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Reynolds, J.W. 2016. (: Acanthodrilidae, Lumbricidae, and Megascolecidae) in the Arkansas Valley Ecoregion (37), USA. Megadrilogica 20(7): 119-124. [paper in...

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EARTHWORMS (OLIGOCHAETA: ACANTHODRILIDAE, LUMBRICIDAE, AND MEGASCOLECIDAE) IN THE ARKANSAS VALLEY ECOREGION (37), USA.

Cite as 2016 Megadrilogica 20(7): 119-124. (June)

John Warren Reynolds

Oligochaetology Laboratory, 18 Broadview Court, Kitchener, ON Canada N2A 2X8, and Research Associate, New Brunswick Museum, Saint John, NB Canada E2K 1E5 (e-mail: [email protected])

ABSTRACT The Arkansas Valley Ecoregion in the United States occurs in portions of two states (Arkansas and Oklahoma). There are 21 species found here, in eight genera and three families (Acanthodrilidae, Lumbricidae and Megascolecidae). Six of the species are native (endemic) to ; all others are European or Oriental (exotic) introductions. A description of the ecoregion and two colour photographs are also included.

Key-words: Arkansas Valley Ecoregion, Oligochaeta, Acanthodrilidae, Lumbricidae, Megascolecidae, earthworms, distribution

RÉSUMÉ

L'écorégion de la Vallée D'Arkansas aux États-Unis se retrouve dans les portions de deux états (Arkansas et Oklahoma). On y retrouve 21 espèces présentes ici dans huit genres et trois familles (Acanthodrilidae, Lumbricidae et Megascolecidae). Six de ces espèces sont indigènes (endémiques) en Amérique du Nord (endemiques), tous les autres sont des introductions (exotiques) européens ou orientaux espèces. Une description de l'écorégion et deux illustrations couleur sont aussi incluses.

Mots-clé: La Vallée D'Arkansas, écorégion, Oligochaeta, Acanthodrilidae, Lumbricidae, Megascolecidae, vers de terre, répartition

RESUMEN La Ecoregión del Valle de Arkansas en los Estados Unidos se extiende en porciones de dos estados (Arkansas y Oklahoma). Se encontraron 21 especies reunidas en ocho géneros y tres familias (Acanthodrilidae, Lumbricidae y Megascolecidae). Seis de las especies son nativas (endémica) en América del Norte, el resto son introducciones (exóticas) europeas, orientales o pantropicales. Se incluye una descripción de la ecorregión y dos fotografías en color.

Key-words: La Ecoregión del Valle de Arkansas, Oligochaeta, Acanthodrilidae, Lumbricidae, Megascolecidae, lombrices de tierra, distribución

ZUSAMMENFASSUNG

Das Arkansastal is eine Ökoregion die inTeilen von zwei US Staaten (Arkansas und Oklahoma) auftritt. Es gibt dort 21 Arten in acht Gattungen und drei Familien (Acanthodrilidae, Lumbricidae und Megascolecidae). Sechs der Arten stammen aus Nordamerika, alle anderen sind europäische, orientalische oder pantropische Einführungen. Eine Beschreibung der Ökoregion und zwei Farbfotografien sind ebenfalls im Artikel enthalten.

Schlüsselworte: Das Arkansastal, Ökoregion, Oligochaeta, Acanthodrilidae, Lumbricidae, Megascolecidae, Regenwürmer, geografische Verteilung. 120 Earthworms in the Arkansas Valley Ecoregion, USA

INTRODUCTION nova Gates, 1977 Native North American species The Arkansas Valley Ecoregion is a relatively Arkansas small area which covers portions of central Arkansas (Pope Co.) and eastern Oklahoma in the United States. A detailed Oklahoma description of the species' locations can be found in (Muskogee Co.) Reynolds (2008) and Reynolds and Damoff (2010). The location of the Arkansas Valley Ecoregion (37) is Diplocardia riparia Smith, 1895 shown in Figure 1 with two photographs in Native North American species APPENDIX, Fig. A1. Arkansas (Johnson, Yell Cos.) Oklahoma (Haskell, Muskogee, Sequoyah Cos.)

Diplocardia rugosa James, 1988 Native North American species Oklahoma (Haskell Co.)

Diplocardia singularis (Ude, 1893) Native North American species Arkansas (Faulkner, Logan, Perry, Yell Cos.) Oklahoma (Haskell, Sequoyah Cos.)

Diplocardia smithi Macnab & McKey-Fender, 1955 Native North American species Oklahoma (Haskell Co.) Fig. 1. The location of the Arkansas Valley Ecoregion (37), USA (modified from Omernik, 1987). [Red lines Diplocardia sylvicola Gates, 1977 are state boundaries.] Native North American species Oklahoma ECOREGION EARTHWORMS (Haskell Co.) The species found in this ecoregion - none collected. will be listed alphabetically under the family, genus, species and counties where the species is present. The GLOSSOSCOLECIDAE - none collected. region of origin is also presented for each species. KOMAREKIONIDAE - none collected. ACANTHODRILIDAE LUMBRICIDAE Diplocardia caroliniana Eisen, 1899 Aporrectodea trapezoides (Dugès, 1828) Native North American species European introduction Arkansas Arkansas (Perry Co.) (Conway, Crawford, Franklin, Johnson, Pope, Oklahoma Pulaski, Yell Cos.) (Sequoyah Co.) Oklahoma (Haskell, Latimer, Sequoyah Cos.) Diplocardia communis Garman, 1888 Native North American species Aporrectodea tuberculata (Eisen, 1874) Arkansas European introduction (Johnson Co.) Arkansas (Franklin Co.) MEGADRILOGICA 121

Aporrectodea turgida (Eisen ,1873) MEGASCOLECIDAE European introduction Amynthas diffringens (Baird, 1869) Arkansas Oriental introduction (Conway, Crawford, Franklin, Perry, Yell Arkansas Cos.) (Crawford, Franklin Cos.) Bimastos beddardi (Michaelsen, 1894) OCNERODRILIDAE - none collected. Native North American species - none collected. Arkansas (Crawford Co.) SPARGANOPHILIDAE. - none collected.

Bimastos longicintus (Smith & Gittins, 1915) DISCUSSION Native North American species There are 21 species reported in the Arkansas Arkansas Valley Ecoregion, in eight genera and three families (Pulaksi Co.) (Acanthodrilidae, Lumbricidae and Megascolecidae). Of the 21 species, 13 are considered native to North Bimastos parvus (Eisen, 1874) America (endemics), seven are European introduced Native North American species species (exotics) and one is an Oriental introduced Arkansas species. In this ecoregion six counties in Oklahoma (Conway Co.) and five counties in Arkansas have no earthworm records. Bimastos tumidis (Eisen, 1874) Native North American species In this relatively small ecoregion there are Oklahoma fewer species than larger regional ecoregions. The (Sequoyah Co.) higher elevation may in part contribute to the balance between native species (13) vis-à-vis introduced species Bimastos zeteki (Smith & Gittins, 1915) (8). In the adjacent ecoregions to the south and east the Native North American species native earthworm species make up more that 50% of Arkansas the species collected. There is a noticeable absence in (Crawford Co.) the species list of introduced European species found in abundance in the north, e.g. Allolobophora chlorotica, Dendrobaena octaedra (Savigny, 1826) Aporrectodea rosea and Lumbricus rubellus. Eisenia European introduction hortensis is found in this ecoregion, but only in one or Arkansas two counties in eight other states (Reynolds and (Crawford, Yell Cos.) Wetzel, 2012). Acknowledgements Eisenia hortensis (Michaelsen, 1890) European introduction I am grateful to Wilma M. Reynolds of the Arkansas Oligochaetology Laboratory, Dr. Frederick W. Kutz (Yell Co.) formerly of the U.S. Environmental Protection Agency, and Dr. Josef H. Görres, University of Vermont for Lumbricus terrestris Linnaeus, 1758 reviewing the manuscript, their comments and European introduction suggestions. The author also wishes to thank Dr. Arkansas Catalina C. de Mischis, Universidad Nacional de (Faulkner Co.) Córdoba, Dr. Jean-Marc Gagnon, Canadian Museum of Nature, and Dr. Josef H. Görres, University of Vermont Octolasion tyrtaeum (Savigny, 1826) for translations of the abstracts and key words. European introduction Arkansas LITERATURE CITED (Crawford, Franklin, Perry Cos.) Generic and species names in brackets that follow citations, below, reflect current nomenclature for those LUTODRILIDAE - none collected. taxa, and thus may be different than nomenclature assigned in the original descriptions (Reynolds and Wetzel, 2016). 122 Earthworms in the Arkansas Valley Ecoregion, USA

Baird, W. 1869. Description of a new species of Michaelsen, W. 1890. Die Lumbriciden Norddeutsch- earthworm (Megascolex diffringens) found in lands. Mitt. Mus. Hamburg 7:1-19. [Eisenia north Wales. Proc. Biol. Soc. London, 1869: hortensis n. sp.] 40-43. [Amynthas diffringens n. sp.] Michaelsen, W. 1894. Die Regenwürmen-Fauna von Dugès, A. 1828. Recherche sur la circulation, la Florida und Georgia. Zool. Jb. Syst. 8(2): respiration et la reproduction des Annélides 177-194. [Bimastos beddardi n. sp.] abranches sétigères. Ann. Sci. Nat. 15(1): 284-336. [Aporrectodea trapezoides n. sp.] Omernik, J.M. 1987. Ecoregions of the conterminous United States. Map (scale 1:7,500,000). Eisen, G. 1873. Om Skandinaviens Oligochaeter. Öfv. Annals Association of American Geographers Vet-Akad. Förh. Stockholm 30(8): 43-56. 77(1): 118-125. [Aporrectodea turgida n. sp.] Omernik, J.M. 1995. Ecoregions: A spatial frame- work for environmental management. Eisen, G. 1874. New Englands och Canadas Phaseolus Pp. 49-62. In: Biological Lumbricider. Öfv. Vet-Akad. Förh. Stock- Assessment and Criteria: Tools for Water holm 31(2): 41-49. [Aporrectodea tuber- Resource Planning and Decision Making. culata n. sp., Bimastos parvus n. sp. and Davis, W.S. and T.P. Simon (eds.) Boca Bimastos tumidus n. sp.] Raton, FL: Lewis Publishers. Eisen, G. 1899. Notes on North-American Omernik, J.M. 2004. Perspectives on the nature and earthworms of the genus Diplocardia. Zool. definition of ecological regions. Environ- Bull. 2(4): 161-172. [Diplocardia caroliniana mental Management 34 (Suppl. 1): S27-S38. n. sp.] Reynolds, J.W. 2008. The earthworms (Oligochaeta: Garman, H. 1888. On the anatomy and histology of a Acanthodrilidae, Lumbricidae, Megascolec- new earthworm (Diplocardia communis gen. idae and Sparganophilidae) of Arkansas, USA et sp. nov.). Bull. Illinois St. Lab. Hist. 3: 47- revisited. Megadrilogica 11(11): 115-130. 77. [Diplocardia communis n. gen. and n. sp.] Reynolds, J.W. and G.A. Damoff. 2010. The earth- Gates, G.E. 1977. More on the earthworm genus worms (Oligochaeta: Acanthodrilidae, Lum- Diplocardia. Megadrilogica 3(1): 1-48. bricidae, Megascolecidae and Spargano- [Diplocardia nova n. sp. and Diplocardia. philidae) of Oklahoma, USA. Megadrilogica sylvicola n. sp.] 13(12): 173-193. James, S.W. 1988. Diplocardia hulberti and D. rugosa, new earthworms (Annelida: Oligo- Reynolds, J.W. and M.J. Wetzel. 2012. Terrestrial chaeta: Megascolecidae) from Kansas. Proc. Oligochaeta (Annelida: ) in North Biol. Soc. Washington 101(2): 300-307. America, including Mexico, Puerto Rico, [Diplocardia. rugosa n. sp.] Hawaii, and Bermuda. III. Megadrilogica 15(8): 191-211. Kinberg, J.G. 1867. Annulata nova. Öfv. Vetensk. Akad. Förh. Stockholm 23: 97-103, 356-357. Reynolds, J.W. and M.J. Wetzel. 2016. Nomenclatura [Amynthas corticis n. sp. (= Amynthas Oligochaetologica – A catalogue of names, diffringens)] descriptions and type specimens. Editio Linnaeus, C. 1758. Systema naturae per regna tris Secunda. URL: naturae, secundum classes, ordines, genera, http://wwx.inhs.illinois.edu/people/ species cum characteribus, differentiis, mjwetzel/nomenoligo (accessed: 5 June 2016). synonymis, locis, Edition decima, reformata. Tom I. Laurentii Salvii, Holmiae, 824 pp. Savigny, J.C. 1826. Analyses des travaux de [Lumbricus terrestris n. gen. and n. sp.] l'Académie Royale des Sciences pendant l'année 1821, partie physique. Cuvier, M. le Macnab, J.A. and D. McKey-Fender. 1955. Studies in Baron G. (ed.) Mém. Acad. Sci. Inst. France the genus Diplocardia (Oligochaeta: Mega- 5: 176-184. [Dendrobaena octaedra n. sp., scolecidae). Wasmann J. Biol. 13(1): 113- Octolasion tyrtaeum n. sp.] 143. [Diplocardia smithi n. sp.] MEGADRILOGICA 123

Smith, F. 1895. A preliminary account of two new Ude, H. 1893. Beiträge zur Kenntnis ausländischer Oligochaeta from Illinois. Bull. Illinois St. Regenwürmer. Z. wiss. Zool. 57: 57-75. Lab. Nat. Hist. 4(5): 138-148. [Diplocardia [Diplocardia singularis n. sp.] riparia]

Smith, F. and E.M. Gittins. 1915. Two new species of Lumbricidae from Illinois. Bull. Illinois St. Lab. Nat. Hist. 10(7): 545-550. [Bimastos longicinctus n. sp. and Bimastos zeteki n. sp.

APPENDIX Arkansas Valley Ecoregion (modified from Omernik, 1987, 1995, 2004; Griffiths et al., 1997)

37. ARKANSAS VALLEY

Arkansas Valley Ecoregion (37) is a synclinal and alluvial valley lying between the Ozark Highlands Ecoregion (39) and the Ouachita Mountains Ecoregion (36). The Arkansas Valley Ecoregion (37) is, characteristically, diverse and transitional. It generally coincides with the Arkoma Basin, an oil and gas province, that developed as sand and mud were deposited in a depression north of the rising Ouachita Mountains during the Mississippian and Pennsylvanian eras. The Arkansas Valley Ecoregion (37) contains plains, hills, floodplains, terraces, and scattered mountains. It is largely underlain by interbedded Pennsylvanian sandstone, shale, and siltstone. Prior to the 19th Century, uplands were dominated by a mix of forest, woodland, savanna, and prairie whereas floodplains and lower terraces were covered by bottomland deciduous forest. Today, less rugged upland areas have been cleared for pastureland or hayland. Poultry and livestock farming are important land uses.

A region of mostly forested valleys and ridges, the physiography of the Arkansas Valley is much less irregular than that of the Boston Mountains to the north and the Ouachita Mountains to the south, but is more irregular than the ecological regions to the west and east. About one fourth of the region is grazed and roughly one tenth is cropland. In the Arkansas Valley, even streams that have been relatively unimpacted by human activities have considerably lower dissolved oxygen levels, and hence support different biological communities, than those of most of the adjacent regions.

The predominant vegetation form is about equal areas of cold-deciduous, broad-leaved forest and needle-leaved evergreen trees. Principal forest cover types are oak-hickory (Quercus-Carya) and loblolly-shortleaf pine (Pinus taeda, Pinus palustris). Species include white, black, bur, post, and blackjack oaks (Quercus alba, Q. velutina, Q. macrocarpa, Q. stellata and Q, marilandica); pignut and mockernut hickories (Carya glabra and Carya tomentosa); and loblolly and shortleaf pines. Oak-gum-cypress forest type is dominant along major river bottoms and includes cottonwood (Populus deltoides), sugarberry (Celtis laevigata), river birch (Betula nigra), and green ash (Fraxinus pennsylvanica).

Annual average precipitation is 1,120 to 1,270 mm. Average temperature is 16 to 17 o C. The growing season lasts 200 to 240 days.

Water quality is generally good and influenced more by land use activities than by soils or geology; average stream gradients and dissolved oxygen levels are lower in the Arkansas Valley (37) than in the Ouachita Mountains (36) or Ozark Highlands (39), whereas turbidity, total suspended solids, total organic carbon, total phosphorus, and biochemical oxygen demand values are typically higher. The Arkansas River is continuously turbid. Summer flow in smaller streams is typically limited or nonexistent. 124 Earthworms in the Arkansas Valley Ecoregion, USA

Fig. A1. The Arkansas Valley Ecoregion (37), USA (from Wikipedia).

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