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Ludwigia Grandiflora (Michx.) Greuter & Burdet Subsp. Hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, an Invasive A

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Ludwigia Grandiflora (Michx.) Greuter & Burdet Subsp. Hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, an Invasive A ISSN 1346-7565 Acta Phytotax. Geobot. 71 (1): 65–71 (2020) doi: 10.18942/apg.201911 Short CommuniCation Ludwigia grandiflora (Michx.) Greuter & Burdet subsp. hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, an Invasive Aquatic Plant in Lake Biwa, the Largest Lake in Japan 1,* 2 3 1 Shinya hieda , yuko kaneko , maSato nakagawa and naohiko noma 1Graduate School of Environmental Science, the University of Shiga Prefecture, 2500 Hassaka-cho, Hikone, Shiga 522-8533, Japan. *[email protected] (author for correspondence); 2Natural Science Laboratory, Department of English and American Literature, Faculty of Letters, Toyo University, 5-28-20 Hakusan, Bunkyo-ku, Tokyo 112- 8606, Japan; 3Research Institute for Biological Sciences, Okayama Prefectural Technology Center for Agriculture, Forestry and Fisheries, 7549-1 Yoshikawa, Kibichuo, Okayama 716-1241, Japan Ludwigia grandiflora (Michx.) Greuter & Burdet is an invasive aquatic plant native to an area ranging from the southeastern United States to South America. In previous reports, L. grandiflorain Lake Biwa has been identified as L. grandiflora subsp. grandiflora, a hexaploid subspecies (2n = 48) or L. grandi- flora sensu lato. Species of Ludwigia sect. Jussiaea, are morphologically very similar. Morphological characters, especially floral characters and cytology (chromosome number) have been used to identify species and subspecies. In our study, the morphological characteristics of L. grandiflora in Lake Biwa substantially corresponded to those of L. grandiflora subsp. hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, a decaploid (2n = 80) subspecies. The chromosome number of L. grandiflorain Lake Biwa was 2n = ca. 80, the same as in L. grandiflorasubsp. hexapetala (2n = 80). Importantly, L. grandiflora in Lake Biwa is not L. grandiflora subsp. grandiflora, but rather L. grandiflora subsp. hexapetala. This re-iden- tification has implications for management in Lake Biwa, since the life history of L. grandiflora subsp. hexapetala has been studied in Europe. Key words: invasive aquatic plant, Lake Biwa, Ludwigia grandiflora subsp. hexapetala Worldwide, there are eight known species in The taxonomic status of the two subspecies the genus Ludwigia L., section Jussiaea (L.) has varied several times, depending on the re- Hoch, W. L. Wagner & P. H. Raven (Nesom & searcher. Based on differences in chromosome Kartesz 2000, Wagner et al. 2007, Armitage et al. number and morphological characteristics (e.g. 2013, Hoch et al. 2015). Ludwigia grandiflora flower size and plant pubescence), Ludwigia uru- (Michx.) Greuter & Burdet, sect. Jussiaea, is native guayensis (Cambess.) H. Hara, was separated from the southeastern United States to South into two species: L. grandiflora and L. hexa- America (Munz 1942, Raven 1963, Hoch et al. petala (Hook. & Arn.) Zardini, H. Y. Gu & P. H. 2015). It includes two subspecies, L. grandiflora Raven (Raven & Tai 1979, Greuter & Raus 1987, subsp. grandiflora, a hexaploid (2n = 48) or, rarely, Zardini et al. 1991a). Nesom & Kartesz (2000) dodecaploid (2n = 96), and L. grandiflora subsp. treated these as subspecies, L. grandiflora subsp. hexapetala (Hook. & Arn.) G. L. Nesom & Kartesz, grandifloraand L. grandiflora subsp. hexapetala, a decaploid (2n = 80) (Raven & Tai 1979, Zardini of a single species because they differ only in et al. 1991a, 1991b, Nesom & Kartesz 2000). An quantitative, intergrading morphological charac- octoploid (2n = 64) hybrid between the two sub- ters. Ward (2012) treated them as two varieties: L. species has been reported (Zardini et al. 1991b). grandiflora var. grandiflora and L. grandiflora 66 Acta Phytotax. Geobot. Vol. 71 var. hexapetala (Hook. & Arn.) D. B. Ward. Environment, Government of Japan 2018). Moreover, in a study of the invasive population in In previous reports, Ludwigia grandiflora in Britain, Armitage et al. (2013) recommended that Lake Biwa was described as L. grandiflorasubsp. the taxa be treated as subspecies because they grandiflora (Kadono 2014) or L. grandiflora sen- differ in chromosome number, but lack clear su lato (Uyemura et al. 2015). Ludwigia grandi- morphological differences. flora is a plastic species, but from our prelimi- Considering the above, we treat the taxa as nary morphological observations we concluded two subspecies within a single species. Species of that L. grandiflora in Lake Biwa was not L. gran- Ludwigia are morphologically very similar diflora subsp. grandiflora. Misidentification of (Ghahramanzadeh et al. 2013). Their identifica- alien species can have serious consequences tion is difficult (Dandelot et al. 2005b), and iden- (Pyšek & Richardson 2010). As the basis for the tification is especially difficult in the absence of management of L. grandiflora in Lake Biwa, its flowers (Ghahramanzadeh et al. 2013). For the correct identification is needed. recognition of subspecies of L. grandiflora, floral In this study, we focused on several morpho- characteristics and the pubescence of the stems logical characters to determine the identity of and leaves have been regarded as significant (Zar- Ludwigia grandiflora in Lake Biwa. To support dini et al. 1991a, Nesom & Kartesz 2000). Addi- our morphological observations, we also deter- tionally, the subspecies differ in chromosome mined the chromosome number. On the basis of number, which was used to support their distinc- the findings, we discuss L. grandiflora in Lake tiveness (Suyama et al. 2008, Kadono & Okamo- Biwa and its infraspecific identity. to 2018). Ludwigia grandiflora has been used as an or- namental (Ruaux et al. 2009), and naturalized Materials and Methods populations have been recorded in the USA (Oka- da et al. 2009) and in Europe (Dandelot et al. Measurement and observation of morphological 2005a, Nehring & Kolthoff 2011, Stiers et al. characters 2014). The invasiveness of L. grandiflorahas eco- Ludwigia grandiflora was first recorded in logical impacts on native aquatic vegetation and Akanoi Bay in southern Lake Biwa in 2009 detritus communities (Stiers et al. 2011). In Ja- (Council to Manage Aquatic Invasive Alien pan, L. grandiflora subsp. grandiflora has been Plants in Lake Biwa 2014). We measured six flo- recorded as being in Wakayama (Naito & Hieda ral characters: length of the sepal, petal, style, 2014) and Hyogo prefectures (Suyama et al. short filament, long filament and ovary (Fig. 2) 2008), and in Lake Biwa in Shiga prefecture (Ka- and evaluated the pubescence of the stems in dono 2014). Ludwigia grandiflora subsp. hexa- plants from the population of L. grandiflora es- petala has been recorded as being in Kagoshima tablished on the eastern coast of the Karasuma Prefecture (Kadono 2014, Kadono & Okamoto Peninsula near the locality where it was first re- 2018). Because L. grandiflora spreads rapidly, its corded. All measurements and observations were dense floating stems in some parts of Lake Biwa made using living materials from 20 emergent cover the water surface (Fig. 1) and cause many stems. The measurements and observations were problems, such as choking of the water channels, then compared with description in the articles us- blocking of sunlight, and interfering with boat ing living materials (Zardini et al. 1991a) and her- navigation and fishing. As a result of these condi- barium specimens (Nesom & Kartesz 2000) and tions, the local government in Shiga Prefecture type specimens examined in K and digital imag- began management activities in 2013. Ludwigia es on the websites E, K and P (Appendix 1). We grandiflora was subsequently included in the List used ImageJ v.1.52a (Schneider et al. 2012), to of Regulated Living Organisms in 2014 under Ja- measure the characteristics of flowers in the digi- pan’s Invasive Alien Species Act (Ministry of the tal images. Characters partially covered by other February 2020 hieda & al. ― Ludwigia grandiflora Subspecies in Lake Biwa 67 Fig. 1. Ludwigia grandiflora in Lake Biwa. A: Dense mats of flowering plants (Shiga Pref., Karasuma Penin- sula, Lake Biwa, June 8, 2014). B: Monospecific stand of flowering plants (Shiga Pref., Karasuma Penin- sula, Lake Biwa, June 1, 2015). Pho- tographs by S. Hieda. Fig. 2. Diagram showing the six floral characters measured in Ludwigia grandiflora. A, Sepal length; B, petal length; C, style length; D, short fila- ment length; E, long filament length; F, ovary length. Fig. 3. Somatic chromosomes (2n = ca. 80) of Ludwigia grandiflora in Lake Biwa. A, A’: sample from Karasuma Peninsula, B, B’: sample from Akanoi Bay. A, B: photographs. A’, B’: drawings of respective preceding photographs. Arrows: satel- lite chromosomes. 68 Acta Phytotax. Geobot. Vol. 71 plant parts were inferred from visible ones. characters (petal length and style length) were within the range of values for L. grandiflora sub- Chromosome count sp. hexapetala. Three other floral characters (se- To determine chromosome numbers, we col- pal length, short filament length and ovary length) lected five stems from the population on the were within the range of values for both subspe- northern coast of Karasuma Peninsula and in cies; long filament length was not within the Akanoi Bay. Since we obtained no accurate chro- range of value for either subspecies. The pubes- mosome count for the population studied mor- cence of the stems corresponded most closely phologically (eastern coast of Karasuma Penin- with the state reported for of L. grandiflora sub- sula), we observed the somatic chromosomes of sp. hexapetala. the population (northern coast of Karasuma Pen- insula and Akanoi Bay)
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