Carabid Beetle Fauna Research Articles

Ecology and Conservation of the Dutch Ground Beetle Fauna - Lessons from 66 Years of Pitfall Trapping

Carl H. Lindroth was the first to present a comprehensive survey with his fundamental work on the Fennoscandian carabid beetles, covering all ecological aspects. The knowledge of the Dutch carabid beetle fauna is also exceptionally good what is especially thanks to Piet den Boer’s initiative and the work of his followers. Apart from the comprehensive book by Hans Turin (2000) on The Netherlands carabid beetles, I only see major publications on parts of a country or certain aspects, respectively, such as Trautner et al. (2017) on carabids of a German Federal State, Baden-Württemberg or Paill (2009) on endemic species in Austria. Turin et al. (1991) analysed the data bank on Dutch carabid beetles on the basis of 1.616 year samples from the period 1953-1983. The present book contains updated and extended analyses of the extraordinary huge data bank on Dutch carabid beetles of 4.363 pitfall trap year samples with more than 3 million specimens covering additionally the period 1984-2018.

Ground beetles (Coleoptera; Carabidae) as an indicator of ongoing changes in forest habitats due to increased water retention.

This study, concerning the epigeic fauna of carabid beetles (Coleoptera; Carabidae), was conducted in the north-east of Poland, in an area which is part of the Dąbrówka Forest Subdistrict and has been included in the “Small water retention program for the Province of Warmia and Mazury in 2006–2015”. The purpose of the study was to assess the impact of the water retention implemented within the framework of the above program on assemblages of ground beetles. These insects are highly sensitive to any anthropogenically induced transformations. This analysis was based on the interactions among the analyzed insects caused by changes occurring in their habitat. During the 3-year study, 5,807 specimens representing 84 species were captured. The water storage had a significant influence on the structure of the Carabidae assemblages. Before the earthworks were constructed for the project, the beetle assemblages had comprised a large group of xerophilous species, whereas after the small retention reservoirs had been created, an increase in the contribution of hygrophilous species was noticed. The results indicate that the retention works cause alterations in the water and environmental conditions of the habitats, and thereby effect changes in the composition of Carabidae assemblages. Moreover, modification in water relations within a habitat causes long-term changes in the structural and functional diversity of the beetles.

Community composition of ground beetles (Coleoptera, Carabidae) in citrus orchard in North-Eastern of Algeria

Carabid beetles are natural enemies of insects pests and are weed seeds predators. They have been often used to study the ecological effects of different agricultural measures, because they are highly sensitive to habitat change. The Carabid beetle fauna is little known in Algeria, especially in agrosystems, thus we undertaken a study of this group of insects in citrus orchard located in the city of Guelma. Carabids were collected by pitfall trapping from december 2017 to may 2018. A total of 16 beetle species and 1072 individuals were recorded. The dominant species were consecutively as follows: Amara subconvexa , Brachinus sclopeta, Phyla rectangulum, Harpalus siculus and Percus lineatus. The data of the species traits as food preference and flying ability show that carnivorous and macropterous species dominate in the study area. The activity of the species was mainly observed during the spring period.

Evolution in Extremis: The When, How, and Why of Hawaiian Carabid Beetles

The Hawaiian Archipelago is the most isolated oceanic island system in the World, separated from the nearest source areas by more than 4000 km. Five independent colonization events have resulted in diversification of a native carabid beetle fauna in excess of 400 known species. This diverse assemblage is disharmonic, with the major radiations restricted to the platynine genus Blackburnia Sharp (139 species), the subgenus Nesocidium Sharp of Bembidion Latreille (21 species), and the moriomorphine genus Mecyclothorax Sharp (239 species). Biogeographical, ecological, and evolutionary attributes of these three radiations are compared in order to determine factors crucial to carabid beetle diversification in this most-isolated situation. Biogeographical attributes include the age of origin of the constituent radiation, the island likely colonized by its common ancestor, and the biological characteristics, where known, of the colonizing ancestors for each independent radiation. Ecological attributes include the amount of habitat specialization undergone during each radiation, taking into account the primordial habitat colonized and the subsequent pattern of occupation of different habitat types during diversification. Evolutionary attributes include brachyptery, body-size evolution, sexual selection, and the evolution of specialized body conformations. It is shown that ecological specialization—i.e., occupation of a diverse array of ecological zones and microhabitats—in concert with reduced dispersal ability brought on by evolution of brachyptery are positively associated with enhanced levels of diversification. Comparing sympatric island faunas, it is shown that the latter operates synergistically with body size, as the smaller-bodied Mecyclothorax beetles speciate much more rapidly than the larger-bodied Blackburnia on Maui and Hawai῾i Island. Nonetheless, small body size does not gaurantee high diversity, as Bembidion beetles attain body sizes similar to Mecyclothorax beetles. Age of origin of a radiation is a subsidiary criterion for diversification given that the Mecyclothorax radiation commenced only 1.2 Ma, whereas it is hypothesized that Blackburnia have been resident in the Hawaiian archipelago for upwards of 28 Ma. Thus especially for Blackburnia we are constrained in our ability to know all of the evolutionary products of the radiation due to extinction of presumably all or nearly all species that occupied the now-sunken islands northwest of the oldest high island of Kauai. We are fortunate to know of several extinct Blackburnia species discovered in lowland subfossil deposits in Kauai, and these species provide crucial information now regarding future patterns of extinction. Sexual selection can be demonstrated for the Bembidion subgenus Nesocidium, and is a likely explanation for genitalic evolution over parts of the Mecyclothorax radiation, but it is not a phenomenon pervasively associated with increased levels of speciation.

Post-fire fauna of carabid beetles (Coleoptera, Carabidae) in forests of the Mordovia State Nature Reserve (Russia)

Post-fire fauna of carabid beetles (Coleoptera, Carabidae) in forests of the Mordovia State Nature Reserve (Russia)

Influence of mowing measures on carabid beetle fauna (Coleoptera: Carabidae) in a post-agricultural area

Background and purpose: Some agricultural practices are considered to be useful tools in biodiversity conservation. Therefore, carabid beetles were collected on post-agricultural fallow land in Western Poland in order to study the impact of mowing treatment. Materials and Methods: Following a before-after-control-impact (BACI) study design over the period of two years (2013 – 2014), standard arrays of pitfall traps were installed on six study sites, of which three were treated by mowing at the beginning of July in the second year of study. The influence of this treatment was analysed statistically with respect to the most frequently collected species, selected ecological traits, as well as the mean individual biomass of the carabid assemblages (MIB). Additionally, Non-metric Multidimensional Scaling (NMDS) was carried out. Results: A total of 1995 individuals belonging to 40 species were collected, with species numbers ranging from 11 to 21 and numbers of individuals ranging from 76 to 278 in the samples. Although some species reacted significantly to the mowing treatment and numbers of individuals of forest species significantly decreased on the treatment sites, in general rather weak effects were observed as a result of the mowing measures. The weak effects of the mowing measures may be explained by the low cutting intensity (only once a year) and differences in environmental conditions between the years of study. Conclusions: The results of the study are assumed to be useful in the context of planning mowing measures in order to conserve biological diversity. Yet, the results also underline the importance of long-term studies.

Body mass distributions along successional gradients in epigeic carabid beetle fauna (Coleoptera: Carabidae)

Background and purpose: Body mass distributions may be related to the stage of succession of a habitat and provide with information for assessing successional processes. Therefore, body mass distributions of carabid assemblages were studied in three research areas, which were post-industrial areas near the city of Belchatow (Central Poland) planted with different tree and shrub species, moist and wet forest stands in the Puszcza Knyszynska forest (Northeastern Poland), and beech stands in the Ruhr valley (Western Germany) in order to analyze the changes in body mass distributions within single assemblages along successional gradients. Materials and Methods: For each carabid assemblage, the mean individual biomass (MIB) as well as MIB standard deviation (SD) and coefficient of variation (CV) were calculated. SD and CV were plotted against the age of study sites and MIB values, respectively. Analyses of Covariance (ANCOVA) were carried out with SD and CV as dependent variables and the age of study sites and MIB as covariates. Results: SD was low at young stages of succession, but increased rapidly and plateaued at advanced stages in beech but not in wet forest stands. Accordingly, CV was low at very young stages of succession, showing a rapid increase and subsequent decrease in the beech stands, whereas in the wet stands it stayed on a constant level. ANCOVA revealed significant differences in SD and CV between the research areas and significant changes with age or MIB, but, with the exception of CV as dependent variable and MIB as covariate, interactions were also significant. Conclusions: The results of the study suggest that data on body mass distributions within single carabid assemblages may be useful in the assessment and comparison of successional stages and processes between different habitat types.

Reassembling a lost lowland carabid beetle assemblage (Coleoptera) from Kauai, Hawaiian Islands

A late Holocene but prehistoric carabid beetle fauna from the lowland Makauwahi Cave, Kauai, is characterised. Seven extinct species – Blackburnia burneyi, B. cryptipes, B. godzilla, B. menehune, B. mothra, B. ovata and B. rugosa, spp. nov. (tribe Platynini) – represent the first Hawaiian insect species to be newly described from subfossil specimens. Four extant Blackburnia spp. – B. aterrima (Sharp), B. bryophila Liebherr, B. pavida (Sharp), and B. posticata (Sharp) – and three extant species of tribe Bembidiini – Bembidion ignicola Blackburn, B. pacificum Sharp and Tachys oahuensis Blackburn – are also represented. All subfossil fragments are disarticulated, with physical dimensions and cladistic analysis used to associate the major somites – head, prothorax and elytra – for description of the new species. The seven new Makauwahi Cave species support recognition of a lowland area of endemism adjoining Haupu, a low-stature 700 m elevation ridgeline in southern Kauai. Four of the extinct Blackburnia are adelphotaxa to extant species currently found at higher elevations in Kauai. Addition of these lowland specialists to the phylogenetic hypothesis undercuts applicability of the taxon cycle for interpreting evolutionary history of these taxa. Two of the extinct species are Kauai representatives in clades that subsequently colonised younger Hawaiian Islands, enhancing support for the progressive biogeographic colonisation of the archipelago by this lineage. And three of the extinct Blackburnia species comprised larger beetles than those of any extant Kauai Blackburnia, consistent with the evolution of island gigantism in the lowland habitats of Kauai.

Changes in carabid beetle fauna (Coleoptera: Carabidae) along successional gradients in post-industrial areas in Central Poland

An inventory of the carabid fauna was carried out in two post-industrial areas in central Poland. The two areas studied were a heap of ash generated by a power station and a colliery spoil heap. In each area sites of different ages were investigated using pitfall traps over an eight year period from 2004 to 2011. At the end of this period each of the youngest sites was as old, or even older, than the next oldest site studied in each area. A pine forest growing on natural soil close to the ash heap was included as a reference study site. Changes were described in terms of the numbers of species and individuals, total biomass, Mean Individual Biomass of Carabi- dae (MIB) and the most frequently collected species. Indirect and direct gradient analyses were carried out in order to determine the environmental basis of the major pattern in variation and analyse the relationships with current environmental parameters. During the eight years of this study 5032 individuals of 84 species were collected. Numbers of individuals decreased significantly with the age of the sites on the ash heap and the MIB values increased significantly on both heaps. Changes in the most frequently collected species were more pronounced on the spoil heap, the study sites on which covered a time span of 26 years, than on the ash heap, for which the period was only 11 years. Characteristic species differed on the spoil and ash heaps and numbers of species were lower on the spoil heap. Despite differences in speed of succession at the sites studied, there was a tendency for the MIB to increase after about 9 to 16 years. Indirect and direct gradients analyses confirmed that the stage of succession (age of the study sites) was an important factor determining the carabid assemblages at the sites studied. The results of this study are important for the restoration and management of post-industrial areas.

The effect of the construction and renovation of a highway bypass in Central Poland on the carabid beetle fauna (Coleoptera: Carabidae)

The aim of this research was to define the effect of the construction and renovation of a highway bypass around the town of Skepe on the environment and assemblages of carabid beetles. This four-year study was based on catching samples of beetles using pitfall traps. The traps were set first in 2008 and later from 2010 to 2012, after the renovation of the road, along roadside verges adjacent to two ca. 100-year-old pine forest stands, wet habitats and fallows of arable fields and inside these habitats about 25 m from the road (control plots). The carabids collected were identified to species and numbers of individuals and dominance values of each of the species were calculated for each study plot and each year. Furthermore, the dominance of the different species of plants was assessed. Numbers of species, numbers of individuals, percentage share of forest species and individuals, and Shannon diversity values of the carabid assemblages were analyzed. Multivariate statistical analyses (ordinations) were carried out using the dominance of the different species of carabids and using the ecological indicator values of plants as environmental factors. The construction of the bypass in 1987 resulted in more diversified carabid assemblages along the roadsides. Carabid fauna along the roadsides differed from that in the control plots in forests and in terms of some parameters from the control plots in wet habitats, but not statistical from the control plots in the fallow areas of arable fields. The renovation of the road 22 years after construction did not result in any significant changes in the characteristic parameters of the carabid assemblages along the roadsides, but gradient analysis indicated that it did have an effect. In terms of the environmental conditions of the habitats based on the plant associations occurring there the roadside habitats differed from the control plots in forests and wetlands. Since the number of sites studied is rather low, the results of this study have to be considered to be preliminary, but may provide research hypotheses for further research on highway roadsides.

Inventory of the carabid beetle fauna of the Gaoligong Mountains, western Yunnan Province, China: species of the tribe Zabrini (Coleoptera, Carabidae).

A ten-year multidisciplinary, multi-national and multi-institutional biodiversity inventory project in the Gaoligong Shan region of western Yunnan Province, China generated more than 35,000 specimens of the beetle (Coleoptera) family Carabidae. In this report, first of a planned series, we focus on diversity in tribe Zabrini. Our study of just over 1300 specimens of zabrine carabids from the project, all in genus Amara Bonelli, found a total of 13 species, all previously described, to occur in the study area, with none of them strictly endemic. We present a key for identification of adults of these species, as well as nomenclatural data, diagnoses, illustrations of dorsal habitus and male genitalia, and information about geographical, altitudinal and habitat distributions within the study area and overall geographical distribution for each species. Distributions of the species within the study area are compared, and broader geographical range patterns are characterized. We also discuss a possible role of the Gaoligong Shan region as one source area for the present-day fauna of the Himalaya and southern edge of the Qinghai-Xizang (Tibetan) Plateau.

Conservation potential for heathland carabid beetle fauna of linear trackways within a plantation forest

Abstract Exploring conservation and connectivity potential of landscape linkages implies studies on the level of population and community structure. We examined carabid beetle communities within a network of open linear trackways embedded in a plantation forest, in a region where conservation interest focuses on heathland associated species. The relative importance of environmental characteristics such as width, orientation, age of surrounding plantations and vegetation height on species associated with grassland, heathland and sandy habitats (GHS species) was explored in 36 trackway elements. A total of 3554 individuals were collected of which 1465 were GHS specialists. Linear trackways supported important carabid beetle diversity including 17 GHS species of which six are nationally scarce within the UK. The age of adjacent plantations was the most important factor determining community composition of carabids with GHS species favouring trackways surrounded by younger plantations. For predicting number and abundance of GHS species within a subset of trackways surrounded by younger plantations, models incorporating sward height in verges showed the strongest relationships. But results were statistically significant only for predicting abundance of the dominant GHS species Harpalus rufipalpis, which favoured wider trackways with lower vegetation and greater cover of bare sand. Current conservation management using forage harvesting of trackways does not improve habitat quality for GHS carabids. Seemingly continuous network of linear linkages is interrupted by sections of less suitable habitat surrounded by older plantations. Conservation potential could be enhanced by widening of trackways and management by physical disturbance of soil and vegetation.

Native and Alien Carabidae (Coleoptera) Share Lanai, an Ecologically Devastated Island

The carabid beetle fauna of Lanai includes nine native Hawaiian species, and four extralimital alien species. The three species of Mecyclothorax Sharp precinctive to Lanai—M. filipes (Sharp), M. mundanus (Sharp), and M. flavipes, new species—are taxonomically revised. Lanai non-native carabid species include Metacolpodes buchanani (Hope) (new island record), Gnathaphanus picipes (MacLeay), G. multipunctatus (MacLeay) (new island record), and Perigona nigriceps (Dejean) (new island record). A key to species of Harpalini introduced to the Hawaiian Islands is provided. Historical records of native Lanai carabid species indicate that they occurred in both summit cloud forest and in riparian corridors of the mesic and dry forest communities. Present-day records indicate restriction of these species solely to the cloud forest zone. The known historical diminution of summit cloud forest establishes a minimum area that has supported persistent populations for several of Lanai's native carabid taxa. Three of the four alien species have been recorded from cloud and mesic forest communities as well as lowland dry forest not known to support native carabid beetles. Currently, native and non-native species exhibit extensive distributional overlap. However, based on fine-scale, field-collection data, this geographic sympatry overstates the extent of ecological interactions between native and non-native taxa. Whereas native species occur in the cloud forest community, non-native species are more broadly distributed in anthropogenically-disturbed situations including the ecologically devastated dry forest community as well as smaller patches of disturbance within the native cloud forest.

Erratum: Harpalus rufipes (Coleoptera: Carabidae) Is not Known from New York State

Hajek et al. (2007) surveyed the carabid beetle fauna of soybean, Glycine max (L.) Merr., Þelds in central NewYork state in 2004 and 2006.We reported capture of one specimen of Harpalus rufipes (DeGeer) from Wayne Co., NY, in 2004. This specimen actually represents Harpalus puncticeps (Stephens). Table 1 in Hajek et al. (2007) should be corrected to show H. puncticeps in Þeld 5, not H. rufipes. The total number of carabid species sampled in this study thus changes to 59. Representatives of all 59 species are deposited in the Cornell University Insect Collection as CUIC voucher lot 1255. Cardenas et al. (2007) report that H. rufipes has expanded its range to near Montreal, QC, Canada, from an initial introduction on Prince Edward Island in 1937. Krinsky andOliver (2001) reported Connecticut records from as far west as NewHaven. However, as of this erratum, these remain the reported western distributional limits of this Old World species in the New World.

Plant and carabid beetle species diversity in relation to forest type and structural heterogeneity

The aim of this study was to evaluate the influence of forest structure (mainly resulting from human uses) and forest type (the identity of the dominant tree species) on biodiversity. We determined the diversity of two taxonomical groups: the understory vegetation and the edaphic carabid beetle fauna. We selected eight types of forest ecosystems (five replicates or stands per forest type): pine (Pinus sylvestris) plantations of three age classes (10, 40 and 80 years since reforestation), an old-growth relict natural pine forest, and four types of oak (Quercus pyrenaica) stands: mature forests with livestock grazing and firewood extraction, mature forests where uses have been abandoned, “dehesa” ecosystems and shrubby oak ecosystems. The results obtained by a global PCA analysis indicated that both tree size and dominant species influenced the ordination of the 40 forest stands. In general, carabids were more sensitive to changes in forest heterogeneity and responded more clearly to the analysed structural variables than the understory vegetation, although the species richness of both groups was significantly correlated and higher in case of oak forests. Pine forest ecosystems were characterised by the lowest species richness for both taxonomical groups, the lowest plant diversity and by the lowest coefficients of variation and, consequently, low structural heterogeneity. As a result, it was very difficult to discriminate the effects of the spatial heterogeneity and the dominant tree species on biodiversity.

Rediscovery of Blackburnia anomala (Coleoptera: Carabidae), in East Maui, Hawai'i, after a 107-Year Hiatus

The highly distinctive and diverse native Hawaiian carabid beetle fauna includes a suite of species not recently observed in nature. These are predominantly historical residents of the mesic Acacia koa forest formation. We report rediscovery of one of these species, Blackburnia anomala (Blackburn), in the shrubland formation near Paliku Cabin, and in koa forest of Kaupo Gap. Prior records of B. anomala are limited to the leeward edges of historical koa forest near Olinda, on the northwestern slope of Haleakala. Rediscovery on the far southeastern side of Haleakala Crater in similar, though conserved, habitats suggests that other long-missing koa associates may persist in similar situations on Haleakala.

The flora and carabid beetle fauna of a mature and regenerating semi-natural oak woodland in south-east Ireland

An investigation of the plant and carabid beetle communities of a mature and adjacent regenerating oak woodland site was carried out from June to September (1999) in County Wexford, south-east Ireland. Phytosociological classification (using 10 m×10 m quadrats) identified the mature site as belonging to the coryletosum sub-association of the Blechno–Quercetum association. Vegetation was recorded in ten 2 m×2 m quadrats at each site within which pitfall traps were placed to sample carabids. Median plant species richness per quadrat was greater in the regenerating site (11) than in the mature site (9.5), although this was not statistically significant. The median species richness of carabids per pitfall trap in the mature site (6) was significantly greater than in the regenerating site (3.5) at P<0.01. There was a significant positive correlation between carabid species richness and leaf litter depth. Further analysis revealed positive correlations between carnivorous/omnivorous ground beetle abundance and litter depth suggesting that availability of prey in leaf litter could be one of the causes for greater carabid species richness and abundances at the mature site. Although the woodland structure is very different at the two adjacent sites investigated, the similarity in plant species composition suggests that the regenerating site will recover in the future.

DISTINCT GROUND BEETLE (COLEOPTERA: CARABIDAE) ASSEMBLAGES WITHIN A NEW YORK STATE WETLAND COMPLEX

Carabid beetle communities were compared for adjacent marsh and bog biotopes at the McLean Bogs Preserve, Tompkins Co., New York by means of pitfall-trap sampling. Though the sampled marsh and bog habitats were only 200 m distant, with the umbrotrophic bog isolated from the marsh/fen complex by a Wisconsin-aged glacial esker of only 7 m ele- vation, the resident wetland carabid species assemblages differed significantly between the sites during the spring and summer seasons. Of 62 species observed in the wetlands, 36 were found exclusively in the marsh biotope, 17 were exclusive to the bog site, and only 9 were found at both sites. This level of wetland habitat fidelity was maintained in spite of potential colonization of the two sites each spring by adult beetles dispersing from overwintering sites in surrounding forest edge habitats. Pitfall sampling found 59 native carabid species, of which 52 were found during a survey conducted from 1916-1925, suggesting that the McLean Bogs Preserve sup- ports locally resident populations for the vast majority of carabid species found during this study. The McLean Bogs Preserve supports geographically southern, peripheral populations of the bog-specialist Platynus mannerheimiiDejean, as well as four marsh/fen-inhabiting species— Trechus crassiscapusLindroth, Bembidion muscicolaHayward, B. praticola Lindroth, and Bra- dycellus semipubescens Lindroth—emphasizing the role this preserve plays in maintaining the distributional ranges of both bog- and marsh-resident taxa. The distinct faunas observed in these two proximate biotopes clearly illustrate that comprehensive conservation of the carabid beetle fauna of northeastern North America should involve preservation of a variety of wetland types.

Introduction and Distributional Expansion of Trechus obtusus (Coleoptera: Carabidae) in Maui, Hawai'i

Trechus obtusus Erichson (tribe Trechini), native to Europe and North Africa and introduced to the Pacific coast of North America, is recorded for the first time from East Maui Island, Hawai'i, based on collections made at Haleakala National Park in September 1998. The species subsequently expanded its distribution to include Polipoli Springs State Recreation Area, East Maui. Range expansion has averaged 3 km per year, based on documented absence of T. obtusus from the Polipoli Springs area in 1998. All Hawaiian individuals are macropterous, even though European and North American populations of T. obtusus are dimorphic for wing configuration, with the brachypterous form most common in long-established populations. The source area for the Hawaiian invasion is hypothesized to be Oregon or the San Francisco Bay area, based on the closest match in the frequency of macroptery between specimens from Hawai'i and those from those mainland areas. Monomorphic macroptery of the Hawaiian populations suggests that the founder population was small, with estimates ranging from as few as 6 individuals to as many as 25, assuming the founding propagule was drawn at random from populations in the western United States. Baseline abundance data are presented for Polipoli Springs State Recreation Area, where T. obtusus co-occurs with seven native Mecyclothorax species (tribe Psydrini), establishing the opportunity for long-term assessment of the impact of the introduced species on the sympatric native fauna. Means to identify T. obtusus in the context of the Hawaiian carabid beetle fauna are presented.

What is the origin of the carabid beetle fauna of dry, anthropogenic habitats in western Europe?

AbstractAimTo study the origin of the carabid beetle fauna of dry, open anthropogenic habitats.LocationVarious parts of Europe.MethodsCollecting by hand in habitats influenced by humans (arable land, sand pits, ruderal places, road verges, meadows) and naturally open habitats such as talus, supralittoral sand dunes, alvar as well as littoral habitats in south and central Scandinavia. Studies made in other parts of Europe were included in the discussion.Results and conclusionThe material comprises about 37,700 specimens. About 8% of the Scandinavian species occurring in anthropogenic habitats may have a littoral origin, about 10% may originate from forests whereas about 9% are euryoecious. A majority of the species (about 66%) originate from primarily open, dry habitats and many of them constitute a steppe or steppe‐like element. They invaded the bare landscape soon after the deglaciation and probably survived in naturally open habitats (supralittoral sand dunes, talus, alvar, old riverbeds) during the postglacial warm period, but some species have a recent origin in Scandinavia.