Research Article Journal of Orthoptera Research 2019, 28(2): 187-193 Response of orthopterans to macroclimate changes: A 15-year case study in Central European humid grasslands ZOLTAN KENYERES!, GABorR TAKACS2, NORBERT BAUER? 1 Acrida Conservational Research L.P., Tapolca 8300, Hungary. 2 Ferté-Hansag National Park Directorate, Sarr6d 9435, Hungary. 3 Department of Botany, Hungarian Natural History Museum, Budapest 1087, Hungary. Corresponding author: Zoltdn Kenyeres (kenyeres@acridabt.hu) Academic editor: Maria-Marta Cigliano | Received 26 February 2019 | Accepted 29 May 2019 | Published 2 October 2019 http://zoobank.org/39E596D 1-EFFF-4E9E-80FA-1E5C94DE31A5 Citation: Kenyeres Z, Takacs G, Bauer N (2019) Response of orthopterans to macroclimate changes: A 15-year case study in Central European humid grasslands. Journal of Orthoptera Research 28(2): 187-193. https://doi.org/10.3897/jor.28.34102 Abstract Orthoptera is a good indicator taxon of macroclimate changes. In our case study, we analyzed data of orthopterans, vegetation, and macrocli- mate collected yearly from 2002 through 2017 in Central European hu- mid grasslands. During the study period, the annual mean temperature increased, while the relative abundance of moderately hygrophilic orthop- teran species decreased significantly. On the other hand, the species rich- ness and diversity of the assemblages increased due, mostly, to an increase of graminicole/thermophilic species. According to our results, the conser- vation of the hygrophilic orthopteran assemblages of Central European humid grasslands under global warming can only be ensured by adequate land management, which can at least mitigate the effects of climate change resulting in the warming and drying of humid habitats. Keywords climate change, Hungary, indicator, landscape management, monitoring, species richness Introduction Global climate change has a significant impact on insect popu- lations and assemblages, both directly (in terms of temperature, precipitation, and seasonal changes) and indirectly (changes in vegetation productivity and quality characteristics, presence and spread of predators and pathogenic organisms) (Ryrholm 2001, Lightfoot 2006, Menéndez 2007). Orthopterans seem to be a good indicator for the monitoring of macroclimate changes, as their distribution, density, and assemblage structure are mostly deter- mined by climatic conditions (Dreux 1962, Wingerden et al. 1992, Racz et al. 1994, Guido and Chemini 2000, Squitier and Capinera 2002, Gardiner and Hassall 2009, Buri et al. 2013, Kenyeres et al. 2018). The mild effects of global climate change include variations in the phenological characteristics of the orthopteran species. Ac- cording to the laboratory test results of Fielding and Defoliart (2010), in the cases of Melanoplus borealis (Fieber) and Melanoplus sanguinipes (Fabricius) a temperature increase of 2, 3, and 4°C re- sults in earlier hatching by 3, 5, and 7 days, respectively. This phe- nomenon was also observed in Hungary between 1958 and 2009: the hatching of early species such as Isophya costata, Isophya stysi, Isophya kraussii, and Isophya camptoxypha, shifted earlier and earlier at a rate of 2-3 days/decade, due to the increasing spring mean temperature (Szabo et al. 2014). Another important phenomenon resulting from global warm- ing is the change in the area boundary of the Orthoptera species according to their cold tolerance (Uvarov 1931, Burton 2001). Eu- ropean species were observed to be expanding to the north due to global warming including, among others, Phaneroptera falcata and Phaneroptera nana (Koéarek et al. 2008), Ruspolia nitidula, Mecoste- thus parapleurus (HoluSa et al. 2007) and Roeseliana roeselii (Gar- diner 2009, Wissmann et al. 2009). The results of the impact of climate change on density of orthopterans are contradictory; there are some localities where an increase has been found while in oth- ers a decrease in density was observed (Bale et al. 2002, O’Neill et al. 2008, Laws and Belovsky 2010). This may be due to the lo- cality-specific reaction of insects to climate change influenced by several circumstances of the habitat (e.g., altitude), nutrition (e.g., polyphagous vs. monophagous) and climatic requirements of the dominant species (Bale et al. 2002). Flightless, habitat-specialist species with a restricted area are particularly vulnerable to the effects of climate change. Their yearly population size, in many cases, responds very sensitively to some climate parameters (Ke- nyeres et al. 2018). In the case of European species, climate change, although to a negligible extent when compared to the above, also influences the composition of the orthopteran assemblages through the in- crease in the chances of survival of the species that overwinter in an imago state (Kiritani 2006). According to our earlier experiences gained in various re- gions of Central Europe, the impact of climate change on the orthopteran assemblages is really pronounced in humid grass- land habitats (Kenyeres and Cservenka 2014). In this case study, we analyzed the data of orthopterans, vegetation, and macrocli- mate collected yearly between 2002 and 2017 under the moni- toring program of the Ferté-Hansag National Park Directorate. Our main questions included: 1) What changes can be seen in JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) 188 the relative abundance of different life forms and eco-types over the 15-year period? 2) Can changes be detected in the relative abundance of species? 3) Is it possible to show relationships be- tween the diversity parameters of orthopteran assemblages and parameters of the macroclimate? We hypothesized that changes in macroclimate (increasing temperature and decreasing precipi- tation) may result in an increase of thermophilic species and in a decrease of hygrophilic species. Material and methods Study area.—The study area (240 ha) (Fig. 1) belongs to the Han- sag mesoregion of the Kisalf6ld macroregion. It is located at an altitude of ~114 maz.s.l. and is characterized by large flatlands. The potential vegetation of Hansag, having previously had hydrological connections with Lake Fert6é, which is 30 km away, is moorlands, fens, and marshlands. The drainage of the Hansag area started with the Romans, but the recent hydrological con- ditions are the result of interventions carried out in the last 100 years. At present, the natural vegetation of the region is restricted to some patches covered by large, mesic grasslands, mosaiced and surrounded by forests, scrubs, and tree plantations. The study area is dominated by intrazonal bog and fen soils. The level of the groundwater is permanently around 1 m. Turf resulting from loosening organic material is about half a meter. The average total duration of annual insolation in the region is 1,900 hours. Mean annual temperature is around 10.1°C. The average annual precipi- tation is 630 mm (Dé6vényi 2010). Experimental design.—The study area included two habitat-mosa- ics: a large area of contacting parcels dominated by humid grass- lands and a smaller one on a comb of the local microrelief with humid and semi-dry grasslands impacted by forest areas (Fig. 1). Seven sampling sites were established as 50x50 m sized quadrats. Data collection was carried out from 2002 to 2017. All quadrats were consistently used: 1-2 were extensively grazed and 3-7 were mowed once a year in early June. Environmental parameters.—Measurements of the main vegetation parameter (average height of the vegetation) were carried out on 3 plots in each sampling site during each orthopteran sampling. The height of the vegetation was measured in cm with the use of a 30 cm wide and 100 cm high white card. The total cover of the vegeta- tion showed only small differences between 90 and 100% cover, so this parameter was not included in the experiment. Regarding the 2002-2005 interval, we used public mac- roclimate data (annual mean temperature and precipitation) from the Hungarian Meteorological Service (www.met.hu), but from 2006 we used detailed daily macroclimate data from the Fert6-Hansag National Park Directorate (coordinates of data collection: 47°42'13.55"N, 17°10'40.43"E). We used the fol- lowing derived parameters as potential background variables: seasonal (winter: December-February, spring: March-May, sum- mer: June-August) annual and mean precipitation; seasonal and annual values of mean, minimum and maximum temperature; seasonal and annual values of mean, minimum and maximum humidity; mean of the monthly active and effective thermic amount (10°C). Orthoptera.—During the study period (2002-2017), sampling of the Orthoptera took place every June, July, August, and September. Z. KENYERES, G. TAKACS AND N. BAUER Fig. 1. Location map of the study area in Hungary. Samplings were carried out by sweep-netting within the 50x50 m sampling sites (altogether 448 samples). Species abundances were recorded by 300 sweeps per sampling site. Sweep-netted samples were identified to species level following Harz (1969, 1975). Sci- entific nomenclature follows Cigliano et al. (2017). The categories defined by Uvarov (1977) and Ingrisch and Kohler (1998) were used for classification of life forms (arbusti- cole: species found in habitats dominated by shrub-sized items; pratinicole: species found in grasslands of tall grass; graminicole: species found in grasslands of short grass; geophilic: species found in grasslands characterized by a high percentage of bare soil; pseu- do-psammophilic: species found in different types of grasslands but usually having a high density in sandy grasslands). The characterization of the climatic requirements of the spe- cies as thermophilic, moderately-thermophilic, mesophilic, mod- erately-hygrophilic, and hygrophilic were assigned based on Varga (1997), Racz (1998), and Ingrisch and Kohler (1998). Statistical analysis. —Samples collected in the same sampling sites in the same year were pooled (the number of pooled samples was 112). The pooled samples were used for calculating assemblage variables and statistical analyses. Shannon diversity, species num- ber, relative abundances of detected species, of life forms, and of species-groups with different climatic requirements were calcu- lated and used as Orthoptera variables in the statistical analyses. The mean values of Orthoptera response variables were calculated for comparison. JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) Z. KENYERES, G. TAKACS AND N. BAUER The Mann-Kendall trend test was used to evaluate temporal trends for both the Orthoptera variables and the macroclimate data. Generalized linear models (response variables: parameters of Or- thoptera showed statistically significant decreasing or increasing trends; predictor variables: macroclimate data) were performed. Canonical correspondence analysis based on Orthoptera species data and environmental parameters were also compiled. All statis- tical analyses were performed using the Past 3.14. software pack- age (Hammer et al. 2001). Results Orthoptera species.—Thirty-four Orthoptera species comprising 11,191 individuals were recorded at the sampling sites. The most prevalent species was Bicolorana bicolor with 1,779 individuals (16%), followed by Chorthippus brunneus with 1,530 individuals (14%), Roeseliana roeselii with 1,317 individuals (12%), Pseudo- chorthippus parallelus with 1,261 individuals (11%), Conocephalus fuscus with 1,084 individuals (10%), Chorthippus mollis with 1,042 individuals (9%), Chorthippus biguttulus with 742 individuals (7%), Euchorthippus declivus with 392 individuals (4%), Stenobothrus linea- tus with 368 individuals (3%), Euthystira brachyptera with 330 indi- viduals (3%), Chrysochraon dispar with 312 individuals (3%), and Mecostethus parapleurus with 194 individuals (2%) (see Appendix 1). Trends in Orthoptera parameters.—During the study a significant decreasing trend in the relative abundance of moderately-hygro- philic species (Fig. 2A), a non-significant increasing in the rela- tive abundance of thermophilic species (Fig. 2B), and a signifi- cant increasing trend in the relative abundance of graminicole species (Fig. 2C) were detected. Species richness also showed a significant increase (Fig. 2D). Of the species recorded, the relative abundance of mesophilic Chrysochraon dispar and thermophilic Euchorthippus declivus increased significantly (Fig. 3A, C), while the moderately-hygrophilic Roeseliana roeselii showed a significant decrease (Fig. 3B). Trends in macroclimate parameters.—A significant increasing trend was seen in the data of annual mean temperature (Fig. 4A); while annual precipitation showed no clear trend (Fig. 4B), mean monthly precipitation showed a non-significant decrease from 2006 to 2017 (Fig. 4C). Effects of macroclimate parameters.—Based on the results of the generalized linear models, annual mean temperature and annual minimum temperature were found to be significant predictors of the relative abundance of graminicole, pratinicole, and thermo- philic species, of the species number, and of the diversity of the as- semblages. The increase of annual mean temperature and annual minimum temperature were found to be significant predictors of a higher relative abundance of graminicole thermophilic species, a lower relative abundance of pratinicole species, and higher or- thopteran species diversity and species richness (Table 1). Also, the relative abundance of graminicole species and the diversity of the assemblages were positively related to mean temperature in summer. Besides, orthopteran diversity and the relative abun- dance of moderately thermophilic and thermophilic species were positively related to the means of the monthly active and effective thermic amount (10°C). Based on canonical correspondence analysis (CCA) ordina- tion, the relative abundance of graminicole species, thermophilic 189 0.25 A Moderately hygrophilic species 9° N 0.15 0.1 Relative abundance 0.05 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 B Thermophilic species Relative abundance C Graminicole species Relative abundance No. species 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 Fig. 2. Decreasing and increasing trends in the relative abundance of some orthopteran parameters (2002-2017). Temporal trends were evaluated by the Mann-Kendall trend test. species, and Euchorthippus declivus were positively correlated with mean temperature in summer (Fig. 5). The relative abundance of Roeseliana roeselii was negatively correlated with the latter pa- rameter. The relative abundance of pratinicole species and Cono- cephalus fuscus were positively correlated to high precipitation in spring (Fig. 5). Species richness was affected by annual mean and minimum temperature and also by monthly active and effective thermic amount (10°C) (Fig. 5). JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) A Chrysochraon dispar Relative abundance Relative abundance 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 20162017 C Euchorthippus declivus Relative abundance 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 Fig. 3. Significant decreasing and increasing trends in the relative abundance of some characteristic Orthoptera species in the stud- ied grasslands (2002-2017). Temporal trends were evaluated by the Mann-Kendall trend test. Discussion Global warming (Bale et al. 2002), which has a complex ef- fect on insect communities, is an existing phenomenon in Central Europe (Anders et al. 2014). It is well known that species-specific ecological requirements make the orthopterans particularly sensi- tive to climate change (Wessely et al. 2017). Between 2002 and 2017, the annual average temperature increased significantly in the humid grasslands of the Carpathian Basin that we studied. No trend was identified in annual rainfall. In the grasslands investi- gated, the relative abundance of moderately hygrophilic orthop- teran species decreased significantly, while the relative abundance of thermophilic species increased. These results suggest that species adapted to cooler climates are more sensitive to climate change (Butterfeld and Coulson 1997). It should be noted that the fluctuation in macrocli- mate data more than the gradual increase in mean temperature (Fig. 4A) may have a stronger negative impact on certain spe- cies in the long run (Easterling et al. 2000). During the study Z. KENYERES, G. TAKACS AND N. BAUER A Annual mean temperature S: 56, Z: 2.476, p=0.013 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 B Annual precipitation 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 C Mean monthly precipitation 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 Fig. 4. Trends in macroclimate parameters on the studied area (2002-2017). Temporal trends were evaluated by the Mann-Ken- dall trend test. period, the relative abundance of graminicole species also in- creased which might be related not only to the macroclimate changes, but also to the eutrophication of the grasslands (Lind et al. 2017). In our study, the effect of macroclimate change was also detectable at species level. The vertical and horizontal area expansion of thermophilic species as a result of global warming which has been described in several areas (Koéarek et al. 2008, Breitenmoser 2015, Kettermann and Fartmann 2018) was also observed in this study. Over the course of the study, some species were found in the sample areas which were not present in the early years, possibly due to their dry-hot climatic requirements (e.g., Calliptamus italicus, Euchorthippus declivus, Omocestus petraeus, and Platycleis affinis). Of the results related to the change in the local relative abundance of the species, a significant decrease in the relative abundance of the moderately-hygrophilic Roeseliana roeselii is remarkable, although the species is represented in the whole dataset with a large number of specimens. In recent decades, the northward expansion of Roeseliana roeselii and JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) Z. KENYERES, G. TAKACS AND N. BAUER 6 Confus 5 4 6) N X & 2 m-ther&ther i] pra © rain-spr# _ 191 Eucdec 4 3 =) - @ *i=-- cee a 3 4 5 s-r MO-ela To ata tem-m ® -1 Roeroe -2 Axis 1 Fig. 5. CCA ordination based on Orthoptera parameters and environmental parameters (Confus: Conocephalus fuscus; Eucdec: Euchor- thippus declivus; gra: graminicole species; mo-ata: mean of the monthly active thermic amount (10°C); mo-eta: mean of the monthly effective thermic amount (10°C); m-ther&ther: moderately-thermophilic and thermophilic species; pra: pratinicole species; rain-spr: rainfall in spring; Roeroe: Roeseliana roeselii; s-r: species richness; tem-m: annual mean temperature; tem-min: annual minimum tem- perature; tem-sum: mean temperature in summer; the: thermophilic species). Table 1. Significant results of GLM testing of macroclimate effects on Orthoptera assemblages (* P<0.05; ** P<0.01; *** P<0.001; Conocephalus fuscus, also occurring in the humid grasslands of Central Europe, was recorded in northern and northwestern data for the mean of the monthly active and effective thermic Europe (Kleukers et al. 1996, Fartmann 2004, Wissmann et amount and precipitation were log transformed). Response variable Predictor variable Estimate St.err. p Roeseliana roeselii Mean temp. in summer -16.634 6.146 ** Euchorthippus declivus Mean temp. in summer 16.701 6.061 ** Graminicole species Mean temp. in summer 14.194 5.102 ** Annual mean temp. 14.467 6.715 ** Annual min. temp. 24.513 8.943 ** Pratinicole species Annual mean temp. -18.041 7.436 * Annual min. temp. -28.779 8.940 ** Diversity Annual mean temp. 163789 53733 7 ** Annual min. temp. 19.768 8.605 * Mean of the monthly active O:698. 03255. ** thermic amount (10°C) Mean of the monthly effective 0.932 0.344 ** thermic amount (10°C) Mean temp. in summer T2809 4:75 a* Precipitation in spring -1.844 0.867 * Species number Precipitation in spring -0.052 0.025 * Annual mean temp. 0.508 0.158 ** Thermophilic species Annual mean temp. T6555 6299. ** Annual min. temp. 22.859 8.486 ** Moderately Mean of the monthly active 0.302. <0y108 ** thermophilic and thermic amount (10°C) thermophilic species Mean of the monthly effective 0.411 0.145 ** thermic amount (10°C) al. 2009). On the one hand, this latter phenomenon confirms the fact that the impact of global warming is more intense in areas closer to the poles (Bale et al. 2002). It is not only recent experiences, but also the results explored in the case of climate changes in earlier geologic epochs (Coope 1970), which indicate that insects do not adapt to the changed conditions in the case of climate change, but follow it by changing their areas. In conclusion, in the Central European humid grasslands studied, the increase in the annual mean temperature most in- tensively affected negatively the relative abundance of moder- ately hygrophilic orthopteran species. The expansion of thermo- philic species could also be observed within the study area (they occupied habitats that were not previously suitable for them). The number of species and diversity of the local orthopteran assemblages was higher as the annual average temperature in- creased. From a conservation point of view, this is not necessarily a positive fact. The orthopteran assemblages of humid grasslands in Central Europe are normally characterized by low diversity, due to the dominance of some hygrophilic and moderately hy- grophilic species. According to our results, the conservation of the main characteristics of the Central European humid grass- lands, under global warming, can only be ensured by adequate land management. JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) 192 Suggestions for adequate land management Due to the causes of global warming, the following sugges- tions for adequate local land management of humid grasslands in Central Europe are suggested: (1) Spatial mosaic grassland man- agement by changing the patches abandoned throughout the sea- son every year. (2) Exclusion of grazing or, at the most, only in an extensive manner during autumn. (3) Abandonment of mowing in extremely dry years with a warm spring (except for patches af- fected by invasive plant species). The above options can result in a mitigating effect of the denser vegetation (Cox and Moore 1980, Schoonhoven et al. 1998), which regulates the microclimate of the humid grasslands. Acknowledgements The authors would like to express their gratitude to the review- ers for their remarks. We are very grateful to Maria Marta Cigliano, Subject Editor of JOR and to Nancy Morris, Editorial Assistant of JOR, for their work with our manuscript. References Anders I, Stagl J, Auer I, Pavlik D (2014) Climate change in Central and Eastern Europe. In: Rannow S, Neubert M (Eds) Managing Protected Areas in Central and Eastern Europe Under Climate Change, Advanc- es in Global Change Research 58, Springer Science+Business Media Dordrecht, 17-30. https://doi.org/10.1007/978-94-007-7960-0_2 Bale JS, Masters GJ, Hodkinson ID, Awmack C, Bezemer TM, Brown VK, Butterfield J, Buse A, Coulson JC, Farrar J, Good JEG, Harrington R, Hartley S, Jones TH, Lindroth RL, Press MC, Symrnioudis I, Watt AD, Whittaker JB (2002) Herbivory in global climate change research: direct effects of rising temperature on insect herbivores. Global Change Biology 8: 1-16. https://doi.org/10.1046/j.1365- 2486.2002.00451.x Breitenmoser S (2015) Expansion de l’'Oedipode automnale Aiolopus stre- pens (Latreille, 1804) (Orthoptera, Acrididae) dans l'Ouest vaudois. Entomo Helvetica 8: 133-139. Buri P, Arlettaz R, Humbert JY (2013) Delaying mowing and leaving un- cut refuges boosts orthopterans in extensively managed meadows: Evidence drawn from field-scale experimentation. Agriculture, Eco- systems and Environment 181: 22-30. https://doi.org/10.1016/j. agee.2013.09.003 Burton JF (2001) The apparent influence of climatic change on recent changes of range by European insects (Lepidoptera, Orthoptera). In: Reemer M, Helsdingen PJ, Kleukers R MJC (Eds) Proceedings of the 13" International Colloquium of the European Invertebrate Survey, Leiden, 2-5 September 2001, 13-22. Butterfeld J, Coulson JC (1997) Terrestrial invertebrates and climate change: physiological and life-cycle adaptations. In: Huntley B, Cramer W, Morgan AV, Prentice HC, Allen JRM (Eds) Past and Future Rapid Environmental Change, Springer, Berlin, 401-412. https://doi. org/10.1007/978-3-642-60599-4_31 Cigliano MM, Braun H, Eades DC, Otte D (2017) Orthoptera Species File. Version 5.0/5.0. http://Orthoptera.SpeciesFile.org Coope GR (1970) Interpretations of Quaternary insect fossils. Annual Review of Entomology15: 97-120. https://doi.org/10.1146/annurev. en.15.010170.000525 Cox CB, Moore PD (1980) Biogeography, an Ecological and Evolutionary Approach. Blackwell, Oxford. Dovényi Z (Ed.) (2010) Magyarorszag kistajainak katasztere. [Cadastre of the Hungarian microregions] MTA Foldrajztudomanyi Kutatdointézet, Budapest. [In Hungarian] Dreux P (1962) Recherches écologiques et biogéographies sur les Or- thoptéres des Alpes Francaises. Théses presentées a la Faculté de Sci- ences de L’Université de Paris. Masson & C* Editeurs, Paris. Z. KENYERES, G. TAKACS AND N. BAUER Easterling DR, Meehl GA, Parmesan C, Changnon SA, Karl TR, Mearns LO (2000) Climate extremes: observations, modeling, and im- pacts. Science 289: 2068-2074. https://doi.org/10.1126/sci- ence.289.5487.2068 Fartmann T (2004) Hydrochorie und warme Jahre - sind das die Griinde fiir die Ausbreitung der Langfliigeligen Schwertschrecke (Conocephalus fuscus) in Ostbrandenburg? Articulata 19: 75-90. Fielding DJ, Defoliart LS (2010) Embryonic developmental rates of north- ern grasshoppers (Orthoptera: Acrididae): Implications for climate change and habitat management. Environmental Entomology 39: 1643-1651. https://doi.org/10.1603/EN09356 Gardiner T (2009) Macropterism of Roesel’s bushcricket Metrioptera ro- eselii in relation to climate change and landscape structure in east- ern England. Journal of Orthoptera Research 18: 95-102. https://doi. org/10.1665/034.018.0110 Gardiner T, Hassall M (2009) Does micro-climate affect grasshopper populations after cutting of hay in improved grassland? Journal of Insect Conservation 13: 97-102. https://doi.org/10.1007/s10841-007- 9129-y Guido M, Chemini C (2000) Response of Orthoptera assemblage com- position to land-use in the southern Alps of Italy. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 73: 353-367. Hammer @, Harper DAT, Ryan PD (2001) PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4: 1-9. Harz K (1969) Die Orthopteren Europas - The Orthoptera of Europe I. Series Ent. 5, The Hague. https://doi.org/10.1007/978-94-017-2511-8 Harz K (1975) Die Orthopteren Europas / The Orthoptera of Europe II. Se- ries Ent. 11, The Hague. https://doi.org/10.1007/978-94-010-1947-7 HoluSa J, Koéarek P, Marhoul P (2007) First sightings of Ruspolia nitidula (Orthoptera: Tettigoniidae) and Mecostethus parapleurus (Orthoptera: Acrididae) after fifty years in the Czech Republic. Articulata 22: 47-51. Ingrisch S, Koéhler G (1998) Die Heuschrecken Mitteleuropas. Die Neue Brehm-Biicherei Bd. 629, Westarp Wissenschaften, Magdeburg. Kenyeres Z, Cservenka J (2014) Effects of climate change and vari- ous grassland management practices on grasshopper (Orthop- tera) assemblages. Advances in Ecology 2014: e601813. https://doi. org/10.1155/2014/601813 Kenyeres Z, Takacs G, Bauer N (2018) Effects of climatic factors on yearly population sizes of Isophya costata (Orthoptera). North-Western Jour- nal of Zoology 14: 13-16. Kettermann M, Fartmann T (2018) Auswirkungen des globalen Wandels auf Heuschrecken - Besiedlung von Steinbriichen der Westfalischen Bucht (NW-Deutschland) durch die Blaufliigelige Sandschrecke. Naturschutz und Landschaftsplanung 50: 23-29. Kiritani K (2006) Predicting impacts of global warming on population dy- namics and distribution of arthropods in Japan. Population Ecology 48: 5-12. https://doi.org/10.1007/s10144-005-0225-0 Kleukers RMJC, Decleer K, Haes ECM, Kolshorn P, Thomas B (1996) The recent expansion of Conocephalus discolor (Thunberg) (Orthoptera: Tettigoniidae) in western Europe. Entomologist’s Gazette 47: 37-49. Koéarek P, HoluSa J, VIk R, Marhoul P, Zuna-Kratky T (2008) Recent expan- sions of bush-crickets Phaneroptera falcata and Phaneroptera nana (Or- thoptera: Tettigoniidae) in the Czech Republic. Articulata 23: 67-75. Laws AN, Belovsky GE (2010) How will species respond to climate change? Examining the effects of temperature and population density on an herbivorous insect. Environmental Entomology 39: 312-319. https:// doi.org/10.1603/EN09294 Lightfoot DC (2006) Climate Change and Rangeland Insects. Beyond Boxes and Arrows, A Workshop on Climate Change and Rangeland Management, Jan. 25-26, 2006, San Carlos, Arizona. Lind EM, La Pierre KJ, Seabloom EW, Alberti J, Iribarne O, Firn J, Gruner DS, Kay AD, Pascal J, Wright JP, Yang L, Borer ET (2017) Increased grassland arthropod production with mammalian herbivory and eu- trophication: a test of mediation pathways. Ecology 98: 3022-3033. https://doi.org/10.1002/ecy.2029 Menéndez R (2007) How are insects responding to global warming? Tijd- schrift voor Entomologie 150: 355-365. JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2) Z. KENYERES, G. TAKACS AND N. BAUER O'Neill BE Zangerl AR, Delucia EH, Berenbaum MR (2008) Longevity and fecundity of Japanese beetle (Popillia japonica) on foliage grown under elevated carbon dioxide. Environmental Entomology 37: 601-607. https://doi.org/10.1603/0046-225X(2008)37[601:LAFOJ B]2.0.CO;2 Racz I (1998) Biogeographical survey of the Orthoptera fauna in central part of the Carpathian Basin (Hungary): Fauna types and community types. Articulata 13: 53-69. Racz I, Szilagyi G, Molnar A (1994) Saskajaras a Hortobagyon. [Locust gradation on the Hortobagy.] II. Kelet-Magyarorszagi erdé, vad- és halgazdalkodasi, természetvédelmi konferencia, Debrecen, 1994. XI. 5-6., 372-380. Ryrholm N (2001) Global warming and the change of butterfly distri- butions: a new opportunity for species diversity or a severe threat (Lepidoptera)? In: Reemer M, Helsdingen PJ, Kleukers RMJC (Eds) Proceedings of the 13 International Colloquium of the European Invertebrate Survey, Leiden, 2-5 September 2001, 7-12. Schoonhoven LM, Jermy T, van Loon JJA (1998) Insect-Plant Biology: From Physiology to Evolution. Chapman & Hall, London. https:// doi.org/10.1007/978-1-4899-3200-6 Squitier JM, Capinera JL (2002) Habitat associations of Florida grass- hoppers (Orthoptera: Acrididae). Florida Entomologist 85: 235- 244. https://doi.org/10.1653/0015-4040(2002)085[0235:HAOFG O]2.0;CO-2 Appendix 1 193 Szabo B, Nagy B, Szentkiralyi F (2014) Egyenesszarnyt rovarfajok (Or- thoptera) larvakeléskezdeteinekhosszu tavu trendjei (1958-2009) az éghajlatvaltozds kévetkezményeként Magyarorszagon. Allattani K6- zlemények 99: 45-59. Uvarov BP (1931) Insects and climate. Transactions of the Entomological Society. London, 79: 1-247. Uvarov BP (1977) Grasshoppers and Locusts, Vol. II. Centre for Overseas Pest Research, London. Varga Z (1997) Trockenrasen im pannonischen Raum: Zusammenhang der physiognomischen Struktur und der florischtischen Komposition mit den Insektenz6nosen. Phytocoenologia 27: 509-571. https://doi. org/10.1127/phyto/27/1997/509 Wessely J, Hiilber K, Gattringer A, Kuttner M, Moser D, Rabitsch W, Schin- dler S, Dullinger S, Ess] F (2017) Habitat-based conservation strategies cannot compensate for climate-change-induced range loss. Nature Climate Change 7: 823-827. https://doi.org/10.1038/nclimate3414 Wingerden WKRE, Kreveld AR, Bongers W (1992) Analysis of species com- position and abundance of grasshoppers (Orth., Acrididae) in natural and fertilized grasslands. Journal of Applied Entomology 113: 138- 152. https://doi.org/10.1111/j.1439-0418.1992.tb00647.x Wissmann J, Schielzeth H, Fartmann T (2009) Landscape-scale expansion of Roesel’s bush-cricket Metrioptera roeselii at the north-western range limit in Central Europe (Orthoptera: Tettigoniidae). Entomologia Generalis 31: 317-326. https://doi.org/10.1127/entom.gen/31/2008/317 Species composition and abundance of the samples pooled per year (LF: life form; EF: ecotype form; arbu: arbusticole; geo: geophilic; gra: graminicole; pra: pratinicole; psps: pseudo-psammophilic; hyg: hygrophilic; mes: mesophilic; m-hyg: moderately-hygrophilic; m-ther: moderately-thermophilic; ther: thermophilic). Taxon LF EF 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 Bicolorana bicolor (Philippi, 1830) pra m-ther 34 159 138 112 157 94 94 71 138 192 128 31 24 158 147 102 Chorthippus brunneus (Thunberg, 1815) pra m-ther 21 130 24 45 50 100 91 96 336 72 206 47 112 #104 62 34 Roeseliana roeselii (Hagenbach, 1822) pra m-hyg 46 176 37 44 60 69 70 126 96 174 82 67 64 102 70 34 Pseudochorthippus parallelus (Zetterstedt, 1821) pra mes 45 8 17 104 58 58 51 144 106 116 120 122 114 152 28 Conocephalus fuscus (Fabricius, 1793) pra hyg 21 60 26 29 24 50 51 52 #152 134 28 89 134 130 58 46 Chorthippus mollis (Charpentier, 1825) pra mes 6 56 0 72 50 64 102 236 31 108 75 72 66 64 £440 Chorthippus biguttulus (Linnaeus, 1758) pra m-ther 4 61 O 22 59 60 58 47 64 96 91 78 49 20 33 Euchorthippus declivus (Brisout de Barneville, 1848) gra ther O 20 144 O 28 28 20 6 4 42 52 14 36 64 46 Stenobothrus lineatus (Panzer, 1796) pra m-ther 2 22 27 26 O 21 22 6 12 47 #%24 #%38 60 15 40 6 Euthystira brachyptera (Ocskay, 1826) pra mes ll 42 O O” "255 "20" “19. 52s “O07 “SO™ “227 F20- 30. 2344 5 0 Chrysochraon dispar (Germar, 1834) pra m-hyg 5 24 3 2 9 6 5 4 13 46 27 18 36 68 26 20 Mecostethus parapleurus (Hagenbach, 1822) pra hyg Zz 0 0 0 0 0 0 0 0 0 5 0 16 47 64 #55 Chorthippus dorsatus (Zetterstedt, 1821) pra mes 12 8 0 8 0 0 0 14 +#O 0 8.9) 1S 28= 925 « =8 12 Decticus verrucivorus (Linnaeus, 1785) pra mes 6 0 0 0 2 3 2 ) Bx, Le BLOF A? 8 9 10 Tettigonia viridissima Linnaeus, 1758 arbu= mes 0 0 8 12 14 12 U 6 2 | 0 0 8 5 4 0 Calliptamus italicus (Linnaeus, 1758) gra ther 0O 0 0 0 Of Thnsitg 39 0 0 2 6 17 4 0 11 Conocephalus dorsalis (Latreille, 1804) pra hyg 0 0 6 2 0 0 0 O> 12% eile Ae pS) SiGe 22 1 0 Omocestus petraeus (Brisout de Barneville, 1856) gra ther 0O 0 fi 3 0 0 G= 113;' £0 Ose se &6 0 Oo Ul 0 Phaneroptera falcata (Poda, 1761) arbu. ther 0 6 0 0 4 3 1 2A 10a, “Oe tO 0 0 0 0 Omocestus haemorrhoidalis (Charpentier, 1825) pra ther O- 21 0 6 0 0 0 0 0 9 5 3 0 2 7 3 Pseudochorthippus montanus (Charpentier, 1825) pra hyg 13 21 0 0 0 0 0 0 0 0 0 0 0 0 0 8 Tetrix subulata (Linnaeus, 1758) geo hyg 0 0 0 0 0 0 0 0 31 #O 3 0 0 0 0 0 Ruspolia nitidula (Scopoli, 1786) pra m-hyg 0 0 0 0 0 0 0 3 0 2 0 11 0 0 =) 5 Chorthippus oschei Helversen, 1986 pra mes 10 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Omocestus rufipes (Zetterstedt, 1821) pra mes 0 0 0 0 0 5 0 0 0 0 5 0 0 0 6 0 Chorthippus dichrous (Eversman, 1859) pra mes 0 0 0 0 0 0 0 0 0 0 0 0 6 0 4 2 Tetrix tenuicornis (Schalberg, 1893) pra (ther; “0; «fe -0 0 0 0 0 0 0 0 0 0 0 0 0 0 Oecanthus pellucens (Scopoli, 1763) pra m-ther 0 0 3 5 0 0 0 0 0 0 0 0 0 0 0 0 Leptophyes albovittata (Kollar, 1833) arbu ther 0 0 0 0 3 0 0 0 0 0 0 0 1 2 0 0 Stethophyma grossum (Linnaeus, 1758) pra hyg 3 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Aiolopus thalassinus (Fabricius, 1781) gra m-ther 0 0 0 0 0 0 0 0 0 0 0 0 6 0 0 0 Stenobothrus nigromaculatus (Herrich-Schaffer, 1840) gra ther 0 0 0 0 0 0 0 0 0 0 2) 0 0 0 0 0 Platycleis grisea (Fabricius, 1781) pra ther 0 0 0 0 0 0 0 0 0 0 0 0 3 0 2 0 Platycleis affinis Fieber, 1853 psps ther 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 1 JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(2)