The fate of an intentional introduction of Formica lugubris to North America from Europe

Red wood ants (Formica s.str.) are not prevalent in the forests of North America, but commonly occur in conifer and mixed conifer forests in northern Europe and Asia. In 1971, a European red wood ant species, Formica lugubris, was intentionally established in a 35‐year‐old predominantly mixed conifer plantation approximately 30 km north of QC, Canada. The purpose of its introduction was to evaluate the potential of this species as a biological control agent against conifer‐defoliating Lepidoptera species. This red wood ant introduction was monitored periodically for about 5 years after establishment, but its long‐term fate has not been reported. We visited this field site in 2005 and found that this species was well established, and we could locate some of the nests that resulted from the original release. We mapped and measured over 100 nests around the site of original release, which ranged from 5 cm in height to over 1 m. We estimated the population of introduced ants to have grown to over 8 million in the last 34 years. Significant clustering of nests suggests that these nests may be one supercolony. F. lugubris has become a dominant understory arthropod in this mixed forest, and is likely to have ecological impacts, including effects at the community and ecosystem level.     

Abundance and distribution of organic mound-building ants of the Formica rufa group in Yellowstone National Park

Red wood ants (Formica rufa group) are ubiquitous in many conifer and mixed‐conifer forests of northern Europe and Asia. In contrast, relatively little is known about the abundance and distribution of the 24 North American F. rufa group species. As ants are important components of most soil invertebrate communities and are considered ecosystem engineers that alter the flow of energy and nutrients through terrestrial systems, it is important to gain information on their distribution and abundance. We conducted a survey for red wood ant mounds in Yellowstone National Park (YNP), Wyoming/Montana, USA, where human disturbance has been kept to a minimum for over 130 years. We found a total of 85 red wood ant mounds (0.11 to 0.17 mounds/ha) on 327 km of roads and 180 km of the hiking trails we surveyed. The occurrence of ant mounds was higher then expected by random distribution at elevations between 1600 and 2400 m, annual precipitation of 250 to 760 mm, middle and late successional lodgepole pine, late successional Douglas fir forest and non‐forested grassland/sagebrush prairie vegetation. Additionally, mounds were clustered in gently sloped not north‐exposed locations and in areas that had not recently burned. Most of the mounds detected were inhabited by Formica obscuripes Forel, which occupied 94% of the mounds sampled. Based on a multi‐criteria binary Geographic Information System model that we developed, we found that ant mounds were to be expected with a high probability in less then 1% of the YNP area. These results together with the detected low density and small size of the red wood ant mounds within the study area suggest that these insects have a much lower impact on invertebrate biodiversity and ecosystem processes, such as forest productivity and carbon and nutrient cycling on the ecosystem scale compared with their counterparts in European or Asian systems.     

The importance of diurnal and nocturnal activity and interspecific interactions for space use by ants in clear‐cuts

1. Full sunlight conditions in open clear‐cuts may limit the activity of ants as soil surface temperatures reach lethal levels. Therefore, differences may be expected between the diurnal and nocturnal activity of ants, and in the interactions between ant species. These predictions, however, have been poorly investigated so far. 2. The circadian activity of ants in clear‐cuts in managed forests in Poland was investigated. Repeated counts of ants were performed during the day and the following night at the clear‐cut edge and in the clear‐cut interior. Interspecific interactions and the effect of plant coverage were also considered. 3. Abundances of Formica fusca Linnaeus and red wood ants were higher during the day, whereas Myrmica were more common at night. Formica fusca, Lasius and red wood ants were more common at the clear‐cut edge than in the interior. Myrmica showed the opposite pattern, but at night, its numbers increased at the edge. Plant coverage positively affected F. fusca and red wood ants. 4. Red wood ants tended to be negatively associated with Lasius, whereas they were neutral for F. fusca. The negative association of red wood ants and Myrmica was stronger during the day compared to night. 5. The time of day was a strong driver of ant activity in the clear‐cuts, whereas the distribution of red wood ants was of lesser importance. It is concluded that circadian activity may substantially contribute to niche separation between coexisting species, therefore, studies performed exclusively during the day cannot reflect the real structure of the community.     

Stand type is more important than red wood ant abundance for the structure of ground‐dwelling arthropod assemblages in managed boreal forests

• 1 The relationships between red wood ants (Formica rufa group) and other ground‐dwelling arthropods were studied in young managed forests stands in Eastern Finland. The main objectives were: (i) to test the influence of stand type (dominant tree species; age: sapling versus pole stage) and numbers of red wood ants on the occurrence of other ground‐dwelling arthropods and (ii) to study the occurrence of red wood ants versus other arthropods on a distance gradient from ant mounds. We used pitfall traps set in 5–14‐year‐old sapling stands and 30–45‐year‐old pole‐stage stands of Scots pine (Pinus sylvestris L.) and birch (Betula spp.) forests.
• 2 Pitfall trap catches of red wood ants did not vary significantly between the forest stand types, although some groups of other arthropods showed clear responses to stand type (e.g. catches of other Formicinae and Gnaphosidae were higher in sapling stands than in pole‐stage stands). The number of red wood ants clearly explained less of the variation in assemblages of other ground‐dwelling arthropods than the forest stand type.
• 3 Red wood ant numbers decreased significantly with distance from the mounds, but the other ground‐dwelling arthropods were insensitive to this gradient or even showed a preference for proximity to ant mounds and high ant activity.
• 4 The results obtained in the present study suggest that wood ants do not have strong effects on several other ground‐dwelling arthropod groups in young managed forests other than in the immediate vicinity of their mounds.

     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

An innate template for enemy recognition in red wood ants

The hypothesis of the innate template for perception and recognition of the enemy image in red wood ants Formica aquilonia Yarr. was tested by initiating conflicts between ants and predatory ground beetles, their competitors for space. Live beetles and their models with different characters were used. In nature, ants respond selectively to such features of competitors as dark coloration, the presence of “outgrowths” (legs, antennae), body symmetry, the rate of movement, and scent. Comparison of behavior of ants from natural colonies and “naive” (laboratory reared) ones showed that ants having no experience of encounters with competitors responded aggressively to an integral and sufficiently realistic enemy image. This suggests that red wood ants possess an innate template for recognition of potential competitors. At the same time, the ability to single out the key features and complete the integral image seems to require accumulation of experience.     

The effect of red wood ant abundance on feeding damage by the pine weevil Hylobius abietis

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Within territory abundance of red wood ants Formica rufa is associated with the body condition of nestlings in the Eurasian treecreeper Certhia familiaris

Studies on individual reproductive success in relation to interspecific competition between distantly related taxa are scarce. We studied whether the abundance of red wood ants Formica rufa -group is related to the breeding habitat selection, fecundity and offspring quality in the Eurasian treecreeper Certhia familiaris, an old-growth forest passerine. The nest-box occupancy data were gathered over a five-year study period, whereas the breeding performance analyses were based on a two-year data set. The abundance of wood ants, measured within 50  m around the nest-boxes, was not related to nest-box occupancy rate, fecundity or the physiological stress of nestlings. In contrast, the abundance of wood ants was negatively related to the offspring quality and life-history traits, such as lowered body mass, subcutaneous fat reserves, and tarsus length at fledging. Our results suggest that exploitative competition between distantly related taxa may have considerable and adverse influences on nestling quality as measured by body mass, subcutaneous, fat and tarsus length. Red wood ants may decrease the fitness of treecreepers as the lower body condition of nestlings has the potential to impair recruitment into the breeding population and, additionally, impair the future reproductive effort.     

Are red wood ants (Formica rufa-group) tectonic indicators? A statistical approach

Recent research in seismically active areas indicated that the spatial distributions of geochemical soil gas anomalies and nests of red wood ants (RWA; Formica rufa-group) are strongly correlated. Here we applied a modified Hough Transform (mHT) and an Iterative Mode Detection (IMD) to RWA nest positions, which we mapped in two Southwest German study areas (Black Forest and Bodanrück), to test for statistical significance of correlation with tectonic features. RWA densities in the seismically active study areas reaching up to 1600 nests/100 ha are about 100× higher than overall values for Northern and Southern Germany. Since the shape of edges of the study area was found to strongly influence the selection of distribution patterns, all subsequent analyses were carried out for circular study areas. Results of the mHT applied on the RWA nests in both study areas clearly showed several modes which correspond to preferred directions. Centres of the modes further processed by IMD transferred into a GIS as RWA prototype lines showed very dominant directions of RWA nest distributions in right lateral strike-slip mode in WNW-ESE resp. NW-SE direction but also in extension direction caused by the recent main stress field in NW-SE resp. NNW-SSE direction. It could thus be clearly shown that the large scale spatial distribution of RWA nests directly reflects significant components of the present day stress field and its accompanying conjugated shear systems: Linear alignments of RWA nests indicate the course of active degassing faults zones and nest clusters indicate area-wide geochemical anomalies respectively crosscut zones of different fault systems. Furthermore, directions of re-activated shear systems, e.g. from Eocene-Oligocene but also from Late Jurassic that had been modified by overprinting due to changes of the main stress field could be identified. Therefore, RWA prototype lines complement and clarify the shear sense and the tectonic regime identified in previous tectonic studies. The high degree of statistical significance of these results will also allow a rating with former contrasting interpretations as to the drivers of RWA distributions.     

Wood ant colouration as an ecological indicator for the level of disturbance in managed coniferous forests

The search for ecological indicators of population well-being in natural and managed ecosystems is a crucial aspect of effective biomonitoring, conservation and nature protection. In long-term monitoring programs environmental stress has a measurable effect on naturally selected traits, such as body shape or size. However, changes in colouration provide information about early warning responses. The red wood ant Formica aquilonia is ecologically and territorially dominant among wood ant species in European boreal coniferous forest, and possesses variable red-brownish melanin-based cuticular colouration. F. aquilonia gynes, which are unfertilized queens, exhibit colour variability on the head, propodeum and abdomen, yet only head colour features allow setting clearly visible and symmetrical (left/right) classes of morphs. We studied phenotypic colour variability, melanisation and fluctuating asymmetry of colour patterns in faces of F. aquilonia gynes from natural (forest interiors) and disturbed (clear-cut zones and forest edges) habitats in Finland. We defined five variable, clearly visible and symmetrical (left/right) colour morphs of gynes' faces. Individuals of totally dark morph were present only in disturbed habitats. General analyses of melanisation degree showed, that gynes from disturbed habitats were significantly darker compared to those from forest interiors. Individuals from the same nest tend to have a similar degree of melanisation. The mean darkness of face was highest among individuals from forest-edge nests and the lowest from forest interior nests. In all habitat types the darkness of face increased with an increase in head width. Increase of cuticular melanisation in F. aquilonia gynes in disturbed habitats could be explained with the theory of thermal melanism and stress-induced immune defence. Although some amount of fluctuating asymmetry among left/right symmetrical colour variations on faces of F. aquilonia gynes was found, it was affected neither by habitat type nor by head width. Melanisation degree of red wood ants have a potency to be used as ecological indicator for the level of disturbance in managed coniferous forests and fluctuating asymmetry of colour variations in ants might be studied furtherly in cases of more severe environmental concerns.     

Behavior of red wood ants (Hymenoptera,Formicidae) during interaction with different symbiont partners

A comparative analysis of the behavior of Formica polyctena Först during interaction with different symbionts (free-living aphids Aphis grossulariae Kalt. and hidden larvae of the sawfly Blasticotoma filiceti Klug) was carried out. Red wood ants demonstrate different levels of functional differentiation in relatively constant groups of foragers collecting honeydew. A deep “professional” specialization with clear division of a number of tasks among foragers was studied in groups of ants tending aphids. Four professional groups of foragers with different tasks were revealed: “shepherds,” “guards,” “transporters,” and “scouts” (or “coordinators”). The groups of foragers caring for sawfly larvae mainly consist of unspecialized ants. Only few ants (about 5%) remain on duty on the fern plant near B. filiceti larvae and protect the food resource from competitors, especially from other ants. In addition, the ants demonstrate simpler behavior while collecting the larval excretion, resembling that at the sugar feeders. On the whole, the behavior of red wood ants is rather flexible. The level of functional differentiation in groups of foragers collecting honeydew is determined not only by the colony size and requirements but by the nature of their interaction with trophobionts, particularly, by the possibility of direct contact.     

Effects of<Emphasis Type="Italic"> Formica</Emphasis> ants on soil fauna-results from a short-term exclusion and a long-term natural experiment

Wood ants (Formica spp.) were hypothesised to affect the composition and greatly reduce the abundance of large-sized soil fauna by predation. This was tested in two ways. Firstly, a 4-year-long experimental study was carried out in a mixed forest. Five ant-free 1.3-m(2) plots were created by fenced exclosures within an ant territory. Five nearby plots had fences with entrances for the ants. In addition, five non-fenced control plots were selected. Soil fauna (e.g. Coleoptera, Diptera larvae, Collembola and Araneae) was sampled during the summers of 1997-2000. The soil fauna was affected by the exclosures but there was no detectable effect of ants on the soil fauna. Secondly, soil fauna was studied within a large-scale natural experiment in which the long-term (30 years) effects of red wood ants could be assessed inside and outside ant territories. This long-term natural experiment revealed no significant effects of ants on the abundance or composition of soil fauna. The results from the two studies indicate that the effects of wood ants on soil fauna are fairly small. The hypothesis that wood ants are key-stone predators on soil fauna could, thus, not be supported.     

Behavioral mechanisms of spatial competition between red wood ants (Formica aquilonia) and ground beetles (Carabidae)

Behavioral aspects of spatial competition between red wood ants (Formica aquilonia) and six mass species of Carabidae were studied in field and laboratory experiments. We showed that red wood ants essentially influence spatial distribution of ground beetles on their common territories. Transplantation experiments suggest that in newly established ants' settlements stronger forms of interrelations arise than in old stable colony. To examine the ability of beetles to avoid collisions with ants we used two experimental techniques. In laboratory, we tested carabids ability to avoid a clash in a Y-shaped labyrinth containing an active tethered ant in one section. In field experiments we compared quantitative characteristics of movements (such as crookedness of individual trajectories, speed of movement, the time spent on stops) for beetles placed close to ants foraging routes and on ant-free plots. All beetles studied displayed a clear tendency to learn, that is, to modity their behavior in order to avoid collisions with ants. Species that exhibited best parameters of learning were closer to ants by their size and characteristic movement, namely, Pterostichus oblogopunctatus and P. magus. Beetles' stereotyped behavioral tactics can be considered universal for avoiding collisions with any subject (for instance, with an ant) of a certain size and speed of movements. A set of tactics in the labyrinth included: (1) attempts to round the ant; (2) turns away after touching the ant with antennae; (3) turns away without a contact; (4) avoidances of a dangerous section; (5) stops near the ant with the antennae hidden. Comparing pairwise difference between four species shows that beetles use species-specific preference for definite combinations of tactics. Effective learning allows carabids to penetrate into ant foraging territory and partly avoide interference competition. It seems that red wood ants are not inclined to learn to avoid collisions with competing carabid species. Instead, they recognize an "enemy's image" and selectively attack relatively small predatory carabids rather than herbivorous species. Experiments with dummy beetles suggest that ants react for several hierarchically organized key characteristics of competitors such as speed of movement, dark color, bilateral symmetry, protuberances (legs, antennae), and smell. Among "professional" groups in ant family, guards and hunters react to beetles aggressively, whereas aphid tenders ignore them. Sophisticated combination of flexible and innate behavioral patterns enables insects to share territories and interact in the mode of relatively mild, spatial, competition instead of predation. Eliciting sets of hierarchically organized features of competitors that govern adjustment of spatial distribution in insects' community enable us to integrate ideas of classic and cognitive ethology.     

RETRACTED ARTICLE: Interspecific competition among ants in the boreal forest: Testing predictions from a linear hierarchical competition model

Competitive interactions often play an important role in local community structure, and particularly so in ant communities. We test predictions derived from a competition model proposed for Scandinavian ant communities by comparing ant nest densities inside and outside fifty randomly-selected territories of competitively dominant wood ants within 30 km of Ume?, northern Sweden. As predicted by the model, competitively-intermediate encounter species, as well as other territorial species, showed complementary occurrences with both studied wood ants, Formica aquilonia Yarrow and F. lugubris Zett.. In contrast, complementary abundances, i.e. lower density of competitively-inferior submissive species in the presence of wood ants, as predicted by the model, was not supported for all submissive species. Of the two studied wood ant species, the nest density of submissive species was negatively correlated only with F. aquilonia. Submissives as a group, as well as Myrmica-species, showed complementary abundances with F. aquilonia, but one Leptothorax and two Serviformica species had higher nest densities in the presence of this competitively-dominant species. We propose that, for Leptothorax, these deviations from the model predictions may be because of limited niche overlap with dominant wood ants, a small worker force and a timid behaviour which does not elicit aggression in wood ants. For the two Serviformica species, we propose a combination of protection against social parasites, inter-specific social control performed by F. aquilonia, and dominance relationships between competitively-inferior submissive species as reasons for the higher nest density inside F. aquilonia territories. Monogyny, and thereby smaller nests, lower worker force, smaller territory (as shown by this study) in F. lugubris, as compared to the polygyny in F. aquilonia, may also help explain the differences in their effects on subordinate species. Our analyses indicate that the linear competition hierarchy model proposed for Scandinavian ants accurately predicts the outcome of interaction between and community composition for dominant territorial and encounter species, but that it needs refinement with respect to the relationship between territorial and submissive species and the resulting community composition. Further studies are needed, especially addressing the complex relationships between these latter groups, and the effects of different competitively-dominant wood ants, to determine the mechanisms determining the outcome of these relationships and to more accurately predict community composition. Received 11 June 2007; revised 11 September 2007; accepted 17 September 2007.     

Ants protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis

• 1 Ants that protect food resources on plants may prey on (or deter) herbivores and thereby reduce damage. Red wood ants (of the Formica rufa group) are dominant ants in boreal forests of Eurasia and affect the local abundance of several herbivorous species.
• 2 The pine weevil Hylobius abietis (L.) is a herbivore that causes severe damage by feeding on the bark of coniferous seedlings within areas of forest regeneration.
• 3 We investigated whether ants can protect conifer seedlings from pine weevil feeding. In a manipulative experiment, ants were attracted to sugar baits attached to spruce seedlings and the damage caused by pine weevils was compared with control seedlings without ant‐baits.
• 4 The feeding‐scar area was approximately one‐third lower on the seedlings with ant‐baits compared with the controls. Besides red wood ants, Myrmica ants were also attracted in high numbers to the ant baits and the relative effects of these species are discussed.
• 5 The results obtained in the present study support the trophic cascade hypothesis (i.e. damage to herbivores is suppressed in the presence of predators). The decreased pine weevil feeding on the baited seedlings was probably a result of nonconsumptive interactions [i.e. the presence of (or harassment by) ants distracting pine weevils from feeding].
• 6 Understanding the role of ants may have important implications for future strategies aiming to control pine weevil damage. For example, maintaining suitable conditions for ants after harvesting stands may be an environmentally friendly but currently unexploited method of for decreasing weevil damage.

     

Efficacy of bifenthrin treatment zones against red imported fire ant

Exclusion of ants, particularly red imported fire ant, Solenopsis invicta (Buren), from homes, nursing facilities, hospitals, and electrical housings is an important strategy in urban and rural pest control. We conducted a laboratory bioassay to determine the repellency of granular bifenthrin (Talstar: rate 2.087 kg of formulated product/92.88 m2 or 4.6 lb formulated product/1000 feet2 or 4.2 g active ingredient/92.88 m2) to S. invicta foragers. In the field, we compared the efficacy of three widths (0.3, 2.0, and 3.0 m) of granular bifenthrin-treated zones at the rate 2.087 kg of formulated product/92.88 m2 and investigated the survival of individual ants successfully crossing the respective zones. Granular bifenthrin was nonrepellent to fire ant foragers in the laboratory. The 2.0- and 3.0-m treatment zones provided 100% protection for 7 wk after treatment and provided a reduction in the number of ants breaching the treated zone compared with the control for the remaining 9 wk of the study. This level of control may be tolerable for homeowners and is, therefore, considered an effective treatment for 15 wk after treatment. Hospitals, nursing homes, and electrical boxes would have to be treated on a monthly or bimonthly to remain ant free.     

Immigration and formation of mixed colonies in red wood ants (Hymenoptera,Formicidae)

Long-term field studies of the composition and spatial structure of settlements of ants of the Formica rufa group were carried out in two regions of Russia (Moscow and Arkhangelsk provinces). Fragmentation of damaged nests followed by reintegration of the fragments is the main way of formation of mixed colonies of ants from different nests (including different species). The principal factor of nest fragmentation is their damage by wild boars, bears, and in some localities, by poachers. The formation of mixed nests and nest complexes with participation of different Formica species was observed. They are formed by joining the ants from several damaged nests or by a colony from a destroyed nest immigrating into an intact one. Regular damage of many nests leads to the formation of broad zones of mixed colonies. The mixed colonies including 2–3 species of wood ants have recently become common. The phenomenon of mixed colonies raises a question as to the relative importance of two basic principles (sociality and specific identity) in the life of ant societies and demonstrates the priority of the social principle.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Interference competition,not predation,explains the negative association between wood ants (Formica rufa) and abundance of ground beetles (Coleoptera: Carabidae)

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