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E Wilcox Juncus2.Pub Watsonia IDENTIFICATION28 : 43–56 (2010) OF JUNCUS × DIFFUSUS AND J. × KERN-REICHGELTII 43 A novel approach to the determination and identification of Juncus × diffusus Hoppe and J. × kern-reichgeltii Jansen & Wacht. ex Reichg. M. WILCOX* 32 Shawbridge St. Clitheroe, BB7 1LZ ABSTRACT specimens on one sheet, No. 2442 – N. Woodhead and two on another sheet No. 4972 The identity of J. inflexus L., J. effusus L. and J. – 2428 - G. Claridge Druce. These specimens conglomeratus L. have had a rather chequered past. have been reviewed in MANCH , with the G. The differences are now relatively well documented C. Druce specimens being attributable to J. and there is less confusion, though there are still effusus and the N. Woodhead specimens are J. misidentifications, particularly in the latter two species. The hybrids Juncus effusus × J. inflexus (J. conglomeratus (only one of which may vaguely × diffusus Hoppe) and J. effusus × J. conglomeratus be called, ‘var. subuliflorus ’) The name ‘var. (J. × kern-reichgeltii Jansen & Wacht. ex Reichg.) laxus ’ appears to be misapplied. However, very present further difficulties in their identification. The lax forms of J. conglomeratus have been found, former hybrid has been accepted for some time, but in which the pedicels are long, up to 8 cm with the latter has always been a difficult plant to identify flower clusters at the distal ends; these are with any certainty. Also, J. × kern-reichgeltii is said particularly found in woodland situations. This to be a fertile hybrid and therefore backcrossing form needs further investigation. Some of these (introgression) is likely with no certain recognition inflorescence types also occur in the hybrid or distinctions that include the potentially introgressed individuals. This article aims to present J. × kern-reichgeltii , though more commonly it information that may be useful in the determination has a compact form of inflorescence as in of these 5 taxa using morphological and anatomical J. conglomeratus . characteristics. In J. × kern-reichgeltii using both The hybrid J. × kern-reichgeltii was morphological and anatomical characters, there is considered on balance not to exist (Stace 1972) evidence for hybrids and introgression; ultimately, and Tweed & Woodhead (1949) reported they these characters provide a more critical could not find evidence for hybrids in their determination of not only the parent taxa, but also of studies. Ascherson & Graebner (1904) had their hybrids. reported J. × kern-reichgeltii (and J. inflexus × J. conglomeratus , for which there is no KEYWORDS : Sclerenchyma, epidermal ridge cells, introgression. evidence at present) from various locations in Britain and Europe. Kriša (1962) considered that J. effusus and J. conglomeratus were just INTRODUCTION end points of the same species. However, Agnew (1968) looked at populations and Within this group of rushes, subgenus Genuini , graphed various characters, such as spathe J. inflexus appears to be the most distinct with length against ridge number, which appeared to stiff glaucous stems that have interrupted pith suggest fertile hybrids existing in populations and lax, suberect branches, (Stace 1997). Two and that ‘introgression’ was occurring with other species, J. effusus and J. conglomeratus J. effusus ; given that fertility was high, this have at times been difficult to separate with a could have equally shown two end points of suggestion that characters overlap (Stace one species or two variable taxa. While 1970b, 1972). This is mainly due to these two Agnew’s (1968) work did suggest a range of species having forms in which the intermediates sometimes based on subjective inflorescence can be effuse or compact in either qualities such as inflorescence colour, though species; J. effusus var. subglomeratus DC a there was no real way of differentiating any compact from and J. conglomeratus var. distinct evidence for hybridity, it could have subuliflorus (Drejer) Asch. & Graebn. having been that one parent was very variable. Also to several stalked heads (Stace 1997). There are see and count the ridges in J. effusus the stems two sheets of in NMW which are labelled J. would have to be dry as it is more or less conglomeratus var. laxus A & G; Four smooth stemmed in life. *e-mail: [email protected] 44 M. WILCOX Fertility in Juncus can be interrupted even in rank and for interspecific hybrids. The hybrid the parental taxa as well as in hybrids often for J. × diffusus is depicted in transverse section different reasons. Juncus hybrids are often on the cover of Stace (1975) and is an excellent considered sterile or with very low fertility, example of an intermediate between the two (Stace 1970a, b, 1972). However, fertility is parents. Thus, while studying the parents of J. known in hybrid rushes ranging from partial × diffusus and this hybrid, two main fertility in some plants such as Juncus anatomical-morphological characteristics were articulatus L. × J. acutiflorus Ehrh. ex Hoffm. noted and these were studied and compared (J. × surrejanus Druce ex. Stace & Lambinon), between the parents and the hybrid. These (Blackstock & Roberts 1986) to almost no characteristics relate to the form, arrangement fertility, e.g. J. balticus Willd. × inflexus , and and type of subepidermal sclerenchymatous J. balticus × effusus ( J. × obotritorum girders, ( SeSgs ) and the shape (and other Rothm.), (both these latter hybrids having characteristics) of the epidermal ridge cells approx <1% fertility and seeds produced in (Ercs ) above these longitudinal strands; the these two taxa do germinate into full sized latter appearing to be useful and novel in the phenotypes, which are ± sterile also – identification of all these taxa. Therefore, the unpublished data M. Wilcox.) Due to this study was extended to J. × kern-reichgeltii as confusing situation, Stace (1975) comments, the characteristics noted may have had a that misidentifications are still not that similar type of inheritance thus possibly uncommon, though identifications are more separating it from the parents and even putative defined today at least for the species (Stace introgressed individuals. The previous 1997). In relation to the hybrids, J. inflexus × difficulties encountered in this latter hybrid J. effusus is relatively widespread being the may account for there being no mention of it most frequently recorded hybrid of the two and therefore a lack of any map in the new (Stace 1997, Preston et al. 2002) and is the Atlas of the British & Irish Flora (Preston et al. only well known hybrid of these two with a 2002) with most records unsubstantiated and few likely to be errors for odd or sterile J. based on field comparisons. This study sheds inflexus (Stace 1975, 1997). The hybrid J. × light on the identification of the parent taxa and kern-reichgeltii on the other hand has always both hybrids but in particular J. × kern- been an uncertain hybrid (Stace 1972) and reichgeltii . It shows a relationship between the today it is said to be difficult to identify, patterns of the subepidermal sclerenchymatous though Stace (1997) now supports the idea that girders and the epidermal ridge cell patterns it occurs sporadically with the parents and also and that these in turn will highlight the hybrids. states that due to its high fertility it is difficult In J. × kern-reichgeltii , it will show that this to determine other than in the field with its hybrid appears to be fertile and that back- parents and that many records are likely to be crossing can be frequent in suitable places erroneous. More recently, O’Mahony (2002) creating a hybrid range, though it appears there provided a more up-to-date key based on field is a more distinct form of the hybrid, (variable observations of external morphological chara- in fertility) which is recognisable from the cteristics. This key is likely to be of some use introgressed individuals. in detecting potential hybrids, but errors are still likely and given there is little conclusive evidence for hybridity due to its fertility, this METHODS key again may be describing large scale varia- tion in one of the parent taxa, namely J. cong- The methods employed in this study are lomeratus due to the presence of ridged stems. relatively simple so that they can be repeated Initially, this project started in the winter of by the amateur botanist with access to a 2006 with the hybrid J. × diffusus , which was compound microscope with approximately investigated to see if there were any ×20–100 magnification; this can be less once characteristics that would help to confirm its familiar with the parts. The photographs identity from any odd forms of the parent taxa, illustrated here may have used magnifications especially from sterile J. inflexus . Stace higher than this but this was for illustrative (1970b), states that the use of micro- purposes. anatomical/morphological features in Juncaceae Material from different populations of is clearly of immense taxonomic value not only approx 60–100 stems was used and other indi- at the level of subgenus but also at the specific vidual specimens from personal collections. IDENTIFICATION OF JUNCUS × DIFFUSUS AND J. × KERN-REICHGELTII 45 Many stems of the parents were also checked. These lines of epidermal ridge cells, (Ercs) A length of stem approximately 3 cm was cut were found to be different from other from about 1–2 cm below the inflorescence. epidermal cells and have no stomata. The shape This material, and if dried, was rehydrated in of the Ercs above the SeSgs was found to be warm distilled water until soft. These lengths different for each species. were then sectioned for transverse and longitudinal sections (TS/s LS/s) each being JUNCUS INFLEXUS : numbered and labelled and kept separate in This species generally stands out as being a glass tubes. A normal shaving razor was used pale somewhat grey-green glaucous rush. The and sections were cut as thin as possible and stem is clearly ridged and rather stiff and has stored initially in the distilled water then later distinctly interrupted pith (Fig.
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