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Isoetes ×Blondeaui Hyb. Nov. (Isoetaceae), a New Hybrid Quillwort from Eastern Québec, Canada

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Isoetes ×Blondeaui Hyb. Nov. (Isoetaceae), a New Hybrid Quillwort from Eastern Québec, Canada Botany Isoetes ×blondeaui hyb. nov. (Isoetaceae), a new hybrid quillwort from eastern Québec, Canada Journal: Botany Manuscript ID cjb-2020-0065.R2 Manuscript Type: Article Date Submitted by the 08-Jun-2020 Author: Complete List of Authors: Brunton, Daniel; Brunton Consulting Services, Sokoloff, Paul; Canadian Museum of Nature, Research and Collections Isoetes ×blondeaui, Isoetes laurentiana, Isoetes echinospora, freshwater Keyword: tidal marsh,Draft St Lawrence River Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/botany-pubs Page 1 of 18 Botany 1 Isoetes ×blondeaui hyb. nov. (Isoetaceae), a new hybrid quillwort from eastern 2 Québec, Canada 3 4 D. F. Brunton1 and, P. C. Sokoloff2 5 1 Beaty Centre for Species Discovery and Botany Section, Canadian Museum of Nature, Ottawa, Ontario K1P 6 6P4, Canada [email protected] 7 2 Beaty Centre for Species Discovery and Botany Section, Canadian Museum of Nature, Ottawa, Ontario K1P 8 6P4, Canada [email protected] 9 10 ABSTRACT 11 The hybrid between diploid Isoetes echinospora and tetraploid I. laurentiana is reported from six sites along 12 the St. Lawrence River near Québec City. Morphological characteristics and distinguishing features are 13 identified and the binomial I. ×blondeaui, hyb. nov. is proposed. All occurrences are in biogeographically 14 significant freshwater tidal marsh habitat. The hybrid is expected to occur in small numbers within most large 15 I. laurentiana populations. Draft 16 17 Isoetes laurentiana, Isoetes echinospora, Canada, Québec, St Lawrence River, freshwater tidal 18 marsh, lycophyte 19 20 The recent discovery of the endemic lycophyte Isoetes laurentiana D.F.Brunton (Isoetaceae) 21 (Brunton et al. 2019) adds to the list of remarkable vascular plant taxa of the freshwater tidal marsh 22 habitat found within an approximately 150 km section of the St. Lawrence River in eastern Québec 23 (QC), Canada (Figure 1). Investigation of that species provided the opportunity to search for hybrids 24 between it and cohabiting taxa. Sterile hybrids are recognized to be important in the evolution and 25 taxonomy of vascular plants (Plume et al 2015; Gianguzzi et al. 2017; Goulet et al. 2017) by 26 confirming the existence of genetic barriers between species pairs. The development of polyploid 27 species through alloploidy within hybrids is also recognized as being particularly important in 28 the evolution of quillwort (Isoetes) taxa (Taylor et al. 1985; Hickey et al. 1989; Brunton and 29 Troia 2018; Schafran et al. 2018; Brunton and Sokoloff 2020). 30 Field investigations for potential hybrids were undertaken within the floristically diverse 31 freshwater tidal marsh habitat of Isoetes laurentiana (Figure 2). This unique habitat is confined to the 32 lower St. Lawrence River and harbours an exceptional number of endemic or near-endemic taxa https://mc06.manuscriptcentral.com/botany-pubs Botany Page 2 of 18 2 33 from various vascular plant groups (Brouillet et al. 2004; Brunton et al. 2019). A global Key 34 Biodiversity Area (KBA), tentatively named The St. Lawrence Fluvial Estuary KBA, has recently 35 been identified here (C. Raudsepp-Hearne, Canadian KBA Co-ordinator, pers. comm., May 2020). 36 While I. laurentiana is locally abundant in this unique habitat, the only other quillwort taxon known 37 there (rarely) is I. echinospora M.Durieu (Brunton et al. 2019). Accordingly, these two taxa were 38 considered the likely putative parents of any hybrid found during these investigations. 39 40 Methods 41 Field investigations focussing on three freshwater tidal marsh Isoetes laurentiana populations 42 were undertaken over three days in mid August 2018 along the St. Lawrence River in eastern 43 Québec. August and September are the peak months for mature spores to be present in Isoetes 44 specimens in eastern Canada (Cody and Britton 1989). One site (Sainte-Petronille, MRC L’lle 45 d’Orleans) was visited twice while the others (Marais-Léon-Provancher Reserve, MNR Portneuf and 46 Saint-Augustin, Communaute de Québec),Draft received a single inspection. Particular attention was paid 47 to specimens demonstrating usually large size relative to their associates, this being a common 48 demonstration of hybrid vigour (Britton 1991; Brunton and Sokoloff 2020). Over 60 herbarium 49 specimens (approximately 250 plants) from this freshwater tidal marsh habitat from CAN, DFB and 50 (especially) MT were also examined in 2018 and 2019 (herbaria acronym of Thiers 2020). 51 For Scanning Electron Microscope (SEM) images produced during the present study, air 52 dried spores were attached to SEM stubs by means of adhesive carbon discs. These were sputter 53 coated with a gold / palladium alloy (Au/Pd) and examined with a 2017 model FEI Apreo SEM (at 54 15 kV and 25 pA, with a working distance of 10 mm and a spot size of 6). 55 The physical characteristics of the Isoetes specimens examined, especially their megaspore 56 form, ornamentation, and size, were viewed through a Leica Wild M3B dissecting microscope at 40× 57 magnification, with the aid of an in-mount graticule (ocular micrometer) for measurements. 58 Megaspore sizes represent the average width (across the equatorial region) of at least 20 (and 59 typically 40) spores. Comparable microspore measurements are based on a minimum of 20 60 longitudinal measurements taken from SEM images of clusters of spores. 61 The determination of ploidy is of great value in identifying suspected Isoetes hybrids 62 (Hickey et al. 1989; Taylor and Hickey 1992; Brunton and Sokoloff 2020). The ploidy level of https://mc06.manuscriptcentral.com/botany-pubs Page 3 of 18 Botany 3 63 the suspected hybrid plants was inferred from that of the putative parents, the latter determined 64 from chromosome counts or flow cytometry investigations reported in Taylor et al. (1993) and 65 Brunton et al. (2019). 66 67 Results 68 The conclusive identification of the hybrid nature of a particular Isoetes plant is usually 69 straightforward, even in old herbarium material and without a living sample from which the ploidy 70 level can be determined. Particularly distinctive characteristics for the detection of Isoetes hybrids 71 have been documented in populations around the world, including misshaped, aborted megaspores 72 (Britton 1991; Taylor et al. 1993) with dense, congested ‘brain-coral-like’ ornamentation (Britton 73 and Brunton 1996), misshaped microspores often including individuals with little or no 74 cytoplasm, and intact microspores and megaspores being of intermediate size and ornamentation 75 vis-a-vis the putative parents (Taylor and Luebke 1988; Musselman et al. 1995; Brunton and 76 Troia 2018). Draft 77 In field investigations in 2018, an apparent hybrid plant with those distinctive morphological 78 characteristics was detected within an Isoetes laurentiana population near Québec City. Ten more 79 plants were detected in I. laurentiana herbarium vouchers in MT collected between 1935 and 1995. 80 These apparent hybrid plants are from six distinct locations (Figure 1). They appear to represent the 81 triploid (2n=3x=33) hybrid of tetraploid I. laurentiana (2n=4x=44) with diploid I. echinospora 82 (2n=2x=22). 83 Figure 3 compares megaspore characteristics (shape, size and ornamentation) typical of the 84 suspected hybrids against those of putative parents Isoetes laurentiana and I. echinospora. The 85 congested ornamentation pattern of densely-packed, loosely interconnected muri (walls) with 86 scattered intervening spines/ narrow tubercles of the hybrid (Figures 3C and 3D) contrasts with the 87 more open, regularly broken-reticulate pattern of I. laurentiana megaspores (Figures 3A and 3B) and 88 the uniformly spiny (echinate) ornamentation of I. echinospora (Figures 3E and 3F). 89 A narrow and often obscure band of short spines is present along the distal (lower) side of the 90 equatorial ridge of the hybrid (Figure 3C), a feature that reflects the spiny equatorial band present in 91 I. laurentiana (Figure 3A) but absent in I. echinospora (Figure 3F). Similarly, the regularly globose https://mc06.manuscriptcentral.com/botany-pubs Botany Page 4 of 18 4 92 shape of the putative parents (Figures 3B and 3F) contrasts with the flattened (lens-shaped) form of 93 the hybrid (Figure 3C). 94 Microspore ornamentation of the hybrid (Figure 4B) is more clearly intermediate between 95 that of a perispore (surface) abundantly covered with broad-based spines as with I. laurentiana 96 (Figure 4A), and that of a plain to sparsely thin-spiny ornamented perispore as with I. echinospora 97 (Figure 4C). 98 Atypically, spore size is not as significant a factor with this hybrid because of the 99 exceptionally small spore size of putative tetraploid parent Isoetes laurentiana which fall within the 100 upper size range typically seen amongst North American diploid Isoetes (Brunton et al. 2019). The 101 megaspores of the hybrid, for example, average 429.14 µm (1 SD 39.89, n=110, five plants), which 102 is less than either I. laurentiana (460 µm - Brunton et al. 2019) or I echinospora (450 µm - Britton 103 and Brunton 1999). 104 Similarly, microspore length apparently does not distinguish itself as a significant 105 distinguishing factor between this hybrid Draftand its putative parents. Limited microspore data suggest a 106 size of 27.9 µm (1SD 1.6 µm, n=50), which is marginally larger than that of I. laurentiana (27.1 µm 107 - Brunton et al. 2019). These data are from only a single specimen, however (F. Coursol et D. 108 Bouchard 95-524, MT 8387) and may represent an aberration. 109 Isoetes laurentiana is unique amongst eastern North American tetraploid Isoetes in having a 110 velum coverage over the sporangium of less than 10%; this is a particularly conspicuous and 111 significant distinguishing character (Brunton et al. 2019). Accordingly, the 14.5 to 33% (n=9, three 112 plants) velum coverage of these hybrid plants is distinctively intermediate between that of I.
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