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Nebria brevicollis

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Nebria brevicollis
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Coleoptera
Suborder:
Adephaga
Family:
Carabidae
Subfamily:
Nebriinae
Genus:
Nebria
Subgenus:
Nebria (Nebria)
Species:
N. brevicollis
Binomial name
Nebria brevicollis
(Fabricius, 1792)

Nebria brevicollis, belonging to the Carabidae family, is a ground beetle. With nearly 500 species and more than 100 subspecies, N. brevicollis is the most diverse genus within the tribe Nebriini. Members of the genus occupy a wide range of habitats.[1] It is a species of ground beetle native to Europe and the Near East, but introduced to the western United States[2] and Canada.[3] The rapid expansion of the beetle in North America is characteristic of an invasive species.[4] Due to the variation in habitat, their diet consists of small arthropods including Collembola, Diptera, earthworms, mites, and spiders.[5] Most prey taken by Nebria brevicollis have been observed to be less than 4 mm in length.[4]

Description

Nebria brevicollis has a body which is dark brown, with reddish-brown on the tibiae, tarsi, palpi and antennae. The pronotum has two lateral setae on either side, and is densely punctuate across the entire base.[3]

This species is most abundant between October and December, then from January through mid-May. It is of modest size, ranging in length from 10–14 mm. It has two sets of wings, protected by a hard shell that is black or dark brown. Only a few beetles have functional flight muscles.[6] No sexual dimorphism has been observed between males and females.[7] They are night-active.[8]

Distribution and habitat

These beetles are plentiful across the globe. In Europe, it is found in all countries and islands except the Azores, the Canary Islands, the Channel Islands, Franz Josef Land, Gibraltar, Madeira, Malta, Monaco, the North Aegean Islands, Novaya Zemlya, San Marino, the Selvagens Islands, Svalbard and Jan Mayen, and Vatican City.[9]

In 2008, it was reported as introduced in western Oregon, U.S.A.,[2] where it has been found in highly disturbed sites as well as in native old-growth forest stands. It has also now been found in Washington State,[10] Northern California, as well as in southern British Columbia, Canada.[3] The rapid expansion of Nebria brevicollis in North America has caused researchers to question whether it meets the criteria of an invasive species. The rate of detection, and the rapid range expansion into Oregon, are characteristics of an invasive species.[4] In addition, its presence has been documented from Europe to North America and from sea beaches to alpine caves. The ecological range these beetles exhibit is unrivaled by any other carabid species in Oregon.[1]

The primary habitat is the cover of shaded leaf litter in deciduous forests; however, it can also be found in disturbed habitat such as parks, gardens, and agricultural lands. In North America, it has found in the summits, forest and meadows of mountains (at over 1200 meters elevation).[10] Unlike other members of Nebria, this species avoids moist areas.[2]

Physiology

Long and large wings protected by the strong exoskeleton of N. brevicollis

Flight

Despite having large and long wings, Nebria brevicollis has a low flight potential, which is a common trait among ground beetles. While all beetles have flight muscles that work antagonistically – when one shortens, the other stretches – only a small percentage are functional. In addition, the beetles with flight potential have a short flight period. The arrested development of the flight muscles, causing the inability to fly, may be an evolutionary choice. Rather than investing energy into the building of flight muscles, these beetles use their energy for the metabolic costs of larval growth.[6]

Dispersers are characterized by their ability to fly as a result of having function fly muscles. Within the N. brevicollis species, dispersers have been observed to form under favorable laboratory condition. Beetles that were fed in excess and kept at an ideal temperature, always emerged with flight muscles. The development of flight muscles took three weeks.[6]

All beetles hatch with flight muscles. Whether the muscles become arrested or not depends on the amount of food available during larval development. Crowding, lack of food, and low temperatures will not produce dispersers. Physiologically, the breaking down of the flight muscles creates more room for the ovaries and the eggs within the abdomen of the beetle.[6]

Genetics

In a study published in 2012, researchers Ikeda, Nishikawa, and Sorta found evidence that the loss of flight within a species promotes beetle diversification. Using carrion beetles, to which the order Coleoptera belongs to, researchers demonstrated higher genetic differentiation in flightless beetle species compared to flight-capable species. Flight ability can facilitate better mating and access to resources. However, the walking ability of N. brevicollis is high and does allow for the colonization of new habitats and exchange between local populations.[6] There is an energetic cost associated with the maintaining of the flight muscles. The evolutionary loss of flight muscles has allowed for beetles to invest energy into other organs, such as those essential for survival and reproduction.[7]

Within flightless species, researchers showed higher genetic differentiation compared to flight-capable species – a result of restricted gene flow between species. They further observed speciation rate within flightless populations to be higher than that of flight-capable ones. Thus, a higher number of genetically distinct lineages can be observed within flightless species. Flightless species is a common pattern among the different families of Coleoptera with Nebria brevicollis likely choosing to use its energy towards larval growth rather than the building of flight muscles.[6]

Behavior

A study found evidence of associative learning in N. brevicollis. Associative learning may lead to increased fitness advantages. The genotype, the physical and social environment, as well as the animal personality, influence one's interactions with the environment. Natural selection may act upon associative learning and personality traits, including boldness, aggressiveness, sociability, and exploratory tendencies.[5]

In 2012, researchers Sih and Del Guidice constructed a model for 'fast' individual cognitive styles and 'slow' individual cognitive styles. Higher expressions of a trait are thought to increase an individual's likelihood for reward or allow for more acquisition of the environment while, simultaneously, increasing one's risk at the cost of accuracy.[11] Greater exploratory behavior may be correlated with increased foraging success.[5] 'Slow' cognitive styles individuals make fewer mistakes; however, are slower to acquire reward and knowledge.[11]

Recent studies regarding N. brevicollis have questioned whether variation in personality traits and associative learning ability are related. The grain beetle Tenebrio molitor and the carabid beetle Pterostichus melanarius have been shown to possess learning ability. Due to the wide range of habitat and diet of N. brevicollis, it has been proposed that learning ability may be favored by natural selection within the species. Researchers demonstrated that N. brevicollis exhibit associative learning ability and demonstrated exploratory behavior as a personality trait of these ground-dwellers.[5]

Varying selection pressures within different habitats has been hypothesized as an explanation for the relationship between personality and cognition. Compared to beetles collected from less urbanized areas, those in urban areas show increased exploratory behavior. In laboratory settings, N. brevicollis females exhibited a large variation in associative learning ability. It is likely that associative learning abilities are specific to one beetle's personality rather than across species population.[5]

As an emerging field of interest, researchers plan to conduct further studies in which a wider range of learning assays are tested to understand the cognition-personality relationships of N. brevicollis.

Life history

Bimodal distribution representing the breeding patterns or Nebria brevicollis

Breeding is influenced by the quantity and quality of the food available. Egg production is influenced by the size of the beetle and diminishing food quantity prolongs the development of the larvae.[4] To rear the fittest offspring, it will avoid conditions of low food supply by following a bimodal breeding curve. It will engage in inactive periods, lasting five to six weeks, prior to a female laying her eggs. The breeding life of females is 27 days. Most eggs take 17 days to hatch.[7]

Breeding and diapause

The ground beetle partakes in sporadic breeding. Adults are active in the early summer, prior to a period of diapause.[7] When active, they build up their food reserves by feeding rapidly. Diapause can be characterized as a period of suspended development. Triggered by environmental cues, metabolic changes occur and the onset of diapause is brought on.[12] There is no locomotion activity and the animals do not feed.[7] In N. brevicollis, the onset of diapause is regulated by the fat content of the beetle's body. The period of diapause lasts 5 to 6 weeks during the months of July and August. Energy for body maintenance during diapause is supplied by the large food reserves accumulated early in the summer. Groups of up to 80 beetles will aggregate together under stones or logs during a period of diapause.[7]

Research regarding the aggregation behavior of Harmonia axyridis, belonging to the same order as N. brevicollis, has demonstrated this social behavior to be advantageous in group living conditions. It can allow for interactivity between individuals, which can allow for further information transfer. Costs, such as increased competition for food, space, and reproduction, may be a consequence of aggregative behavior; however, the advantages associated with group living outweigh the costs for H. axyridis and likewise other beetles.[13]

Following diapause, the breeding season begins. Various theories have been brought forth concerning the diapause in N. brevicollis. It may allow beetles to avoid conditions of low food supply during the summer.[7] Conditions of low food supply can negatively influence both the number of eggs produced and the viability of the eggs.

Low feeding levels produce small beetles, and thus unfavorable conditions during larval development.[14] Secondly, it is hypothesized that diapause allows for the development of the gonads. Fat reserves are depleted for gonad maturation.[7]

Predation

The abundance of beetles in Oregon may have deleterious effects on other carabid species. It is possible that the increase in prevalence of Nebria brevicollis is causing population declines within other carabid species. Increased competition for food between species may emerge, especially between species with similar diets and breeding practices. However, more research regarding the interactions between Nebria brevicollis and other carabid taxa must be conducted before concluding that negative impacts have occurred.[4]

Predation of Nebria brevicollis on non-carabid species may also have deleterious effects. The Fender's blue butterfly is considered to be endangered. During diapause, the butterflies are at increased risk of predation by Nebria brevicollis. Both species share similar habitats, as the occurrence of N. brevicollis has been found at sites where the butterflies exist.[4]

N. brevicollis has also been documented to kill, consume, and scavenge the slug Deroceras reticulatum.[15] Also known as the gray field slug, it is found in the Pacific Northwest and worldwide.[16] N. brevicollis killed more injured individuals than healthy individuals. However, there was a stronger preference in scavenging for dead slugs than those that were alive. Slugs produce mucus as a defense mechanism at the onset of a beetle attack. However, deceased slugs no longer produce mucus. They are therefore a more suitable prey, as they require a reduced handling time.[15] A reduced handling time decreases the energy a beetle must invest to kill a prey.

Nebria brevicollis at various stages of infection by Erynia nebriae

In north-western Germany, it was observed by researchers that an entomopathogenic fungus, Erynia nebriae was preying upon N. brevicollis [12]. Although Nebria brevicollis is widely considered to be solely carnivorous, multiple small studies made by enthusiasts have proven that many Nebria brevicollis will resort to eating various types of fungi that can be found in the soil they live on/around.[17]

The fungus infects the beetle, which results in a swollen abdomen. Observation concluded that low temperatures neither killed nor impeded the epizootic of the fungus. While more research regarding the relationship between fungus and beetle must be conducted, it is possible that the fungus manipulates the behavior of the beetle to their advantage. However, the strong exoskeleton of most carabid species may serve as a defense mechanism against fungi predation.[12]

Interaction with humans and conservation

Composition of the soil on which Nebria brevicollis walks

The use of ground beetles (Coleoptera, Carabidae) to study negative consequences caused by human activities has been shown to be an useful bioindicator. Habitat alterations due to urbanization, crop and forest management, overgrazing by domestic livestock, and soil pollution can be studied using ground beetles.[18] Possible effects environmental pollution may have on the habitat of the beetles includes changing the pH value of the soil, as well as altering its sodium and calcium content. Physiologically, pollution has been observed to cause physiological and behavioral changes.[18] Research has confirmed the property of the soil can shape carabid communities, including Nebria brevicollis.[19] Changes in species number or abundance can be correlated with habitat alterations caused by humans. Changes in the reproductive ability of ground-dwelling beetles may also serve to highlight the negative consequences of environmental pollution.

While carabids are useful bioindicators, more research must be conducted.[18]

References

  1. ^ a b Kavanaugh, David H.; et al. (16 June 2021). "Phylogeny of the supertribe Nebriitae (Coleoptera, Carabidae) based on analyses of DNA sequence data". ZooKeys (1044): 41–152. Bibcode:2021ZooK.1044...41K. doi:10.3897/zookeys.1044.62245. PMC 8222211. PMID 34183875.
  2. ^ a b c Kavanaugh, David H.; LaBonte, James R. "Discovery of Nebria brevicollis (Fabricius) (Coleoptera: Carabidae: Nebriini), a European Ground Beetle, Established in the Willamette Valley, Oregon" (PDF). Proceedings of the California Academy of Sciences. 59 (9): 481–488. Retrieved 19 November 2023.
  3. ^ a b c McGregor, Robert R.; Goulet, Henri; LaBonte, James R. (18 December 2020). "First western Canadian records of Nebria brevicollis (Coleoptera: Carabidae) and establishment of populations in Coquitlam, British Columbia, Canada". The Canadian Entomologist. 153 (2): 237–243. doi:10.4039/tce.2020.72. ISSN 0008-347X. S2CID 232233993.
  4. ^ a b c d e f LaBonte, James R. (16 November 2011). "Nebria brevicollis (Fabricius, 1792) in North America, benign or malign? (Coleoptera, Carabidae, Nebriini)". ZooKeys (147): 497–543. Bibcode:2011ZooK..147..527L. doi:10.3897/zookeys.147.2119. PMC 3286253. PMID 22371674.
  5. ^ a b c d e Harris, Ciaran; Liedtke, Jannis; Drees, Claudia; Schuett, Wiebke (1 November 2020). "Exploratory behaviour is not related to associative learning ability in the carabid beetle Nebria brevicollis". Behavioural Processes. 180: 104224. doi:10.1016/j.beproc.2020.104224. ISSN 0376-6357. PMID 32828809. S2CID 221198469.
  6. ^ a b c d e f Nelemans, M. N. E. (1 July 1987). "Possibilities for flight in the carabid beetle Nebria brevicollis (F.)". Oecologia. 72 (4): 502–509. Bibcode:1987Oecol..72..502N. doi:10.1007/BF00378974. ISSN 1432-1939. PMID 28312510. S2CID 20103414.
  7. ^ a b c d e f g h Penney, Margaret M. (1966). "Studies on Certain Aspects of the Ecology of Nebria brevicollis (F.) (Coleoptera, Carabidae)". Journal of Animal Ecology. 35 (3): 505–523. Bibcode:1966JAnEc..35..505P. doi:10.2307/2488. JSTOR 2488 – via JSTOR.
  8. ^ Harris, Ciaran; Liedtke, Jannis; Drees, Claudia; Schuett, Wiebke (1 November 2020). "Exploratory behaviour is not related to associative learning ability in the carabid beetle Nebria brevicollis". Behavioural Processes. 180: 104224. doi:10.1016/j.beproc.2020.104224. ISSN 0376-6357. PMID 32828809. S2CID 221198469.
  9. ^ "Nebria (Nebria) brevicollis (Fabricius, 1792)". 2.6.2. Fauna Europaea. 29 August 2013. Archived from the original on 13 November 2013. Retrieved 13 November 2013.
  10. ^ a b LaBonte, James (16 November 2011). "Nebria brevicollis (Fabricius, 1792) in North America, benign or malign? (Coleoptera, Carabidae, Nebriini)". ZooKeys (147): 527–543. Bibcode:2011ZooK..147..527L. doi:10.3897/zookeys.147.2119. ISSN 1313-2970. PMC 3286253. PMID 22371674.
  11. ^ a b Sih, Andrew; Del Giudice, Marco (5 October 2012). "Linking behavioural syndromes and cognition: a behavioural ecology perspective". Philosophical Transactions of the Royal Society B: Biological Sciences. 367 (1603): 2762–2772. doi:10.1098/rstb.2012.0216. ISSN 0962-8436. PMC 3427552. PMID 22927575.
  12. ^ a b c Keller, Siegfried; Hülsewig, Thorben (5 January 2018). "Amended description and new combination for Entomophthora nebriae Raunkiaer, (1893), a little known entomopathogenic fungus attacking the ground beetle Nebria brevicollis (Fabricius, 1792)". Alpine Entomology. 2: 1–5. doi:10.3897/alpento.2.22136. ISSN 2535-0889.
  13. ^ Durieux, Delphine; Fassotte, Bérénice; Deneubourg, Jean-Louis; Brostaux, Yves; Vandereycken, Axel; Joie, Emilie; Haubruge, Eric; Verheggen, François J (2015). "Aggregation behavior of Harmonia axyridis under non-wintering conditions". Insect Direct. 22 (5): 670–678. Bibcode:2015InsSc..22..670D. doi:10.1111/1744-7917.12144. PMID 24889907. S2CID 1954451 – via PubMed.
  14. ^ Nelemans, M.N.E. (1986). "On the Life-History of the Carabid Beetle Nebria Brevicollis (F.)". Netherlands Journal of Zoology. 37 (1): 26–42. doi:10.1163/002829687X00026.
  15. ^ a b Mair, J.; Port, G. R. (May 2001). "Predation by the carabid beetles Pterostichus madidus and Nebria brevicollis is affected by size and condition of the prey slug Deroceras reticulatum". Agricultural and Forest Entomology. 3 (2): 99–106. doi:10.1046/j.1461-9563.2001.00093.x. ISSN 1461-9555. S2CID 84793192.
  16. ^ "Biology and Life Cycle of the Gray Field Slug". agsci.oregonstate.edu. 18 January 2016. Retrieved 26 February 2024.
  17. ^ "Nebria brevicollis (Fabricius, 1792)". www.gbif.org (in Arabic). Retrieved 26 February 2024.
  18. ^ a b c Avgin, Sakine Serap; Luff, Martin Leslie (2010). "Ground beetles (Coleoptera: Carabidae) as bioindicators of human impact". Munis Entomology & Zoology. 5 (1): 209–215.
  19. ^ Vician, Vladimír; Svitok, Marek; Michalková, Eva; Lukáčik, Ivan; Stašiov, Slavomír (1 August 2018). "Influence of tree species and soil properties on ground beetle (Coleoptera: Carabidae) communities". Acta Oecologica. 91: 120–126. Bibcode:2018AcO....91..120V. doi:10.1016/j.actao.2018.07.005. ISSN 1146-609X. S2CID 92140437.
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