Research Article Journal of Orthoptera Research 2021, 30(1): 35-41 Conservation possibilities of lsophya costata (Orthoptera: Tettigoniidae: Phaneropterinae) based on frequency, population size, and habitats ZOLTAN KENnyeRES!, NorBERT BAUER? 1 Acrida Conservational Research L.P., Tapolca, Hungary. 2 Department of Botany, Hungarian Natural History Museum, Budapest, Hungary. Corresponding author: Z. Kenyeres (kenyeres@acridabt.hu) Academic editor: Alina Avanesyan | Received 2 October 2020 | Accepted 5 January 2021 | Published 26 April 2021 http://zoobank.org/93247656-GE5F-44BD-91 70-69360F1 6B3AE Citation: Kenyeres Z, Bauer N (2021) Conservation possibilities of Isophya costata (Orthoptera: Tettigoniidae: Phaneropterinae) based on frequency, population size, and habitats. Journal of Orthoptera Research 30(1): 35-41. https://doi.org/10.3897/jor.30.59262 Abstract Isophya costata Brunner von Wattenwyl, 1878, commonly called the Keeled Plump Bush-cricket, is an endemic Natura 2000 species in the Car- pathian Basin and is included in the IUCN Red List of Threatened species. We used extensive data collection from Hungary retrieved between 2004 and 2019 from 700 sampling sites spread over an area of 12,700 km? to examine the occurrence of the species in different regions in grasslands of similar structure but different origin, naturalness, and character. The results confirmed that I. costata currently occurs with the highest num- ber of populations and highest density in regularly mowed, mesophilic hayfields rich in dicotyledonous plants (Arrhenatheretalia). The species also appears in smaller numbers in grasslands adjacent to hayfields, such as wetland meadows (Molinion coeruleae), marsh meadows (Deschamp- sion caespitosae, Alopecurenion pratensis), and edge habitats dominated by herbaceous plants. However, the results show that the extension of these habitats has a negatively significant correlation with both the occurrence of the species and its density. Isophya costata occurs in steppe meadows much less frequently than in mesophilic hayfields. The species is endemic to the Pannonian Steppe, and the key to their conservation is by main- taining stocks of hayfields in the species’ area of distribution. According to this study, overseeding of mowed grasslands leads to the decline of the species. To preserve I. costata, it is necessary to eliminate trampling in its areas of occurrence (prohibition of grazing) and encourage late-season mowing adapted to the phenology of the species (not as early as mid-July) or, if this is not feasible, mosaic-type treatment leaving unmown patches (e.g., 1/3 of the plot). Keywords Carpathian Basin, endemic species, grassland character, hayfields, mow- ing, overseeding Introduction Isophya costata Brunner von Wattenwyl, 1878 is an endemic, postglacial-steppe relict species (Varga 1995) found in the Car- pathian Basin. It is a highly protected species of community im- portance (Natura 2000) in Hungary, and as such, it is included in the IUCN Red List of Threatened species (Chobanov et al. 2016). The species has long been the focus of research because of its hab- itat loss due to cultivation, abandonment of mowing, etc. (Nagy 1974) that has led to the Hungarian Red Book classifying it as an endangered species (Rakonczay 1989). I. costata is mostly found in Hungary (Nagy and Racz 2014). Based on previous studies, I. costata may have been a characteristic species of loess meadows and closed-steppe meadows rich in dicotyledonous plants that later adapted to hayfields rich in dicotyledonous species, simi- lar to their previous habitats but formed under anthropogenic influence. These hayfields have become the primary habitat of I. costata (Kenyeres et al. 2004, Bauer and Kenyeres 2006, Kenyeres et al. 2009). Thanks to recent systematic research new occurrence data have been revealed - not only in Hungary, but also in Slovakia (Nuhlickova et al. 2017), Serbia (Sz6vényi and Szekeres 2011, Ivkovi¢ and Horvat 2020), Austria (Zuna-Kratky et al. 2017), and Romania (lorgu et al. 2008). In the present study, the data collection covers about a quar- ter of the territory of Hungary and took place between 2004 and 2019. We sought to answer the following questions: (1) What re- gional differences can be detected in the frequency of the occur- rence of I. costata among the regions within the study area? (2) Are there any differences in terms of the frequency of occurrence of the species among its potential habitats with a similar structure but different species composition, origin, and landscape history? (3) Can the effects of the condition and use of the studied grass- lands (regularly mowed, overseeded, shrubbing, humid, semi-dry, dry, etc.) on the occurrence of the species be revealed? (4) Does the density of the species correlate with the local surface cover of the studied habitat types? Methods Between 2004 and 2019, we examined the presence/absence, density, and habitat requirements of I. costata (Figs 1, 2) at 700 sites (12,700 km?) in northwestern Hungary. The study area was divided into 12 regions (Fig. 3) according to landscape and bioge- ography. The CORINE Landcover habitat structure of the regions and the distribution of the studied grasslands among habitat types are shown in Fig. 4 and Table 1. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) Z. KENYERES AND N. BAUER Fig. 2. Known distribution of Isophya costata in the Carpathian Basin at the level of ETRS quadrates (circle: presence data of the species; red circle: presence data of this study; triangle: absence data of this study). JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) Z. KENYERES AND N. BAUER 37 Fig. 3. Sub-areas of the study area (A, B.) (1: Alpokalja, 2: Gyér Basin, 3: Komarom-Esztergom Plain, 4: Pannonhalma Region, 5: Sopron- Vas Plain, 6: Marcal Basin, 7: Kemeneshat, 8: Bakonyalja, 9: Southern Bakony, 10: Balaton Uplands, 11: Eastern Bakony, 12: Mezéféld), sampling sites (red points), and the most typical studied habitat-types: C. Typical hayfield rich in dicotyledonous plant species; D. Overseeded hayfield poor in dicotyledonous plant species; E. Loess steppe; F. Slope steppe. Table 1. Extent (mean+SE) and habitat types of the grasslands containing the study quadrates (N=700). Habitat type Red beds Rich fens Mesotrophic wet meadows Hayfields Overseeded hayfields Calcareous rocky steppes Slope steppes on stony soils Closed forest steppe meadows Closed steppes on loess Uncharacteristic mesic grasslands Uncharacteristic dry and semi-dry grasslands Range in hectare (mean+SE) 3.6440.64 14.32+3.52 6.7440.67 6.40+0.97 13.2043.41 4.03+0.45 3.59+0.62 2.29+40.47 0.68+0.27 2.864+0.42 5.20+0.86 N Sweep-netting, direct observation, and acoustic detection were performed at the studied 10x10 m quadrates for the dura- tion (30 minutes). All detected individuals were recorded. Based on the number of individuals registered in the study quadrates, we determined the presence/absence data and the density of the species per square meter related to the habitat patch concerned. During sampling, we recorded the type of the habitat according to the Hungarian General National Habitat Classification System (A- NER): red bed, rich fen, mesotrophic wet meadow, hayfield/over- seeded hayfield, calcareous rocky steppe, slope steppe on stony soils, closed forest steppe meadow, closed steppe on loess, unchar- acteristic mesic, and dry and semi-dry grassland. We also recorded the naturalness (natural, pseudo-natural: anthropogenic hayfields with good naturalness, shrubby, disturbed, and overseeded) and microclimatic conditions (humid: directly affected by water and JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) 38 Regions of the studied area Mezéfold Ji i Komarom-Esztergom Plain Jj fi Gy6r Basin |i | Marcal Basin il Sopron-Vas Plain Kemeneshat SI (i cotoryal2 I Pannonhalma Region i ij Balaton Uplands ee | | | Apotal: [iT castern Sokory TS I Southern Bakony I i Z. KENYERES AND N. BAUER CORINE habitat types mm Broad-leaved forest Complex cultivation patterns Coniferous forest Construction sites Continuous urban fabric Discontinuous urban fabric Dump sites Fruit trees and berry plantations Green urban areas Industrial or commercial units Inland marshes Land principally occupied by agriculture, with significant areas of natural vegetation Mineral extraction sites Mixed forest Natural grasslands i | Non-irrigated arable land Pastures Peat bogs Road and rail networks and associated land Sparsely vegetated areas Sport and leisure facilities Transitional woodland-scrub Vineyards Water bodies Water courses Fig. 4. Relative frequency of CORINE habitat types in the studied sub-areas. characterized by water supply for at least a part of the year; meso- philic: closed, semi-dry grasslands; dry: xerophilous grasslands with open soil surfaces). We determined the size of the habitat type corresponding to the quadrates studied based on data collected in the field by hand- held computer (Trimble Juno3B) with the use of QGIS 2.16 soft- ware (QGIS Development Team 2016). A basic database containing the records of all sites (N = 700) was used for analyses, but a database containing the relative fre- quency (rel. freq.) of the recorded variables at the ETRS quadratic level was also generated (N = 78). Furthermore, we determined the sub-areal (12 regions) indicators for the presence/absence of the species (Ns: number of sampling sites with presence/absence of I. costata, and Rf: relative frequency of presence of I. costata). The relationships between the presence/absence of the species and the presence/absence of the habitat types and habitat characteristics studied were examined by Generalized Linear Models (GLM) with binomial distribution. Division of the habitats with the presence of I. costata among habitat characteristics recorded (habitat types, naturalness, structure, and microclimate) were described by bar charts. The paired t-test was used to evaluate statistical differences among the variables. In order to analyse the relationship between the density of I. costata and environmental data at the European Terrestrial Reference System (ETRS) quadrate scale (in which the dependent variable was the mean density of I. costata; the habitat variables were mean of the patch size; and the relative frequencies (rel. freq.) of the recorded habitat types were categorized accord- ing to in the Habitat Classification System (A-NER), to their natu- ralness and to their microclimatic conditions; see above), single tests of GLM (Poisson distribution) were used. Overdispersion was not detected in the variables. Statistical procedures were per- formed using PAST 2.16 (Hammer et al. 2001) and CANOCO 4.5 (Braak and Smilauer 2002) software packages. Results Regional differences.—Of the 700 study sites, the presence of the species was detected at 280 locations. The frequency of the species showed significant regional dif- ferences (Fig. 5). I. costata proved to be particularly frequent (rel. freq. of positive cases: 0.63) in the Balaton Uplands sub- area (267 study sites), presumably because of the richness of potential habitats in the region. Among the studied quadrates, the proportion of hayfield habitat type was high (rel. freq.: 0.42), but slope steppes on stony soils (rel. freq.: 0.15), rich fens (rel. freq.: 0.15), and uncharacteristic dry and semi-dry grasslands (rel. freq.: 0.15) habitat types were also significantly present. Of the grasslands studied, the proportion of pseudo- natural (rel. freq.: 0.55) and mesophilic (rel. freq.: 0.59) patch- es was very high. Roughly a third of the 112 study sites showing the presence of the species were in the Eastern Bakony region (rel. freq.: 0.29). In this region, most of the grasslands studied were calcareous rocky steppes habitat type (rel. freq.: 0.80) with a significant share of uncharacteristic dry and semi-dry grassland habitat types (rel. freq.: 0.15). Most sites included in the study were edaphic dry grasslands (rel. freq.: 0.76) with a dry microclimatic character (rel. freq.: 0.98). Also, we found the presence of the species in about a third of the 72 study sites in the Marcal Basin region (rel. freq.: 0.32). The majority of the examined patches (rel. freq.: 0.51) belonged to the rich fens habitat type, with a significant number of hayfields and overseeded hayfields (rel. freq.: 0.32 and 0.10). In terms of char- acter, the grasslands studied were pseudo-natural (rel. freq.: 0.81), humid (rel. freq.: 0.58), or semi-dry (rel. freq.: 0.32). Among the regions with less than 10% of the total sam- ple, I. costata was found in about a third of the study sites. This JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) Z. KENYERES AND N. BAUER fi presence [§ absence Sampling sites GyB B SV K KE PR 0.00 0.00 0.00 BU EB MB_~ SB M Al 063 0.29 0.32 030 033 039 015 020 0.11 Relative frequency Fig. 5. Presence or absence and relative frequency of presence of Isophya costata in the studied regions (BU: Balaton Uplands; EB: Eastern Bakony; MB: Marcal Basin; SB: Southern Bakony; M: Mez6foéld; Al: Alpokalja; GyB: Gy6ér Basin; B: Bakonyalja; SV: Sop- ron-Vas Plain; K: Kemeneshat; KE: Komarom-Esztergom Plain; PR: Pannonhalma Region). includes Southern Bakony (no. of samples: 57, rel. freq.: 0.30), Mez6féld (no. of samples: 36, rel. freq.: 0.33), and in the Alpokal- ja region (no. of samples: 23, rel. freq.: 0.39). From a microcli- matic point of view, the grasslands under review were dry (rel. freq.: 0.68) and semi-dry (rel. freq.: 0.30) in all three regions. In the sampling quadrates of the Southern Bakony, uncharacteristic dry and semi-dry grasslands (rel. freq.: 0.28), hayfields (rel. freq.: 0.26), rich fens (rel. freq.: 0.28), and slope steppes on stony soil (rel. freq.: 0.26) habitats occurred in a roughly balanced propor- tion. Among the grasslands examined, pseudo-natural patches (rel. freq.: 0.40) were dominant, but the proportion of disturbed (rel. freq.: 0.30) and shrubby (rel. freq.: 0.25) habitats were also high. Closed forest-steppe meadows (rel. freq.: 0.31) and closed steppes on loess habitat types (rel. freq.: 0.31) were dominant in the studied grasslands of the Mezéfdld region, but hayfields (rel. freq.: 0.26) also occurred (rel. freq.: 0.17). Among the grasslands examined, shrubby (rel. freq.: 0.33), pseudo-natural (rel. freq.: 0.22), disturbed (rel. freq.: 0.22), and edaphic patches (rel. freq.: 0.22) were present in a balanced proportion. In the Alpokalja re- gion, in addition to the dominant hayfields (rel. freq.: 0.26), rich fens (rel. freq.: 0.13), and overseeded hayfields (rel. freq.: 0.13) habitat types also occurred. Of the grasslands studied, pseudo-nat- ural patches (rel. freq.: 0.61) were dominant, but the proportion of overseeded (rel. freq.: 0.13) habitats was also high. In the other sub-areas, either the species was not detected or the frequency of its occurrence was very low. Of these, the Bakony- alja should be highlighted, where most of the grasslands exam- ined belonged to the pseudo-natural, but rather dry, microclimate occurrences of the hayfield (rel. freq.: 0.80) habitat type. However, the number of samples (corresponding to the occurrence of poten- tial I. costata habitats) was low in this region. The other sub-areas can be divided into two groups. The examined patches of the Gy6r Basin (N = 52), Kemeneshat (N = 29) and Sopron-Vas Plain (N = 37) predominantly belonged to overseeded hayfield habitat type (rel. freq.: 0.55-0.68), the character of which was determined by overseeding (rel. freq.: 0.55-0.70) and dry stock climate (rel. freq.: 0.59-0.83). The dominant habitat types in the Komarom-Eszter- 39 gom Plain (N = 8) and Pannonhalma Hills (N = 2) regions can be classified as diverse: rich fens (rel. freq.: 0.38-0.50), overseeded hayfields (rel. freq.: 0.00-0.38), uncharacteristic dry and semi-dry grasslands (rel. freq.: 0.00-0.25), and closed steppes on loess (rel. freq.: 0.00-0.50). In terms of character, both humid (rel. freq.: 0.38-0.50) and dry (rel. freq.: 0.50-0.63) grasslands were deter- mined to be in high proportion in these regions, and also in terms of naturalness, several types occurred: pseudo-natural (rel. freq.: 0.13-0.50), shrubbing (rel. freq.: 0.00-0.50), disturbed (rel. freq.: 0.00-0.50) and overseeded (rel. freq.: 0.00-0.70). Habitat requirements.—Based on the data as a whole, more than half (51.1%) of the sampled quadrates with the presence of the species were hayfields rich in dicotyledonous plant species (Fig. 6). Additional occurrences were distributed among nine dif- ferent habitat types, of which the proportion of marsh meadows (12.9%) and calcareous rocky steppes (12.5%) were significant. In terms of habitat character (Fig. 6), most occurrences were found in regularly mowed hayfields (64.6%) showing good natu- ralness or edaphic steppes (14.6%). The proportion of disturbed or shrubby grasslands among positive samples was 12.1% and 7.14%, respectively, and the presence of I. costata was rarely ob- served on overseeded hayfields (1.4%). In terms of microclimate, 66.4% of the grasslands where the species occurred were semi- arid, 27.9% were dry, and only 5.7% were humid. GLM (Table 2) of the data on all study quadrates showed a sig- nificant positive correlation between the presence of I. costata and the presence of hayfields rich in dicotyledonous plant species. Sig- nificant negative relationships were seen between the occurrence of I. costata and the presence of overseeded hayfields, mesotrophic wet meadows, and uncharacteristic dry and semi-dry grasslands. Density.—The density of I. costata in the study quadrates was 0.24 individuals/m? (mean + 0.11). There was no significant correlation between species density and size of the habitat patch (p=0.609, R2=0.00037). GLM showed (a) a significant positive correlation (p=0.004, R’=0.301) between the mean density of I. costata at the ETRS quadrate level and the frequency of hayfields rich in dicotyledon- ous plant species (Fig. 7) and (b) a significant negative correlation (p<0.001, R?=0.462) between the mean density of I. costata at the ETRS quadrate level and the frequency of grasslands characterized by humid microclimate (Fig. 7). Discussion Our results show that, although the Keeled Plump Bush-crick- et can be found in several areas on steppe meadows, loess grass- lands, and other grassland types (weedy humid grasslands, marsh meadows, etc.), their highest density populations live on regularly mowed hayfields rich in dicotyledonous plant species, confirm- ing our earlier statement (Bauer and Kenyeres 2006). According to this study, the species rarely occurs in overseeded hayfields or, if it does, only in very small numbers. This may be related to the fact that overseeding is usually done with grass species, leading to the decline of dicotyledonous plant species cover (Blackmore and Goulson 2014). Also, the soil surface is regularly damaged by overseeding (e.g., harrowing, ploughing) (Cardarelli et al. 2020), which is fatal in terms of the reproduction of I. costata, which lays its eggs in the upper-most layers of the soil (Nagy and Racz 2014). Intensive mowing of the whole plot is also pervasive, which can lead to a further reduction of the cover of dicotyledonous plant JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) 40 7) Bo) 2 = > & x= Relative frequency in the dataset FH Calcareous rocky steppes FH Mesotrophic wet meadows FH Uncharacteristic dry and semi-dry grasslands FH Closed forest steppe meadows HUncharacteristic mesic grasslands H Slope steppes IH Overseeded hayfields H Rich fens Examined habitat types Character of grassland naturalness and structure Character of grassland microclimate -- Regularly mowed » FH Semi-dry » Relative frequency in the dataset b L 6 © i a fH Disturbed > H{Shrubbbed & HOverseeded 0 H Humid 0 Character of the examined habitat types Fig. 6. Bar charts (+SE) of the studied habitat types with a pres- ence of Isophya costata and bar charts (+SE) of the studied habitats (characterized by different naturalness, structure, and microcli- mate) with a presence of Isophya costata. species to a level that is no longer able to meet the dicotyledon- ous nutritional requirements of I. costata (Orci et al. 2007); in this situation, the grasslands become unsuitable for maintaining the species’ populations (Kenyeres et al. 2004). The presence of I. costata in hayfields rich in dicotyledonous plant species and the absence of the species in overseeded mowed grasslands was also reflected in the sub-areas of the study. In the Balaton Uplands sub-area, which showed the highest frequency of the species, the prevalence of large hayfields rich in dicotyle- donous plant species was substantial. The Balaton Uplands sub- area is also characterized by the occurrence of steppe grasslands, which is a priori favourable for the species (Varga 1995). In addi- tion to the above, the extent of potential habitats for I. costata in the Balaton Uplands has increased due to the impact of humans, such as through deforestation and the establishment of grasslands maintained by mowing. Typically, extensive mowing creates large anthropogenic hayfields rich in dicotyledonous plant species with a physiognomy very similar to steppes. However, according to our results, I. costata occurs with a much lower frequency in steppe grasslands, and in sub-areas dominated by that habitat type (e.g., Eastern Bakony), than in hayfields. These grasslands have a simi- lar structure to hayfields but are less advantageous to I. costata in terms of protection from both predators (Kruess and Tscharn- tke 2002, Gardiner and Haines 2008) and weather challenges, as these grasslands have a different structure and shorter grass (Ke- nyeres et al. 2018). Z. KENYERES AND N. BAUER Table 2. Results of GLM with binomial distribution of presence/ absence data of Isophya costata and the studied habitat types, with indication of p-, R-values and estimates (grassland types examined using more than five quadrates and with a mean larger than 1 hectare were used). Isophya costata Presence of p R est. Rich fens ns. -0.053— -57.81 Mesotrophic wet meadows 0.043 -0.471 -251.44 Hayfields <0.001 2.358 -270.28 Overseeded hayfields <0.001 -2.775 -184.61 Calcareous rocky steppes ns. -0.464 -241.94 Slope steppes on stony soils ns. -0.623 -97.71 Closed forest steppe meadows n.s. 0.172 -75.84 Uncharacteristic mesic grasslands ns. -3.319 = -75.69 Uncharacteristic dry and semi-dry grasslands <0.001 -1.331 -170.63 The other sub-areas characterized by typical, but fewer, occur- rences of the species (Southern Bakony, Mez6féld, Bakonyalja, Marcal Basin) have a similar landscape to the Balaton Uplands sub-area. I. costata can also be considered frequent in the Alpok- alja sub-area (Vienna Basin), where loess grasslands and steppe meadows rich in dicotyledonous plant species (assumed to be the original habitat of the species) were present before the landscape was changed by human activities (Bauer and Kenyeres 2006, Ke- nyeres et al. 2009). Based on this extensive study, I. costata is also extremely rare in the intermediate, lowland areas (Gy6r Basin, Sopron-Vas Plain, Kemeneshat, and Komdarom-Esztergom Plain) The potential habitats of I. costata in these areas, dominated by arable lands, have drastically changed, and most of the remain- ing grasslands in these areas are located in depressions and, thus, are dominated not by semi-dry hayfields, but by marsh meadows with a humid microclimate that are less suitable for the ecologi- cal requirements of I. costata. According to the results of previous (Kenyeres et al. 2017, 2018) and the present study, grasslands with humid climates negatively correlate with the occurrence of the species and its density. Conservation possibilities. From the perspective of species protec- tion, it is essential to highlight that trampling, fires, and mow- ing in spring and early summer are a threat to I. costata due to its phenological characteristics and low mobility. The species places its eggs 1-2 cm deep in the soil, making them easily destroyed by passing fires. Among grassland management procedures, inter- ventions involving soil damage seriously endanger a population’s survival; ploughing and overseeding are lethal, but harrowing, with only a few centimeters of soil damage, can also significantly endanger the success of the species. In areas where I. costata occurs, the prohibition of grazing and tourism that cause trampling is justified; in hayfields, the applica- tion of late mowing (beginning of July at the earliest) or, if this is not feasible, mosaic-type treatment leaving unmown patches (e.g., 1/3 of the plot) is recommended (Kenyeres et al. 2017, 2018). The latter is justified by the fact that from the second half of May, which is the typical mowing period of these meadows, the spe- cies is in its adult state when reproduction and egg-laying take place. Also, in the distribution area of I. costata, mowed grasslands are often located between intensively used arable lands, and it is important for their conservation to reduce the use of pesticides in this area (Ivkovic and Horvat 2020). JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) Z. KENYERES AND N. BAUER y = 0,19915x+0,21697 Average density of /sophya costata 0.24 0.36 0.48 0.60 0.72 0.84 0.96 1.08 Frequency of hayfields rich in dicotyledonous plant species y = -0,363x+0,96308 Average density of Isophya costata 0.0 0.12 0.24 0.36 0.48 0.60 0.72 0.84 0.96 Frequency of grasslands characterized by humid microclimate Fig. 7. Results of GLM testing of the effect of the presence of hay- fields rich in dicotyledonous plant species and of the presence of grasslands characterized by a humid microclimate on the density of Isophya costata (based on data in ETRS quadrat, N=78). Acknowledgements The authors would like to express their gratitude to Ming Kai Tan and Ionut Stefan Iorgu for their remarks. We are also grateful to Tony Robillard, Editor-in-Chief of JOR, Alina Avanesyan, Sub- ject Editor of JOR, and Nancy Morris, Editorial Assistant of JOR for their work with our manuscript. 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