673975),
Gryllomorpha dalmatina (e.T47695140A74518522), Gryllotalpa gryllotalpa (e.T68380889A74521302), Gryllus
bimaculatus (e.T68382951A74519241), Gryllus campestris (e.T68382951A74519241), Leptophyes punctatissima
(e.T44709622A74526293. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T44709622A74526293.en),
Phaneroptera falcata (e.T68451243A74526336), Phaneroptera nana (e.T68451285A74526741), Polysarcus denticauda
(e.T68459646A74529234. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68459646A74529234.en), Tylopsis
lilifolia (e.T68485724A74529809). The IUCN Red List of Threatened Species, 2016. (accessed 15 January 2018).
Hochkirch, A., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. & Szövényi, G. (2016)
Myrmeleotettix maculatus (e.T16084559A74495466), Omocestus rufipes (e.T16084488A74508968), Psophus stridulus
(e.T16084568A74509860), Stenobothrus lineatus (e.T16084422A74514438). Stenobothrus nigromaculatus
(e.T16084451A74515226), Stenobothrus stigmaticus (e.T16084454A74515452), Stenobothrus rubicundulus
(e.T16084507A74515247. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16084507A74515247.en.),
Stethophyma grossum (e.T16084611A74516611), Tetrix bipunctata (e.T68469349A74530155). The IUCN Red List of
Threatened Species, 2016. (accessed 08 April 2017).
Hochkirch, A., Willemse, L.P.M., Rutschmann, F., Chobanov, D.P., Krištín, A., Presa, J.J., Szövényi, G. & Kleukers, R. (2016)
Arcyptera fusca. The IUCN Red List of Threatened Species, 2016, e.T16084432A72594321. (accessed 10 April 2017).
Hochkirch, A., Willemse, L.P.M., Rutschmann, F., Danielczak, A., Chobanov, D.P., Kleukers, R., Krištín, A., Presa, J.J. &
Szövényi, G. (2016) Eumodicogryllus bordigalensis (e.T68378608A74518073), Oecanthus pellucens
(e.T68444291A74520428). The IUCN Red List of Threatened Species, 2016. (accessed 15 January 2018).
Hochkirch, A., Willemse, L.P.M., Szövényi, G., Rutschmann, F., Presa, J.J., Krištín, A., Kleukers, R. & Chobanov, D.P. (2016)
Bicolorana bicolor (e.T64550306A6967917), Conocephalus dorsalis (e.T68280729A74535339), Conocephalus fuscus
(e.T68280913A74535358), Decticus verrucivorus (e.T68288600A74536544), Meconema thalassinum
(e.T68427417A74540428. DOI: http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T68427417A74540428.en),
Metrioptera brachyptera (e.T68427614A74540433), Pholidoptera griseoaptera (e.T68451642A74621940), Platycleis
albopunctata (e.T68451982A74622396), Roeseliana roeselii (e.T68468465A74623999), Ruspolia nitidula
(e.T68468487A74624078), Tettigonia cantans (e.T68484522A74624729), Tettigonia caudata (e.T68485021A74624838),
Tettigonia viridissima (e.T68485046A74624843). The IUCN Red List of Threatened Species, 2016. (accessed 15 January
2018).
Hochkirch, A., Willemse, L.P.M., Szövényi, G., Rutschmann, F., Presa, J.J., Krištín, A., Kleukers, R. & Chobanov, D.P. (2016)
Chorthippus vagans (e.T16084357A74251494), Chrysochraon dispar (e.T16084524A74251505), Dociostaurus genei
(e.T16084544A74253488), Euchorthippus declivus (e.T16084407A74491942. DOI: http://dx.doi.org/10.2305/
IUCN.UK.2016-3.RLTS.T16084407A74491942.en), Euthystira brachyptera (e.T16084369A74492646),

94 · Zootaxa 4533 (1) © 2018 Magnolia Press SKEJO ET AL.


Gomphocerippus rufus (e.T16084377A74493482), Tetrix ceperoi (e.T68469367A74530260), Tetrix subulata
(e.T68469394A74530607), Tetrix tenuicornis (e.T68469416A74530618), Tetrix undulata (e.T68469441A74531235, DOI:
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2016. (accessed 08 April 2017).
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