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Article

Immediate Response of Carabids to Small-Scale Wildfire Across a Healthy-Edge-Burnt Gradient in Young Managed Coniferous Forest in Central Europe

by
Václav Zumr
*,
Jiří Remeš
and
Oto Nakládal
Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague, Kamýcká 129, Praha 6–Suchdol, 165 00 Prague, Czech Republic
*
Author to whom correspondence should be addressed.
Fire 2024, 7(12), 436; https://doi.org/10.3390/fire7120436
Submission received: 20 October 2024 / Revised: 20 November 2024 / Accepted: 25 November 2024 / Published: 26 November 2024
(This article belongs to the Special Issue Integrated Vulnerability of Forest Systems to Wildfire: Implications on Forest Management Tools. VIS4FIRE Project)
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Abstract

:
Wildfire is a type of disturbance that plays a critical role in affecting forest ecosystems. Wildfires also have a significant effect on shaping arthropods communities. Carabids (family Carabidae) are often used as a bioindicator group of altered biocenoses. Methods: For carabid sampling, pitfall traps were used in three habitats, healthy-edge-burnt, fifteen days after the suppression of wildfire. Seven traps were evenly placed on each transect. In total, twenty-one traps were used for the study. Aim of the study: (i) evaluate the overall diversity of carabids, (ii) sex change and distribution within the studied habitats, (iii) dynamics of pyrophilous carabids. Results: In total, 1051 individuals within 42 species were recorded. The total number of species was higher in the edge and burnt habitats and differed from the healthy habitat. The abundance of carabids did not differ significantly across the three habitats. However, the healthy habitat exhibited both lower species numbers and abundance. Communities, species richness, and diversity indices were similar in the edge and burnt habitats, while the healthy habitat had lower species richness, diversity indices, and more homogenized communities. The overall sex ratio was nearly equal, with females comprising 519 individuals (49.4%) and males 532 individuals (50.6%), showing nonsignificant differences among study habitats. Among the nine most dominant species, a general trend of female dominance was observed. Many species showed different patterns in sex distribution in relation to the study habitats. Pyrophilous species accounted for the majority of individuals, comprising 55% of all carabids in the burnt habitat, predominantly represented by Pterostichus quadrifoveolatus. The rare species Sericoda quadripunctata was observed infrequently with only twenty-three individuals recorded. These two species are highly correlated, potentially indicating their near-habitat requirements. Males of pyrophilous species in general colonize the area in the earliest post-fire period. Conclusions: The immediate response of carabids to forest wildfire is significant, primarily influencing species richness and communities. While wildfire did not affect overall sex distribution, it shaped interspecies sex distribution across the study habitats.

1. Introduction

Wildfire is one of the important factors influencing the character and form of forest landscapes [1] especially in the maintenance of open forests [2]. Fires also alter the underground environment, altering its physical, chemical, and overall biological composition [3,4]. The frequency of fires has increased with human activities since the Holocene [5], primarily driven by settlement and land-use changes [6]. As a result, the western Palearctic is one of the most fire-affected regions in the world [7]. While fire is no longer used as a tool for landscape settlement management, human activity—mainly negligence—plays a primary role in the occurrence of forest fires [8]. Drier seasons may also contribute to creating environments more susceptible to wildfire [9]. Over the past 20 years, scientific research on wildfires has increased rapidly [10]. However, while climate change has received significant attention, the impact of wildfires on biodiversity remains underexplored [10], despite their crucial role in shaping biodiversity [11]. Wildfire creates unique natural environments that cannot be replicated by other forms of disturbance or management [12,13]. For this reason, controlled burning is often mentioned or used as a conservation tool [14]. Unlike wildfires, prescribed fires affect smaller areas and have lower severity [14], which can limit their effectiveness in supporting biodiversity of specific groups of fire-adapted species [15,16]. These factors may contribute to the broader pattern of insect decline observed globally [17].
The family Carabidae is common and widespread globally, with around 35,000 species [18]. In the Czech Republic, 517 species and subspecies are currently known, of which 171 (33%) are registered in the Czech Red List [19]. The carabids are a widely studied group of beetles [20], also recognized for their bioindicative value due to their ecological diversity, functional traits, and role as pest and weed regulators, all of which demonstrate environmental characteristics [20,21,22]. For example, carabids are often used to assess forest ecosystems [23,24], agricultural landscapes [25], and post-industrial areas [26]. They are also valuable for studying broader ecological changes resulting from long-term impacts on landscapes [21,22], as well as the effects of drought [27,28] and fire [16]. Carabids are among the most commonly studied groups of invertebrates in relation to wildfires [16]. Carabids include several species called the pyrophilous group “sensu stricto”, which are strongly adapted to fires. These adaptations include heightened sensitivity to smoke, high reproductive capacity, and the ability to fly over long distances [29]. In contrast, the “sensu lato” group is not directly adapted to fire by bionomic traits but is associated with open habitats [29]. This group often comprises smaller macropteran species, as larger brachypteran species are more severely affected by fire [13].
The Central European region is not strongly influenced by wildfires, unlike boreal forests or the Mediterranean region [30,31,32]. However, scientific interest in pyroentomology within Central Europe is increasing [33,34,35,36]. Fewer studies have addressed the direct response of carabids to fire [37]. Research has typically focused on temporal responses, ranging from annual [13,38,39,40] to multiyear lags [36,41,42,43]. This can significantly affect inferences regarding the behavior of carabids, and particularly pyrophilous species of carabids, which are very rare and often rapidly disappear from burnt areas [44]. Their detection can be virtually impossible over large areas after fire [13], potentially due to years of fire protection [45]. Conversely, research on biodiversity and the response of invertebrates to wildfire is limited due to the infrequency of such events [46] and the unpredictability of fire dynamics. Therefore, any study supporting pyrophilous insect knowledge is needed [29]. Additionally, topics such as the distribution and sex differentiation of carabids in relation to fire remain largely unexplored. Colonization by the sexes may provide clues to the bionomics of carabids and may provide insight into whether there is a sex difference in habitat colonization activity. Colonization of burnt areas can be massively driven by one sex of the carabids. Especially in pyrophilous species, this phenomenon is unexplored; colonization of freshly burnt areas may be driven by a high number of females associated with a high reproductive capacity strategy to maximize reproduction on unstable burnt areas, as described by Bell [29]. A potential bias in sex ratios in burnt areas, compared to unburnt conditions, necessitate direct assessment to verify this effect. The objectives of this study were to: (i) assess the immediate impact of fires on carabid communities, (ii) investigate their sex distribution, and (iii) investigate dynamics of pyrophilous carabids.

2. Materials and Methods

2.1. Study Site

The study was performed in the Czech Republic (Central Europe) in the municipal forest of the town of Horní Cerekev. The area is located at an altitude of 600 m above sea level, with an average temperature of 8 °C. The studied stands are typical acidic, commercial, even-aged coniferous forest stands, primarily consisting of Picea abies and Pinus sylvestris. This small forest complex covers an area about of 15 ha. On 30 April 2024, a negligent fire occurred during railway maintenance activities, affecting approximately 3.5 ha of the forest stands, which were divided by a railway into two parts. The fire was extinguished within a few hours and mainly affected clearings and young coniferous stands up to 20 years of age.
The fire had the character of a fast surface fire in the clearings. In the young stands, it manifested as both a surface and crown fire, resulting in the mortality of all affected trees. The spatial distribution of traps was influenced by the limited size of the fire and the age of the affected stands. For an appropriate comparison of fire effects, one part of the affected area was selected for beetle sampling, where young conifer stands (<20 years of age) were present. The study design was based on established recommendations [47]. The trap layout followed a transect pattern (Figure 1). Each transect contained seven pitfall traps spaced ten meters apart, with thirty meters between each row (transect). In total, twenty-one traps (7 × 3) were deployed for the study.

2.2. Sampling Beetle

Beetle sampling began on 15 May 2024, 15 days after the fire was extinguished, and data collection was completed on 23 September 2024. Pitfall traps were used for beetle sampling, as this method is widely recognized for collecting general epigeic organisms [47] and is very informative, especially for ground beetles [48]. Each trap consisted of four semi-transparent plastic components (Figure 2): a large container that kept its shape when buried in the soil and a smaller catch container with preservative fluid into which the funnel exited. The funnel of the trap was aligned with the ground to minimize the capture limitations for small species and thus potentially underestimate the overall results. Each trap was covered with a roof to prevent washout by rain, evaporation of the preservative fluid, and fallout of plant debris into the trap [47]. These aspects can strongly negatively affect the data obtained [48]. A metal roof was chosen to reduce evaporation on the exposed, dark surface of the post-fire ground, where high temperatures do not arise as they do under transparent roof [49]. The preservative solution used was vinegar (acetic acid 8%). The traps were replenished regularly every 2–3 weeks with fresh preservative fluid. Due to decreased beetle activity, the final replenishment and removal of traps was performed after an interval of more than one month. A total of six sampling sessions were conducted during the study period. Each captured adult of the family Carabidae was identified by the authors, in particular by co-author and carabid specialist Oto Nakládal. Additionally, each adult was classified by sex (male/female) based on external morphological characteristics, in particular the enlarged tarsal segment of the front legs and/or the genitalia.

3. Data Analysis

Analyzing the overall diversity was performed at the alpha, beta, and gamma diversity levels. All analyses were calculated based on each trap considered as an independent unit. To evaluate the number of species and abundance (α diversity), the analytical method used was a generalized linear χ2 model using a Poisson distribution for species numbers and negative binomial for abundance. This procedure was also used to evaluate sex differences in their abundance within the study habitats. The following R packages were used for data analysis: “glmmtbk” [50] and “car” [51]. For multiple comparisons, the “emmeans” package [52] was applied with post hoc Tukey HSD test, and differences were visualized using the “multcompView” package [53].
The carabid assemblage composition (β diversity) was evaluated by non-metric multidimensional scaling (NMDS), a recommended robust technique for indirect gradient analysis [54]. The NMDS analysis was conducted using the metaMDS function from the “vegan” package [55], with a “bray” dissimilarity matrix. To evaluate differences between carabid communities, a PERMANOVA global test was employed [56]. This analysis was also performed using the “adonis2” function in the “vegan“ package, and for multiple comparisons, the “pairwise.adonis” function with Bonferroni adjustment was used.
γ diversity was estimated using the sample coverage method as described by Chao and Jost [57], along with Hill numbers (q = 0 for species richness; q = 1 for the exponential of Shannon’s entropy index; and q = 2 for the inverse of Simpson’s concentration index), as proposed by Chao et al. [58]. The Chao and Jost [57] method provided a simplified estimation of species richness with lower sampling effort, calculated in the “iNEXT” package [59]. The estimation of significant differences was made using a conservative method based on non-overlapping confidence intervals [60].
A comparison of the sex distribution of the studied carabids was made based on the dominant species. Since many of the species recorded had only a few individuals, a sex comparison would not be relevant. Dominance was calculated based on Boháč [61] and was calculated as the percentage composition of the total number in the community (representation of populations in the total number of individuals). D = N/s where N is the number of individuals of a species and s is the total number of individuals. The dominance of each species was expressed in classes that corresponded to the following percentage ranges: eudominant species (ed) more than 10%; dominant species (d) 5–9.99%; subdominant species (sd) 2–4.99%; redundant species (r) 1–1.99%; sub-redundant species (sr) less than 1%. The total number of dominant species (including eudominant, dominant, and subdominant) in our study was nine. Analysis and figure preparations were created in R version 4.3.1 [62], and figures were plotted with “ggplot2“ [63].

4. Results

A total of 1051 individuals of carabids were captured during the study and classified into 42 species. Overall, the nine most dominant species accounted for 87% (913 individuals) of all individuals captured. The pyrophilous species Pterostichus quadrifoveolatus was the most abundant species. The list of all species recorded in the study is shown in Table 1.

4.1. Overall Diversity

Total number of species captured in traps varied across the studied habitats (χ2 = 18.6; p < 0.001). The greatest number of species per trap was recorded in the edge habitat, followed by the burnt habitat, with no difference between these two habitats (Figure 3A). However, no difference was found in the abundance (χ2 = 1.9; p = 0.38). The greatest variation in abundance was in the burnt habitat (Figure 3B). Carabid communities differed significantly between the habitats (Figure 4, F = 4.9; R2 = 35.6; p < 0.001), with the least variation observed in the healthy habitat. The main differences in communities were found between healthy habitats and both edge and burnt habitats (Table 2). A similar pattern was observed in gamma diversity (Figure 5), with edge and burnt habitats exhibiting the highest species richness and diversity, while the healthy habitat had the lowest richness and diversity.

4.2. Sex Distribution

The overall sex distribution of carabids was evenly represented, with females accounting for 519 individuals (49.4%) and males for 532 individuals (50.6%). This balanced sex distribution was also consistent across the studied habitats. The sex ratios in each habitat type—burnt habitat (average: F = 28.7, M = 32.7), edge habitat (F = 28.7, M = 27.3), and healthy habitat (F = 16.7, M = 16)—along with the total counts of each sex across all recorded carabid species, are presented in Table 1. Among dominant species, a general trend was observed indicating an over-representation of females in six species, with only three species having a higher abundance of males. Differences among sex distribution of dominant species across study habitats is shown in Figure 6. This shows a relatively high variability in the numbers of carabid males and females and the habitats studied. The movement activity (catches per day) of dominant carabids over the study period between males/females and study habitat is shown in Figure 7.

4.3. Pyrophilous Carabids

Two pyrophilous carabid species (P. quadrifoveolatus, S. quadripunctata) were recorded with 392 individuals (i.e., 37% of the overall number). Mainly males of the recorded pyrophilous carabid species inhabited the initial post-burn stage (Figure 7). P. quadrifoveolatus accounted for 30% of the total number of carabids in the edge habitat and 50% in the burnt habitat. S. quadripunctata was exclusively found in the burnt habitat, representing 5% of the total number of carabids in this habitat. Overall, pyrophilous species constituted the majority (55%) of the abundance in the burnt habitat. Both species were recorded together only in the burnt habitat. A strong relationship was found for the recorded pyrophilous species in the burnt habitat (Spearman r = 0.815, p = 0.025) (Figure 8).

5. Discussion

5.1. Carabid Assemblages

The reaction of carabids to the forest wildfire was very significant, with an observed increase in species richness compared to the previous stand condition (healthy habitat). A similar finding was reported by Zumr et al. [13] in healthy habitats, but their study did not confirm a positive relationship between forest fires and species richness when comparing different types of disturbance (burnt vs. unburnt). This discrepancy may be attributed to the affinity of a number of species for specific habitat or disturbance types, which could result in similar species richness across these conditions [66]. The edge habitat exhibited the highest species richness. As an ecotone, it represents a transitional area where a mixture of conditions from both burnt forest and healthy undisturbed forest combines. This environment provides optimal foraging and microclimatic conditions for more different species. The increased species richness of the edge habitat was due to the presence of species typical of the adjacent forest, in addition to edge-preferring species [67]. Differences in carabid communities between habitats were statistically significant. Specifically, communities in healthy (fire-unaffected) habitats differed significantly from those in burnt and edge habitats. This suggests that fire creates different community relationships in these conditions, as with a study from Heliölä et al. [68], which found carabid communities in the edges were more similar to forest and community than found in the clearcuts. In contrast to a large-scale wildfire [13], the decrease in carabid abundance following the small-scale fire in this study was not as pronounced. Although a reduction in abundance was observed, it was not statistically significant, potentially indicating lower fire intensity. For instance, Mason et al. [16] reported that prescribed burns did not result in significant negative effects on carabid abundance, as is often observed with wildfires. This could indicate that prescribed fires typically exhibit lower severity, as mentioned by Hiers [14]. The area affected by fires may also play a significant role, as indicated by findings of Zumr et al. [13]. In general, brachypteran species are the most affected in large-scale fires, but they may be able to colonize the area more rapidly following smaller-scale fires. As the area of the fire increases, colonization becomes even more difficult. Movement activities of carabids in the context of forest wildfires is an unexplored area. Therefore, studies examining carabid movement in forest or agricultural ecosystems were used as analogs for discussion. Anderson et al. [69] studied small macropteran carabids and found that long-distance movement among carabid beetles was rare, as most beetles moved about 5 m/day in the agricultural landscape. Studies examining the movement of carabids (mainly genus Carabus) across forest, edge, and fallow habitats were performed by, e.g., Růžičková and Veselý [70] and Sklodowski [71]. These studies generally observed very low movement activity across habitats, typically on a scale of a few dozen meters. For instance, Sklodowski [71] documented daily movement distances of up to 4 m in the species Pterostichus niger and Carabus. hortensis. In addition to their vulnerability to fire and low mobility, larger carabid species are limited in food supply in burnt areas. Typically, large carabids feed on snails or earthworms [18], which are invertebrates generally associated with moist environments and are not typically found in fully open burnt habitats. This may also suggest distinct movement patterns among different carabid species within ecotones [71]. Wetter conditions are more favorable for these groups, which is why forest moisture has been shown to have a positive effect on the abundance of carabids [43].

5.2. Sex Distribution

Evaluation of the sex proportion of captured individuals for eudominant, dominant, and subdominant species in each habitat showed that, for the pyrophilous species P. quadrifoveolatus, the burnt site was colonized at an increased rate by males immediately after this site was formed. Subsequently, the sex ratio gradually evened out over the season. P. quadrifoveolatus responded in a similar pattern in the edge habitat. A similar trend was observed for another pyrophilous species, S. quadripunctata, where males were more frequently captured at the early post-fire stage in burnt habitat. Similarly, Bell et al. [44] also observed primary colonization mainly by males. However, in our study, the sex ratio reversed very quickly, and females predominated during the season. However, the total number of this species captured in our study was low. Among the large brachypteran carabids of the genus Carabus (C. violaceus and C. hortensis) that were unable to fly to the freshly burnt habitat and, therefore, colonized it by ground movement, both male and female individuals were represented. The results suggested that the newly formed burnt site was an unsuitable habitat for these carabids. Despite the relatively quick re-establishment of the vegetation on the burnt site, their access across the edge habitat remained limited, as they were captured in much larger numbers in the healthy forest. Carabus scheidleri lives in various types of mainly open habitats [72]. The results indicated that the source of colonization by both sexes was other, less intensively burnt parts within the burnt habitat itself, rather than the surrounding healthy forest, where captures of this species were minimal. Longeard et al. [73] found that even minimal preserved vegetation on a burnt site can significantly enhance recolonization, including the less-mobile species. In freshly burnt habitats, smaller macropteran carabids showed a balanced sex ratio or slight female dominance; however, due to the low abundance of captured individuals, this pattern should be interpreted with caution. Differences in capture rates between males and females do not necessarily indicate an imbalanced sex ratio in the population. One possible source of the sex ratio imbalance is the higher movement activity of males during the reproductive period. Another factor may be the longer lifespan of females, as they persist to lay eggs, whereas males often disappear from the population shortly after copulation. Additionally, slight differences in habitat requirements for egg-laying by females and foraging by adults of both sexes may also contribute. Monitoring sex ratios of specific species over time and space may reveal more about species’ responses than simple species abundance measures.

5.3. Pyrophilous Carabids

In our study, we captured two species of carabids, which are considered obligatory pyrophilous due to their adaptations for searching and living in burnt forest sites [29]. However, only 23 individuals of S. quadripunctata were recorded with seven pitfall traps in burnt habitat. This scarcity is likely due to several facts: long-term fire suppression across European landscapes [45], low fire severity [40], and the uneven spatiotemporal distribution of fires [74]. For example, Zumr et al. [13] collected only four individuals (24 traps) of S. quadripunctata after a megafire in Central Europe during the first year after the fire. Similarly, a recent European survey by Aleksanov et al. [38] recorded only seven individuals (several dozen traps) over two years of post-fire forest surveys in 2021. However, in the past, this species appeared to be much more abundant in burnt forest sites. For example, Blažej [33] recorded 756 (3 traps) individuals of S. quadripunctata in the same state of the Czech Republic during the first year after a forest wildfire in 2006. Gongalsky et al. [37] observed several hundred individuals (30 traps) in the boreal forests of Sweden in the first year after the forest wildfire in 1999. Similarly, Süda et al. [75] documented nearly four hundred (20 traps) of Sericoda in Estonia after the first year after a wildfire in 2006. Moreover, other earlier studies focusing on this group of Sericoda reported many thousand individuals (24, 400, several hundreds), depending on the type of fire studied (prescribed, wildfire) in boreal forests of North America between 2000 and 2002 [12,39,40]. This may be explained by the fact that boreal forests in North America are frequently and extensively affected by forest fires [30], which may create more suitable conditions for obligatory pyrophilous species. The second species recorded, P. quadrifoveolatus, is thought to have pyrophilous adaptations, such as larger wings compared to the related P. oblongopunctatus, which may enhance its ability for mobile colonization of post-fire forest stands [29,76]. P. quadrifoveolatus is often referred to as a secondary pyrophilous carabid beetle; high fire severity providing environmental sterility is less important for this species compared to S. quadripunctata, which is primarily limited by the sterile conditions of the burnt soil that impact egg survival [44]. However, we found, given the close correlation of these two species, it is possible to expect them to seek similar characteristics of burnt sites, and P. quadrifoveolatus may benefit from higher soil sterility. S. quadripunctata was observed to inhabit exclusively the central areas of the burnt site, avoiding the edges. P. quadrifoveolatus showed no such preference in our study but needs a burnt habitat to live, so it occurred on both the burnt and the edge habitat that was also affected by fire. Similar behavioral patterns were reported by Gongalsky et al. [37]. Interestingly, P. quadrifoveolatus seems to be a more resilient species, persisting in burnt sites for longer periods and in relatively high numbers, with trap captures often reaching hundreds of individuals [33,38]. The maximum occurrence of this species is likely to be within seven years after a fire [43].

6. Conclusions

Fire is a part of the natural cycles within forest ecosystems. In our study, we assessed the immediate response of carabids to forest fire. Overall, fire had a positive effect on the species richness of the studied beetles, supporting the hypotheses that fire can enhance landscape biodiversity. We also observed an overall neutral effect of studied habitats on sex ratio of carabids. However, many species showed different patterns in sex distribution in relation to the study habitats. In particular, pyrophilous males mainly colonized the newly burnt habitats during the early post-fire period. This finding may help to better understand the bionomics of the generally poorly studied pyrophilous carabids [29]. Notably, the low number of individuals of the pyrophilous species, mainly S. quadripunctata, may indicate a severe decline in this species in central Europe. Further studies of forest fires are needed from a pyroentomological perspective, as the unpredictability and strong spatiotemporal distribution of forest fires pose challenges to research in this scientific discipline.

Author Contributions

Conceptualization, V.Z., O.N. and J.R.; data curation, V.Z. and O.N.; formal analysis, V.Z.; funding acquisition, J.R.; investigation, V.Z. and O.N.; methodology, V.Z. and O.N.; project administration, J.R.; resources, V.Z.; visualization, V.Z.; writing—original draft, V.Z.; writing— review and editing V.Z., O.N. and J.R. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by grant No. QK23020008, funded by the Ministry of Agriculture of the Czech Republic.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The raw data supporting the conclusions of this article will be made available by the authors on request.

Acknowledgments

We are grateful to Markéta Macháčová, who proofread the language of this manuscript. We would also like to thank the town Mayor and Vice Mayor of city Horní Cerekev, for the permission to conduct this research in municipal forests. Finally, we are grateful to three anonymous reviewers for their valuable comments.

Conflicts of Interest

The authors declare no conflicts of interest.

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Fire 07 00436 g001
Figure 1. Location of the study area with the distribution of forest stands affected by the wildfire. Points indicate the location of pitfall traps. The photos show the typical appearance of the study stands.
Figure 1. Location of the study area with the distribution of forest stands affected by the wildfire. Points indicate the location of pitfall traps. The photos show the typical appearance of the study stands.
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Figure 2. A pitfall trap installed in the ground, with dimensions of its parts (cm).
Figure 2. A pitfall trap installed in the ground, with dimensions of its parts (cm).
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Figure 3. Number of species (A) and individuals (B) per trap of carabid beetles. Boxes indicate the interquartile range 1–3Q, the solid line in the box represents the median, and the error lines are min–max values. The letters above the bars indicate differences.
Figure 3. Number of species (A) and individuals (B) per trap of carabid beetles. Boxes indicate the interquartile range 1–3Q, the solid line in the box represents the median, and the error lines are min–max values. The letters above the bars indicate differences.
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Figure 4. Non-metric multidimensional scaling (NMDS) shows differences in carabid communities between study habitats. Ellipses indicate 95% confidence interval. (*) Indicate sampling units. Solid points are centroids of carabid communities.
Figure 4. Non-metric multidimensional scaling (NMDS) shows differences in carabid communities between study habitats. Ellipses indicate 95% confidence interval. (*) Indicate sampling units. Solid points are centroids of carabid communities.
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Figure 5. Sample coverage-based rarefaction and extrapolation of species richness (q = 0), the exponential of Shannon’s entropy index (q = 1), and the inverse of Simpson’s concentration index (q = 2). Solid points indicate observed sample coverage with the number of recorded species. Dashed lines indicate extrapolation to 100% sample coverage. Shaded area represents 95% confidence intervals.
Figure 5. Sample coverage-based rarefaction and extrapolation of species richness (q = 0), the exponential of Shannon’s entropy index (q = 1), and the inverse of Simpson’s concentration index (q = 2). Solid points indicate observed sample coverage with the number of recorded species. Dashed lines indicate extrapolation to 100% sample coverage. Shaded area represents 95% confidence intervals.
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Figure 6. Numbers of individuals per sex (F—female, M—male) among dominant carabid species recorded in the study in three habitats.
Figure 6. Numbers of individuals per sex (F—female, M—male) among dominant carabid species recorded in the study in three habitats.
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Figure 7. Dominant carabid activity (catches per day over sampling period; events) categorized by sex and study habitats (represented by colored lines). The number of events indicates the intervals of trap exposition and days after the end of wildfire. (1. = 15.5–28.5 (15–28); 2. = 28.5–18.6 (28–49); 3. = 18.6–6.7 (49–67); 4. = 6.7–25.7 (67–86); 5. = 25.7–13.8 (86–105); 6. = 13.8–23.9 (105–146).
Figure 7. Dominant carabid activity (catches per day over sampling period; events) categorized by sex and study habitats (represented by colored lines). The number of events indicates the intervals of trap exposition and days after the end of wildfire. (1. = 15.5–28.5 (15–28); 2. = 28.5–18.6 (28–49); 3. = 18.6–6.7 (49–67); 4. = 6.7–25.7 (67–86); 5. = 25.7–13.8 (86–105); 6. = 13.8–23.9 (105–146).
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Figure 8. Relationship between two pyrophilous carabid species recorded in the burnt habitat. The shaded area represents the 95% confidence interval.
Figure 8. Relationship between two pyrophilous carabid species recorded in the burnt habitat. The shaded area represents the 95% confidence interval.
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Table 1. The list of carabids and their sex (F—female, M—male) recorded across study habitats by the largest number. Indicator Species Analysis (IndVal) approach [64], “indicspecies” package was employed with the “multipatt” function [65]. Results of IndVal (** p < 0.01; + p< 0.1) within affiliation of species to habitats. Dominance in carabid recorded community (D) following Boháč [61]. Undrelined: eudominant (ed) >10%; dominant (d) 5–9.99%; subdominant (sd) 2–4.99% and other redundant (r) 1–1.99%; sub-redundant (sr) < 1%.
Table 1. The list of carabids and their sex (F—female, M—male) recorded across study habitats by the largest number. Indicator Species Analysis (IndVal) approach [64], “indicspecies” package was employed with the “multipatt” function [65]. Results of IndVal (** p < 0.01; + p< 0.1) within affiliation of species to habitats. Dominance in carabid recorded community (D) following Boháč [61]. Undrelined: eudominant (ed) >10%; dominant (d) 5–9.99%; subdominant (sd) 2–4.99% and other redundant (r) 1–1.99%; sub-redundant (sr) < 1%.
Burnt Edge Healthy
DFM∑FM∑FM∑∑Total
Pterostichus quadrifoveolatus (Letzner, 1852)ed951552504673119 369
Carabus violaceus (Linnaeus, 1758)ed437412061463682150
Carabus hortensis (Linnaeus, 1758)ed415231538174966 **109
Bembidion lampros (Herbst, 1784)d302353182947 100
Pterostichus niger (Schaller, 1783)d13114175221772460
Notiophilus biguttatus (Fabricius, 1779)sd9101953813102350
Pterostichus oblongopunctatus (Fabricius, 1787)sd5611107171 129
Carabus scheidleri (Panzer, 1799)sd52796151 123
Sericoda quadripunctata (De Geer, 1774)sd14923+ 23
Carabus auronitens (Fabricius, 1792)r 3312113+16
Poecilus versicolor (Sturm, 1824)r 118311 ** 12
Amara curta (Dejean, 1828)r415246 11
Microlestes minutulus (Goeze, 1777)r448+123 11
Trechus quadristriatus (Schrank, 1781)r1 12 244811
Harpalus laevipes Zetterstedt, 1828 sr2 24261 19
Harpalus rufipes (DeGeer, 1774) sr415123 119
Poecilus cupreus (Linnaeus, 1758)sr246 33 9
Notiophilus palustris (Duftschmid, 1812)sr123224 7
Bembidion mannerheimii C.R. Sahlberg, 1827sr 314+ 4
Carabus linnei (Panzer, 1810)sr 2244
Calathus fuscipes (Goeze, 1777) sr 11112 3
Notiophilus aestuans (Dejean, 1826) sr112 11 3
Pterostichus melanarius (Illiger, 1798)sr 111123
Pterostichus strenuus (Panzer, 1796) sr 123 3
Abax parallelepipedus (Piller & Mitterpacher, 1783)sr 11 11 2
Badister lacertosus (Sturm, 1815) sr 2 2 2
Calathus melanocephalus (Linnaeus, 1758)sr1 1 11 2
Demetrias monostigma (Samouelle, 1819)sr 1 11 12
Epaphius secalis (Paykull, 1790) sr 112 2
Agonum spp.sr 1 1 1
Amara familiaris (Duftschmid, 1812)sr 1 11
Amara plebeja (Gyllenhal, 1810)sr1 1 1
Bembidion humerale Sturm, 1825 sr 1 1 1
Bradycellus harpalinus (Audinet-Serville, 1821) sr 1 1 1
Carabus glabratus (Paykull, 1790)sr 111
Carabus granulatus (Linnaeus, 1758)sr1 1 1
Cychrus caraboides (Linnaeus, 1758)sr 11 1
Dromius fenestratus (Fabricius, 1794)sr 11 1
Leistus ferrugineus (Linnaeus, 1758)sr 11 1
Nebria brevicollis (Fabricius, 1792)sr 11 1
Stomis pumicatus (Panzer, 1796) sr 11 1
Trichotichnus laevicollis (Duftschmid, 1812) sr 11 1
Total ∑ 2012294302011913921171122291051
Table 2. Pairwise comparison of carabids communities within non-metric multidimensional scaling. Significant differences are in bold. (Bonferroni adjustment).
Table 2. Pairwise comparison of carabids communities within non-metric multidimensional scaling. Significant differences are in bold. (Bonferroni adjustment).
FR2p. Adjusted
Healthy vs. edge5.5660.3170.003
Healthy vs. burnt8.1840.4050.003
Edge vs. burnt1.9840.1420.177
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Zumr, V.; Remeš, J.; Nakládal, O. Immediate Response of Carabids to Small-Scale Wildfire Across a Healthy-Edge-Burnt Gradient in Young Managed Coniferous Forest in Central Europe. Fire 2024, 7, 436. https://doi.org/10.3390/fire7120436

AMA Style

Zumr V, Remeš J, Nakládal O. Immediate Response of Carabids to Small-Scale Wildfire Across a Healthy-Edge-Burnt Gradient in Young Managed Coniferous Forest in Central Europe. Fire. 2024; 7(12):436. https://doi.org/10.3390/fire7120436

Chicago/Turabian Style

Zumr, Václav, Jiří Remeš, and Oto Nakládal. 2024. "Immediate Response of Carabids to Small-Scale Wildfire Across a Healthy-Edge-Burnt Gradient in Young Managed Coniferous Forest in Central Europe" Fire 7, no. 12: 436. https://doi.org/10.3390/fire7120436

APA Style

Zumr, V., Remeš, J., & Nakládal, O. (2024). Immediate Response of Carabids to Small-Scale Wildfire Across a Healthy-Edge-Burnt Gradient in Young Managed Coniferous Forest in Central Europe. Fire, 7(12), 436. https://doi.org/10.3390/fire7120436

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