Open AccessArticle

Climate Change Influences on Central European Insect Fauna over the Last 50 Years: Mediterranean Influx and Non-Native Species

Attila Haris

^1,*

Zsolt Józan

¹,

Péter Schmidt

¹,

Gábor Glemba

²,

Bogdan Tomozii

György Csóka

⁴

Anikó Hirka

⁴

Peter Šima

⁵ and

Sándor Tóth

⁶

Rippl-Rónai Museum, H-7400 Kaposvár, Hungary

Hungarian Ornithological and Nature Conservation Society, H-1121 Budapest, Hungary

“Ion Borcea” Museum of Natural Sciences Complex, 600043 Bacău, Romania

⁴

Forest Research Institute, University of Sopron, H-3232 Mátrafüred, Hungary

⁵

Koppert s.r.o., Komárňanská cesta 13, 940 01 Nové Zámky, Slovakia

⁶

Bakony Museum of Hungarian Natural History Museum, H-8420 Zirc, Hungary

Author to whom correspondence should be addressed.

Ecologies 2025, 6(1), 16; https://doi.org/10.3390/ecologies6010016

Submission received: 2 December 2024 / Revised: 4 February 2025 / Accepted: 5 February 2025 / Published: 13 February 2025

(This article belongs to the Special Issue Feature Papers of Ecologies 2024)

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Abstract

The Central European fauna, in the last decades, has been undergoing a strong transformation due to four main factors: the retreat of living organisms that require cool and wet habitats, the proliferation of organisms that thrive in warm and dry conditions, the northward migration of Mediterranean organisms, and the gradual establishment of plants and animals with tropical origins. In this study, we detail the changes in the orders Diptera, Hymenoptera, and Lepidoptera and analyze the establishment of non native insects and the northward migration of Mediterranean insect species. The transformation towards a Mediterranean-type fauna is prominently indicated by the population in total abundance increase of xerothermic Aculeata, bee flies (Bombyliidae), and horse flies (Tabanidae). Additionally, groups that require cool and wet ecological conditions, particularly hoverflies (Syrphidae), but also sawflies (Tenthredinidae) and tachinid flies (Tachinidae), have shown a notable decline. In nocturnal moths, we observe a decrease in species richness in certain areas, as well as frequent outbreaks in populations of some less climate-sensitive species. Some species of butterflies are less sensitive to the current extent of climate change, exhibiting significant population in total abundance growth under protected conditions. However, most of the previously sporadic and rare butterfly species have proven to be climate-sensitive, unable to achieve significant population in abundance growth even under strict nature conservation. In recent decades, the influx from Mediterranean regions and the establishment of tropical non native insect species have turned at an exponential rate. We have reviewed the presence of alien insect species, recording 803 alien insect species in our region; 298 of them have arrived in the past quarter-century, with a significant proportion (54%) originating from tropical and Mediterranean regions.

Keywords:

climate change; insects; alien species; Hymenoptera; Diptera; Lepidoptera; Mediterranean influx; Central Europe

1. Introduction

“If you put a frog in a pot and slowly turn up the heat, it won’t jump out. Instead, it will enjoy a nice warm bath until it is cooked to death. We humans seem to be doing pretty much the same thing”.
(Jeff Goodell (The Water Will Come, 2010)

Jeff Goodell’s renowned quote aptly illustrates the gradual yet destructive process of climate change. Global warming, a critical aspect of climate change, profoundly impacts biodiversity and ecosystems worldwide. Insects, which play crucial roles in various ecological processes such as pollination, decomposition, and serving as food sources for other animals, are particularly affected. This paper focuses on the changes in the fauna of three major insect orders, Lepidoptera, Hymenoptera, and Diptera in Central Europe, driven by the ongoing shifts in climate patterns, additionally, we discuss all groups of the alien, introduced insects (Figure 1) and the Mediterranean influx.

Previous studies have demonstrated that climate change can lead to shifts in species richness and population densities of insects. For instance, research on European butterflies has documented northward range shifts and changes in abundance patterns. Parmesan and his colleagues [1] found that 63% of 35 non-migratory European butterfly species had shifted their ranges northward by 35–240 km over the 20th century. This shift is attributed to rising temperatures, which create more favorable conditions in previously cooler regions. Similarly, the phenology of many insect species is being altered by climate change. For example, the flight periods of butterflies and moths are occurring earlier in the year. Roy and Sparks [2] observed that the first appearance dates of 35 British butterfly species advanced by an average of 2.3 days per decade from 1976 to 1998.

Diptera, including many species of flies, are also experiencing changes in their populations in total abundance due to climate change. For example, the distribution of the common housefly (Musca domestica Linnaeus, 1758) has been influenced by temperature changes, with populations in abundance increasing in regions that have become warmer. Additionally, climate change can affect the prevalence of disease vectors such as mosquitoes [3,4]. Researchers suggested that rising temperatures and altered precipitation patterns could expand the range of mosquitoes that transmit diseases like malaria and dengue fever. In Hymenoptera, a similar phenomenon may be observed. Ornithologists have noted that European bee-eaters (Merops apiaster Linnaeus, 1758) are expanding northward following their prey animals [5]. Central Europe, with its diverse habitats ranging from lowland forests to alpine meadows, provides a unique setting to study these changes. The region has already experienced noticeable shifts in climate, with increasing average temperatures and altered precipitation patterns. These climatic changes are expected to continue, posing challenges for the conservation of insect biodiversity.

The present work is a continuation of our monograph titled “Changes in Population Densities and Species Richness of Pollinators in the Carpathian Basin” [6]. The conclusion of this work is the basic assumption of our entire work. During the writing of that article, we realized that instead of a pollination crisis in our region, a strong Mediterranean-type transformation is taking place here in Central Europe.

In this study, we concluded that although a pollination crisis is moderately present in our region, particularly for certain groups, the more concerning issue is the gradual Mediterranean-type transformation of the fauna. In this article, we examine this transformation in detail. We will discuss the adaptation of individual insect species and their established ecotypes, examining how each ecotype responds to changing environmental conditions. We also explore the transformation of fauna in terms of agricultural plant protection, forest protection, and nature conservation.

The study area is Central Europe, with the Carpathian Basin at its heart (Figure 2 and Figure 3). Of the 11 biogeographic regions in Europe, three are present here: the Pannonian, Continental, and Alpine regions [7].

Global warming is merely one aspect of the broader set of contemporary issues termed “polycrisis”. This includes climate change, habitat destruction, loss of natural vegetation and fauna, overpopulation, degradation of agricultural land leading to food crises, dwindling potable water supplies triggering global migration, energy crises, and the depletion of raw materials critical to the economy. Collectively, these issues herald a global collapse process, threatening economic, commercial, political, social, and ultimately, cultural stability. These climate-related challenges can trigger or worsen other crises listed above. The complex interplay between climate change and these other factors creates a web of interconnected problems that reinforce and amplify one another, making it difficult to address any single crisis in isolation.

Fauna is fundamentally determined by vegetation [7]; hence, changes in fauna are driven by changes in vegetation through ecological networks. The interaction between climate change and vegetation is complex. Climate change and warming favor the expansion of the steppe zone while adversely affecting montane beech and other woodlands. Concurrently, afforestation mitigates the impact of climate change. These processes have been studied extensively in our region [9,10]: researchers mapped the changes in vegetation under climate change, and by using the REMO model, they concluded that forest restoration could reduce surface temperatures by up to 0.7 °C and may increase precipitation by more than 10%.

Key aspects of faunal change include the decline of abundances of insects adapted to cool climates, the expansion of insects adapted to warm, dry, or occasionally warm-humid climates, the northward spread of Mediterranean insects, and the promotion of invasive species. Insects have short generation times and high reproductive rates, making them more likely to respond rapidly to climate change compared to plants and vertebrates. Possible responses include changes in phenological patterns, habitat selection, and the expansion and/or contraction of geographic ranges. These rapid responses can trigger processes with potentially devastating consequences for crop production and food security [11]. Karuppaiah and Sujayanad classified these responses into changes in insect population dynamics, migration, development and reproductive biology (phenological changes, nutrient cycling, etc.), diapause, stage survival, growth rate, and species voltinism [12].

The indirect effects of climate change are examined in a UFZ study [13], including the re-emergence of mosquito-borne diseases like dengue fever or malaria, the occurrence of multiple generations of pests due to extended reproductive periods, increasing damage, and the likelihood of forest fires in boreal areas such as Canada. The increased prevalence of certain insect species can also serve as an indicator of climate change [14].

The temporal movements of different insect groups at varying altitudes exhibit diverse patterns depending on climate change and the ecological needs of specific groups [15]. In the present work, we will also focus on these patterns, using the Carpathian Basin as a representative sample area for Central Europe.

The climate of Central and Eastern Europe is characterized by a general rise in temperature (Figure 4), particularly pronounced in the Alps. Overall, temperatures have risen significantly by approximately 1.2 °C during the 20th century, with seasonal increases of 1.1 °C in spring, 1.3 °C in summer, 1.2 °C in autumn, and 1.3 °C in winter. However, the frequency of cold days and nights has decreased over the past 40 years.

There is no overall trend in precipitation across the region, but regional specificities must be considered. For instance, precipitation increased by about 9% in the northwest, while it decreased similarly in the southeast. In Hungary, precipitation has generally decreased by 11% since 1901, particularly since the 1970s. Droughts are common in late spring/early summer and late autumn [16]. Hungary, due to its low-lying position in the Carpathian Basin, is particularly affected by a negative water balance. The Tisza, the region’s second-largest river, has been strictly regulated since 1846. Consequently, the river acts as a drainage channel for the vast volumes of water from the surrounding mountains, further drying out the Carpathian Basin. In 2019, the volume of water flowing into the lower areas of the Carpathian Basin was 112 km³, while the volume flowing out was 117 km³ [17,18]. Global warming is disrupting this balance, leading to increased droughts in recent years.

In Romania, the temperature has increased by about 0.5 °C since 1901, with up to 1 °C in the southeastern region. Significant summer warming has been observed since the 1970s, particularly in the Northeast and Southwest. Winter warming has been steady, leading to the warmest winter in 2006/2007 with a +6 °C anomaly. Precipitation does not show a uniform long-term trend, but there are regional variations, with some areas experiencing increases and others decreases. Extreme precipitation events show a varied pattern.

In Slovakia, after 1990, the average air temperatures have increased compared to the period 1951–1990. The number of tropical (≥30 °C) and supertropical days (≥35 °C) has significantly increased. Dry periods have become more frequent due to rising temperatures and high potential evapotranspiration, leading to a significant decrease in soil moisture. Global warming in Slovakia has resulted in an increased average annual air temperature of 1.1 °C over the last 100 years (1901–2000) with noticeable warming in the last two decades. This climate shift has led to new plants and animals migrating from the south while native species are forced to move northward or to higher altitudes, sometimes leading to extinction. Severe drought episodes lasting several weeks are becoming almost annual events, with rainfall occurring mainly as short, intense showers. In Central and Eastern Europe, the mean annual temperature is projected to increase between 1 °C and 3 °C by mid-century and up to 5 °C by the end of the century. However, one-way projections do not account for the phenomena of action-reaction and climatic feedback [17].

Figure 4. Average annual temperatures of three Central European countries between 1970 and 2023. Green: Hungary, orange: Romania, red: Slovakia. Climatic data from World Bank Portal [19].

Our paper synthesizes current research on the impacts of global warming on Lepidoptera, Hymenoptera, and Diptera in Central Europe. By examining changes in distribution, phenology, and population dynamics, we aim to understand the broader ecological consequences and inform conservation strategies. Particular attention is given to the influx of insect species from the south, as well as the introduction and establishment of alien insects, often referred to as insect globalization, within the context of the current warming climate. The findings underscore the urgency of addressing climate change to preserve the intricate web of life that sustains our ecosystems.

2. Materials and Methods

2.1. Diptera, Hymenoptera

Our research is based on 3 to 5 decades of systematic collection results from the 1970s and 1980s onwards. The collecting area covers the Carpathian Basin (Pannonian Basin) region, the core region of Central Europe. Approximately 4000 collection points were utilized, with 30–35 collection days per year, depending on the weather conditions. The collected material is partly stored in the Natural History Museum in Zirc, the insect collection of the Rippl-Ronai Museum in Kaposvár, and the Hungarian Natural History Museum. The collection method used was grass netting. For the analysis of individual trends, changes in the number and species richness of the groups defined by each ecotype were examined. The ecotypes of Aculeate were defined as euryecious intermediate, euryecious eremophilic, euryecious hylophilic, stenoeceous eremophilic, and stenoeceous hylophilic, following Pittioni [20,21]. The ecotypes of Diptera were defined following Hagen [22]. Bumblebees were categorized according to their climatic sensitivity based on the monograph by Rasmont and colleagues (“Climatic Risk and Distribution Atlas of European Bumblebees”) [23] and compared with our own recording experience. The above-mentioned researchers established the ecotypes based on the empirical method. The basis of the applied and generally widespread ecotypes is the regular collection or comparison and indexing of the ecological parameters of the distribution area (studying large museum collections).

We provided a list of insects that have not been collected in recent decades, i.e., those whose numbers have fallen below the detection level. These species were studied in parallel in the lower parts of the Carpathian Basin and at high altitudes of 1200–1800 m above sea level (Tatras, Carpathians, Slovakia, Romania) to determine whether high altitudes can provide suitable habitats for their survival [24,25,26,27,28,29,30,31,32,33,34]. While the moth data are unpublished, Hymenoptera and Diptera data are published in part [35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51,52,53,54,55,56,57,58,59,60].

2.2. Lepidoptera

2.2.1. Rhopalocera, Butterflies

The two primary sources of data on butterflies are the butterfly collection of the Rippl-Rónai Museum in Kaposvár and the Danube-Ipoly National Park. The museum’s collection consists of approximately 50,000 specimens from nearly 300 sites across the Carpathian Basin, regularly collected with 30–35 field days annually. However, as the collection is selective, it does not provide reliable data on common butterfly species. Levente Ábrahám systematically collected specimens of 33 species over 30 years, offering a reliable picture of these species’ changes over time. Ecotypes by habitat have only occasionally been applied to diurnal butterflies by various authors, and this is also true for nocturnal butterflies, so they remain to be developed further [61,62].The ecological usability and limitations of museum butterfly data are discussed by several authors [63,64,65,66]. They consider museum collections as indispensable historical and ecological resources for comparing climate change and present conditions. However, Davis [64] argues, “Museum specimens were not collected with the purpose of estimating population aboundance trends and thus can exhibit spatiotemporal and collector-specific biases that can impose severe limitations to using NHC data for evaluating population trajectories”. With the help of collectors, rare and sporadic butterflies were systematically collected, and only those were analyzed for quantitative data. The trend of decline for common species cannot be described from this collection of data alone.

The Danube-Ipoly National Park, covering about 600 km² in the middle of the Carpathian Basin, is an excellent source of butterfly data for analyzing these butterfly species. As a nature reserve, it minimizes disturbances such as pesticide use (partly banned or restricted in some areas), creation of ecological corridors between core areas instead of habitat fragmentation, systematic eradication of invasive plants, and adaptation of the mowing regime to phenological stages. These measures allowed for studying the impact of climate change by eliminating most disturbances. The increase in the butterfly population was not only expected but also logical. By filtering out factors causing the decline in butterflies, it became clearer which species were sensitive to climate change and which were more adaptive.

Annual data collections ran from early March to the end of October, usually twice a month, in the first and third weeks. On average collection days, a net of three hours was spent in the field, not including travel time between sites. Regular survey areas included Csomád: Magas-hegy, Fót: Somlyó-hegy, and Csömör: Ősláp, upon the request of the Danube-Ipoly National Park. This 10-year data set from regular surveys covering all butterfly species is unique in Central Europe and an outstanding source for ecological-historical studies, allowing tracking changes even in common butterfly species.

Here, we employed the Pollard transect method and periodically cross-checked using the MRR (Mark-Release-Recapture) method to reduce the possibility of double-counting specimens. The procedure followed that of Nowicki et al. [67], but in our case, the duration of the Pollard transect was extended to 60 min instead of 30 min. Additionally, avoiding double counting was more feasible for species that protect their territories (e.g., Papilio machaon Linnaeus, 1758 (swallowtail butterfly)) or where distinct individual lesions were visible during sightings.

The proportion of captured and mounted butterflies was 10%. These were species that were difficult to identify, and several specimens were photographed from multiple angles (wing back, color) for accurate identification, allowing the relative proportions of similar species to be determined. Annual results showed considerable fluctuation depending on weather factors; thus, pooling several years of data in the analysis helped mitigate the impact of annual weather variations. The climatic sensitivity categorization of each species was based on the monograph by Settele and colleagues (Climatic Risk Atlas of European Butterflies) [68] and compared with our own recording experience. Values of Species Temperature Index (STI), Species Precipitation Index (SPI), Habitat Suitability Index (HSI) are based on the papers of Schweiger et al.; Mora, A. et al. and Kudrna et al. [69,70,71].

2.2.2. Nocturnal Macrolepidoptera

The species richness and number of nocturnal moths were analyzed using light trap material from faunistic research conducted in the Carpathian Basin. Notably, this includes material from the national forest light trap network in Hungary. Light trap material was collected from 41 sites across the Carpathian Basin and identified by experts such as Csaba Szabóky, Csaba Gáspár, Lajos Kovács, Katalin Leskó, Levente Ábrahám, Ákos Uherkovich, Zoltán Varga, and Péter Schmidt, among others. A single trapping point was consistently included over several decades, and changes in light sources were also considered.

Methodological changes were necessary for nocturnal butterfly studies. Since 2014, UV LED light traps have been used, replacing the black light UV 20 W (used between 1990 and 2010) and the Jermy-type light trap with a 125 W mercury vapor lamp (used till 1990 circa). These different light sources had varying selectivities [72,73,74]. Consequently, we calculated a long-term trend from 1970 and a separate trend from 2014 to draw reliable conclusions. Additionally, an apparent trend reversal in the 1980s was observed and calculated separately for each species as a control. Specific analyses of Hymenoptera, Lepidoptera, and alien insects are in Supplementary Materials A–C.

From the light trap data, we selected only those series that covered six months of intensive daily collection, typically from April to October and, in some cases, from May to November. The traps operated continuously during these periods. Other shorter or incomplete trap data were excluded from the analysis. The number of sampling points was 24, and several locations (such as the Drava Plain or the Zselic Mountains) were resampled after 40 years. To differentiate surveys from various locations in the same year, letters of the alphabet were used as follows: Felsötárkány: 1970a, 1971a, 2020a, 2021b, 2022c Tompa: 1970b, 2020b, 2021c, 2022b Sopron: 1970c, 202c, 2021d, Gilvánfa: 1970d, 1973b, Sellye: 1971c, Magyarszombatfa: 1975, 1976a, Kisvaszar: 1973a, Mike: 1976b, Vásárosbéc: 1977, 1978a, Bőszénfa: 1978b, 1979c, 1980c, Almamellék: 1979a, 1980a, Palé: 1979b, 1980b, 1981b, Lipótfa 1986, 1987, Jósvafő: 1990, 1998, 2005, Aggtelek: 2000, 2001, Répáshuta: 2015, 2016, 2017, 2018, 2019a, Sellye: 2019b, 2020d, 2021a, 2022a, Ropoly: 2019c, 2020e (Supplementary Material B).

The ecotypes of moths were discussed partially following the work of Sage and Utschick [61], where the authors ranked different habitats on a scale from 0 to 10. Additionally, the classification by Kanarskyi, Y. et al. [62] was employed, assigning ecotypes as follows: U (ubiquists), M1 (grassland mesophiles), M2 (seminemoral mesophiles), M3 (nemoral mesophiles), H2 (nemoral hygrophiles), X1 (grassland xerothermophiles), and X2 (seminemoral xerothermophiles).

2.3. Mathematical Methods and Data Standardization

Over the years, a significant amount of data has been collected: approximately 300,000 specimens of nocturnal moths, 50,000 specimens of butterflies, 40,000 specimens of Diptera, and 60,000 specimens of Hymenoptera. For Hymenoptera, Diptera, and butterflies, data from individual years were aggregated into 3, 5, and 6-year groups to smooth out population fluctuations caused by annual weather variations. Nocturnal moths were the exception, with annual data series being sequentially analyzed. We employed trend analysis to track changes, modeling most changes using linear and exponential trends. We plotted the trend line and the trend equation, determining the linear trend coefficient of determination (r² value). The coefficient of determination of the linear trend and the trend’s steepness, expressing the rate of change and its direction (positive or negative). The coefficient of determination of linear trend (r²) reveals whether our data follows a trend or is dominated by non-trend factors, potentially indicating random changes. The r² value ranges from 0 to 1, with higher values indicating a stronger trend influence, such as climate change or gradual habitat loss and degradation. We calculated the r² using the sum of squares method. The dominance relationships at the beginning and end of the studied period were determined using the Ranking Method or Borda Count method, eliminating the overwhelming effect of outbreaks while appropriately calculating their magnitude and frequency relative to the locations where the occasional outbreaks occurred.

To measure the impact of climate change, we used the coefficient of determination (tR²) value for the temperature change (it differs from the r², which measures the fiting to the linear model as it is described above). To differentiate between the two types of coefficients, the coefficient of determination for the linear trend (trend-model for the changes) is marked with r², while the coefficient of determination related to the change of temperature (see Figure 4),depending on where the data comes from: Hungary, Slovakia, or Romania) is marked with tR² (t for the change of temperature). Here (with the determination coefficient of the temperature change), we compared population size changes and fluctuations with climatic data only for major groups. Other obvious methods, like ANOVA, cannot be applied because during the extensive data collection spanning up to six decades, concepts such as climate change and the threats posed by polycrisis components to biodiversity and pollination crises were not even conceived at that time. Consequently, the last 4–5 decades’ millions of additional influencing ecological data were not collected or quantified for biotic, abiotic, and anthropogenic environmental factors. However, the coefficient of determination excellently demonstrates how much climate change can play a role in the growth, decline, or even extinction of specific groups in abundance. While the coefficient of regression r² helps us to decide whether the changes of abundances or species richness follow a trend or not, at the same time, the coefficient of determination, tR² value helps us quantify the impact of climate change on population dynamics, showing how significant the role of climate change is in driving population growth in total abundance, decline, or extinction. In the analysis, approximately 900 insect species were analyzed, but in the family and group statistics there are data for 5 thousand insect species or even more (all species that were captured during the 30–50 years of investigations).

2.4. Non Native and Invasive Insect Species, Mediterranean Influx

With climate change, species that were previously not members of our natural fauna are appearing. Additionally, species introduced by humans, either intentionally or unintentionally, are becoming more prevalent. Mediterranean newcomers and alien species are treated separately. The northward spread of Mediterranean insect species is directly attributable to climate change, while the establishment, spread, and reproduction of adventitious species from warmer regions may be facilitated by mild winters.

As a starting point, we utilized the monograph titled “Alien Terrestrial Arthropods of Europe” [75], which lists 330 insect species from the Carpathian Basin. This number was more than doubled by processing additional works. Insects of alien origin, whether deliberately or accidentally introduced by humans, and species that have migrated northward from the Mediterranean to Central Europe in the last two decades, were analyzed separately. The temporal appearance, taxonomic composition, and zoogeographic division of non-native species were depicted on graphs.

Looking back over the last 10,000 years, we evaluated changes that occurred in each historical period, comparing them with the rate of change of the fauna in our present time, and derived mathematical relationships. We looked back 10,000 years, this time, at the end of the Ice Age, ref. [76] the climate had stabilized, and present conditions gradually emerged. Therefore, the first three human-mediated introductions of Phthiraptera species and subspecies in the Palaeolithic and earlier are omitted, as subsequent insect movements induced by climate changes completely redrew the fauna during the interglacials. It is challenging to determine the precise introduction date of alien species. Thus, we rely on the date of the first sighting, which can only be indicative. For large, conspicuous species (e.g., Antheraea yamamai), this is a reliable guide. In other cases, there may be a preceding period of incubation or breeding, or the problem may be due to a lack of specialists. Currently, the published date of first sightings is constantly being revised by the growing GBIF database [77], which contains numerous unpublished old records for a wide variety of animal species. Additionally, archaeology has recently played a significant role in detecting the establishment dates of invasive species, extending these dates back to centuries or millennia [78]. Our detailed data on the invasive and non-native insects of the Carpathian Basin can be found in Supplementary Material C, which is based partly on our own published investigations and partly on papers by various experts from around the world [79,80,81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109,110,111,112,113,114,115,116,117,118,119,120,121,122,123,124,125,126,127,128,129,130,131,132,133,134,135,136,137,138,139,140,141,142,143,144,145,146,147,148,149,150,151,152,153,154,155,156,157].

Predicting the emergence and expansion of non-native insects or the contraction or vertical movement of native species northward in response to climate change is currently not possible. Numerous predictions exist (for example, [22,71]) based on specific climatic models [158], which predict rapid or slow warming depending on atmospheric pollution. More recent predictions, based on the impact-versus-impact principle with a very high uncertainty factor (e.g., cooling due to the collapse of Atlantic Meridional Overturning Circulation (AMOC) [159,160], have emerged. It must be acknowledged that interference with the climate triggers chaotic processes, creating extreme and highly variable conditions and rendering these predictions outdated. Extreme weather variability is at least as burdensome, if not more so, on wildlife than continuous warming.

The definition of non-native, alien, and invasive insect species is based on the 1999 US Executive Order #13112 quoted by Ziska et al. [161], which defines “invasive” species and “alien” species for federal agencies as follows: “Alien” species means, with respect to a particular ecosystem, any species, including its seeds, eggs, spores, or other biological material capable of propagating that species, that is not native to that ecosystem. This term also includes non-indigenous or exotic species. In contrast, “invasive or noxious species” means an alien insect species whose introduction does or is likely to cause economic or environmental harm or harm to human or animal health. We may add that an animal may become an invasive species through deliberate introduction, e.g., the harlequin lady beetle (Harmonia axyridis Pallas, 1773).

3. Results

3.1. Diptera

3.1.1. Horse-Flies, Tabanidae

Horse-flies are primarily xerothermic. Therefore, the Tabanidae population abundance has shown an increase (Table 1 and Figure 5). We saw a significant decline only at the Band-eyed Tabanus bromius Linnaeus, 1758 (Brown Horse-fly). A moderately significant decline was experienced at Haematopota pluvialis (Linnaeus, 1758) (Common Horse-fly). Among these, Haematopota pluvialis is the most common hylophilous species in our region. When examining trends in detail, the vast majority of species have shown an increase, which is not surprising given that most members of this group are xerothermic.

In the long term, since the larvae require aquatic or semi-aquatic environments for development, species richness and population density are expected to decrease as these environments become more scarce. In the last part of the time series, these signs are already visible. Thus far, many species in this group have benefited from climate change.

While there is a close correlation between the ecotypes of individual species, it is not absolute. We can say that species showing stronger growth trends, with one exception, all have xerothermic ecological requirements in their adult form. The number of Tabanidae species exhibiting a decline is very small, with only two species showing moderate or strong decreasing trends. One of the most common species, the hylophilous Haematopota pluvialis, also showed a strong increase till the end of the 1990s; after this, the population density of this species dropped. Overall, the gradually rising temperature, with its low coefficient of determination (tR² = 0.0467), has a very slight influence (approximately 5%) on the quantitative changes in all Tabanidae populations in total abundance. We attribute this to the fact that the group, as shown in the table, is divided into two main ecological types: one group is moisture-loving (sylvicol), while the other is specifically xerotherm. Thus, the two ecotypes cancel each other’s effects. Among the sylvicol forest species, Haematopota pluvialis is the dominant one and has also suffered the most drastic decline. We specifically examined how much influence the warming may have on this decline. The obtained Coefficient of Determination (tR²) related to the temperature trend is 0.2346, indicating that while climate change is an important factor in the abundance decrease, it alone does not fully explain the observed decline in numbers.

The ecotypes identified in the literature through field research and the observed reactions, while not exhaustive, show overall good agreement. The classification of species that exhibit abundance growth is nearly all xerothermic, with only two exceptions. Of those horse-flies where we observed an abundance decline, two-thirds are classified as silvicol and hylophilous. The reasons for the lack of complete agreement could be multiple, such as incorrect determination of ecological requirements or ecotypes in the literature, changes in ecological requirements, adaptation processes, or some degree of drift during fieldwork, which is more or less unavoidable.

Overall, the response of Tabanidae species to climate change is twofold: while warming has favored xerothermic species in the adult stage, it is also evident that the larvae are dependent on water. Seven species have not been collected in the Carpathian Basin for 20 years or more. Among these, three species appear to find refuge in higher elevations of the region, such as the Tatras or the Carpathians. These species are Pangonius pyritosus (Loew, 1859), Hybomitra aterrima (Meigen, 1820), and Hybomitra montana (Meigen, 1820).

3.1.2. Hoverflies, Syrphidae

This group can be characterized by a significant decline in numbers, regardless of ecotypes (Table 2 and Figure 6). Among the studied Diptera families, this group shows the strongest decline. The likely reason is the moisture-dependent lifestyle of the larvae. However, this explanation is not entirely satisfactory, as the larvae of Tabanidae also require aquatic or semi-aquatic environments but do not exhibit the same intense decline as hoverflies.

The overall picture becomes even more interesting and disheartening when we examine whether the mountainous regions of the Carpathian Basin provide refuge for hoverflies that have disappeared from lower-lying areas or have fallen below detectable levels. A significant number of Syrphidae species have not been collected in this century, although they were once sporadically found throughout the Carpathian Basin. Among the four studied groups, 89 species disappeared in the last 2 decades, with the vast majority, 50 species, belonging to the Syrphidae. We investigated whether high-altitude habitats serve as refuges for these hoverflies. Of the 89 Diptera species, only 14 have been found in the last 24 years in the Tatras or the Carpathians, representing a mere 16%. The changes in hoverfly abundances are strongly negatively correlated with the gradually rising temperature of our environment, as they are known to be moisture-dependent insects. This is further supported by the high coefficient of determination obtained from the results (Coefficient of Determination tR² = 0.6768).

Almost all ecotypes have seen a rapid decline in numbers. However, we observed a strong decline in the number of silvicol and intermediate species, while the decline in xerothermic species was moderate or showed a balance. The abundances of two hoverfly species from the lower regions has declined to such an extent that the decrease is better approximated by an exponential trend rather than a linear one, these 2 species are Sphaerophoria scripta (linear r²: 0.73, exponential r²: 0.76) and Syrphus torvus (linear r²: 0.58, exponential r²: 0.72). Interestingly, the only species showing a significant increase is Anasimyia contracta Claussen & Torp, 1980, an Atlantic moisture-loving species. This indicates that while climate change greatly influences population dynamics, it is not the sole factor, and we are facing highly complex processes. Additionally, a significant decline in the population abundance of the Syrphidae family is not surprising, as it was expected based on previous research. However, it is surprising that the species showing the most pronounced declines are classified as euryecious. This is a consequence of the fact that these hoverflies were widespread and common before the effects of climate change became apparent. This significant decline of species historically classified as “euryecious” also highlights the necessity for the reevaluation of the ecotypes of this group.

3.1.3. Bee Flies, Bombyliidae

We examined all common or frequent species within the approximately 65 species of Bombyliidae. With few exceptions, almost all species have shown an intense increase in numbers or have remained stable over the past 40 years (Table 3 and Figure 7). Bee flies likely include some montane, forest species, which are among the few within the family that are sensitive to climate change. Their distribution supports this, as they are most common between the latitudes of 48–52 degrees in Western Europe (GBIF, [77]).

Three Bombyliidae species have not been detected in the Carpathian Basin for 20 years or more: Bombylius trichurus Wiedemann, 1818; Bombylius quadrifarius Loew, 1855; and Bombylius ambustus Pallas & Wiedemann, 1818. The disappearance of these southern species is certainly not due to climate change. Overall, Bee flies are clearly the winners of the prevailing temperature increase caused by climate change so far. When we examine the proportion of total population abundance changes attributed to warming, we find a very strong 57% share (Coefficient of Determination (tR² = 0.5744).

3.1.4. Tachinidae

No growth trends were observed for any species (Table 4 and Figure 8). However, the decline is not as pronounced as in the Syrphidae species (for comparison, see also Table 5). On the other hand, several species, including Linnaemya frater (Rondani, 1859), Meigenia dorsalis (Meigen, 1824), Exorista larvarum (Linnaeus, 1758), and Tachina fera (Linnaeus, 1761), have shown specific trends: the general trend is decline but displayed significant increase in the last five years of investigation.

These species primarily have silvicol (forest-dwelling, cooler microclimate-preferring) ecological requirements (12 species) or are intermediate in terms of temperature (6 species), with only one xerothermic species (Table 4). Based on this, most Central European Tachinidae species are sensitive to climate change. Among the 27 species not collected for over 20 years, only seven are known to still exist in the high mountains surrounding the Carpathian Basin (and 17 of the 50 hoverfly species) (Table 6). These include Cadurciella tritaeniata (Rondani, 1859), Catharosia albisquama (Villeneuve, 1932), Ceranthia tristella (Herting, 1966), Chetoptilia puella (Rondani, 1862), Eloceria delecta (Meigen, 1824), Ligeriella aristata (Villeneuve, 1911), and Winthemia bohemani (Zetterstedt, 1844), primarily of Scandinavian distribution. The species Amelibaea tultschensis (Brauer & Bergenstamm, 1891) and Bithia acanthophora (Rondani, 1861) are rare mountainous species with records from the Italian Alps. The species Anthomyiopsis plagioderae (Mesnil, 1972) is of Atlantic distribution, while Vibrissina debilitata (Pandellé, 1896) is of Scandinavian and Atlantic distribution. The species Aphria xyphias (Pandellé, 1896), Campylocheta latigena (Mesnil, 1974), Conogaster pruinosa (Meigen, 1824), Phytomyptera abnormis (Stein, 1924), Estheria acuta (Portschinsky, 1881), Gonia bimaculata (Wiedemann, 1819), Heraultia albipennis (Villeneuve, 1920), Minthodes pictipennis (Brauer & Bergenstamm, 1889), and Psalidoxena transsylvanica (Villeneuve, 1929) have always been extremely rare in our region (GBIF database, [77]). In the case of the Tachinidae family, global warming and population changes are negatively correlated. However, the impact of temperature increase is small, which is not surprising given their parasitoid lifestyle. According to the literature (detailed in the next chapter), the population fluctuations of Tachinidae species are unpredictable and require extensive further research. Regarding climate change, we can currently say that it influences them, but there are other unexplored factors that predominantly affect them. At the group level, the Coefficient of Determination of the temperature increase is tR² = 0.2205.

3.2. Hymenoptera

3.2.1. Bumblebees

Upon preliminary observation, it is evident that bumblebee abundances are experiencing a significant and marked decline across both the lower regions of the Carpathian Basin and mountainous areas, with high linear regression coefficient (r²) values from 2005 (Table 7). Bombus terrestris (Linnaeus, 1758) (Buff-tailed Bumblebee) and Bombus lapidarius (Linnaeus, 1758) (Red-tailed Bumblebee) dominate the lower elevations, while Bombus lucorum (Linnaeus, 1761) (White-tailed Bumblebee) prevails at higher elevations. Cuckoo bumblebees are rare at lower altitudes, whereas they are primarily found in mountainous areas. As cuckoo bumblebees are parasites of social bumblebees, it is evident that their numbers also decline alongside those of their host bumblebees. At lower elevations, two Mediterranean-origin bumblebee species, Bombus haematurus Kriechbaumer, 1870 and Bombus argillaceus (Scopoli, 1763), are replacing the native fauna. Bombus haematurus may eventually become the dominant species at lower altitudes. According to our observations, Bombus haematurus queens terminate their diapause relatively early, in the first half of March. The populations peak in April, and the last individuals have been observed in the first third of July. This phenological pattern is not typical in our region for any bumblebee species sharing the typical habitat with B. haematurus, except B. pratorum. With warming, flowers bloom earlier. Thus, B. haematurus was better able to track and exploit this new opportunity compared to other Bombus species, thereby gaining an advantage. This opened up the possibility of northward expansion.

Bombus argillaceus did not show significant abundance at higher altitudes. In general, while the Bombus fauna is more abundant in high mountains, a similar declining trend is occurring, albeit with some delay, in the low mountain regions. In high mountainous regions, highly specialized species, such as Bombus gerstaeckeri F. Morawitz, 1881—already a rare sight in the Carpathians—are particularly vulnerable to the impacts of climate change, which can negatively affect the habitats of their preferred plant species.

Climatic Risk Categories align with the observed decline in bumblebee populations in total abundances. Species that have declined below detection levels are categorized as extreme high risk (HHHR) at low latitudes, where the impacts of climate change are most pronounced. Conversely, the two species exhibiting population increases in abundances are classified as high risk (HR) and low risk (LR) in the literature [22]. A different trend is observed at high altitudes: species experiencing the most severe declines do not fall into the most vulnerable categories. These findings underscore that while climate change significantly impacts bumblebee population dynamics, it is not the sole influencing factor.

In the Carpathians, we have surveyed bumblebees over the past decades. In the Eastern Carpathians, certain species, such as Bombus soroeensis (Fabricius, 1776), appear to be thriving in the more humid glades within forested areas. Conversely, species like Bombus wurflenii Radoszkowski, 1860, which typically inhabit the transition zone between the upper coniferous forest and the subalpine region, have become less common, likely due to increased exposure to direct climatic fluctuations. Additionally, Bombus pyrenaeus Pérez, 1879 has not been observed in recent years, despite extensive efforts to locate it.

Meteorological data for the Eastern Carpathians indicate a significant rise in temperature and solar radiation accompanied by a marked decline in humidity at higher altitudes, particularly in the subalpine zone. These climatic changes are likely driving the observed shifts in Bombus species distribution and abundance. At high altitudes, after a general decline since 2005, B. lucorum appears to have stable populations. Despite the general decline, the higher altitudes have helped preserve the richness of the local fauna up until this phase of warming (Table 7 and Figure 9). While we observed a 30% reduction in species richness at lower altitudes, at higher altitudes, we captured the same number of species among the regularly monitored species at the beginning (19) and the end (19) of the study. Furthermore, to this day, the higher altitudes continue to help maintain the populations of certain mountain species, as discussed below. The high-altitude regions have played a particularly important role in the preservation of cuckoo bumblebees. While 80% of cuckoo bumblebee species have disappeared from lower-altitude regions, all species have been successfully collected continuously in high-altitude regions (see Table 7). Thus, high mountains have a prominent role in the preservation of cuckoo bumblebees. We can conclude that the decline of cuckoo bumblebees accounts for the majority of the disappearance of all Bombus species. Another species exhibiting notable success is the Common Carder Bumblebee: Bombus pascuorum (Scopoli, 1763), which can frequently be found up to the subalpine zone in the Ceahlău Mountains, after its decline in the last two decades. Alongside B. pascuorum and B. soroeensis, Bombus campestris (Panzer, 1801) is also commonly encountered at elevations reaching up to 1750 m. However, the impact of climate change is evident on oligolectic species such as B. gerstaeckeri, which forages primarily on Aconitum spp., plants that thrive in shaded, cool, and moist environments. Despite extensive surveys, we have not been able to locate this species in recent years, suggesting a potential decline in its population abundance linked to habitat changes driven by climate fluctuations. Some less frequent species, such as Bombus confusus Schenck, 1861, Apple Bumblebee: Bombus pomorum (Panzer, 1805), and the Large Garden Bumblebee: Bombus ruderatus (Fabricius, 1775), tend to expand or persist in very abundance in lower mountainous areas (though this should be closely monitored in the coming years to ensure it is not accidental).

Of particular interest is the adaptability of B. haematurus, which appears more inclined to spread along forest margins, parks, and dry open areas, compared to B. argillaceus. Based on our observations, B. argillaceus is found predominantly within deciduous forests and parks, suggesting differing ecological tolerances between these two species. The maximum altitude for B. haematurus in Romania in recent years was 780–800 m (Valea Uzei—Râmeț) in the Apuseni Mountains [162]. For B. argillaceus, the highest recorded altitude was 852 m (around Gheorghieni, Harghita) [162]. Additionally, B. argillaceus was observed in a valley north of the Făgăraș Mountains (Sâmbăta de Sus) at approximately 770 m. In the Eastern Carpathians, it was recorded at a lower altitude of around 680 m (Ticoș, Neamț County). These observed abundance trends highlight the need for continued and focused monitoring in the coming years to better understand the long-term impacts of climate change and habitat modifications on Bombus species. Examining the impact of gradually rising temperatures on bumblebee population dynamics, we observe an interesting pattern: the effect differs between lowland and highland regions (correlating with different species compositions and dominance relations). However, an increase in environmental temperature negatively affects population size at all elevations. In low altitudes, climate change exceptionally strongly impacts bumblebee populations, with the most extreme highest values observed here: tR² = 0.9006. Conversely, in highland regions, this effect is significantly moderated, tR² = 0.5045 due to more balanced temperature conditions. The population in abundance of four Bombus species only from the lower regions has declined to such an extent that the decrease is better or similarly approximated by an exponential trend rather than a linear one, these species are Bombus lapidarius (linear r²: 0.71, exponential r²: 0.71), Bombus ruderarius (linear r²: 0.42, exponential r²: 0.66), Bombus pauscoum (linear r²: 0.51, exponential r²: 0.71) and Bombus hortorum (linear r²: 0.42, exponential r²: 0.66).

3.2.2. Aculeata

The two main groups under examination display radical ecological differences. The suborder Symphyta typically exhibits higher species richness in northern Europe, while the Aculeata group reaches its peak species richness in southern regions. For instance, Finland records 690 Aculeata species and 769 Symphyta species, a ratio of approximately 1:1. In contrast, Hungary reports 615 Symphyta species and 1400 Aculeata species, yielding a ratio of about 2:1. Turkey presents an even more pronounced disparity with 370 Symphyta species compared to 3382 Aculeate species, a ratio nearing 10:1 [163,164,165,166,167,168]. Consequently, the Mediterranean transformation of Central European fauna is most evident through the proliferation of Aculeata species and the gradual decline of Symphyta species. The subsequent figures (Figure 10, Figure 11, Figure 12 and Figure 13) illustrate this process, partially based on our previous paper [169].

The Supplementary Material A summarizes the changes in the 241 most common Aculeata species within the deeper areas of the Carpathian Basin, categorized by their ecotypes: eurytherm xerophile, stenotherm hygrophile (sh), eurytherm intermediate (ei), eurytherm hygrophile (eh), and stenotherm xerophile (sh). Given that approximately 85% of these species fall into the intermediate and xerophilic groups, it follows that gradual warming would lead to a significant increase in individual numbers. Several groups, however, have experienced substantial declines in abundance. One such group includes the cool and humid-adapted bees, primarily from the genera Alysson and Didineis, and to a lesser extent, some Andrena, Megachile, and Osmia. Another affected group is the bumblebees (Bombus spp.), which warrant separate discussion due to their economic importance and unique responses. Additionally, numerous Crabronidae species inhabiting wet marshes or wooded areas are also in decline. In this way, the families and genera of the Aculeata respond differently to climate change. We can say that up to the current extent of warming, the reaction is positive, except for two genera, namely Crabro Fabricius, 1775 and Crossocerus Lepeletier & Brullé, 1834 (Table 8).

Of the approximately 1480 Aculeata species that once inhabited the lower Carpathian Basin, 209 species have not been recorded for 20 years or more. These have been categorized into three groups. Species that have found refuge in the higher elevations of the Tatras or the Carpathians are indicated in grey in Table 9. Species absent due to rarity have been excluded from our list. The remaining species have disappeared from our region but have been recorded in Scandinavia, the northern parts of Western Europe, the Atlantic region, and the Alps. The primary distribution of these species serves as an indicator of climate sensitivity, suggesting that these species may have shifted their southern range limit further north, thereby reducing their abundance below the detection limit in our region. The Coefficient of Determination of the temperature trend (tR²) value for all Aculeata (in terms of changes in total abundance) is 0.5071, including hylophilous and xerothermic groups in total. It means that this group is dominantly eremophilous-xerothem, which is the reason Aculeata reaches its highest species richness and population densities in the Mediterranean, tropical, and subtropical areas (see details in the discussion part). We specifically examined the determination coefficient (tR²) of the temperature trend for hylophilous species to determine the extent to which the effect of average temperature increase on the population abundance decline of cold and moisture-loving species can be justified. We found a strong negative correlation: thermal det coeff: tR² = 0.752 and linear regr. det. coeff.: r² = 0.715.

3.2.3. Sawflies, Symphyta

Due to their northern distribution, it is not surprising that climate change affects this group significantly. Using Malaise trap methods, we have successfully demonstrated this decline [6], as reflected in Figure 13 and Figure 14. Regular data from sweep net collections have been available since the 1960s. Although this method collects few specimens (300–700 per year). Thus, the decline in collected specimens is masked by variations in different habitats, yet the decreasing trend in species that were once common is still noticeable (Table 10).

Most Tenthredo species show a strong declining trend, as does the once-common Pachyprotasis rapae (Linnaeus, 1767). It is not surprising that the number of specimens in the Argidae family, which is primarily characteristic of the Mediterranean (and also the Afrotropical region), is increasing. Among Symphyta, several agricultural and horticultural pests belong to this family. Our data indicate that the current rate of warming has positively impacted the Athalia rosae (Linnaeus, 1758) (Turnip Sawfly) and two Dolerus species that occasionally damage cereals and grass crops. The drastic decline in the number of Athalia ancilla Serville, 1823, began before climate change became noticeable. Additionally, it appears that climate change is also unfavorable for the Cephus pygmeus (Linnaeus, 1767) (Wheat Stem Sawfly).

The mountainous regions surrounding the Carpathian Basin still provide refuge for previously sporadic species [6].

The measured Coefficient of Determination of the temperature trend (negative trend) (tR²) value for all Symphyta (in terms of changes in total abundance) is 0.8212, which is a very high value. It is closely related partly to the fact that sawflies reach their highest abundances in the northern regions and at high altitudes (see details in the Discussion part). For sawflies, the Malaise trap method (see the table in our previous article [6]) was re-evaluated here. The p-value is very low, almost equal to zero. It is 3.129 × 10⁻¹⁰ using the Pearson method. The p-value indicated by the Spearman method is 0.0006799. This indicates a very strong statistical correlation between the two series (abundance change and temperature change). The difference between the two methods is natural, as the Pearson method measures the linear relationship, while the Spearman method measures rank correlation.

One series represents the annual total number of caught Symphyta, and in the absence of local temperature recordings, the annual average temperature of Slovakia (where the Malaise trap surveys were conducted).

3.3. Lepidoptera

3.3.1. Butterflies, Rhopalocera

Table 11 and Table 12 show the changes in the number of individuals of certain species of butterflies between 1980 and 2023. The correlation between the Species Temperature Index (STI), Species Precipitation Index (SPI), Habitat Suitability Index (HSI) values, and changes in the number of individuals of each butterfly species is very weak for HSI (0.075), STI (0.038), and SPI (0.058), but stronger for Climate Risk Category (CRC) values. When the CRC (1980–2009) values are ranked from 1 to 6 (1: HHHR—extreme high risk, 6: PR—potential risk), and the most resistant species are assigned a value of 6, the correlation coefficient of 0.35 indicates a significant connection (Table 11). Conversely, the correlation between the calculated CRC (2016–2023) values (Table 12) and the strength of the change in the number of individuals (lin x correlation coefficient) in the national park is only 0.034, indicating no real correlation. This suggests that the relationship between the decrease in the number of butterfly species and climate change exists and, in our case, is of medium strength (Table 11). On the other hand, the increase in more or less climate-resistant species is a result of nature conservation measures rather than climate change (Table 12 and Table 13).

Where we have observed a strong increase in certain species, we can infer that these species have shown resilience to the extent of climate change to date, but other conditions were necessary for their increase, such as protected habitat, a network of core areas and ecological corridors, prohibited or strictly controlled pesticide use, and mowing regimes adapted to the species’ lifestyle. Therefore, sensitivity to climatic conditions is becoming more pronounced over time and is strongly reflected in the numbers of many butterfly species. Furthermore, the reproduction of species less sensitive to climatic variation requires the aforementioned conditions.

These findings are described in more detail below. Less climate-sensitive species have experienced a strong increase in the last 10 years, as listed in Table 11 and Table 14. In contrast, the populations of sensitive species have declined since the mid−1980s (e.g., Polyommatus daphnis (Denis & Schiffermüller, 1776) (Meleager’s Blue), Lycaena hippothoe (Linnaeus, 1761) (Purple-edged copper); or mostly from the early 90s Zerynthia polyxena (Denis & Schiffermüller, 1775) (Southern Festoon), Melitaea diamina (Lang, 1789) (False Heath Fritillary), Heteropterus morpheus (Pallas, 1771) (Large Chequered Skipper), Apatura iris (Linnaeus, 1758) (Purple Emperor), Nymphalis antiopa (Linnaeus, 1758) (Camberwell Beauty) and Aglais urticae (Linnaeus, 1758) (Small Tortoiseshell) or have suffered from an intense decline in numbers from the second half of the 1990s onwards (e.g., Brenthis hecate (Denis & Schiffermüller, 1775) (Twin-spot Fritillary) (Table 12, Table 13 and Table 15). Among the listed species, climate change plays a major role in the decline of almost all, with the exception of N. antiopa, which is classified as LR (low risk) and only slightly sensitive to climate change.

Aglais urticae, once common until the late 1980s and then extremely rare by the early 1990s, is now absent from most records (Figure 15). Since the early 2000s, it has been largely absent from surveys, now a distinct rarity in the deeper areas of the Carpathian Basin.

The other group includes species that have been resistant to climate change thus far, showing a significant increase in density since the early 2010s (Table 12 and Table 13). This increase occurred only under conservation protection, excluding other factors affecting their natural abundance but not mitigating the impact of climate change. Species such as Maniola jurtina (Linnaeus, 1758) (Meadow Brown), Minois dryas (Scopoli, 1763) (Dryad), Melanargia galathea (Linnaeus, 1758) (Marbled White), Brintesia circe (Fabricius, 1775) (Great Banded Grayling) (Figure 16), Polyommatus icarus (Rottemburg, 1775) (Common Blue), Aricia agestis (Denis & Schiffermüller, 1775) (Brown Argus) and Coenonympha glycerion Borkhausen, 1788 (Chestnut Heath) were notable for both the intensity of the increase in numbers (slope of the trend line) and the strength of the trend (r²), alongside several other species (see Table 12). The species that declined in the 1990s and 2000s have not been able to increase their numbers despite conservation efforts, suggesting that climate change has most likely played a critical role in their decline.

At the family level, gossamer-winged butterflies (Lycaenidae) have been the most affected by climate change: our studies show that most species are below detection levels, their population density compensated by some common species, while populations in total abundance of rare and sporadic species have drastically reduced. A similar pattern is seen in the Nymphalidae family, where the decline of the vast majority of species was offset by a few species formerly belonging to the Satyridae, listed above.

Overall, climate change has had a negative impact from a conservation perspective for a significant proportion of species. Rare or sporadic butterfly species have drastically declined since the 1990s. Conservation measures have ensured that, with the exception of a few species, butterflies have not yet disappeared from the Carpathian Basin. In protected habitats, significant reproduction has been achieved, but this increase predominantly affects species that were already common. The vast majority of species that were once rare or sporadic are expected to fall victim to climate change in Central Europe, and conservation measures alone will not be sufficient to protect them.

Under optimal, protected conditions, only certain species from the families Papilionidae (Figure 17), Nymphalidae, and Lycaenidae were able to increase their numbers, ensuring an upward trend for these families (Figure 13). However, families like Hesperidae and Pieridae did not show a positive trend even under these conditions (Table 12, Table 14 and Table 15). The number of species collected in each period showed a slight downward trend (Table 13 and Table 14). This decrease is not only local. In the last 20 years, seven species have disappeared (or fallen below the detection limit), while another three species are considered historically extinct, even before climate change. These are: Carcharodus lavatherae (Esper, 1783) (Marbled Skipper), Coenonympha tullia (Müller, 1764) (Large Heath) and Melanargia russiae (Esper, 1783) (Esper’s Marbled White). Nine of the 10 species find habitat in the mountain ranges surrounding the Carpathian Mountains (Tatras, Carpathians) up to a given extent of climate change (Table 16). The species Polyommatus damon (Denis & Schiffermüller, 1775) (Damon blue) has not been collected in the lower regions of the Carpathian Basin for 10 years, so it, like the previous species, persists in higher regions in our region. Overall, the temperature related Coefficient of Determination (tR²) value = 0.1088 (Kaposvár, Museum, selected sporadic and rare species) and tR² value = 0.5884 (Danube.Ipoly National Park, all butterfly species). This value suggest, although the climatic factor is well represented, however, decline of butterflies is a polycausal phenomenon and the climatic change manifests indirectly (for instance, we assume that, some butterflies (overwintering in imaginal phase) starts to fly during mild late winters and then be killed by subsequent frosts (due to irregular weather conditions). However, it is only an assumption that requires further evidence and investigations. The other values have a strong good correlation, but this is very misleading, and this is where the importance of the rational evaluation of statistical results comes in. This increase is primarily due to conservation measures, not climate change.

3.3.2. Nocturnal Macrolepidoptera

Similar to the vast majority of local faunistic data and regional reports, we found a decline in numbers, but it was so slight that it was not significant, and no significant r² value was measured (Table 17 and Figure 14). Compared to the 1970s, that slight decrease in the number of specimens caught (Figure 18) turned into a slight increase from 2014. At the same time, the decline in species richness (number of species below the detection limit in a given area) is decreasing slightly but showing a real trend (r² = 0.14) (Table 17 and Figure 19). By family, the Noctuidae (owlet moths) are the most sensitive in terms of numbers, and the decline in this group has not stopped even in the last decade. Other families, especially the Geometridae, have experienced a less significant decline in numbers compared to 1970. The Coefficient of Determination of thermal sensitivity (tR²) is 0.0051. It is practically zero. This is due to the fact that circa a quarter of the species show positive responses, another quarter show negative responses, while the majority of species remain in the neutral range in terms of aboundance changes. For nocturnal moths, the p-value (related to change of abundance of the total group) is very high, 0.7279 for the Pearson method and 0.6007 for the Spearman method. This also reinforces the evaluation of the two types of correlation coefficients. Specifically, it indicates that the very distinct response of nocturnal moths to climate change is not reflected in the change in population size but rather in parameters assessed in this article (such as changes in dominance).

In terms of moth species captured by individual traps (Figure 20), local species depletion has been most pronounced in the Geometridae family. Meanwhile, in the Noctuidae and Erebidae families, species diversity captured by individual traps has increased (Table 18). A strong increase in the total number of species studied was observed in Peribatodes rhomboidaria (Denis & Sch., 1775), Noctua pronuba (Linnaeus, 1758), Phyllodesma tremulifolia (Hübner, 1810), Paracolax tristalis (Fabricius, 1794), Hyloicus pinastri (Linnaeus, 1758), Zanclognatha lunalis (Scopoli, 1763), Euphyia biangulata (Haworth, 1809), Catocala hymenaea (Denis & Schiffermüller, 1775), Eilema sororcula (Hufnagel, 1766) and Hypomecis punctinalis (Scopoli, 1763). These are believed to be the most adaptive to the extent of climate change to date. The most sensitive species were Chiasmia clathrata (Linnaeus, 1758), Smerinthus ocellata (Linnaeus, 1758), Phragmatobia fuliginosa (Linnaeus, 1758), Pseudeustrotia candidula (Denis & Schiffermüller, 1775), Spilosoma lubricipeda (Linnaeus, 1758), Ascotis selenaria (Denis & Schiff., 1775), Rivula sericealis (Scopoli, 1763), Mamestra brassicae (Linnaeus, 1758) and Arctia caja (Linnaeus, 1758) (Supplementary Material B).

From a practical point of view, certain agricultural pests, Agrotis segetum (Denis & Schiffermüller, 1775), Agrotis exclamationis (Linnaeus, 1758), and Lacanobia oleracea Linnaeus, 1758) have not been sensitive to climate change to date and have been found to be tolerant of climate change. Population fluctuations of these agricultural pests were close to equilibrium. Moderately significant declines were observed in abundances of the Xestia c-nigrum (Linnaeus, 1758), and Hyphantria cunea (Drury, 1773), significant declines in population aboundances of the Cabbage Moth: Mamestra brassicae (Linnaeus, 1758) and moderate increases in population aboundances of Autographa gamma (Linnaeus, 1758) and Helicoverpa armigera (Hübner, 1808). The latter two species are migrant moths from the south (Supplementary Material B). Helicoverpa armigera has produced such outbreaks in certain areas in recent years that the population growth in aboundance can be approximated with an exponential trend with the same r2 value as the linear trend.

Among forest pests, an increase was observed in Pheosia tremula (Clerck, 1759) and Panolis flammea Denis & Schiffermüller, 1775. Nycteola asiatica (Krulikowsky, 1904), Agrotis vestigialis (Hufnagel, 1766), Bupalus piniaria (Linnaeus, 1758), Leucoma salicis Linnaeus, 1758, Clostera anastomosis (Linnaeus, 1758), Euproctis chrysorrhoea (Linnaeus, 1758), Thaumetopoea processionea (Linnaeus, 1758), Malacosoma neustria (Linnaeus, 1758) and Lymantria dispar (Linnaeus, 1758) did not show any trend-like variation in positive or negative directions, i.e., they were not affected by the warming up to date. In practice, this means that climate change has had a positive or neutral effect on these forest pests. The increasing prevalence of outbreaks can be examined in Supplementary Material B.

In the beginning and end of the 50 years studied, all but 5 of the top 20 species with the highest abundance were replaced, which means that climate change has rearranged the dominance of each species almost completely (Table 16). If we take the 20 most common species, Miltochrista miniata (J. Förster, 1771), Orthosia cruda Lempke, 1964, Lithosia quadra (Linnaeus, 1758), Eilema sororcula (Hufnagel, 1766) and Polypogon tentacularia (Linnaeus, 1758) have remained in the dominant 20 species, as they did in the 1970s and 1980s as 30–50 years later in the 2010s and 2020s.

Codes of Table 19: 0 = xerothermophilic species (burning, semi-dry grassland, dry fallow land, heaths, gravel pits, quarries); 1 = ubiquitous open land species and cultural followers in fields, meadows, gardens, parks and on forest edges; caterpillars almost exclusively on grasses and herbs; 2 = ubiquitous species and cultural followers in open land and forest biotopes; caterpillars on grasses, herbs or shrubs; 3 = species mostly in tree-covered areas; common in forests, but also in gardens or parks with good tree cover; caterpillars predominantly on grasses, herbs or shrubs; 4 = like group 3, but caterpillars predominantly on hardwood and coniferous wood; 5 = like group 3, but more oriented towards softwood floodplains; caterpillars predominantly on softwood; 6 = forest species; caterpillars mostly on wood, rarely on herbs or grasses; 6a = ubiquitous forest species, 6b = coniferous forest species, 6c = deciduous forest species; 7 = species of moist forests and open wetlands; caterpillars predominantly on algae, lichens and liverworts; 8 = as in group 7, but caterpillars predominantly on deciduous wood (mainly hygrophilous floodplain, marsh and moor forest inhabitants; 9 = as in group 7, but caterpillars on grasses and herbs (mainly sedge meadows and reed beds inhabitants); 10 = typical floodplain forest species of larger river valleys; caterpillars predominantly on soft deciduous wood or clematis).

Regarding changes in dominant moth species, we observed the following: no extremely xerophilic species dominated in any period. The extreme hygophilous species, Rivula sericealis (Scopoli, 1763) and Caradrina morpheus (Hufnagel, 1766), have been eliminated from the top 20 species. The proportion of species native to open grasslands has declined, indicating their sensitivity to climate change. Their place has understandably been taken by other species in more closed habitats, leading to a shift in favor of species living in habitats more protected from climate exposure.

Table 16 lists species that have been missing for 20 years or more. Species marked with an X are still found in the higher regions of the Carpathian Basin (Tatras, Carpathians). Most nocturnal lepdoptera species from low-lying regions disappeared several decades ago, before the 1980s. Only three species from the list are suspected to have been affected by climate change: Cosmotriche lobulina (Denis & Schiffermüller, 1775), Eublemma pannonica (Freyer, 1840), and Syngrapha ain (Hochenwarth, 1785).

3.4. Non Native Insects and Mediterranean Influx

3.4.1. Non Native Species

The most comprehensive work on our region and Europe to date is the Biorisk series monograph titled “Alien Terrestrial Arthropods of Europe” [75], which lists about 330 insect species from the studied region. Through deep research, we have expanded this list to 803 insect species by carefully reviewing faunistic reports and the literature. Our chart shows that the influx of alien species is accelerating, and the dynamics of the influx of established alien insects can be described by an exponential equation (Figure 21).

The first alien insect species likely appeared in the Carpathian Basin with Homo erectus, as parasites living on the human body. The first major invasion wave occurred in the Neolithic era when humans brought various livestock, such as sheep or goats, along with the first invasive arthropods of veterinary importance. The introduction of agriculture also brought the first storage pests from the Eastern Mediterranean [78]. During antiquity, with Roman conquests, the first exotic animals, like peacocks and pheasants, likely introduced new arthropods (their lice, ectoparasits); also those associated with the European rabbit, which was bred by the Romans.

Medieval records of invasive insects primarily document locust swarms, with the earliest record from 1191. The 1346 plague, which reached the Carpathian Basin in 1349, is often attributed to invasive rats and their fleas. Transoceanic insect invasions likely began with maritime exploration and geographic discoveries, bringing exotic animals (like guinea pigs) and food and fodder plants (tomatoes, peppers, corn, tobacco, etc.). However, early entomological literature, such as the richly illustrated manuscript “Mira Calligraphiae Monumenta” from 1590, did not depict any alien insect species.

After Linnaeus, researchers in our region turned their focus toward insects. The first description of an invasive species from our area, published by Mátyás Piller and Lajos Mitterpacher, was Epauloecus unicolor (Piller & Mitterpacher, 1783), a foreign warehouse pest from the Dermestidae family. As international trade increased, so did the number of reports and descriptions of new alien species. This accumulation has become exponential over the past quarter-century, with climate change playing a significant role, according to the literature.

In terms of distribution, approximately half of the non-native insect species came from the Eastern Palaearctic (144 species, 18%) and North American regions (Nearctic, 135 species, 17%). We will not discuss these in more detail, as their spread was already ensured earlier due to climatic similarities, and the main driver here is the increasing logistics. The milder winters have contributed to the influx of Mediterranean, tropical, and subtropical species, accelerating their invasion (Figure 22) and allowing many species to survive and proliferate to detectable levels.

The majority of incoming species, approximately 38% (Figure 23), belong to the order Hemiptera (true bugs, cicadas, aphids, planthoppers, leafhoppers, assassin bugs, scale insects, bed bugs, and shield bugs), with scale insects constituting the majority (83 species). Due to their small size (especially as overwintering larvae), they can easily evade plant health checks, whether on propagating material, ornamental plants, or the skins of imported fruits. Some have only been able to reproduce in greenhouses until now, but increasingly, thanks to milder weather, they can survive outdoors, such as Ceroplastes ceriferus (Fabricius, 1798), which has been detected in the open field. Whether it can overwinter in our region remains to be seen, but it is likely. Originally, Nezara viridula (Linnaeus, 1758) also occurred exclusively in greenhouses, but it has now become a significant outdoor pest.

The second largest group is beetles (Figure 23) due to their very high diversity in nature. Many beetle species are warehouse pests and are spread worldwide through transportation, making the original habitats of many species difficult to trace. An example of an invasive beetle is the Harmonia axyridis (Pallas, 1773) (Asian Lady Beetle), introduced as a biological control agent, which is now displacing native ladybird species. Another alien ladybird species, Cryptolaemus montrouzieri Mulsant, 1853, is polyphagous and may threaten non-damaging non-target fauna.

Other beetle species associated with crops and seeds include Megabruchidius dorsalis (Fahraeus, 1839), Megabruchidius tonkineus (Pic, 1904), Murmidius ovalis (Beck, 1817), and Bruchidius siliquastri Delobel, 2007. Beetles associated with woody plants include Lyctus brunneus (Stephens, 1830), Rhynchophorus ferrugineus (A. G. Olivier, 1791), Xylosandrus germanus (Blandford, 1894), Apate monachus Fabricius, 1775, and Coccotrypes dactyliperda (Fabricius, 1802). These beetles have been introduced into European ports and easily enter the Carpathian Basin, acting as forest or crop pests. Dermestes haemorrhoidalis Küster, 1852, is known for infesting mummified human corpses.

Certain tropical and subtropical species, such as Tuta absoluta (Meyrick, 1917), Hipoepa fractalis (Guenée, 1854), Monema flavescens Walker, 1855, Phthorimaea operculella (Zeller, 1873), Spodoptera littoralis (Boisduval, 1833), Apomyelois ceratoniae (Zeller, 1839), and Cacyreus marshalli Butler, 1897, may have increased in prevalence due to climate change. According to internet sources, Cacyreus marshalli has already reached Zagreb. This non-native butterfly lives on Pelargonium and is a synanthropic species, similar to many invasive organisms that spread during the Neolithic period. Notably, the colonization of Paysandisia archon (Burmeister, 1879), a member of the Castniidae family (Lepidoptera) previously unknown in our region, is significant. The larvae of this large moth of South American origin can overwinter inside the ornamental plants, and some of its host plants can survive outdoors in the Carpathian Basin climate.

In the 2000s, two heat-demanding tropical and northern Mediterranean cockroach species, Periplaneta australasiae (Fabricius, 1775) and Planuncus tingitanus (Bolívar, 1914), became widespread in our region.

Economically, Hierodula tenuidentata Saussure, 1869 (Southeast Asian Mantis), poses a minimal threat but may carry nature conservation risk. Dragonflies, due to their excellent flying ability, can naturally colonize northern areas from the Mediterranean. However, one species, Trithemis arteriosa (Burmeister, 1839), was introduced by humans from Cyprus through the import of aquatic plants, likely arriving with dragonfly larvae.

Among the alien species, Xenylla uniseta Da Gama, 1963, was the first springtail recorded in the Carpathian Basin. Within the primitive insect groups, Ctenolepisma longicaudatum (Escherich, 1905), a relative of the European silverfish, Lepisma saccharinum Linnaeus, 1758, is noteworthy.

Since 2004, alien hymenopteran species permitted for agricultural use (Ministerial Regulation on Biological Control, No 89/2004, FVM Decree) have been spreading. Some species, such as Torymus sinensis Kamijo, 1982, introduced for biological control of Dryocosmus kuriphilus Yasumatsu, 1951 (chestnut gall wasp), are now found in the wild [149].

Among Diptera, larvae Hermetia illucens (Linnaeus, 1758) have been used in organic waste processing, with potential genetic modifications to expand their waste consumption range. Rhynchomicropteron nudiventer Papp, 1982, described by Hungarian researcher László Papp from India, was caught in Hungary over three decades after its description. Clogmia albipunctatus (Williston, 1893) became a mass species after 2012, frequently found in restrooms and sanitary facilities, gaining hygienic significance. In hospitals, as a passive vector of various bacteria, it poses a risk to patients when colonizing operating theaters.

Aedes albopictus (Skuse, 1894) (Tiger Mosquito) is a vector of arboviral pathogens such as West Nile virus, yellow fever virus, encephalitis, dengue fever, Chikungunya fever, Zika virus, and several nematodes (e.g., Dirofilaria immitis).

3.4.2. Mediterranean Influx

According to our results, of the 803 alien insect species counted so far (Supplementary Material C), 22% are of Mediterranean origin, and 21% are from one of the tropical regions. The quantity of species imported from the Nearctic (17%) and temperate Asian (Eastern Palaearctic) region (18%) is also significant (Figure 24). The most difficult thing to decide about Mediterranean invasive species is whether they have arrived in Central Europe naturally as a result of climate change (we are neighbours of the Mediterranean region) or whether they have been affected by human intervention. There are a few cases where human conscious or accidental transport is documented. For example, several Mediterranean species have been introduced and spread for biological control purposes, such as Coenosia attenuata Stein, 1903, Diptera, Muscidae (predator fly), Orius laevigatus (Fieber, 1860), Heteroptera, and Anthocoridae (predator bug).

It is quite likely that it has been introduced by the Juniper Ermine Moth: Argyresthia trifasciata Staudinger, 1871, Pyralidae, (with dried food) or Phyllonoricter millierella Staudinger, 1871, Gracillariidae (with propagules of the Celtis australis). It is not surprising then that there are species that have been introduced both naturally and artificially, i.e., more than once, such as the Ovalisia festiva (Linnaeus, 1767) (Cypress Jewel Beetle).

Here, we focus mainly on those species, from the last quarter of a century alone, that may have spread without human intervention, simply following weather trends.

In this group, there is so-called internal dispersal, where the range of a species is limited to the Carpathian Basin, and within this range, it can naturally spread from the southern countries to the northern countries. Thaumetopoea pityocampa (Denis & Schiffermüller, 1775) (Notodontidae) was recorded in Hungary as early as 1977, but it reached Slovakia by 2011.

The northern limit of the distribution of the Mediterranean Pseudocephaleia praeteritorum (Semenov, 1934) (Pamphiilidae) has been Hungary, from where it has been known for a long time, but was collected in Slovakia for the first time in 2019. Hyles vespertilio (Esper, 1779) first entered our region as a migrant in 2020 and has been observed several times since then. In such cases, the area boundary fluctuates depending on climatic conditions. A good example is Choreutis nemorana (Hübner, 1799) (Choreutidae), which feeds on fig leaves and appeared in 1955 in the southern Carpathian Basin, receded and reappeared in 2011 due to warming. But it also includes Tenthredo costata Klug, 1817 (Tenthredinidae), a species common in the Mediterranean and Anatolian regions, sometimes dominant in some places, which occurred sporadically in Hungary in the 1970s, then disappeared and was recaptured on 8 June 2016 at the Nagybajom woodcutter. These kind of species can be identified by the fact that they are frequent or even common in the Mediterranean, while they are rare in our country and absent in areas further north, since we are on the border of their distribution area, sometimes they disappear sometimes appear as the “border” of their area is fluctuating. There is relatively little information on the insect species that move northwards, partly because they do not receive the same attention as agricultural and forest pests and partly because there are very few specialists collecting, identifying, and publishing them. Based on the available data for the Carpathian Basin, the following can be said about the last quarter of a century. Ectobius vittiventris (Costa, 1847) is a species of cockroach that is distributed northwards and is otherwise harmless. Of the two species of praying mantis, Ameles spallanzania (Rossi, 1792) has reached the central parts of the Carpathian Basin, while Iris oratoria (Linnaeus, 1758) has conquered the eastern part of the Carpathian Basin. A Neuroptera, Sisyra iridipennis Costa, 1884 and three Mediterranean dragonfly species, the Erythromma lindenii (Selys, 1840), Selysiothemis nigra (Vander Linden, 1825) and Trithemis annulata (Palisot de Beauvois, 1807), also reached the north. The host plants of three scale insects of Mediterranean origin, Aonidia lauri (Bouché, 1833), Carulaspis minima (Signoret, 1869), and Dynaspidiotus britannicus (Newstead, 1896), can also survive in the wild and may have spread spontaneously or been introduced by plants. The same is true for Livilla variegata (Löw, 1881), Trioza alacris (Flor, 1861) and Uroleucon telekiae (Holman, 1965). The Mediterranean Acrosternum heegeri Fieber, 1861, which is a polyphagous bug, has been on the path to invasion, and its appearance in the Carpathian Basin is likely to have been by introduction, but spontaneous colonization can not be excluded. Entomobrya unostrigata Stach, 1930 is a springtail of Mediterranean origin. Silba adipata McAlpine, 1956 (Black Fig Fly) has been introduced into our region as a pest of figs this year, as its food plant has recently been able to ripen to sweetness in increasing areas throughout the Carpathian Basin. Anacridium aegyptium (Linnaeus, 1764) (Egyptian locust) has been introduced into our region only occasionally, as stray specimens, but has been seen from time to time for a few years. The two leafroller moths, Cydia interscindana (Möschler, 1866) and Cacoecimorpha pronubana (Hübner, 1799), may have been introduced directly from the Mediterranean region or even by plant detour from elsewhere. Helicoverpa armigera (Hübner, 1808), for example, was extremely rare in Hungary until the mid-1980s. In a quarter of a century, the more than 20 traps of the Forestry Light Trap Network caught only 4 specimens. After that, it became increasingly common, with some traps now catching thousands of the species annually.

Ovalisia festiva (Linnaeus, 1767) (Buprestidae) has been introduced to Hungary twice (Figure 25). A natural immigration into the Barcs Juniper Woodland Nature Conservation Area was followed about 10 years later by the destruction of evergreen ornamental plants by another population introduced by Italian and Dutch imports. While several mud-dauber wasps species of tropical origin and one Mediterranean species (Sphecidae: Sceliphron madraspatanum (Fabricius, 1781) Sceliphron curvatum (F. Smith, 1870) and Sceliphron caementarium (Drury, 1773)) invaded the Carpathian-Basin, the native species, including the most common Sceliphron destillatorium (Illiger, 1807), were largely suppressed. (Supplementary Material C). The accelerating rate of influx of insect species of Mediterranean origin is shown in Figure 26.

4. Discussion

4.1. Diptera

4.1.1. Bombyliidae

In the literature, decades-old Bombyliiidae data are not available; however, the available ecological observations support our findings. Bombyliid species are typically and most frequently encountered in arid areas and constitute a high percentage of the diversity of flies in the more or less desert regions of the earth [170]. Adult bee flies are frequently the primary pollinators of a wide variety of flowering plants, particularly in arid environments [171]. Their hosts are typically insects from warm, dry areas such as Acrididae, Formica spp., Myrmeleontidae, Tenebrionidae, Sphecidae, as well as Noctuidae, Pyralidae, Gelechiidae, Scarabaeidae, etc. 63% of the world’s species, including the most common ones, parasitize xertolerant hosts like bees and wasps, locusts, and grasshoppers [172]. Since these animals mostly have xerothermic ecological requirements, we have not found a decrease in their population numbers according to the current extent of climate change. At present, they are among the winners of climate change. No similar data sets have been published by others. Herrera studied the population dynamics of pollinators in the Mediterranean for over 20 years [173]. Among the observed species, a Bombyliidae genus, Hemipenthes (Bombyliidae), exhibited a significant increase in aboundance.

4.1.2. Horse-Flies, Tabanidae

There is no time-series analysis available in the literature for this group, yet the work of Havkenberget and colleagues [174] is interesting from this perspective because they describe and prove that both species richness and individual density are dependent on different ecotypes. Studying the Tabanid fauna on both sides of the Velika Kapela mountain range (Croatia), they found that species occupied either the southern slope (Mediterranean zone) or the northern slope (Continental zone) according to their ecotype and similarly occupied different altitudinal zones. In our study, the Tabanids responded differently to changing climatic conditions rather than to slopes and altitudinal zones, depending on the ecotypes they belong to. Herczeg, T. and colleagues [175] also examined ecological and climatic factors in terms of Tabanid distribution and proliferation. They studied six species and observed that the 31–35 °C range was when flight activity was most vigorous, i.e., the traps captured the most individuals. As a warmth-loving species, we also observed increasing captures in most species following the warming trend. Haematopota subcylindrica Pandellé, 1883 is a moisture-loving forest species [37]. In our case, the population size of this species showed a downward trend, while other xerothermic species showed a decline according to their ecological requirements. Climatic changes do not justify the decrease in the population size of Tabanus bromius Linnaeus, 1758. This decline was first observed in Great Britain, where it was found that it is now a local species in Britain that has declined substantially, possibly as a result of the widespread ploughing and destruction of meadows [176].

4.1.3. Hoverflies, Syrphidae

Hallmann et al. [177] found a decrease of the number of individuals of hoverflies of 89% and a decrease in species richness of 23% over a 26-year period (1989–2014) for a river valley in Germany. Barendregt et al. [178] reported similar results, a decrease of 80% in the number of individuals of hoverflies over 40 years (1982–2021) and 44% of the species over 43 years (1979–2021) for hoverflies in a forest in the Netherlands. Concerning migratory hoverflies, Gatter et al. [179] report a decline of individuals of 90–97% over 40–50 years (based on transect counts 1970–2019 resp. Malaise traps 1980–2019). Finally, comparing the period 2008–2022 with 1900–1969, Reemer et al. [180] found as many species of hoverflies with a declining trend (147 species) as with an increasing trend (146 species). These studies describe a strong decline of both the number and the species richness of hoverflies since 1980 in western Europe. The current (1993–2017) extinction rate for hoverflies is estimated to be nearly five times higher than the historical rate (1942–1992) [181]. Longer warmer and drier periods will harm most hoverflies, but especially the aphidophagous larvae of zoophagous species because aphids are highly sensitive to drier conditions. In addition, there may be a disconnect in phenological timing between aphid and hoverfly life cycles. Warmer winters can make it harder to overwinter successfully in a dormant state, particularly if breaking diapause requires a cold spell. Also, potentially, warmer winters could promote fungal infections or increase predation during the winter [182].

4.1.4. Tachinid Flies, Tachinidae

Zeegers [183] examined six Tachinidae species in the Netherlands for climate sensitivity. The tachinids Brullaea ocypteroidea Robineau-Desvoidy, 1863, Entomophaga exoleta (Meigen, 1824), Istocheta longicornis (Fallén, 1810), Phasia aurigera (Egger, 1860), Senometopia intermedia (Herting, 1960), and Thecocarcelia acutangulata (Macquart, 1850) are recorded for the first time for the Netherlands, bringing the number of Tachinid species recorded from the country to 331. Most of these species have a southern distribution in Europe, recently extending northwards due to climate change. Similarly, many Tachinid species exhibit this northward spread: Four of these species show recent expansions in their range. The discovery of Leucostoma abbreviatum Herting, 1971, near Templin provides the first records for this species in the province of Brandenburg, and Elomya lateralis (Meigen, 1824) has been found for the first time in Sachsen-Anhalt, Brandenburg, and Mecklenburg-Vorpommern. Both species were known only from the south and have now been found in northeast Germany, more than 600 km north of the previous limit of their distribution. Two other bug parasitoids, Phasia barbifrons (Girschner, 1887) and Phasia aurigera (Egger, 1860), are currently expanding their range north from southern and central Germany [184]. Many species of Tachinid flies possess relatively large host ranges, and the frequency of parasitism by tachinids exhibits no relationship with climatic variability among the data sets that estimated their frequency. Thus, the population changes in the total abundance of Tachinids are unpredictable [184]. Our findings also support this uncertainty. While many species show a steady decline, we observed a significant increase at the end of the study period compared to the beginning in species such as Linnaemya frater (Rondani, 1859), Meigenia dorsalis (Meigen, 1824), and Exorista larvarum (Linnaeus, 1758). The family should not be treated as a single ecological unit. Each species has different ecological needs, and various ecotypes are present within the family. Despite the declining trend, we see that most species have been able to adapt to the changed environmental conditions. We must also acknowledge that under changing conditions, the population size of the Tachinidae species is indeed unpredictable (Table 8 and Table 9). Thus, our detailed data sets reinforce the opinion of Stireman and colleagues [185].

4.2. Hymenoptera

4.2.1. Bumblebees

A remarkable observation was made by Biella et al. [186], which explained why, contrary to expectations, bumblebee numbers are decreasing in high-mountain areas: “In high-mountain areas, climate is changing faster than the global average. This warming is harmful because it accelerates the metabolisms of ectothermic organisms and increases the activity of harmful fungi and parasites, impacting survival and fecundity in different taxa, including cold-adapted bumblebees.” In the Italian Alps, an increase was observed [187] in bumblebee numbers at altitudes of 1400–2000 m a.s.l. from 1994 onwards. They studied four species: Bombus alpinus (Linnaeus, 1758), Bombus mendax Gerstaecker, 1869, Bombus monticola Smith, 1849, and Bombus mucidus Gerstaecker, 1869, none of which are typical of our region. This may explain why our results align with those of Biella et al. [186].

In Hymenoptera, such as bees and wasps, climate change has been linked to changes in population densities and community composition. Bumblebee populations in total abundance in North America and Europe have been declining, partly due to climate-induced habitat loss and changes in floral resources [188,189]. Kerr and colleagues [188] highlighted that bumblebee species are failing to track warming temperatures, leading to range contractions at their southern limits without corresponding expansions at their northern limits. Arnóczkyné et al. [190] reported a general decline in Hungary, citing an increase in 7 species compared to 1953. From the 1980s onwards, we observed an increase in 6 species [6], but since 2005, only two Mediterranean species have shown an upward trend: Bombus haematurus Kriechbaumer, 1870, and Bombus argillaceus (Scopoli, 1763). This trend of these species is observed by other researchers from our region [191,192,193,194,195,196].

The drastic decline of bumblebees in both number and diversity in Central Europe is described in detail by Kosior et al. [197]. The typical mountain species of the Carpathians were also detected by Šima and Smetana [198] from Slovakia, similarly from our own results from the Carpathians, but the only 527 specimens (20 species) observed during the 12-year research period (2007–2019), suggest that the bumblebee population in total abundance has significantly decreased not only in the Romanian Carpathians, but also in the northern mountains of the Carpathian Basin, although high altitudes still hold a rich bumblebee fauna in relatively higher density than frequently found in the low altitudes, this is especially true for cuckoo bumblebees. From Slovakia, we have a report on the spreading of Bombus semenoviellus Skorikov, 1910. Apparently, the East-West spread of insects is not attributed to the effects of global warming. However, as the authors [199] write, certain adaptations to relatively dry and warm biotopes have been observed in this species, showing a degree of plasticity in ecological adaptations.

4.2.2. Aculeata

Similar data are scarce in the literature. In the absence of long-term data series, various models are employed to understand the relationship between Hymenoptera and climate change. These models are based on findings such as differential species responses, the necessity for wild bees to shift to higher latitudes due to temperature increases, and the importance of examining climate change effects on habitat quality [200]. Predictions are often derived from data sets of few species, obtained from the GBIF database using the SDM software packages. However, these models have limitations, as they use baselines such as RCP 4.5 and RCP 8.5 (Representative Concentration Pathways), which assume continuous warming without accounting for counter-effects like the potential shutdown of the Gulf Stream, leading to drastic cooling. Therefore, computer-based projections of Hymenoptera expansions are not always accurate, as climate change is inherently chaotic, with numerous feedback mechanisms and effects.

In the absence of extensive data, the impact of climate change on Hymenoptera can be estimated through direct observation of their activity. For example, Raider et al. [201] studied 20 species of American wild bees, comparing their temperature-dependent activity to that of Apis mellifera, thereby establishing a heat tolerance ranking for some wild bee species.

To our knowledge, only Belgium has produced a similar dataset [202]. Despite the different climatic conditions (Belgium—Atlantic, Central Europe—continental), the dynamics of wild bee populations in both regions show a similar upward trend in aboundance (Compare Figure 10 and Figure 27).

Megachilid species such as Megachile flavipes Spinola, 1838, Megachile sicula (Rossi, 1792), Coelioxys coturnix Pérez, 1884, Megachile minutissima Radoszkowski, 1876, and Osmia submicans F. Morawitz, 1870, have been studied [203], It is recognized that many species within the family Megachilidae are sensitive to warming, including several euryecious hylophyllous species. However, predicting the distribution of these species 80 years in advance is precarious, as climate change processes are not unidirectional. This complexity is why the term “climate change” is now preferred over “global warming”, reflecting the intricate feedbacks that challenge our ability to make reliable long-term predictions. On the other hand, the observation of the authors well agrees our results with hylophilous Megachilid bees. The work of Tryjanowski et al. [204] is a crucial complementary publication, as it addresses common species that are underrepresented in our data sets. They found no significant trend in the abundance of Vespula germanica (Fabricius, 1793). Climate change has not had an impact on the species’ populations in aboundance so far. This species is apparently resistant to climate change up to this date. Eusocial Vespidae were studied by Pawlikowski and Pawlikowski [205] between 1981 and 2000. Overall, the wasp populations in total abundance proved to be stable in this period. Specifically, Vespula vulgaris (Linnaeus, 1758) showed a significant increase over 20 years, from 713 to 1556 individuals; Vespula rufa (Linnaeus, 1758) increased from 8 to 23; and Dolichovespula media (Retzius, 1783) from 0 to 12. Dolichovespula sylvestris (Scopoli, 1763) showed a decrease from 11 to 6, and the Vespa crabro Linnaeus, 1758 from 212 to 182. Vespula germanica numbers stagnated at 11,540 versus 11,352 specimens. However, the European hornet has definitely increased its populations in total abundance in deciduous forests, particularly oak mixt forests, in Romania, based on our observations in the Moldova and Dobrogea regions. Overall, apart from a few forest species, climate change has had a beneficial or neutral impact on social wasps.

The observed increase in cuckoo wasps may be attributed more to the rise in their hosts than to a direct response to climate change. This is supported by Verheyde [206], who stated that after more than 80 years of the vespid wasp Euodynerus dantici (Rossi, 1790) being present, its main parasitoid, the cuckoo wasp Chrysis sexdentata (Christ, 1791), was reported for the first time in both the Netherlands (2019) and Belgium (2022). They mentioned that the main reason for these unexpected reports was the densification of available host populations driven by climate change over the past decade.

The increase in the number of Aculeata shows trends opposite to those observed in several Western European countries, particularly Germany [207]. While warming is evidently favorable for most Aculeata, it does not account for the drastic decline in the German Hymenoptera population in total abundance. This discrepancy may be explained by the fact that, for instance, in Hungary, pesticide use reached 7 kg of active ingredients per hectare of agricultural area by 1989. Following the political turn, pesticide use dropped to just 1.4 kg in 1995 [208].

4.2.3. Sawflies, Symphyta

Overall, sawflies have suffered a significant decline in our region in the last decades, being mainly a northern, Scandinavian group, with their greatest species richness in the Scandinavian areas. It can also be observed that species of the Carpathian Basin, once sporadic in the more marshy regions, now below the detection limit, find shelter in high altitudes, in contrast to species of the Diptera and Aculeata, where vertical dispersal is more limited [6]. Among the genera that produce growth, there are mainly those that reach the highest species numbers in the tropics and Mediterranean regions, these are species of the family Argidae [209]. In the Mediterranean and Anatolian regions, species of the family Megalodontesidae and Tenthredo distinguenda (R. Stein, 1885) are also the commonest in the Mediterranean region with an Anatolian distribution either [210]. The most common Dolerus species are warm-loving to some extent but are beginning to decline or even disappear in the Mediterranean region [211]. A large part of the literature deals with the climatic sensitivity of species of the families Diprionidae and Pamphiliidae [212,213]. Species of these families are not present in our region in numbers sufficient to allow their population increase or decline in total abundance to be assessed statistically. Their outbreaks have been reported from time to time, but we have not observed this during our fieldwork. Migration within the Carpathian Basin has also been observed in this group. The northernmost distribution of Pseudocephaleia praeteritorum (Semenov, 1934) was previously in Hungary, but it was found in Slovakia in 2019 [214].

4.3. Lepidoptera

4.3.1. Butterflies, Rhopalocera

Parmesan and his colleagues [1] studied the northward shift in the areal distribution of butterfly species in the British Isles. Our results are challenging to compare due to the minimal number of common species. Northward shifts have been observed in Parnassius mnemosyne (Linnaeus, 1758), Glaucopsyche alexis (Poda, 1761), Boloria selene (Denis & Schiffermüller, 1775), Argynnis niobe (Linnaeus, 1761), and Apatura iris (Linnaeus, 1758). The distributions of Lycaena virgaureae (Linnaeus, 1758) and Limenitis reducta (Staudinger, 1901) have not significantly changed. Brenthis ino (von Rottemburg, 1775), however, has spread southwards. In Central European conditions (Table 11 and Table 12), the abundance of Boloria selene increased significantly. The abundance of the Argynnis niobe is relatively stable but fluctuating. In contrast, Glaucopsyche alexis, Brenthis ino, and Apatura iris have significantly declined, aligning with Parmesan’s findings in Central Europe [1]. The populations in abundance of Lycaena virgaureae and Limenitis reducta showed stability. Our results corroborate Parmesan et al.’s conclusions [1] regarding the climate sensitivity of butterfly species. Exceptions include Boloria selene, which is classified in the lowest climate sensitivity category (PR) [68], and species of the Papilionidae family, including Parnassius mnemosyne (Linnaeus, 1758) (Figure 28), which appear stable in our region, with some species showing an increase.

According to the Environmental Statistics and Reporting team at the Department for Environment, Food & Rural Affairs [215], of the 26 butterfly species surveyed, 7 species increased, 12 decreased, and the rest remained statistically unchanged. Similarly, in our results, of the 33 species surveyed, 6 species increased, 15 decreased, and the rest remained unchanged (Table 11). The two studies had few common species due to differences in zoogeography. Among the 5 common species, Boloria selene exhibited opposite trends, increasing in the Carpathian Basin and decreasing in the British Isles. Apatura iris decreased in the Carpathian Basin, while their numbers increased dramatically in the British Isles. Aglais urticae (Linnaeus, 1758) declined in all areas, but drastically in the Carpathian Basin. Papilio machaon (Linnaeus, 1758) remained stable everywhere. Euphydryas aurinia (Rottemburg, 1775) increased in the Carpathian Basin and remained stable in the British Isles. In 2008, Panigaj and Panigaj [216] re-surveyed the butterfly fauna of Temnosmrečinská dolina in the High Tatras. According to their results, fifteen of the originally recorded species were not found; however, the occurrence of six new species was recorded. Similar studies are reported by our team [6] based on the field studies of the Hungarian Natural History Museum and independent authors. A 35–68% decrease in species abundance was detected in different areas (Simontornya, Bátorliget, Drávasík). The problem with these studies is that the collection method and frequency were different, but at that time, there were no other sources. In the Kaposvár collection, we have regular records of 33 species made by the same collector, Dr. Levente Ábrahám, over three decades (Table 11). Here, we have seen that out of the initial 33 butterfly species, the density of 6 species has fallen below the detection level. At the level of the total Carpathian Basin, 9 species have fallen below the detection level in the last 24 years, and no detections of these species have been made in the lower parts of the Carpathian Basin since then.

Beyond the Central European situation, in a broader perspective, we observe the following: Settele et al., in their monograph Climatic Risk Atlas of European Butterflies [68], predict a decline of various butterfly species by 2050 and 2080, respectively, and a shift to northern areas based on the situation in the year 2000. We are now halfway through the first period. Obviously, the decline of a large number of butterfly species seems to be the realization of the authors’ predictions. As stated in Parmesan, C.; Ryrholm, N.; Stefanescu, C. et al.: “Mean global temperatures have risen this century, and further warming is predicted to continue for the next 50–100 years [1]”. The basic assumption of these and hundreds of similar predictions is that climate change equals global warming. However, climate change does not imply a general rise in temperature but rather weather disturbances, strong warming in some places, strong cooling in others, and ultimately the collapse of global ocean transport systems. Rather, abrupt regional cooling and gradual global warming may occur simultaneously. In fact, greenhouse warming is one of the destabilizing factors that makes abrupt climate change more likely. According to a 2002 report by the US National Academy of Sciences (NAS), available evidence suggests that abrupt climate changes are not only possible but likely in the future, with potentially large impacts on ecosystems and societies. In extreme cases, indefinite warming could turn into the opposite (glacial) due to a breakdown of thermohaline circulation [159,160].

Numerous papers study the changes in the population abundances of particular butterfly species. Birch, R. J.; Markl, G.; and Gottschalk [217] discuss the adaptation and population growth in abundance of Brintesia circe (Fabricius, 1775) (Figure 12) in the context of climate change. The increase in Minois dryas (Scopoli, 1763) under suitable conditions (nature conservation) is also well documented [218]. We have observed similar trends in the Danube-Ipoly National Park area. Hudák [219] noted an increase in the following species: Satyrium pruni (Linnaeus, 1758), Maculinea teleius (Bergstrasser, 1779), Aricia agestis (Denis & Schiffermüller, 1775), Libythea celtis (Füssly, 1782), Melitaea phoebe (Denis & Schiffermüller, 1775), Apatura ilia (Denis & Schiffermüller, 1775), Coenonympha arcania (Linnaeus, 1761), Minois dryas (Scopoli, 1763), and Brintesia circe (Fabricius, 1775). This largely aligns with our observations, but other species with strong population abundance growth in our area are not included in his work, as they were the most common species even in the past. Consequently, their population changes are rarely investigated.

Fox et al. [220] examined 58 Butterfly species from two aspects: the change in abundance and the change in distribution. We only examined the change in abundance. On the one hand, we examined 33 selected species similarly to the method of British authors) (Table 11). On the other hand, the total butterfly fauna of a given area (National Park), which is 111 species, was investigated by our team as well. In this case, the area was limited, but the number of species covered the total species richness of a given area (about 600 km²). Our data are not comparable because the aim of British researchers is to determine the status and change of the entire butterfly fauna, i.e., to examine the influence of all environmental factors. In contrast, we tried to eliminate all other factors influencing the number of individuals (like farming, pesticides, landscape destruction, isolation of habitats, etc.) and focus only on the effect of climate change. This is the reason that our results are different. Instead of the general decline of abundance, we observed an outbreak of common and frequent butterflies (these butterflies proved to be more or less climate change tolerant), and mostly decline of rare and sporadic butterflies which are sensitive for climate changes.

4.3.2. Moths, Nocturnal Macrolepidoptera

According to the prediction of Uhl and colleagues [221], climate change, with hot summers, might exceed the larval temperature optima and drought reduces food plant quality. Increasing frequency and severity of temperature and drought extremes due to climate change, therefore, might amplify insect decline in the future. Therefore, summer-developing larvae seem particularly sensitive. Changes in insect diversity caused by climate change might not be relevant in the Mediterranean area only, where summer heat might exceed the species’ physiological optima, but also in Central Europe, where optima will likely be exceeded in the future.

Our results also show some decline in Central Europe, but this decline is not significant at this time just because the decline of numerous species is compensated by increased frequency of outbreaks of some common species, also due to the gradually warming climate (see Table 17 and Figure 18). As for future projections, we are cautious to make any statement. Although the trends are clear, we should not forget the counter-effects, such as the possible shutdown of the Atlantic Meridional Overturning Circulation (AMOC) or Gulf Stream, which could lead to significant cooling.

In Sweden, Betzholtz and colleagues [222] studied Catocala nupta Linnaeus, 1767, Noctua interjecta Hübner, 1803, Noctua interposita (Hübner, 1790), Mythimna albipuncta Gaede, 1916, Eucarta virgo (Treitschke, 1835), Watsonalla binaria (Hufnagel, 1769), Pseudeustrotia candidula (Denis & Schiffermüller, 1775) and Eupithecia pulchellata Stephens, 1831. According to their results, the abundances of range-expanding moths in southeastern Sweden has increased both in abundance and species richness over 16 years, and their potential for rapid population density growth seems to be favored by a warming climate. We studied two of these species in Central Europe, with stable populations of Catocala nupta (Linnaeus, 1767) and a sharp decline of Pseudeustrotia candidula (Denis & Schiffermüller], 1775), apparently due to climatic differences between these 2 regions.

According to Sparks et al. [223], the number of migratory Lepidoptera (moths and butterflies) species reported annually at a site in the south of the UK has been steadily increasing. This rise is strongly linked to rising temperatures in Southwestern Europe. It is anticipated that further climate warming in Europe will increase the number of migratory Lepidoptera reaching the UK, necessitating urgent attention to the consequences of this invasion. In our region, as it is not on the border of the distribution of these species, they do not show migratory behaviour. However, we have studied the variation in the number of individuals of certain migratory species. In our area, the increase in the number of species is not very pronounced, but the expansion of one species of migrant moth, namely Helicoverpa armigera, and its increase causes significant damage to agricultural crops, and here it is the main concern among migrant moths. Other migratory moth species, such as Hyles livornica Esper, 1780, and Acherontia atropos Linnaeus, 1758 did not show an increasing trend. For Daphnis nerii Linnaeus, 1758, there is a definite positive trend (number of observations: 2014: 1 specimen, 2018: 8 specimens, 2019: 8 specimens, 2020: 13 specimens, 2021: 3 specimens, 2023: 23 specimens, 2024: 25 specimens) [224].

On the other hand, the increased population density predicted by the British authors [223], in the form of densifying gradients (Table 17, Figure 14 and Figure 15), have already arrived in our region in the form of reduced species diversity and increased frequency of outbreaks.

In Central-Europe, Szabó, Árnyas, and Varga [225] measured an increasing trend in the number of individuals between 1997 and 2004. According to their conclusion, arid weather conditions provided an ideal environment for the proliferation of moth species living in deciduous forests prone to outbreaks. The highest species richness was measured in the last year of their study series. Varga and colleagues [226], from 2019 to 2022, showed a decreasing trend in both species’ numbers and number of individuals. Furthermore, Szentkirályi et al. from Várgesztes experimental site [227], between 1962 and 1999, published a significant decrease in both parameters and observed outbreaks of species susceptible to outbreak in dry, drought years. According to the observation of Uherkovich [228], there was a significant decline in species diversity, which was a clear consequence of the rapid climate change of the last decades. There was also a decrease in the overall abundance of macrolepidoptera, which was clearly seen through his recent samplings from the 2020s. In the Subcarpathian region, there was no visible impoverishment of the general species diversity within a 100-year retrospective [59]. Szentkirályi and his colleagues [227,229] reviewed the results of a 50-year light trap network. Their results proved that there was a definitive, significant decreasing trend of moth assemblages in time series at certain trap stations but not at all stations.

Hufnagel and Sipkay [230] analyzed data from 8 light traps between 1962 and 2006. When examining the total number of moth species and individuals per year, they found that the number of species captured per year remained relatively stable despite a significant decline in the number of individuals. We extended this examination, including a significant number of additional traps, covering the period between 1970 and 2022. The key difference was that the decline in the number of individuals was hardly observable and not statistically significant (although our data detected the same decline in the early 2000s). Concurrently, the number of species caught by each trap decreased (Table 18 and Figure 20). Post-2006, the warming trend persisted, leading to the disappearance of some species from the light traps (their numbers fell below the detection limit locally). Simultaneously, several authors have observed the periodic outbreaks of certain nocturnal moth species (species prone to gradation) [225,226,227,229].

In our results, the lower Carpathian Basin over 50 years contains the main conclusions of the above articles. There is a decline in numbers, but this decline is not significant in the long term (r² = 0.01), as it is more or less compensated by the outbreaks of some common species due to the increasing frequency of dry periods since the beginning of the warming period in the 1980s, (r² = 0.13). This increase in the number of individuals is unfortunately associated with a decrease in species diversity per light trap. At the same time, the number of species that have disappeared from the entire Carpathian Basin during the last 20 years is small (Table 16).

In addition, we have long data series from Great Britain. According to the Butterfly Conservation Organization [231], the total abundance of larger moths caught in the RIS light-trap network in Britain decreased by 33% over 50 years (1968–2017). Losses were greater in the southern half of Britain (39% decrease) than in the northern half (22%). It is very different from the situation in Central Europe. Our experience confirms the results of the Ukrainian and Hungarian authors [59,225,226,227,229], as more frequent outbreaks during the warming period compensate for the decrease in abundance; furthermore, the largest number of moth captures in 50 years was taken in 2022 (Table 17 and Figure 14 and Figure 15). We also detected regional species impoverishment per trap. (Table 18 and Figure 20). but this was not associated with species impoverishment across the whole region (Table 16). On the other hand, the replacement of dominant species was the strongest indicator of climate change (Table 19).

4.4. Alien and Invasive Species, Mediterranean Influx

The most comprehensive study to date on invasive species in our region (and Europe) is the monograph “Alien Terrestrial Arthropods of Europe” by Roques and colleagues [75], published in the Biorisk series. This study analyzes approximately 330 insect species from our area, the Carpathian Basin, up to 2009. Building on this work and thoroughly reviewing the relevant literature, we expanded this to 803 insect species. One part of the remaining 473 species has been introduced or invaded since 2009, and the other part was identified through a more in-depth examination of the literature, having been omitted from the mentioned work. We compared our findings with similar research results from nearby and distant regions.

Solarz and colleagues [232] in Poland assessed 60 non-native plant species and 58 animal species. For seventy-nine species, they evaluated climate change as a factor that increases the likelihood of future introduction, establishment, and spread. Climate change increased the number of high-risk invasive alien species from 38 to 63. Species from warmer parts of the world were the most sensitive to climate change. We examined 803 insect species based on their origin and time of appearance, with a particular focus on the climate change situation. Of the 803 incoming species, 174 are from tropical areas. Among these 174, 74 tropical species (43%) entered the Carpathian Basin in the last quarter-century (Figure 22), along with 86 Mediterranean-origin species (Figure 26). This high rate of influx and establishment in recent decades is largely driven by climate warming. This is supported by Gutierrez and colleagues’ monograph on the influx of alien insect species and its interpretation from an agricultural perspective, stating that whether the introduction of a new species becomes invasive or not partly depends on the biological and physical characteristics of the habitat into which it is introduced. These habitat characteristics are largely influenced by climate, which is altered by human activities [233].

Gutierrez and Ponti [233] report a phenomenon in North America similar to the Mediterranean influx we experience, using examples of four pests: Cochliomyia hominivorax (Coquerel, 1858), Pectinophora gossypiella (Saunders, 1844), Ceratitis capitata (Wiedemann, 1824), and Bactrocera oleae (Rossi, 1790). These species first appeared in California and are spreading northward and southward toward Mexico due to temperature changes. Their findings suggest that the primary driver of northward spread is temperature increase, while southward spread is facilitated by adequate moisture levels. Consequently, their forecasts indicate that the arid regions of Arizona and northern Mexico are more protected, whereas the subtropical areas of California and western and southern Mexico are more susceptible to infestations. Since desertification in Central Europe is currently limited to small areas but increasing (Kiskunság: Homokhátság in Hungary), future studies on the effects of desertification would be important. However, due to a lack of financial resources, these studies were stopped for a while (Fülöpháza, Kiskunság National Park, light trap project).

According to the IUCN’s forecast [234], the number of alien insect species will increase by 36% between 2005 and 2050. However, in our region, we have surpassed this prediction, with the number of insect species alone increasing by 54% over the past 24 years compared to all previous years. Of these, 86 are Mediterranean, and 74 are tropical species (Figure 21). Thus, in this century, out of the 298 incoming species, 160 (54%) have arrived from warm (Mediterranean, tropical) regions, indicating that climate change is a major driver of this influx.

Out of the 1390 alien insect species established in Europe [75], our current findings show that 803 live in the Carpathian Basin, which is 58% of the 1390 species reported from Europe. Specifically, this includes 164 of the 398 non-native Coleoptera species (41%) established in Europe, 303 of the 318 Hemiptera species (95%), 88 of the 297 Hymenoptera species (30%), 99 of the 97 recorded Lepidoptera species (102%), and 65 of the 98 Diptera species (66%). The total insect influx in Europe can be described by the curve 15.40 exp (0.50x) r² = 0.87, while in our region, it is not so steep: 5.86 exp (0.49x) r² = 0.84. With a linear approximation, the intensity of alien species influx in Europe is 67.75 (lin. x coefficient, slope of the line) r² = 0.86, whereas it is more moderate in the Carpathian Basin, with an intensity of 36.82 (lin. x coefficient) r² = 0.77.

From a biogeographic perspective, the two studies are not fully comparable because we separately treated the Mediterranean-origin species, as different drivers (including climate change-induced influx) primarily govern them. Additionally, instead of Asia, we separately considered the Eastern Palaearctic and Oriental regions, as these are distinct units from climatic and zoogeographic points of view. It is evident that for a more accurate assessment of the influx rate, comparing 2024 data with European data from 2010 is not advisable. It indicates the necessity of a deeper analysis of the European situation and the update of the data of the European monograph.

The influx of alien insect species in other regions also currently shows an exponential trend. In Eastern Russia, the equation for the influx trend is quadratic: AS = exp (−5.73 + 0.000002536 × Years²) or e − 5.73 + 0.000002536 × Years² (xsquared) [235].

Similar data are reported from China [236], where we observe an exponential increase again. The influx of alien species correlates with China’s GDP. This correlation is also evident when it is split up by provinces, as the increase in freight transport and GDP are closely linked. In Central Europe, this is not necessarily the case, as the region does not show similar economic growth compared to China; however, it is still highly susceptible to the influx of alien insect species (Supplementary Material C). It is evident that alien species enter through the ports and rail yards of economically stronger EU states and then spread rapidly across the continent. Therefore, in Europe, there is no such strong correlation between economic development and the influx of alien species. The ranking of European countries by non-native insect species is as follows: Italy, France, Great Britain, Spain, and the Czech Republic [75]. Globally, the most heavily infested countries and regions by infection of non native insect species are Cuba, Great Britain, France, Taiwan, Florida, Texas, California, Hawaii (USA), Queensland, and Victoria (Australia) [237].

Another significant driver of the establishment of alien insects is the influx of alien plant species. Plants that have been established or introduced in Europe show an exponential trend similar to that of insects [238]. In this context, our region is positioned in the middle of the ranking (at the boundary of the upper and lower 50%).

5. Conclusions

The entomofauna of Central Europe, including the Carpathian Basin, is undergoing transformation due to four main factors: the retreat of insect species that require cool and wet habitats, the proliferation of insect species that thrive in warm and dry conditions, the northward migration of Mediterranean insect species, and the gradual establishment of introduced insects.

The proportion of insect species arriving from tropical and Mediterranean regions has increased significantly, with their numbers rising by 58% over the past 24 years compared to all previous years. Of the 298 insect species that have arrived in this century, 160 (54%) are from warm regions (Mediterranean and tropical), including 86 Mediterranean and 74 tropical insect species. This indicates that climate change is a major driver of transformation.

Among native insect species, hoverflies (Syrphidae) have suffered the most significant abundance decline, consistent with European and global trends. The population of Tachinidae species has also decreased, although less markedly than hoverflies. Conversely, species from xerothermic families, such as beeflies (Bombyliidae) and horse-flies (Tabanidae), have shown population increases up to the current extent of warming. However, since horsefly larvae are associated with wet or aquatic environments, this trend is expected to stop if warming continues.

In terms of climate change, different insect species exhibit different patterns based on their reactions. For certain species, climate change not only indicates spontaneous colonization but also an increased frequency of outbreaks, making these insects capable of showing an exponential trend. Such is the case with Helicoverpa armigera, which has become a serious insect pest due to this characteristic. Extremely xerothermic insect species, like members of the Philantidae family, have shown a continuous upward trend. Since the Anatolian and Middle Eastern regions are the most optimal for these insects in terms of population size and species richness, if the warming trend continues, we can expect further growth. For moderately xerothermic insects, we can assume (though not yet proven) that the current rate of warming may have already exceeded their optimal level. These types are summarized in Figure 29. For insect species showing a declining trend, a few insect species in two groups (Bombus species and hoverfly species) responded with a decrease in population size that can be described by an exponential trend. The linear decline in the abundances of many insects (hylophilous and silvicolous species) is a logical consequence of the warming trend. For other insects (e.g., Aglais urticae), their population size was not directly affected by temperature change; we believe that their decline or disappearance from their original range is due to the increasingly extreme climate. The reactions of these types can be seen in Figure 30.

The warming climate has proven to be favorable for most of the Aculeata Hymenoptera, which is logical, given that they achieve their highest species diversity and population densities in the Mediterranean region. Alongside horse-flies and bee flies, the increase of this group clearly demonstrates the Mediterranean transformation. However, several groups have shown significant declines, including bumblebees, Megachilid bees, and certain Crabronidae species that require cool and wet habitats (forests and marsh meadows). The abundance decline of bumblebee species is more complex and likely only partially caused by warming. This issue requires further investigation.

In the case of butterflies, a decline in species richness is observable in certain areas. Some previously common species have proven resistant to the current extent of climate change, showing intense reproduction under protected conditions. However, most previously sporadic and rare species have been unable to adapt to the changed climate conditions, resulting in a decline in their population numbers, frequently even falling below the detection level regionally. The most benefited are Papilionidae. Papilionidae are climate tolerant (only up to the present measure) and benefited from the economic collapse (no money for pesticides), especially Iphiclides podalirius and Papilio machaon. They were rare during the 80s and are now frequent and locally even common. However, the majority of butterflies, which were originally sporadic or rare, have become even rarer. We assume (no evidence) that some of them, which overwinter as imagos, start to fly too early, and the late frosts kill them. Earlier frequent and common butterfly species have increased because nature conservation has also helped them to maintain and increase their abundances, and they seem to be euryecious. One exception is Aglais urticae, which was also very frequent earlier but has now nearly disappeared.

The population in total abundance of nocturnal moths has entered a growth phase over the past 10 years. This growth phase has been accompanied by a decrease in species diversity. This population growths are caused by the more frequent outbreaks of some common moths.

Only a smaller proportion of insect species find refuge in higher regions. According to our current findings, this includes 20 out of 87 Diptera species, 32 out of 71 Aculeata Hymenoptera species, and 15 out of 23 Lepidoptera species, which have not been collected in the lower regions for over 20 years. We do not and cannot attempt to make forecasts, as several predictions suggest that current trends may reverse, and our climate may become even more chaotic if ocean currents potentially collapse. Finally, we, as individuals, when we can do nothing more, should never lose hope in the cohesive power of communities, even in such challenging circumstances.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/ecologies6010016/s1, Supplementary Materials A–C.

Author Contributions

Hymenoptera, Diptera, Lepidoptera, non-native species, analysis A.H. (Attila Haris); Aculeata, Z.J.; Diptera, S.T.; bumblebees, Aculeata, P.Š.; bumblebees, Aculeata, Dipetra in part B.T.; Lepidoptera, P.S.; Lepidoptera, G.G.; Invaziv and non native species, Lepidoptera, G.C.; Invaziv and non native species, Lepidoptera, A.H. (Anikó Hirka); statistical analysis, A.H. (Attila Haris); resources, All authors; data curation, All authors; writing—original draft preparation, A.H. (Attila Haris). All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not Applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Supporting data are available: 1. University of Sopron, Forest Research Institute, Mátraháza, Hungary; 2. Digital Archives of Rippl-Rónai Museum, Kaposvár, Hungary.

Acknowledgments

Authors express their grateful thanks to Vladimír Smetana (Tekov Museum, Levice, Slovak Republic), Levente Ábrahám (Rippl-Rónai Museum, Kaposvár, Hungary), Ákos Uherkovich, (Janus Pannonius Museum Pécs, Hungary), Attila Huber (Aggtelek National Park, Aggtelek, Hungary) and Zoltán Varga (University of Debrecen, Debrecen, Hungary) for their advises and generous supports, data provisions and advises. We are also grateful to Ádám Gór (Büro Geyer und Dolek, Wörthsee, Germany) for the permission to publish his photo.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. The Graphocephala fennahi Young, 1977 (Rhododendron Leafhopper), native to North America, were first found on the leaves of Rhododendron catawbiense in Hungary in 2012. It well illustrates the role of alien insect species in the transformation of our fauna (photo: György Csóka).

Figure 2. Biogeographic map of the investigated area. Pannonian (brown), Continental (green), and Alpine biogeographic regions (violet). (source: EEA [7]).

Figure 3. Hydrogeological map of the Carpathian Basin (source: [8]).

Figure 5. Long-term trend of horse-flies (Tabanidae) in the Carpathian Basin.

Figure 6. Long-term declining trend of hoverflies (Syrphidae) in the Carpathian Basin.

Figure 7. Long-term increasing trend of xerotherm bee flies (Bombyliidae) in the Carpathian Basin, winners of the warming climatic conditions.

Figure 8. Long-term trend of tachinid flies (Tachinidae) in the Carpathian Basin. General declining trend with strong fluctuation probably typical for parasitoids.

Figure 9. Long-term trend of species richness of Bumblebees in the high (blue) and low (red) elevations of the Carpathian Basin.

Figure 10. Long-term trend of wild bee families in the Carpathian Basin.

Figure 11. Long-term trend of Bembecidae, Psenidae, Sphecidae and Pemphedronidae in the Carpathian Basint.

Figure 12. Long-term trend of Chrysididae, Philantidae and Crabronidae in the Carpathian Basin.

Figure 13. Long-term trend of numbers of specimens of sawflies (Symphyta) by Malaise trap method based on Haris et al., 2024 [6]. Sawflies are primarily distributed in northern and montane regions, mostly inhabiting moderately cool, rainy areas.

Figure 14. Long-term trend of species richness of sawflies (Symphyta) by Malaise trap method based on Haris et al., 2024 [6].

Figure 15. Aglais urticae (Linnaeus, 1758) (Small Tortoiseshell) was once common until the late 1980s. In the last decades, it nearly disappeared from the deeper region of the Carpathian Basin (photo: Ádám Gór).

Figure 16. Brintesia circe (Fabricius, 1775) (Great Banded Grayling) has successfully resisted climate change so far (photo: Péter Schmidt).

Figure 17. Some butterflies, like Iphiclides podalirius (Linnaeus, 1758) (Scarce Swallowtail), have even been able to increase their abundance in Central Europe (Photo: Gábor Glemba).

Figure 18. Changes in nocturnal macrolepidoptera populations in total abundance between 1970 and 2022.

Figure 19. Changes in nocturnal macrolepidoptera populations in total abundance between 2014 and 2022.

Figure 20. Changes in species richness of nocturnal macrolepidoptera between 1970 and 2022.

Figure 21. Timeline of non-native insects in the Carpathian Basin from the Neolithic period till 2024.

Figure 22. Timeline of the introduction of tropical insect species.

Figure 23. Division of non-native insects according to their taxonomic groups.

Figure 24. Division of non-native insects according to their origin.

Figure 25. Ovalisia festiva (Linnaeus, 1767) (Cypress Jewel Beetle) had double colonization: natural and introduction with ornamental plants (photo: György Csóka).

Figure 26. Timeline of the influx of Mediterranean insect species.

Figure 27. Long-term trend of wild bees in Belgium’s aboundances (after Duchenne et al. [202]).

Figure 28. The Parnassius mnemosyne (Linnaeus, 1758) (Clouded Apollo) appears stable in our region (photo: Gábor Glemba).

Figure 29. Different types of increase of insect populations in abundance as a response to climate change (yellow: exponential growth of invasive insect, red: moderate population growth after the temperature optimum, blue: continuous growth of abundance of xerotherm insect).

Figure 30. Different types of decline of insect abundances as a response to climate change (yellow: exponential decline of highly sensitive insects of climate change, red: gradual decline of population in abundance of hylophilous insects, blue: disappearance of climate change-sensitive butterfly species).

Table 1. Long-term trend of horse-fly (Tabanidae) species (abbreviations: hy: hylophilous, si: silvicolous, and xe: xerothermic).

Tabanidae (Horse-Flies)	1980–84	1985–89	1990–94	1995–99	2005–09	2010–14	lin. x coeff.	r²	diff.	Ecotype
Hybomitra pilosa (Loew, 1858)	2	8	6	9	38	16	4.66	0.45	8.00	hy
Haematopota scutellata (Olsufje et al., 1964)	13	6	9	14	25	17	2.34	0.44	1.31	xe
Philipomyia graeca (Fabricius, 1794)	1	4	5	4	39	15	4.97	0.42	15.00	xe
Tabanus bovinus (Linnaeus, 1758)	8	13	20	14	81	40	10.23	0.38	5.00	xe
Tabanus autumnalis (Linnaeus, 1761)	29	18	14	28	67	41	6.31	0.38	1.41	xe
Heptatoma pellucens (Fabricius, 1777)	16	4	4	11	42	21	4.17	0.30	1.31	xe
Therioplectes gigas (Herbst, 1787)	5	1	3	5	34	14	4.17	0.30	2.80	xe
Silvius alpinus (Scopoli, 1763)	5	0	1	1	31	9	3.22	0.26	1.80	hy
Hybomitra solstitialis (Meigen, 1820)	0	1	1	5	30	2	2.89	0.22	NA	xe
Philipomyia aprica (Meigen, 1820)	17	4	3	16	34	15	2.66	0.20	0.88	xe
Hybomitra confiformis (Chvála & Moucha, 1971)	4	3	2	7	35	0	2.31	0.11	0.00	xe
Chrysops caecutiens (Linnaeus, 1758)	14	16	13	33	83	41	10.17	0.05	2.93	si
Chrysops viduatus (Fabricius, 1794)	74	20	29	59	90	48	3.24	0.05	0.65	si
Haematopota italica Meigen, 1804	19	114	227	42	85	58	−2.2	0.00	3.05	hy
Atylotus rusticus (Linnaeus, 1767)	116	13	13	40	72	61	−2.03	0.01	0.53	xe
Haematopota pluvialis (Linnaeus, 1758)	189	228	205	760	161	70	−6.88	0.03	0.37	hy
Chrysops relictus (Meigen, 1820)	62	35	94	67	55	23	−4.63	0.12	0.37	si
Haematopota subcylindrica (Pandellé, 1883)	14	8	34	7	4	1	−2.97	0.22	0.07	hy
Tabanus bromius (Linnaeus, 1758)	212	128	187	134	116	62	−23.97	0.71	0.29	xe

Table 2. Long-term trend of hoverfly (Syrphidae) species (abbreviations: eur: euryecious [wide ecological tolerance], hyg: hylophilous, sil: silvicolous, and xer: xerothermic).

Syrphidae (Hoverflies)	1980–1984	1985–1989	1990–1994	1995–1999	2000–2004	2005–2010	lin. x coeff.	r2	diff.	Ecotype
Sphaerophoria scripta (Linnaeus, 1758)	5421	2681	600	916	107	341	−937.31	0.73	0.06	eur
Episyrphus balteatus (De Geer, 1776)	3289	2432	928	2446	198	341	−569.26	0.68	0.10	eur
Melanostoma mellina (Linnaeus, 1758)	2016	1694	527	555	129	238	−387.34	0.83	0.12	eur
Syrphus torvus (Osten Sacken, 1875)	640	185	91	86	42	78	−92.68	0.58	0.12	sil
Eristalis arbustorum (Linnaeus, 1758)	2043	801	548	536	150	296	−305.71	0.70	0.14	eur
Pipizella viduata (Linnaeus, 1758)	724	866	141	315	91	117	−148.17	0.68	0.16	eur/sil
Syrphus vitripennis Meigen, 1822)	1842	673	740	393	62	336	−277.43	0.70	0.18	eur
Eristalis tenax (Linnaeus, 1758)	2990	1539	422	422	202	559	−461.89	0.65	0.19	eur
Scaeva pyrastri (Linnaeus, 1758)	531	151	165	129	33	112	−71	0.58	0.21	eur
Syrphus ribesii (Linnaeus, 1758)	895	329	88	282	54	190	−118.74	0.52	0.21	eur
Myathropa florea (Linnaeus, 1758)	237	230	82	96	27	58	−42.57	0.79	0.24	eur/sil
Syritta pipiens (Linnaeus, 1758)	1096	982	679	460	99	290	−197.09	0.89	0.26	eur
Cheilosia variabilis (Panzer, 1798)	810	243	128	252	63	218	−96.46	0.46	0.27	sil
Eristalis pertinax (Scopoli, 1763)	356	354	110	165	52	101	−60.74	0.73	0.28	eur
Cheilosia impressa (Loew, 1840)	270	273	106	175	53	95	−40.89	0.70	0.35	eur
Cheilosia soror (Zetterstedt, 1843)	170	160	88	71	28	69	−26.23	0.77	0.41	sil/xer
Eupeodes luniger (Meigen, 1822)	111	54	56	142	22	49	−9.14	0.15	0.44	eur
Volucella pellucens (Linnaeus, 1758)	159	32	49	52	33	78	−11.4	0.20	0.49	sil
Platycheirus albimanus (Fabricius, 1781)	373	226	189	274	75	190	−36.66	0.48	0.51	eur
Chrysotoxum cautum (Harris, 1776)	135	66	293	229	78	81	−8.51	0.03	0.60	sil
Eristalis interrupta (Poda, 1761)	248	206	109	123	88	158	−22.57	0.47	0.64	eur
Eumerus tricolor (Fabricius, 1798)	42	6	32	11	19	30	−1.2	0.03	0.71	sil
Volucella zonaria (Poda, 1761)	25	0	16	8	93	22	7.31	0,17	0.88	sil
Baccha elongata (Fabricius, 1775)	67	35	79	66	60	59	0.63	0.01	0.88	sil
Xylota segnis (Linnaeus, 1758)	104	102	150	152	31	96	−7.17	0.09	0.92	sil
Cheilosia scutellata (Fallén, 1817)	46	96	71	68	36	45	−5.37	0.20	0.98	sil
Epistrophe nitidicollis (Meigen, 1822)	59	79	134	163	47	61	−1.63	0.00	1.03	sil
Helophilus pendulus (Linnaeus, 1758)	195	192	241	654	107	211	6.8	0.00	1.08	hyg
Epistrophe eligans (Harris, 1780)	80	48	143	189	31	91	1.43	0.00	1.14	sil
Baccha perexilis (Harris, 1776)	65	34	38	49	43	79	3.08	0.11	1.22	sil/hyg
Lejops vittatus (Meigen, 1822)	118	40	49	29	76	152	7.37	0.08	1.29	hyg
Paragus tibialis (Fallén, 1817)	16	6	23	3	12	22	0.8	0.03	1.38	xer
Merodon constans (Rossi, 1794)	29	32	52	45	19	40	0.26	0.00	1.38	xer
Paragus finitimus (de Tiefenau, 1971)	18	9	45	16	20	44	3.83	0.22	2.44	xer
Anasimyia contracta (Claussen & Torp, 1980)	13	24	19	66	30	52	7.43	0.46	4.00	hyg
Anasimyia lineata (Fabricius, 1787)	33	167	74	178	76	135	9.74	0.10	4.09	hyg
Merodon nigritarsis (Rondani, 1845)	5	28	166	110	33	51	5.4	0.03	10.20	xer

Table 3. Long-term trend of bee-fly (Bombyliidae) species (members of this group are categorized as xerothermic).

Bombyliidae (Bee Flies)	1980–84	1985–89	1990–94	1995–99	2000–04	2005–09	2010–14	2016–20	lin. x coeff.	r²	diff.
Hemipenthes velutina (Meigen, 1820)	4	2	9	8	8	12	13	14	1.6	0.86	3.50
Conophorus virescens (Fabricius, 1787)	9	32	19	45	38	57	72	60	7.9	0.83	6.67
Bombylius nubilus (Mikan, 1796)	1	1	0	6	9	6	13	11	1.8	0.8	11.00
Bombylius medius (Linnaeus, 1758)	7	7	9	14	10	34	34	70	7.7	0.73	10.00
Lomatia sabaea (Fabricius, 1781)	3	0	15	3	18	13	23	38	4.39	0.73	12.67
Exoprosopa jacchus (Fabricius, 1805)	1	2	7	1	8	27	23	18	3.46	0.67	18.00
Exoprosopa minos (Meigen, 1804)	2	0	0	1	3	13	19	12	2.45	0.67	6.00
Villa hottentotta (Linnaeus, 1758)	20	6	72	33	36	70	73	99	10.54	0.65	4.95
Anthrax ricardoi (Greathead, 2003)	0	0	0	0	3	2	4	2	0.51	0.62	NA
Bombylius canescens (Mikan, 1796)	8	6	15	27	13	32	19	33	3.29	0.59	4.13
Bombylius discolor (Mikan, 1796)	13	14	8	41	41	23	31	91	8.04	0.55	7.00
Bombylius cinerascens (Mikan, 1796)	13	33	19	24	28	51	27	79	6.33	0.53	6.08
Bombylius fulvescens (Meigen & Wiedemann, 1820)	7	20	15	21	9	23	27	27	2.23	0.51	3.86
Exoprosopa capucina (Fabricius, 1781)	3	0	3	1	6	3	10	6	0.9	0.48	2.00
Anthrax trifasciatus (Meigen, 1804)	5	2	12	20	6	21	15	17	1.93	0.43	3.40
Anthrax anthrax (Schrank, 1781)	5	1	9	0	1	9	10	18	1.63	0.43	3.60
Exhyalanthrax muscarius (Pallas & Wiedemann, 1818)	0	0	0	1	0	1	0	2	0.19	0.38	NA
Bombylius analis (Olivier, 1789	0	5	8	4	1	5	2	31	2.26	0.31	NA
Bombylius fimbriatus (Meigen, 1820)	8	31	18	40	41	13	26	52	3.2	0.27	6.50
Triplasius pictus (Panzer, 1794)	2	5	3	4	3	10	21	4	1.36	0.27	2.00
Bombylisoma nigriceps (Loew, 1862)	2	1	0	4	3	4	14	2	0.9	0.26	1.00
Micomitra stupida (Rossi, 1790)	1	0	0	2	0	0	8	1	0.45	0.17	1.00
Phthiria pulicaria (Mikan, 1796)	5	1	11	4	1	8	35	4	1.8	0.16	0.80
Apolysis szappanosi (Papp, 2005)	0	1	0	0	1	0	5	0	0.25	0.13	NA
Bombylius posticus (Fabricius, 1805)	6	4	30	24	9	19	20	15	1.13	0.09	2.50
Hemipenthes morio (Linnaeus, 1758)	67	42	90	254	31	78	89	124	4.46	0.02	1.85
Bombylius major (Linnaeus, 1758)	54	156	105	160	63	26	28	199	0.49	0	3.69
Bombylella atra (Scopoli, 1763)	23	39	22	53	3	45	36	7	−1.28	0.03	0.30
Hemipenthes maura (Linnaeus, 1758)	12	3	1	7	4	3	11	1	−0.4	0.05	0.08
Bombylius venosus (Mikan, 1796)	27	2	3	13	9	1	6	11	−1.21	0.12	0.41

Table 4. Long-term trend of tachinid fly (Tachinidae) species (abbreviations: eur: euryecious [wide ecological tolerance], hy: hylophilous, sil: silvicolous, and xe: xerothermic).

Tachinidae (Tachinids)	1980–84	1985–89	1990–94	1995–99	2000–04	2005–09	2010–2014	lin. x coeff.	r²	Change	Exotype
Linnaemya frater (Rondani, 1859)	39	234	17	22	16	75	NA	−13.40	0.09	1.92	sil
Meigenia dorsalis (Meigen, 1824)	31	120	21	21	7	54	NA	−6.40	0.09	1.74	eur
Exorista larvarum (Linnaeus, 1758)	59	228	82	31	35	100	NA	−12.00	0.12	1.69	sil
Tachina fera (Linnaeus, 1761)	135	346	299	117	59	201	NA	−20.37	0.12	1.49	sil
Blondelia nigripes (Fallén, 1810)	83	283	85	63	18	111	NA	−19.34	0.16	1.34	xer
Gymnosoma dolycoridis (Dupuis, 1961)	78	195	82	49	52	93	NA	−11.06	0.15	1.19	eur
Actia crassicornis (Meigen, 1824)	73	569	152	57	9	83	NA	−39.29	0.22	1.14	sil
Peleteria varia (Fabricius, 1794)	72	111	69	17	40	75	NA	−7.14	0.17	1.04	sil
Gymnosoma clavatum (Rohdendorf, 1947)	33	69	31	34	11	32	NA	−5.02	0.25	0.97	sil
Gymnosoma rotundatum (Linnaeus, 1758)	106	208	153	42	56	95	NA	−17.80	0.29	0.90	sil
Athrycia trepida (Meigen, 1824)	81	80	33	44	18	62	NA	−7.71	0.32	0.77	eur
Nemoraea pellucida (Meigen, 1824)	86	246	125	26	32	57	NA	−25.31	0.33	0.66	sil
Thelaira solivaga (Harris, 1780)	14	36	3	4	12	8	NA	−2.88	0.20	0.57	sil
Phryno vetula (Meigen, 1824)	47	97	43	51	8	26	NA	−10.40	0.42	0.55	eur
Phasia pusilla (Meigen, 1824)	88	121	42	24	14	39	NA	−16.69	0.57	0.44	sil
Phasia hemiptera (Fabricius, 1794)	138	124	87	7	22	57	NA	−22.60	0.63	0.41	sil
Chetogena filipalpis (Rondani, 1859)	17	18	18	5	8	6	NA	−2.80	0.69	0.35	eur
Compsilura concinnata (Meigen, 1824)	135	50	231	44	31	47	NA	−19.54	0.22	0.35	eur
Solieria vacua (Rondani, 1861)	44	53	31	3	13	7	NA	−9.51	0.74	0.16	sil

Table 5. Long-term trend of various Diptera families.

	1980–84	1985–89	1990–94	1995–99	2000–04	2005–09	2010–14	2014–19	lin. x coeff.	r²	Change
Syrphidae	35,119	22,381	11,520	18,504	5694	11,755	NA	NA	−1812.49	0.86	0.2
Bombilidae	530	533	569	615	452	701	928	980	63.79	0.65	1.8
Tabanidae	978	1046	977	1387	1387	1614	968	NA	136.93	0.84	0.6
Tachinidae	6145	14,339	6701	3479	1906	6793	NA	NA	−523.31	0.37	0.2

Table 6. Previously sporadically occurring Diptera species, but not collected for at least 20 years, in the low-lying regions of the Carpathian Basin. Gray background and x-mark: species that still occur in the higher regions and the northern parts.

Species	High alt.	Species	High alt.
Bombyliidae		Callicera rufa (Schummel, 1842)
Bombylius trichurus (Wiedemann, 1818)		Callicera spinolae (Rondani, 1844)
Bombylius quadrifarius (Loew, 1855)		Chalcosyrphus curvipes (Loew, 1854)
Bombylius ambustus (Pallas & Wiedemann, 1818)		Cheilosia bracusi (Vujic & Claussen, 1994)
		Cheilosia brunnipennis (Becker, 1894)
Tabanidae		Cheilosia hypena (Becker, 1894)
Pangonius pyritosus (Loew, 1859)	x	Cheilosia insignis (Loew, 1857)
Hybomitra arpadi (Szilády, 1923)		Cheilosia melanopa (Zetterstedt, 1843)	x
Hybomitra aterrima (Meigen, 1820)	x	Cheilosia melanura (Becker, 1894)	x
Hybomitra expollicata (Pandellé, 1883)		Cheilosia pictipennis (Egger, 1860)
Hybomitra montana (Meigen, 1820)	x	Cheilosia sahlbergi (Becker, 1894)	x
Hybomitra nigricornis (Zetterstedt, 1842)		Cheilosia subpictipennis (Claussen, 1898)
Hybomitra tarandina (Linnaeus, 1758)		Chrysogaster basalis (Loew, 1857)	x
		Cliorhina pachymera (Egger, 1858)	x
Tachinidae		Epistrophe obscuripes (Strobl, 1910)
Amelibaea tultschensis (Brauer & Bergenstamm, 1891)		Eristalis vitripennis (Strobl, 1893)
Anthomyiopsis nigrisquamata (Zetterstedt, 1838)	x	Eumerus hungaricus (Szilády, 1940)
Anthomyiopsis plagioderae (Mesnil, 1972)		Eumerus longicornis (Loew, 1855)
Aphria xyphias (Pandellé, 1896)		Eumerus ruficornis (Meigen, 1822)
Besseria dimidiata (Zetterstedt, 1844)	x	Eumerus sabulosum (Fallén, 1817)	x
Besseria melanura (Meigen, 1824)	x	Eumerus tauricus (Stackelberg, 1952)
Bithia acanthophora (Rondani, 1861)		Eupeodes lucasi (Marcos-García & Láska, 1983)
Blepharomyia pagana (Meigen, 1824)	x	Hammersmidtia ferruginea (Schummel, 1834)
Cadurciella tritaeniata (Rondani, 1859)		Helophilus affinis (Wahlberg, 1844)
Campylocheta latigena (Mesnil, 1974)		Lejota ruficornis (Zetterstedt, 1843)
Catharosia albisquama (Villeneuve, 1932)		Melanogaster curvistylus (Vujić-Stuke, 1998)
Ceranthia tristella (Herting, 1966)		Melanostoma dubium (Zetterstedt, 1837)
Chetoptilia puella (Rondani, 1862)		Milesia crabroniformis (Fabricius, 1775)	X
Conogaster pruinosa (Meigen, 1824)		Orthonevra tristis (Loew, 1871)
Phytomyptera abnormis (Stein, 1924)		Paragus medeae (Stanescu, 1991
Eloceria delecta (Meigen, 1824)		Paragus punctulatus (Zetterstedt, 1938)
Estheria acuta (Portschinsky, 1881)		Pipiza fenestrata (Meigen, 1822)
Gonia bimaculata (Wiedemann, 1819)		Pipizella pennina (Goeldlin de Tiefenau, 1974)
Heraultia albipennis (Villeneuve, 1920)		Platycheirus complicatus (Becker, 1889)
Ligeriella aristata (Villeneuve, 1911)		Platycheirus immarginatus (Zetterstedt, 1849)
Minthodes pictipennis (Brauer & Bergenstamm, 1889)		Platycheirus jaerensis (Nielsen, 1971)
Psalidoxena transsylvanica (Villeneuve, 1929)		Platycheirus nielseni (Vockeroth, 1990)
Siphona confusa (Mesnil, 1961)	x	Platycheirus perpallidus (Verrall, 1901)
Siphona ingerae (Andersen, 1982)	x	Rhingia austriaca (Meigen, 1830)
Therobia leonidei (Mesnil, 1965)	x	Scaeva albomaculata (Macquart, 1842)
Vibrissina debilitata (Pandellé, 1896)		Sphaerophoria shirchan (Violovitsh, 1957	x
Winthemia bohemani (Zetterstedt, 1844)		Sphiximorpha binominata (Verrall, 1901)
Syrphidae		Syrphus nitidifrons (Becker, 1921)
Brachyopa panzeri (Goffe, 1945)		Syrphus sexmaculatus (Zetterstedt, 1838)	x
Brachyopa vittata (Shummel, 1834)		Trichopsomyia joratensis (Goeldlin de Tiefenau, 1997)
Brachypalpus chrysites (Egger, 1859)	x	Xylota coeruleiventris (Zetterstedt, 1838)
Callicera macquarti (Rondani, 1944)

Table 7. Number of individuals of various bumblebee and cuckoo bumblebee species between 2005 and 2023 in the low and high altitudes of the Carpathian Basin (CRC: Climate Risk Categories: LR: low risk, R: risk, HR: high risk, HHR: very high risk, HHHR—extreme high risk). The gray color separates cuckoo bumblebees from bumblebees.

Taxon/Data from the Low Altitudes	2005–2009	2010–2014	2015–2019	2020–2023	lin. x coeff.	r²	CRC
Bombus confusus (Schenck, 1861)	2	0	0	0	NA	NA	HHHR
Bombus subterraneus (Linnaeus, 1758)	0	0	0	0	NA	NA	HHHR
Bombus pomorum (Panzer, 1805)	0	0	0	0	NA	NA	HHHR
Bombus terrestris (Linnaeus, 1758)	944	888	479	205	−262.6	0.94	HR
Bombus lapidarius (Linnaeus, 1758)	835	291	481	45	−218.0	0.71	HHR
Bombus pascuorum (Scopoli, 1763)	692	944	401	284	−176.7	0.59	R
Bombus hortorum (Linnaeus, 1761)	173	357	106	11	−73.7	0.42	HR
Bombus ruderarius (Müller, 1776)	174	151	56	13	−57.8	0.95	HHR
Bombus humilis (Illiger, 1806)	75	7	32	16	−15.2	0.42	HHR
Bombus lucorum (Linnaeus, 1761)	41	104	31	7	−17.5	0.30	HR
Bombus hypnorum (Linnaeus, 1758)	31	18	12	7	−7.8	0.94	HR
Bombus ruderatus (Fabricius, 1775)	18	11	0	0	−6.5	0.90	HR
Bombus sylvarum (Linnaeus, 1761)	77	98	24	65	−11.0	0.21	HHHR
Bombus muscorum (Linnaeus, 1758)	6	1	1	2	−1.2	0.42	HHR
Bombus pratorum (Linnaeus, 1761)	1	28	18	2	−0.7	0.00	HR
Bombus haematurus (Kriechbaumer, 1870)	80	93	103	116	11.8	0.99	HR
Bombus argillaceus (Scopoli, 1763)	4	28	8	31	6.1	0.33	LR
Bombus vestalis (Geoffroy, 1785)	60	49	55	13	−13.5	0.67	HR
Bombus rupestris (Fabricius, 1793)	12	56	6	0	−8.6	0.19	HHR
Bombus bohemicus (Seidl, 1838)	3	29	21	0	−1.7	0.02	HHR
Bombus barbutellus (Kirby, 1802)	5	1	0	0	−1.3	0.79	HHR
Bombus campestris (Panzer, 1801)	2	4	1	0	−0.9	0.46	HHR
Total specimens	3235	3158	1835	817	−857.7	0.91	NA
Total species	20	19	18	14	−1.9	0.87	NA
Taxon Data from the High Altitudes	2005–2009	2010–2014	2015–2019	2020–2023	lin. x coeff.	r²	CRC
Bombus lucorum (Linnaeus, 1761)	357	80	210	63	−75.2	0.51	HR
Bombus pascuorum (Scopoli, 1763)	210	115	167	54	−41.6	0.64	R
Bombus hortorum (Linnaeus, 1761)	86	48	19	32	−19.1	0.72	HR
Bombus pratorum (Linnaeus, 1761)	71	27	34	20	−14.6	0.69	HR
Bombus lapidarius (Linnaeus, 1758)	87	42	50	44	−12	0.55	HHR
Bombus terrestris (Linnaeus, 1758)	44	20	12	9	−11.3	0.85	HR
Bombus sylvarum (Linnaeus, 1761)	35	27	11	7	−10	0.95	HHHR
Bombus ruderarius (Müller, 1776)	30	22	8	10	−7.4	0.85	HHR
Bombus humilis (Illiger, 1806)	21	13	15	5	−4.6	0.81	HHR
Bombus confusus (Schenck, 1861)	12	0	3	1	−3	0.5	HHHR
Bombus ruderatus (Fabricius, 1775)	10	5	0	2	−2.9	0.74	HR
Bombus subterraneus (Linnaeus, 1758)	11	5	7	2	−2.5	0.73	HHHR
Bombus pomorum (Panzer, 1805)	8	2	0	1	−2.3	0.68	HHHR
Bombus hypnorum (Linnaeus, 1758)	14	12	10	8	−2	1	HR
Bombus argillaceus (Scopoli, 1763)	0	0	0	0	NA	NA	LR
Bombus muscorum (Linnaeus, 1758)	0	0	0	0	NA	NA	HHR
Bombus haematurus (Kriechbaumer, 1870)	0	0	0	0	NA	NA	HR
Bombus rupestris (Fabricius, 1793)	41	53	23	35	−4.8	0.75	HHR
Bombus campestris (Panzer, 1801)	101	85	55	25	−25.8	0.98	HHR
Bombus vestalis (Geoffroy, 1785)	9	3	5	2	−1.9	0.63	HR
Bombus bohemicus (Seidl, 1838)	157	34	70	40	−31.5	0.51	HHR
Bombus barbutellus (Kirby, 1802)	46	28	14	22	−8.6	0.67	HHR
Total specimens	1350	621	713	382	−281.2	0.77	NA
Total species	19	18	17	19	−0.1	0.02	NA

Table 8. Long-term trends of various Aculeata genera (above) and families (below) between 1988 and 2023.

Genera	1988–93	1994–99	2003–08	2012–17	2018–23	lin. x coef.	r²	Changes
Sceliphron	15	18	28	17	64	9.7	0.56	4.3
Cerceris	287	367	536	548	1084	177.5	0.81	3.8
Gorytes	72	74	180	247	209	44.7	0.78	2.9
Oxybelus	245	250	348	447	659	102.5	0.89	2.7
Priocnemis	33	33	127	68	83	13.5	0.26	2.5
Hedychrum	157	162	207	287	372	56.5	0.91	2.4
Crossocerus	195	1430	1064	288	370	−79.2	0.06	1.9
Chrysis	481	496	758	1089	903	143.7	0.75	1.9
Scolia	68	65	49	107	126	15.8	0.60	1.9
Diodontus	316	541	883	503	552	43.4	0.11	1.7
Ectemnius	241	245	540	305	255	8.8	0.01	1.1
Crabro	32	32	35	17	34	−1.1	0.07	1.1
Ammophila	87	78	101	106	82	1.8	0.06	0.9
Megascolia	0	0	0	1	3	0.7	0.72	NA
Families	1988–93	1994–99	2003–08	2012–17	2018–23	lin. x coef.	r²	Changes
Chrysididae	612	954	1454	1824	1707	306.0	0.89	2.8
Bembicidae	82	124	250	324	278	59.2	0.82	3.4
Psenidae	89	357	280	196	274	20.9	0.11	3.1
Philanthidae	338	419	581	600	1163	183.1	0.80	3.4
Sphecidae	130	157	220	308	345	58.1	0.97	2.7
Crabronidae	2110	5235	5503	3698	4405	305.3	0.13	2.1
Pemphredonidae	190	441	396	232	309	2.9	0.00	1.6
Halictidae	2225	3320	4524	4122	6633	961.8	0.86	3.0
Colletidae	1132	1538	1817	1834	2660	335.2	0.89	2.3
Andrenidae	1447	1518	2615	1573	2295	175.1	0.27	1.6
Megachilidae	1234	1421	1237	1522	1860	135.3	0.69	1.5
Apidae	1148	1143	1489	1354	2066	168.7	0.76	0.7
Aculeata total	10,681	18,010	22,516	19,167	25,740	3127.5	0.77	2.4

Table 9. Previously sporadically occurring Aculeata species, but not collected for at least 20 years, in the low-lying regions of the Carpathian Basin. Gray background and x-mark: species that still occur in the higher regions and the northern parts.

Family	Species	H. alt.	Family	Species	H. alt.
Chrys.	Cleptes semicyaneus (Tournier, 1879)		And.	Andrena fuscipes (Kirby, 1802)
Chrys.	Elampus constrictus (Förster, 1853)		And.	Andrena gallica (Schmiedeknecht, 1883)
Chrys.	Hedychrum chalybaeum (Dahlbom, 1854)		And.	Andrena nuptialis (Pérez, 1902)	x
Chrys.	Chrysis immaculata (Du Buysson, 1898)		And.	Andrena tridentata (Kirby, 1802)
Chrys.	Chrysura rufiventris (Dahlbom, 1854)	x	And.	Bombus mesomelas (Gerstäcker, 1869)	x
Chrys.	Stilbum cyanurum (Förster, 1771)	x	And.	Bombus cullumanus (Kirby, 1802)	x
Vesp.	Polistes semenowi (Morawitz, 1889)	x	Apid.	Bombus wurflenii (Radoszkowski, 1860)	x
Vesp.	Dolichovespula norwegica (Fabricius, 1781)	x	Apid.	Bombus quadricolor (Lepeletier, 1832)	x
Vesp.	Ancistrocerus ichneumonideus (Ratzeburg, 1844)		Apid.	Habropoda zonatula (Smith, 1854)	x
Vesp.	Ancistrocerus scoticus (Curtis, 1826)	x	Apid.	Thyreus orbatus (Lepeletier, 1841)
Vesp.	Delta unguiculatum (Villers, 1789)	x	Apid.	Thyreus truncatus (Pérez, 1883)	x
Vesp.	Microdynerus exilis (Herrich-Schäffer, 1839)	x	Apid.	Nomada baccata (Smith, 1844)
Pomp.	Priocnemis cordivalvata (Haupt, 1926)	x	Apid.	Nomada conjungens (Herrich-Schäffer, 1839)	x
Pomp.	Agenioideus apicalis (Vander Linden, 1827)		Apid.	Nomada moeschleri (Alfken, 1913)	x
Pomp.	Arachnospila alvarabnormis (Wolf, 1965)		Apid.	Nomada mutica (Morawitz, 1872)	x
Pomp.	Arachnospila rufa (Haupt, 1927)		Apid.	Nomada noskiewiczi (Schwarz, 1966)	x
Pomp.	Arachnospila sogdianoides (Wolf, 1964)		Apid.	Nomada obscura (Zetterstedt, 1838)
Pomp.	Episyron rufipes (Linnaeus, 1758)		Apid.	Nomada obtusifrons (Nylander, 1848)	x
Pomp.	Evagetes alamannicus (Blüthgen, 1944)		Apid.	Nomada opaca (Alfken, 1913)
Pomp.	Evagetes subglaber (Haupt, 1941)		Apid.	Nomada pulchra (Arnold, 1888)	x
Pomp.	Ferreola diffinis (Lepeletier, 1845)		Coll.	Colletes floralis (Eversmann, 1852)
Crab.	Diodontus medius (Dahlbom, 1844)		Hal.	Lasioglossum kussariense (Blüthgen, 1925)	x
Crab.	Mimesa crassipes (A. Costa, 1871)		Hal.	Lasioglossum damascenum (Pérez, 1910)	x
Crab.	Mimumesa beaumonti (van Lith, 1949)		Hal.	Lasioglossum intermedium (Schenck, 1870)
Crab.	Passaloecus brevilabris (Wolf, 1958)		Hal.	Lasioglossum podolicum (Noskiewicz, 1925)	x
Crab.	Passaloecus borealis (Dahlbom, 1844)	x	Hal.	Lasioglossum rufitarse (Zetterstedt, 1838)
Crab.	Pemphredon montana (Dahlbom, 1845)	x	Meg.	Pseudoanthidium alpinum (Morawitz, 1874)	x
Crab.	Pemphredon morio (Vander Linden, 1829)	x	Meg.	Coelioxys alatus (Förster, 1853)	x
Crab.	Psenulus chevrieri (Tournier, 1889)		Meg.	Megachile nigriventris (Schenck, 1870	x
Crab.	Psenulus fulvicornis (Schenck, 1857)		Meg.	Megachile pyrenaea (Pérez, 1890)	x
Crab.	Harpactus tumidus (Panzer, 1801)		Meg.	Hoplitis lepeletieri (Pérez, 1879)
Crab.	Crossocerus heydeni (Kohl, 1880)		Meg.	Hoplitis loti (Morawitz, 1867)
Crab.	Crossocerus walkeri (Shuckard, 1837)		Meg.	Hoplitis mitis (Nylander, 1852)	x
Crab.	Ectemnius borealis (Zetterstedt, 1838)	x	Meg.	Hoplitis papaveris (Latreille, 1799)	x
			Meg.	Hoplosmia scutellaris (Morawitz, 1868)	x
			Meg.	Osmia mustelina (Gerstäcker, 1869)	x
			Meg.	Melitta wankowiczi (Radoszkowski, 1891)	x

Table 10. Long-term trend of sawfly species between 1960 and 2022.

Taxon	1960–66	1967–73	1971–75	1986–98	1990–92	1996–99	2009–12	2019–22	lin x coef.	r²
Tenthredo mesomela (Linnaeus, 1758)	59	35	27	32	11	2	9	0	−7.61	0.88
Athalia glabricollis (Thomson, 1870	35	44	20	0	0	0	3	0	−6.07	0.68
Dolerus haematodes (Schrank, 1781)	4	7	3	2	0	0	1	0	−0.82	0.66
Tenthredo „arcuata” spec.gr.	84	99	2	4	7	0	22	1	−11.54	0.49
Tenthredopsis tarsata (Fabricius, 1804)	25	11	29	7	2	0	18	1	−2.68	0.34
Tenthredopsis nassata (Linnaeus, 1767)	21	8	24	0	6	10	11	1	−1.92	0.3
Cephus pygmeus (Linnaeus, 1767)	57	23	49	11	4	14	23	31	−3.50	0.22
Tenthredopsis litterata (Geoffroy, 1785)	20	5	3	0	6	2	8	4	−1.12	0.2
Tenthredo omissa (Foerster, 1844)	6	4	3	0	8	1	5	0	−0.42	0.12
Tenthredo zonula (Klug, 1817)	46	9	1	21	9	0	34	2	−2.36	0.11
Pachyprotasis rapae (Linnaeus, 1767)	34	32	25	39	16	10	47	13	−1.67	0.1
Macrophya blanda (Fabricius, 1775)	10	3	21	20	0	0	17	1	−0.90	0.06
Arge pagana (Panzer, 1798)	19	0	1	1	3	0	16	3	−0.39	0.05
Tenthredo atra (Linnaeus, 1758)	13	16	57	0	7	0	32	5	−1.67	0.04
Tenthredo vespa (Retzius, 1783)	19	11	0	0	17	0	20	2	−0.68	0.03
Tenthredopsis stigma (Fabricius, 1798)	33	10	3	4	6	0	19	19	−0.71	0.02
Tenthredo solitaria (Scopoli, 1763)	3	6	28	3	1	2	22	0	−0.25	0,00
Ametastegia glabrata (Fallén, 1808)	3	2	0	1	1	8	2	2	0.20	0.04
Macrophya ribis (Schrank, 1781)	2	0	6	2	18	10	6	0	0.52	0.04
Pachynematus annulatus (Gimmerthal, 1834)	0	0	1	0	0	0	6	0	0.37	0.08
Stethomostus fuliginosus (Schrank, 1781)	10	10	1	6	4	27	17	7	1.07	0.1
Pristiphora pallidiventris (Fallén, 1808)	0	0	0	4	0	0	7	4	0.82	0.27
Allantus cinctus (Linnaeus, 1758)	5	1	4	3	2	7	8	6	0.60	0.35
Tenthredopsis ornata (Serville, 1823)	0	0	0	0	0	0	8	4	0.81	0.44
Tenthredo amoena (Gravenhorst, 1807)	10	13	0	0	4	0	30	0	0.23	0.00
Tenthredo marginella (Fabricius, 1793)	13	18	0	2	9	0	14	19	0.35	0.01
Dolerus gonager (Fabricius, 1771)	14	6	18	23	1	3	21	17	0.35	0.01
Tenthredo temula Scopoli, 1763	21	10	16	10	6	0	44	14	0.82	0.02
Athalia lugens (Klug, 1815)	3	3	1	0	4	21	4	1	0.65	0.06
Tenthredo campestris (Linnaeus, 1758)	0	0	0	23	9	0	11	5	0.90	0.07
Arge nigripes (Retzius, 1783)	10	1	1	6	2	0	12	10	0.57	0.08
Tenthredo bifasciata rossii (Panzer, 1803)	13	10	9	5	12	0	31	13	1.01	0.08
Arge ochropus (Gmelin, 1790)	7	5	2	0	0	0	16	9	0.75	0,11
Arge berberidis Schrank, 1802	5	2	1	2	2	0	12	5	0.56	0,13
Arge melanochra (Gmelin, 1790)	37	19	9	8	12	0	101	40	4.86	0.13
Megalodontes plagiocephalus (Fabricius, 1804)	4	0	1	4	0	1	18	5	1.11	0.21
Macrophya duodecimpunctata (Linnaeus, 1758)	2	4	18	8	18	13	46	5	2.69	0.21
Tenthredo distinguenda (R.Stein, 1885)	4	0	0	0	4	0	21	8	1.63	0.31
Monophadnus pallescens (Gmelin, 1790)	6	7	1	36	6	8	41	25	3.50	0.31
Tenthredopsis sordida (Klug, 1817)	9	10	19	9	12	2	37	42	3.79	0.41
Arge enodis (Linnaeus, 1767)	7	1	2	0	15	8	56	26	5.25	0.46
Arge cyanocrocea (Forster, 1771)	10	4	6	3	6	0	37	58	5.79	0.47
Eutomostethus ephippium (Panzer, 1798)	32	20	31	34	31	18	62	82	6.17	0.47
Athalia rosae (Linnaeus, 1758)	9	22	21	47	12	18	93	247	23.54	0.51
Aglaostigma aucupariae (Klug, 1817)	9	16	5	13	12	10	56	49	5.88	0.54
Dolerus puncticollis (Thomson, 1871)	1	5	7	27	3	10	40	35	4.74	0.56
Dolerus nigratus (O.F. Müller, 1776)	11	8	2	54	15	28	75	101	11.95	0.66
Aglaostigma fulvipes (Scopoli, 1763)	10	15	6	28	11	68	74	113	14.11	0.76

Table 11. Long-term trend of selected butterfly species between 1980 and 2009 (CRC: Climate Risk Categories: PR: potential risk, LR: low risk, R: risk, HR: high risk, HHR: very high risk, HHHR—extreme high risk), STI: Species Temperature Index, SPI: Species Precipitation Index, HSI: Habitat Suitability Index.

Species	1990–94	1995–99	2000–04	2005–09	lin. x coeff.	r²	STI	SPI	HSI	CRC	SD	MEAN
Euphydryas maturna (Linnaeus, 1758)	8	17	28	23	3.94	0.68	7.64	715.62	0.046	LR	8.96	15
Boloria selene (Denis & Schiffermüller, 1775)	13	0	12	27	3.2	0.41	6.93	793.83	0.577	PR	9.31	11
Carcharodus flocciferus (Zeller, 1847)	2	2	3	4	0.31	0.31	9.92	844.02	0.174	R	1.05	3
Phengaris nausithous (Bergstrasser, 1779)	1	5	17	27	2.66	0.25	8.39	771.69	1.000	HHHR	9.81	13
Iphiclides podalirius (Linnaeus, 1758)	4	1	1	9	0.66	0.17	10.87	780.33	0.105	LR	3.08	4
Euphydryas aurinia (Rottemburg, 1775)	13	3	14	18	1.62	0.15	9.53	853.12	0.102	PR	7.82	12
Lycaena dispar (Haworth, 1803)	9	10	6	16	0.69	0.09	9.34	706.61	0.408	R	4.32	10
Parnassius mnemosyne (Linnaeus, 1758)	29	25	5	33	1.57	0.06	8.79	803.6	0.354	HR	11.70	21
Pieris ergane (Geyer, 1828)	3	0	16	8	0.63	0.04	11.69	826.43	0.408	HHR	5.72	7
Lycaena virgaureae (Linnaeus, 1758)	7	0	6	7	0.16	0.03	7.27	757.44	0.408	HR	2.64	5
Libythea celtis (Füssly, 1782)	3	0	3	1	0.09	0.02	12.19	814.02	0.707	HHR	1.22	2
Neptis sappho (Pallas, 1771)	12	14	6	6	0.03	0	9.4	741.16	0.408	HHR	4.83	9
Limenitis reducta (Staudinger, 1901)	0	3	2	0	−0.06	0.01	11.07	863.96	0.577	R	1.21	1
Chazara briseis (Linnaeus, 1764)	2	0	6	1	−0.11	0.01	10.29	782.68	0.333	HR	2.58	2
Papilio machaon (Linnaeus, 1758)	6	3	2	9	−0.31	0.04	9.28	737.43	0.053	PR	2.79	6
Maculinea teleius (Bergstrasser, 1779)	19	21	38	25	−1.17	0.05	8.6	798.04	1.000	HHR	9.85	30
Argynnis niobe (Linnaeus, 1761)	0	0	2	2	−0.2	0.06	8.5	782.2	NA	PR	1.52	2
Scolitantides orion (Pallas, 1771)	27	16	2	0	−1.48	0.07	8.98	796.79	NA	HHR	10.25	10
Spialia orbifer (Hübner, 1823)	6	3	9	6	−0.37	0.09	12.1	710.83	NA	HR	2.32	7
Brenthis ino (von Rottemburg, 1775)	0	0	18	1	−1.88	0.13	6.86	741.56	NA	HR	9.81	8
Colias chrysotheme (Esper, 1777)	23	1	4	3	−2	0.16	9.42	642.2	NA	HHHR	9.24	8
Glaucopsyche alexis (Poda, 1761)	3	3	1	7	−0.97	0.21	9.59	763.99	0.065	PR	3.95	5
Phengaris arion (Linné, 1758)	11	2	0	0	−1.46	0.34	8.64	802.03	0.500	R	4.71	4
Zerynthia polyxena (Denis & Schiffermüller, 1775)	3	4	0	12	−3.34	0.36	10.67	777.33	0.500	HR	10.40	11
Melitaea diamina (Lang, 1789)	2	1	11	3	−4.4	0.47	8.03	817.02	0.126	HR	12.06	10
Heteropterus morpheus (Pallas, 1771)	3	1	2	4	−2.4	0.53	9.52	764.79	0.250	HR	6.15	6
Iolana iolas (Ochsenheimer 1816)	5	0	0	0	−1.09	0.54	11.18	777.18	0.707	HHHR	2.76	2
Apatura iris (Linnaeus, 1758)	0	1	4	0	−1.77	0.54	8.51	774.96	0.316	HHR	4.52	4
Brenthis hecate (Denis & Schiffermüller, 1775)	14	0	9	1	−2.51	0.55	10.62	794.77	0.408	HHR	6.35	8
Lycaena hippothoe (Linnaeus, 1761)	3	4	0	4	−4.03	0.59	6.45	761.59	0.289	R	9.83	8
Nymphalis antiopa (Linnaeus, 1758)	0	0	1	1	−1.82	0.6	7.61	742.04	0.192	LR	4.41	3
Polyommatus daphnis (Denis & Schiffermüller, 1776)	5	9	0	0	−2.71	0.74	9.58	773.06	0.129	HR	6.05	6
Aglais urticae (Linnaeus, 1758)	6	2	3	2	−2.71	0.77	8.12	781.52	0.707	R	5.78	7
Total	242	151	231	260	−21.89	0.35	NA	NA	NA	NA	69.28	259

Table 12. Long-term trend of selected butterfly species between 2016 and 2023 (Duna-Ipoly National Park) (CRC: Climate Risk Categories: PR: potential risk, LR: low risk, R: risk, HR: high risk, HHR: very high risk, HHHR—extreme high risk), STI: Species Temperature Index, SPI: Species Precipitation Index, HSI: Habitat Suitability Index.

Species	2016–19	2020–21	2022–23	lin. x coeff.	r²	STI	SPI	HSI	CRC	SD	MEAN
Maniola jurtina (Linnaeus, 1758)	109	225	351	121	0.99	9.85	797.53	0.289	PR	121.03	228
Minois dryas (Scopoli, 1763)	20	101	168	74	0.99	9.52	807.1	0.200	HR	74.11	96
Aphantopus hyperantus (Linnaeus, 1758)	7	56	133	63	0.98	7.9	770.27	0.063	HR	63.52	65
Pieris brassicae (Linnaeus, 1758)	12	15	20	4	0.97	NA	NA	NA	PR	4.04	16
Parnassius mnemosyne (Linnaeus, 1758)	6	23	63	28.5	0.95	8.79	803.60	0.354	HR	29.26	31
Polyommatus coridon (Poda, 1761)	11	32	89	39	0.93	9.31	801.21	1.000	HR	40.36	44
Melanargia galathea (Linnaeus, 1758)	61	122	394	166.5	0.88	9.71	782.60	0.107	R	177.29	192
Brintesia circe (Fabricius, 1775)	11	25	157	73	0.82	11.07	796.98	0.204	HR	80.56	64
Plebejus argus (Linnaeus, 1758)	63	100	101	19	0.77	8.61	778.56	0.069	PR	21.66	88
Polyommatus icarus (Rottemburg, 1775)	168	170	352	92	0.76	9.07	789.28	0.144	PR	105.66	230
Apatura iris (Linnaeus, 1758)	0	0	2	1	0.75	8.51	774.96	0.316	HHR	1.15	1
Aricia agestis (Denis & Schiffermüller, 1775)	10	7	228	109	0.74	10.16	773.49	0.102	PR	126.74	82
Coenonympha glycerion (Borkhausen, 1788)	80	188	186	53	0.74	8.06	736.54	0.258	R	61.78	151
Scolitantides orion (Pallas, 1771)	2	0	33	15.5	0.7	8.98	796.79	0.333	HHR	18.50	12
Lasiommata megera (Linnaeus, 1767)	61	51	138	38.5	0.65	10.39	775.99	0.120	PR	47.61	83
Plebejus argyrognomon (Bergstrasser, 1779)	57	117	106	24.5	0.59	9.51	766.48	0.224	HHR	31.94	93
Vanessa atalanta (Linnaeus, 1758)	18	9	44	13	0.51	9.07	785.78	0.316	PR	18.18	24
Colias croceus (Geoffroy, 1785)	4	2	8	2	0.43	10.69	798.28	0.041	LR	3.06	5
Issoria lathonia (Linnaeus, 1758)	138	46	272	67	0.35	9.33	748.91	0.408	HR	113.65	152
Cupido alcetas (Hoffmannsegg, 1804)	3	0	7	2	0.32	10.81	844.04	0.183	HR	3.51	3
Plebeius idas (Linnaeus, 1761)	30	135	66	18	0.11	6.68	789.00	0.041	PR	53.36	77
Glaucopsyche alexis (Poda, 1761)	5	3	6	0.5	0.11	9.59	763.99	0.065	PR	1.53	5
Pieris rapae (Linnaeus, 1758)	73	42	86	6.5	0.08	NA	NA	NA	PR	22.61	67
Coenonympha pamphilus (Linnaeus, 1758)	138	69	165	13.5	0.07	8.96	793.06	0.096	PR	49.51	124
Coenonympha arcania (Linnaeus, 1761)	105	194	119	7	0.02	9.04	772.28	0.169	R	47.86	139
Spialia orbifer (Hübner, 1823)	1	32	4	1.5	0.01	12.1	710.83	0.224	HR	17.10	12
Maculinea teleius (Bergstrasser, 1779)	0	6	0	0	0	8.6	798.04	1.000	HHR	3.46	2
Polyommatus daphnis (Denis & Schiffermüller, 1776)	0	1	0	0	0	9.58	773.06	0.129	HR	0.58	0
Vanessa cardui (Linnaeus, 1758)	17	1	14	−1.5	0.03	9.04	770.51	0.083	PR	8.50	11
Lycaena dispar (Haworth, 1803)	5	0	4	−0.5	0.04	9.34	706.61	0.408	R	2.65	3
Colias hyale (Linnaeus, 1758)	36	77	10	−13	0.15	8.37	730.95	0.204	HR	33.78	41
Papilio machaon (Linnaeus, 1758)	36	7	22	−7	0.23	9.28	737.43	0.053	PR	14.50	22
Brenthis hecate (Denis & Schiffermüller, 1775)	4	0	2	−1	0.25	10.62	794.77	0.408	HHR	2.00	2
Leptidea sinapis (Linnaeus, 1758)	91	149	31	−30	0.26	9.11	770.01	0.144	PR	59.00	90
Anthocharis cardamines (Linnaeus, 1758)	231	87	136	−47.5	0.42	8.3	778.77	0.101	PR	73.21	151
Neptis sappho (Pallas, 1771)	9	13	0	−4.5	0.46	9.4	741.16	0.408	HHR	6.66	7
Aglais io (Linnaeus, 1758)	37	9	15	−11	0.56	8.84	787.83	0.236	R	14.74	20
Zerynthia polyxena (Denis & Schiffermüller, 1775)	20	1	2	−9	0.71	10.67	777.33	0.500	HR	10.69	8
Colias chrysotheme (Esper, 1777)	1	1	0	−0.5	0.75	9.42	642.20	0.408	HHHR	0.58	1
Melitaea diamina (Lang, 1789)	1	0	0	−0.5	0.75	8.03	817.02	0.126	HR	0.58	0
Lycaena hippothoe (Linnaeus, 1761)	1	0	0	−0.5	0.75	6.45	761.59	0.289	R	0.58	0
Cupido minimus (Füssly, 1775)	9	0	0	−4.5	0.75	8.76	817.26	0.105	R	5.20	3
Iphiclides podalirius (Linnaeus, 1758)	61	41	41	−10	0.75	10.87	780.33	0.105	LR	11.55	48
Libythea celtis (Füssly, 1782)	99	44	41	−29	0.79	12.19	814.02	0.707	HHR	32.65	61
Pyrgus malvae (Linnaeus, 1758)	190	151	80	−55	0.97	8.74	765.57	0.192	PR	55.77	140

Table 13. Long-term trend of number of individuals of butterfly families between 2013 and 2023 (Duna-Ipoly National Park).

Family	2013–19 (36 Days)	2020–21 (31 Days)	2022–23 (35 Days)	lin. x coeff.	r²	SD	MEAN
Papilionidae	119	72	128	4.5	0.02	30.07	106
Hesperiidae	325	292	202	−61.5	0.93	63.66	273
Pieridae	587	543	458	−64.5	0.97	76.55	523
Lycaenidae	545	815	1265	360.0	0.98	363.73	875
Nymphalidae	1294	1593	2670	688.0	0.90	198.62	1519
Total	2870	3315	4723	926.5	0.92	967.31	3636

Table 14. Long-term trend of number of species richness of butterfly families between 1980 and 2009 low-lying region of the Carpathian Basin.

Number of Captured Species (1980–2009) per 100 Days	1980–84	1985–89	1990–94	1995–99	2000–04	2005–09	lin x coeff.	r²	SD	MEAN
Papilionidae	4	4	4	4	3	4	NA	NA	0.41	4
Hesperiidae	13	12	13	11	10	13	−0.23	0.11	1.26	12
Pieridae	12	15	13	10	14	14	0.11	0.01	1.79	13
Lycaenidae	40	35	37	30	33	29	−1.94	0.62	4.20	34
Nymphalidae	50	52	47	42	49	51	−0.26	0.02	3.62	49
Total	119	118	114	97	109	111	−2.40	0.31	8.02	111

Table 15. Long-term trend of number of species richness of butterfly families 2016 and 2023 (Duna-Ipoly National Park).

Family	2016–19 (30 Days)	2020–21 (30 Days)	2022–23 (30 Days)	lin. x coeff.	r²	SD	MEAN
Papilionidae	4	4	4	NA	NA	0.00	4
Hesperiidae	10	10	10	NA	NA	0.00	10
Pieridae	15	13	12	−1.50	0.96	1.53	13
Lycaenidae	25	23	25	NA	NA	1.15	24
Nymphalidae	37	33	36	−0.50	0.06	2.08	35
Total	91	83	87	−2.00	0.25	4.00	87

Table 16. Previously sporadically occurring Lepidoptera species, but not collected for at least 20 years, in the low-lying regions of the Carpathian Basin. Species that still occur in the higher regions are marked by an x-mark.

Species	High Altitudes	Species	High Altitudes
Hesperiidae		Lasiocampidae
Carcharodus lavatherae (Esper, 1783)	x	Cosmotriche lobulina (Denis & Schiffermüller, 1775)	x
Pieridae		Geometridae
Colias myrmidone (Esper, 1781)	x	Eucrostes indigenata (Villers, 1789)	x
Pieris mannii (Mayer, 1851)	x	Lomaspilis opis (Butler, 1878)
Pieris bryoniae (Hübner, 1806)	x	Erebidae
Lycaenidae		Eublemma pannonica (Freyer, 1840)	x
Lycaena helle (Denis & Schiffermüller, 1775)	x	Noctuidae
Polyommatus damon (Denis & Schiffermüller, 1775)	x	Syngrapha ain (Hochenwarth, 1785)	x
Nymphalidae		Mesotrosta signalis (Treitschke, 1829)
Nymphalis vau-album (Denis & Schiffermüller, 1775)	x	Hyppa rectilinea (Esper, 1788)	x
Lasiommata petropolitana (Fabricius, 1787)	x	Fabula zollikoferi (Freyer, 1836)
Coenonympha tullia (Müller, 1764)	x	Photedes captiuncula (Treitschke, 1825)	x
Melanargia russiae (Esper, 1783)		Polia serratilinea (Ochsenheimer, 1816)
		Divaena haywardi (Tams, 1926)
		Oxytripia orbiculosa (Esper, 1800)
		Saturniidae
		Saturnia spini (Denis & Schiffermüller, 1775)

Table 17. Long and short-term trends of a number of individuals of nocturnal macrolepidoptera families between 1970–2022 and 2014–2022 completed with average, median values and determination coefficient for linear trend. For the explanation of the letters following the years, see the Materials and Methods section.

Families	1970a	1970b	1970c	1970d	1971a	1971b	1973a	1973b	1975	1976a	1976b	1977	1978a	1978b	1979a	1979b	1979c	1980a	1980b
Geometridae	720	11,723	1409	10,538	1167	1979	4513	7299	901	1749	1378	1013	1217	3367	2853	2954	2723	1669	1088
Noctuidae	1913	5431	1169	7987	1676	1259	5897	7599	3301	6560	5872	3481	3077	4855	1032	3560	4571	1059	2488
Lasiocampidae	19	15	115	269	45	21	211	393	126	206	81	94	45	93	42	43	55	54	35
Saturniidae	2	0	0	4	0	0	4	12	56	75	16	29	25	23	1	2	44	0	1
Sphingidae	16	5	38	182	2	0	218	161	237	570	205	142	81	167	15	202	142	12	87
Drepanidae	13	7	15	660	42	2	141	674	108	153	119	28	47	129	34	43	49	22	18
Notodontidae	21	321	64	1828	42	303	485	2160	319	598	278	99	191	808	50	233	403	22	85
Erebidae	580	7020	785	6150	1604	909	4845	12,153	989	2864	3259	884	735	3123	1501	1668	2298	1498	1283
Nolidae	6	158	153	1031	12	55	185	529	30	151	142	20	43	41	91	153	68	109	71
Thyatridae	11	545	18	543	29	103	92	502	18	109	151	57	40	415	66	26	166	62	25
Total	3318	25,228	3770	28,830	4683	4610	16,606	31,002	6115	13,104	11,501	5847	5533	13,045	5693	8901	10,681	4511	5191
Families	1980c	1981a	1981b	1986	1987	1990	1998	2000	2001	2005	2014	2015	2016	2017	2018	2019a	2019b	2019c	2020a
Geometridae	862	2552	1398	2573	1831	2517	2809	3924	2217	2265	570	1073	838	693	1105	1021	525	3578	766
Noctuidae	2817	1065	1990	3605	4094	5060	4729	NA	NA	6974	4619	2053	2587	2359	2862	5236	459	1972	1276
Lasiocampidae	64	42	47	221	55	163	200	364	471	119	18	6	34	60	104	160	3	31	67
Saturniidae	21	8	2	24	11	3	16	1	0	4	2	0	2	6	3	22	0	29	5
Sphingidae	125	10	95	215	85	252	170	288	353	238	78	73	169	193	164	213	2	18	103
Drepanidae	81	31	39	74	29	77	58	158	115	39	NA	NA	NA	NA	NA	NA	36	195	40
Notodontidae	302	35	92	376	96	142	290	102	77	280	197	153	182	134	184	166	36	284	124
Erebidae	1235	1709	1409	2415	1939	3245	1891	NA	NA	1683	1313	3240	2527	1899	3552	2694	683	3355	1330
Nolidae	33	135	73	143	110	48	73	NA	NA	37	7	58	52	24	8	14	58	94	18
Thyatridae	139	235	31	76	39	61	74	53	38	61	NA	NA	NA	NA	NA	NA	46	205	36
Total	5688	5839	5203	9786	8308	11,607	10,323	12,732	11,422	11,725	6914	6876	6450	5488	8121	9638	1842	9761	3775
Families	2020b	2020c	2020d	2020e	2021a	2021b	2021c	2021d	2022a	2022b	2022c	lin. x coeff.1970	r²	lin. x coeff.2014		r²	AVG	MD
Geometridae	2050	1047	1951	5336	1996	1121	3838	1056	2053	5916	1265	−79.7	0.15	113.9		0.22	1908	1749
Noctuidae	4842	1818	1127	3142	715	2597	7099	1664	1103	18,135	2089	−45.5	0.08	−26.4		0	2633	3481
Lasiocampidae	110	56	49	84	34	49	100	23	52	510	81	−1.8	0.04	0.76		0.01	67	55
Saturniidae	27	13	28	60	18	5	3	10	5	21	54	NA	NA	NA		NA	16	4
Sphingidae	130	111	70	18	81	88	147	113	35	417	176	−0.54	0	−3.13		0.07	99	142
Drepanidae	69	52	106	152	208	141	43	53	36	132	65	−1.77	0.02	−2.35		0.02	91	43
Notodontidae	292	208	106	659	208	159	341	226	223	951	490	−8.64	0.06	8.06		0.11	225	278
Erebidae	6623	1271	2891	3486	1402	2918	5369	1423	1252	14,262	1586	−17.9	0.01	16.73		0	2679	1604
Nolidae	335	77	195	142	129	35	708	36	56	3624	42	−1.91	0.01	12.47		0.15	129	91
Thyatridae	133	85	113	262	59	33	165	58	104	597	134	−2.33	0.04	−0.74		0.01	108	66
Total	14,755	4757	6613	13,341	4750	7146	17,911	5676	5020	44,700	5982	−113	0.06	136.67		0.03	9424	6914

Table 18. Long and short-term trends of species richness of nocturnal macrolepidoptera families between 1970–2022 and 2014–2022 completed with average, median values and determination coefficient for linear trend. For the explanation of the letters following the years, see the Materials and Methods section.

Family	1970a	1970b	1970c	1970d	1971a	1971b	1971c	1973a	1973b	1975	1976a	1976b	1977	1978a	1978b	1979a	1979b	1979c
Geometridae	114	107	147	176	120	68	143	132	139	100	122	105	120	124	127	126	134	133
Noctuidae	106	102	120	163	118	61	156	159	151	143	164	133	164	160	153	98	150	162
Lasiocampidae	6	3	7	13	5	2	11	10	13	11	12	9	10	10	9	9	9	11
Saturniidae	1	0	0	1	0	0	2	1	3	2	4	3	2	2	3	1	2	3
Sphingidae	2	1	5	10	2	0	11	8	10	7	10	9	11	9	9	6	10	9
Drepanidae	3	2	4	5	3	1	5	6	5	6	6	13	12	12	6	9	10	6
Notodontidae	11	11	10	25	9	14	24	24	20	27	26	24	25	25	25	9	21	22
Erebidae	37	33	39	57	42	24	54	51	54	40	39	49	47	44	52	47	52	48
Nolidae	3	8	5	8	4	2	7	5	7	5	7	5	5	4	4	5	6	3
Thyatiridae	4	4	5	6	4	2	6	7	6	3	6	7	6	7	7	4	6	6
Moths total	287	271	342	464	307	174	419	403	405	346	399	356	401	396	396	309	393	404
Family	1980a	1980b	1980c	1981a	1981b	1986	1987	1990	1998	2000	2001	2005	2014	2015	2016	2017	2018	2019a
Geometridae	109	122	103	122	125	148	144	90	120	117	121	136	115	143	127	127	149	134
Noctuidae	103	136	145	96	132	163	165	155	183	NA	NA	207	73	75	72	79	91	89
Lasiocampidae	8	7	8	6	9	11	9	9	10	12	9	8	8	3	9	9	10	9
Saturniidae	0	1	2	1	2	4	1	1	2	1	0	2	1	0	1	2	2	2
Sphingidae	5	7	8	5	9	10	8	7	7	10	8	10	9	8	9	9	9	10
Drepanidae	8	10	7	8	11	12	15	6	7	5	5	6	7	11	7	9	10	10
Notodontidae	9	19	20	10	14	23	23	27	27	15	17	27	17	17	17	17	17	19
Erebidae	49	44	44	42	44	55	54	53	58	NA	NA	63	29	34	33	33	43	43
Nolidae	5	7	4	5	7	9	8	3	5	NA	NA	6	3	3	2	2	3	1
Thyatiridae	3	5	6	3	5	6	8	4	4	NA	NA	2	NA	NA	NA	NA	NA	NA
Moths total	294	350	348	293	349	435	428	356	425	377	378	469	266	300	281	288	337	321
Family	2019b	2019c	2020a	2020b	2021a	2021b	2021c	2021d	2022a	2022b	2022c	lin. x coeff. 1970	r²	lin. x coeff. 2014	r²	AVG	MD
Geometridae	61	103	101	113	110	91	74	77	107	66	41	−0.77	0.17	−2.25	0.18	117	121
Noctuidae	36	108	86	113	89	153	128	125	95	126	58	−0.99	0.11	2.43	0.20	124	127
Lasiocampidae	1	7	7	9	7	8	4	3	6	7	2	−0.06	0.09	−0.20	0.13	8	9
Saturniidae	0	1	1	1	1	2	1	4	2	2	3	NA	NA	NA	NA	NA	NA
Sphingidae	5	2	6	6	4	9	9	8	3	8	3	0.01	0	−0.22	0.18	7	8
Drepanidae	1	5	3	5	3	5	2	4	3	2	3	−0.03	0.01	−0.45	0.54	6	6
Notodontidae	10	15	11	14	14	16	17	22	14	18	12	−0.07	0.03	−0.07	0.01	18	17
Erebidae	27	48	42	48	39	50	39	39	52	37	20	−0.12	0.03	−0.18	0.01	44	44
Nolidae	2	10	10	9	11	5	7	2	6	6	2	0	0.00	0.20	0.09	5	5
Thyatiridae	5	4	5	5	4	5	4	5	4	3	2	−0.03	0.07	−0.2	0.46	5	5
Moths total	142	304	268	324	277	344	285	289	295	276	147	−2.02	0.14	−2.16	0.04	334	342

Table 19. Changes in dominant Lepidoptera species over a 50-year period (ecotype codes according to Kanarskyi, Y. et al. [62] was employed, assigning ecotypes as follows: U (ubiquists), M1 (grassland mesophiles), M2 (seminemoral mesophiles), M3 (nemoral mesophiles), H2 (nemoral hygrophiles), X1 (grassland xerothermophiles), and X2 (seminemoral xerothermophiles), numeric ecotype codes are detailed above). Species that maintained their dominance in both time intervals are marked with a grey background.

Rank	1970–1981	Ecotype [62]	Ecotype [61]	2005–2022	Ectype [62]	Ecotype [61]
1	Xestia c-nigrum (Linnaeus, 1758)	M2	1	Paracolax tristalis (Fabricius, 1794)	M3	6c
2	Phragmatobia fuliginosa (Linnaeus, 1758)	M2	1	Eilema sororcula (Hufnagel, 1766)	M3	7
3	Miltochrista miniata (Forster, 1771)	M3	7	Miltochrista miniata (Forster, 1771)	M3	7
4	Rivula sericealis (Scopoli, 1763)	M1	9	Hypomecis punctinalis (Scopoli, 1763)	M3	6a
5	Chiasmia clathrata (Linnaeus, 1758)	M1	1	Eilema complana (Linnaeus, 1758)	M2	6c
6	Orthosia cruda (Denis & Schiffermüller, 1775)	M3	4	Cyclophora annularia (Fabricius, 1775)	M2	3
7	Orthosia gothica (Linnaeus, 1758)	U	3	Lithosia quadra (Linnaeus, 1758)	M3	7
8	Pseudeustrotia candidula (Denis & Schiffermüller, 1775)	M2	2	Orthosia cruda (Denis & Schiffermüller, 1775)	M3	4
9	Conistra vaccinii (Linnaeus, 1761)	M2	4	Polypogon tentacularia (Linnaeus, 1758)	H2	2
10	Lithosia quadra (Linnaeus, 1758)	M3	7	Pelosia muscerda (Hufnagel, 1767)	H2	7
11	Caradrina morpheus (Hufnagel, 1766)	H2	9	Eilema depressa (Esper, 1787)	M3	6a
12	Lomaspilis marginata (Linnaeus, 1758)	M3	5	Zanclognatha lunalis (Scopoli, 1763)	X2	3
13	Timandra comae (Schmidt, 1931)	U	2	Eilema lurideola (Zincken, 1817)	M2	7
14	Ceramica pisi (Linnaeus, 1758)	M1	3	Idaea aversata (Linnaeus, 1758)	M2	3
15	Eilema sororcula (Hufnagel, 1766)	M3	7	Helicoverpa armigera (Hübner, 1808)	U	1
16	Polypogon tentacularia (Linnaeus, 1758)	H2	2	Euphya biangulata (Haworth, 1809)	M3	4
17	Spilosoma lubricipeda (Linnaeus, 1758)	M2	2	Herminia tarsicrinalis (Knoch, 1782)	M3	3
18	Tholera decimalis (Poda, 1761)	M1	2	Athetis furvula (Hübner, 1808)	X1	0
19	Mythimna pallens (Linnaeus, 1758)	M1	1	Colocasia coryli (Linnaeus, 1758)	M3	4
20	Ascotis selenaria ([Denis & Schiffermüller], 1775)	U	1	Orthosia cerasi (Fabricius, 1775)	M3	6a

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Haris, A.; Józan, Z.; Schmidt, P.; Glemba, G.; Tomozii, B.; Csóka, G.; Hirka, A.; Šima, P.; Tóth, S. Climate Change Influences on Central European Insect Fauna over the Last 50 Years: Mediterranean Influx and Non-Native Species. Ecologies 2025, 6, 16. https://doi.org/10.3390/ecologies6010016

AMA Style

Haris A, Józan Z, Schmidt P, Glemba G, Tomozii B, Csóka G, Hirka A, Šima P, Tóth S. Climate Change Influences on Central European Insect Fauna over the Last 50 Years: Mediterranean Influx and Non-Native Species. Ecologies. 2025; 6(1):16. https://doi.org/10.3390/ecologies6010016

Chicago/Turabian Style

Haris, Attila, Zsolt Józan, Péter Schmidt, Gábor Glemba, Bogdan Tomozii, György Csóka, Anikó Hirka, Peter Šima, and Sándor Tóth. 2025. "Climate Change Influences on Central European Insect Fauna over the Last 50 Years: Mediterranean Influx and Non-Native Species" Ecologies 6, no. 1: 16. https://doi.org/10.3390/ecologies6010016

APA Style

Haris, A., Józan, Z., Schmidt, P., Glemba, G., Tomozii, B., Csóka, G., Hirka, A., Šima, P., & Tóth, S. (2025). Climate Change Influences on Central European Insect Fauna over the Last 50 Years: Mediterranean Influx and Non-Native Species. Ecologies, 6(1), 16. https://doi.org/10.3390/ecologies6010016

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