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Komarekiona

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Komarekiona
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Annelida
Clade: Pleistoannelida
Clade: Sedentaria
Class: Clitellata
Order: Opisthopora
Suborder: Crassiclitellata
Family: Komarekionidae
G. E. Gates, 1974
Genus: Komarekiona
G. E. Gates, 1974
Species:
K. eatoni
Binomial name
Komarekiona eatoni
G. E. Gates, 1974

Komarekiona eatoni, or the Kentucky earthworm, is a species of nearctic Annelid found in the southwestern United States, especially near the Appalachian Mountains.[2][3][4][5][6]. Komarekionidae is one of the few endemic Megadrile families in North America north of Mexico.[7]

It is the only species in the family Komarekionidae[8] and the genus Komarekiona, though some sources have combined it into the family Hormogastridae.[9] Other sources place it with Ailoscolecidae.[10][11] However, later sources have pointed out critical differences between Ailoscolecidae and Komarekionidae.[12]

The genus and family (Komarekion-) are named in honor of a Director of Tall Timbers Research Station and Land Conservancy.[13]

Range

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Due to the Last Glacial Maximum during the Pleistocene, there are very few endemic species of earthworm in North America. The range of the species is thus further south, as are most earthworm species on the continent.[14]

A map of the glaciated regions of North America during the Last Glacial Maximum
K. eatoni is the only species of its family to survive the Last Glacial Maximum

The Kentucky earthworm has been found in several states in the southeast of the United States. It ranges through Kentucky,[15][16] Illinois,[17][18] Georgia,[19] Tennessee,[20] Virginia,[21] Indiana,[22] and North Carolina.[23] So far it has only been found in the unglaciated portions of Indiana.[22]

A map of southeastern US states with counties highlighted in Illinois, Indiana, Kentucky, Virginia, Tennessee, North Carolina, and Georgia
A map of the counties where K. eatoni has been documented

Its physiographic regions are the Appalachian Highlands, Southern Piedmont, and Ridge-and-Valley Appalachians.[6]

Phylogeny

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11,000 years ago, during the Late Wisconsin stage of the Pleistocene glaciation, most species of earthworms perished.[14] Komarekiona eatoni is believed to be the only species of its family to have survived.[24]

In phylogenetic analyses, Komarekiona forms a clade with Sparganophilus.[25][26] The two families diverged an estimated 69.57-110 Mya.[27][28]

Morphology

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K. eatoni worms are less than 110 millimetres (4.3 in) long and have a diameter of 5–6 millimetres (0.20–0.24 in). Specimens have between 86-163 segments.[29]

Specimens east of the Great Smoky Mountains are known to reproduce sexually, as a "brilliant spermatozoal iridescence on the male funnels and in the spermathecal coagulum" has been noted. However, those west of the mountains lack spermathecae and testes and have rudimentary seminal vesicles. This indicates that the western organisms reproduce asexually. The author noting this discovery labeled this mutagenesis with a different definition than the accepted definition of the term, citing the Kentucky earthworm as a prime example: "A permanent change in the method of reproduction by an organism, as when a portion of the range is isolated from the main gene pool or centre of origin; ex. nearctic earthworm Komarekiona eatoni."[29]

Biology

[edit]

The Kentucky earthworm is endogeic, meaning it lives in the soil and makes horizontal burrows.[4][30] Other sources note the species as epiendogeic worm.[26] The species is easily found and dug up with a small hand pick "from the surface centimetres of the soil profile."[29] Others have noted its presence in the mineral soil.[31] It has frequently been found in formalin extraction sampling efforts.[29]

The species prefers less acidic environments.[32] It is associated with soil high in organic matter that are mesic habitats.[29][26][6] Its preferred soil great groups are Dystrochrepts and Hapludults.[6] The worms are commonly found among "leaf litter, under debris, logs, and rocks etc."—areas that notably hold onto precipitation longer[29] It is also found along roadsides, in sinkholes, and near sewage installations.[33]

The Kentucky earthworm seems to breed during the winter.[29][34] This likely has to do with the increase of precipitation in its regions during winter months.[29] They appear to go into aestivation during the drier parts of the year, unlike many other earthworm species in their range.[29]

Its ovaries occur collectively.[35] K. eatoni specimens found east of the Great Smoky Mountains are amphimictic and those west of the mountains are parthenogenetic.[36][2]

It has at least 143,281 contigs and one extracellular CYGB globin gene.[37][38]

Associated species

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It has been associated with Tsuga canadensis, Rhododendron maximum, and Liriodendron tulipifera. However, R. maximum has been found to create drier soils and phenolic repellants and toxins that may keep worms like K. eatoni away.[31] It tends to be found in forest regions of Loblolly-Shortleaf pines, Oak-Hickory, and Oak-Pine trees.[6]

It has also been associated with vermivorous snakes, as there is overlap in the ranges of the worm and Carphophis amoenus, Diadophis punctatus, and Thamnophis sirtalis.[39]

Conservation

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The Kentucky earthworm is susceptible to disturbance and competition with non-native species.[40] It survives in minimally disturbed second growth forests.[16] In fact, it has been found only to occur in minimally disturbed areas.[41]

In North Carolina, the species was considered endangered.[42] In 2013, the species was found to be vulnerable.[43] This is the last time it was assessed for a conservation status by the International Union for Conservation of Nature.[44]

References

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  1. ^ Blakemore, R. (2014). "Komarekiona eatoni". IUCN Red List of Threatened Species. 2014: e.T11049A21416036. doi:10.2305/IUCN.UK.2014-1.RLTS.T11049A21416036.en. Retrieved 2 January 2025.
  2. ^ a b Reynolds, John Warren (2020). Earthworms in American Ecoregions. LAP LAMBERT Academic Publishing. p. 433. ISBN 9786202808934.
  3. ^ Reynolds, John Warren (May 2017). "Earthworms (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Megascolecidae, Ocnerodrilidae and Sparganophilidae) in the Ridge and Valley Ecoregion (67), USA". Megadrilogica. 22 (5): 111–123.
  4. ^ a b Reynolds, John Warren (January 2018). "First Earthworm (Annelida: Oligochaeta) Species' Collections in Canada and the Continental United States". Megadrilogica. 23 (1): 1–50.
  5. ^ Reynolds, John Warren; Wetzel, Mark Julian (March 2004). "Terrestrial Oligochaeta (Annelida:Clitellata) in North America North of Mexico". Megadrilogica. 9 (11): 82.
  6. ^ a b c d e Reynolds, John Warren (April 2011). "The Earthworms (Oligochaeta: Acanthodrilidae, Eudrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Lutodrilidae, Megascolecidae, Ocnerodrilidae, Octochaetidae and Sparganophilidae) of Southeastern United States". Megadrilogica. 14 (9–12): 238, 301.
  7. ^ Edwards, Clive A. (2022). Biology and Ecology of Earthworms. Norman Q. Arancon (4th ed.). New York, NY: Springer. p. 68. ISBN 978-0-387-74943-3.
  8. ^ Edwards, C. A., ed. (2004). Earthworm ecology (2nd ed.). Boca Raton, Fla: CRC Press. p. 68. ISBN 978-0-8493-1819-1.
  9. ^ Glasby, Christopher; Biriukova, Olga; Martin, Patrick; Dyne, Geoffrey; Utevsky, Serge; Wilson, Robin (2024-09-27), "Annelida – diagnoses, descriptions and keys to family-level taxa", ARPHA Preprints, vol. 5, pp. e137961, doi:10.3897/arphapreprints.e137961, retrieved 2025-01-01
  10. ^ Sims, R. W. (1980). "A classification and the distribution of earthworms, suborder Lumbricina (Haplotaxida: Oligochaeta)". Bulletin of the British Museum (Natural History). Zoology. 39 (2): 103–124.
  11. ^ Jiangping Qiu; Bouché, Marcel (1998), Révision des taxons supraspécifiques de Lumbricoidea [Revision of Lumbricoidea supraspecific taxa] (in French), vol. 3, doi:10.13140/RG.2.2.24269.49127, retrieved 2025-01-05
  12. ^ Rouse, Greg; Pleijel, Fredrik (2006). Reproductive biology and phylogeny of Annelida. Enfield: Science Publ. pp. 322–324. ISBN 978-1-57808-313-8.
  13. ^ Reynolds, John Warren (2017). "Earthworms (Oligochaeta: Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Megascolecidae, Ocnerodrilidae and Sparganophilidae) in the Piedmont Ecoregion (45), USA". Megadrilogica. 22 (4).
  14. ^ a b Reynolds, John Warren (2024), Kooch, Yahya; Kuzyakov, Yakov (eds.), "Earthworms in Canadian Forest Regions Revisited", Earthworms and Ecological Processes, Cham: Springer Nature Switzerland, pp. 33–93, doi:10.1007/978-3-031-64510-5_2, ISBN 978-3-031-64510-5, retrieved 2025-01-06
  15. ^ Reynolds, John Warren (June 2008). "The Earthworms (Oligochaeta: Acanthodrilidae, Komarekionidae, Megascolecidae, and Sparganophilidae of Kentucky, USA". Megadrilogica. 12 (5): 83.
  16. ^ a b Kalisz, P. J.; Powell, J. E. (October 2000). "Invertebrate Macrofauna in Soils Under Old Growth and Minimally Disturbed Second Growth Forests of the Appalachian Mountains of Kentucky". The American Midland Naturalist. 144 (2): 297–307. doi:10.1674/0003-0031(2000)144[0297:IMISUO]2.0.CO;2. ISSN 0003-0031.
  17. ^ Wetzel, Mark Julian; Reynolds, John Warren (December 2021). "A Preliminary Inventory of Earthworms (Annelida, Clitellata) of the Nachusa Grasslands Area, Lee and Ogle Counties, Illinois, ILLINOIS, USA". Megadrilogica. 26 (8): 91–125.
  18. ^ Webb, Donald W.; Taylor, Steven J.; Krejca, Jean K. (1993). "The biological resources of Illinois caves and other subterranean environments". Illinois Natural History Survey Center for Biodiversity: 31.
  19. ^ Reynolds, John Warren (December 2015). "A Checklist by Counties of Earthworms Oligochaeta: Acanthodrilidae, Udrilidae, Komarekionidae, Lumbricidae, Maegascolecidae, Ocnerodrilidae, and Paragnophilidae) in Georgia, USA". Megadrilogica. 19 (6): 85–114.
  20. ^ Reynolds, John Warren (2016). "A Cheklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Megascolecidae and Sparganophilidae) in Tennessee, USA". Megadrilogica. 19 (11).
  21. ^ Reynolds, John Warren (1994). "Earthworms of Virginia (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Megascolecidae, and Sparganophilidae)". Megadrilogica. 5 (8).
  22. ^ a b Reynolds, John Warren (January 1994). "The distribution of the earthworms (Oligochaeta) of Indiana: A case for the post quarterly introduction theory for megadrile migration in North America". Megadrilogica. 5 (3): 30.
  23. ^ Reynolds, John Warren (May 1994). "Earthworms of North Carolina (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Megascolecidae, Ocnerodrilidae and Sparganophilidae)". Megadrilogica. 5 (6).
  24. ^ Gates, G. E. (1976). "On earthworm ovaries and their importance in Megadrile Systematics I.". Megadrilogica. 2 (12).
  25. ^ Jamieson, B.; Tillier, S.; Tillier, A.; Justine, J.; Ling, E. Y.; James, S.; Mcdonald, K.; Hugall, A. (2002-12-27). "Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28S) and mitochondrial (12S, 16S) rDNA". Zoosystema. S2CID 56327366.
  26. ^ a b c James, Samuel W.; Davidson, Seana K. (2012-08-06). "Molecular phylogeny of earthworms (Annelida : Crassiclitellata) based on 28S, 18S and 16S gene sequences". Invertebrate Systematics. 26 (2): 213–229. doi:10.1071/IS11012. ISSN 1447-2600.
  27. ^ Helsem, Stian Aleksander (June 2021). "Divergence time estimates for several phylogenies of Annelida (Lophotrochozoa)" (PDF). University of Oslo Department of Biosciences.
  28. ^ Erséus, Christer; Williams, Bronwyn W.; Horn, Kevin M.; Halanych, Kenneth M.; Santos, Scott R.; James, Samuel W.; Creuzé des Châtelliers, Michel; Anderson, Frank E. (September 2020). "Phylogenomic analyses reveal a Palaeozoic radiation and support a freshwater origin for clitellate annelids". Zoologica Scripta. 49 (5): 614–640. doi:10.1111/zsc.12426. ISSN 0300-3256.
  29. ^ a b c d e f g h i Reynolds, John Warren (1977). "The earthworms of Tennessee (Oligochaeta). III Komarekionidae with notes on distribution and biology". Megadrilogica. 3 (4).
  30. ^ "Earthworm Ecology". www.earthwormsoc.org.uk. Archived from the original on 20 December 2024. Retrieved 2025-01-01.
  31. ^ a b Boettcher, S. E.; Kalisz, P. J. (1991). "Single-Tree Influence on Earthworms in Forest Soils in Eastern Kentucky". Soil Science Society of America Journal. 55 (3): 862–865. Bibcode:1991SSASJ..55..862B. doi:10.2136/sssaj1991.03615995005500030040x. ISSN 1435-0661.
  32. ^ Boerner, R. E J. (April 2006). "Unraveling the Gordian Knot: Interactions among vegetation, topography, and soil properties in the central and southern Appalachians". The Journal of the Torrey Botanical Society. 133 (2): 321–361. doi:10.3159/1095-5674(2006)133[321:UTGKIA]2.0.CO;2. ISSN 1095-5674.
  33. ^ Parker, Sybil P., ed. (1982). Synopsis and classification of living organisms. Vol. 2. New York: McGraw-Hill. p. 54. ISBN 978-0-07-079031-5.
  34. ^ Belato Costa, Flavia Ariany (2019). "Diversidade e evolução de hemoglobinas extracelulares em Metazoa" [Diversity and evolution of extracellular hemoglobins in Metazoa.]. sucupira-legado.capes.gov.br (in Portuguese). Archived from the original on 1 January 2025. Retrieved 2025-01-01.
  35. ^ Jamieson, B. G. M. (December 1988). "On the Phylogeny and Higher Classification of the Oligochaeta". Cladistics. 4 (4): 367–401. doi:10.1111/j.1096-0031.1988.tb00520.x. ISSN 0748-3007. PMID 34949062.
  36. ^ MisirlioğLu, Mete; Reynolds, John Warren; Stojanović, Mirjana; Trakić, Tanja; Sekulić, Jovana; James, Samuel W.; Csuzdi, Csaba; DecaëNs, Thibaud; Lapied, Emmanuel; Phillips, Helen R. P.; Cameron, Erin K.; Brown, George G. (2023-03-15). "Earthworms (Clitellata, Megadrili) of the world: an updated checklist of valid species and families, with notes on their distribution". Zootaxa. 5255 (1): 417–438. doi:10.11646/zootaxa.5255.1.33. ISSN 1175-5334.
  37. ^ Belato Costa, Flavia Ariany (2019). "Diversidade e evolução de hemoglobinas extracelulares em Metazoa" [Diversity and evolution of extracellular hemoglobins in Metazoa]. Universidade Federal do Rio de Janeiro (in Portuguese).
  38. ^ Belato, Flávia A; Coates, Christopher J; Halanych, Kenneth M; Weber, Roy E; Costa-Paiva, Elisa M (October 2020). Betran, Esther (ed.). "Evolutionary History of the Globin Gene Family in Annelids". Genome Biology and Evolution. 12 (10): 1719–1733. doi:10.1093/gbe/evaa134. ISSN 1759-6653. PMC 7549130. PMID 32597988.
  39. ^ Gray, Brian S. "Distribution of native and exotic earthworms in the eastern US: implications for the ecology of vermivorous snakes". Bulletin of the Chicago Herpetology Society. 45 (5): 73–86.
  40. ^ Mark Timothy Thogerson (1997). Life histories and population dynamics of three earthworm species (Oligochaeta:Lumbricidae) in a northern Michigan hardwood forest (Masters thesis). Michigan State University. doi:10.25335/yqks-cy60.
  41. ^ Kalisz, Paul J. (1993). McKnight, Bill N. (ed.). Native and Exotic Earthworms in Deciduous Forest Soils of Eastern North America. Indiana Academy of Sciences. pp. 93–100. ISBN 978-1-883362-00-3.
  42. ^ McGaugh, M. Houston; Genoways, Hugh H. (1976). "State laws as they pertain to scientific collecting permits". Museology. 2: 51.
  43. ^ "IUCN Red List: Table 7: Species changing IUCN Red List Status (2013-2014)" (PDF). 2014. Archived (PDF) from the original on 28 Feb 2024.
  44. ^ Blakemore, R. (2014). "Komarekiona eatoni". IUCN Red List. Archived from the original on 7 Jan 2025.