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Gametophyte Life-History Dominance of Chondrus Crispus (Gigartinaceae

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Gametophyte Life-History Dominance of Chondrus Crispus (Gigartinaceae NRC Publications Archive Archives des publications du CNRC Gametophyte life-history dominance of Chondrus crispus (Gigartinaceae, Rhodophyta) along the Atlantic coast of Nova Scotia, Canada McLachlan, Jack L.; Blanchard, Wade; Field, Christopher; Lewis, Nancy I. This publication could be one of several versions: author’s original, accepted manuscript or the publisher’s version. / La version de cette publication peut être l’une des suivantes : la version prépublication de l’auteur, la version acceptée du manuscrit ou la version de l’éditeur. For the publisher’s version, please access the DOI link below./ Pour consulter la version de l’éditeur, utilisez le lien DOI ci-dessous. Publisher’s version / Version de l'éditeur: https://doi.org/10.4490/algae.2011.26.1.051 ALGAE, 26, 1, pp. 51-60, 2011-03-15 NRC Publications Record / Notice d'Archives des publications de CNRC: https://nrc-publications.canada.ca/eng/view/object/?id=1b199bcd-c661-471d-abb6-d24921df9708 https://publications-cnrc.canada.ca/fra/voir/objet/?id=1b199bcd-c661-471d-abb6-d24921df9708 Access and use of this website and the material on it are subject to the Terms and Conditions set forth at https://nrc-publications.canada.ca/eng/copyright READ THESE TERMS AND CONDITIONS CAREFULLY BEFORE USING THIS WEBSITE. L’accès à ce site Web et l’utilisation de son contenu sont assujettis aux conditions présentées dans le site https://publications-cnrc.canada.ca/fra/droits LISEZ CES CONDITIONS ATTENTIVEMENT AVANT D’UTILISER CE SITE WEB. Questions? Contact the NRC Publications Archive team at [email protected]. If you wish to email the authors directly, please see the first page of the publication for their contact information. Vous avez des questions? Nous pouvons vous aider. Pour communiquer directement avec un auteur, consultez la première page de la revue dans laquelle son article a été publié afin de trouver ses coordonnées. Si vous n’arrivez pas à les repérer, communiquez avec nous à [email protected]. Research Article Algae 2011, 26(1): 51-60 DOI: 10.4490/algae.2011.26.1.051 Open Access Gametophyte life-history dominance of Chondrus crispus (Gigartinaceae, Rhodophyta) along the Atlantic coast of Nova Scotia, Canada Jack L. McLachlan1,a, Wade Blanchard2, Christopher Field2 and Nancy I. Lewis1,* 1National Research Council Canada, 1411 Oxford St., Halifax, NS B3H 3Z1, Canada 2Department of Statistics, Dalhousie University, Halifax, NS B3H 4J1, Canada adeceased December 2010 Similar to other species of Gigartinaceae Chondrus crispus has an alternation of perennial, isomorphic gametophytic and sporophytic generations. As these two generations co-exist independently within populations and obtain their re- sources in a similar manner, intraspeciic competition is expected. In populations within the southern Gulf of St. Law- rence, fronds of both generations of C. crispus occur in similar numbers. This equivalency can be related to substratum instability, where the population is dynamic with a high turn-over rate of genets. These observations support a stochas- tic hypothesis to account for distribution of gametophytes and sporophytes in this area. Along the Atlantic coast of Nova Scotia, where the substratum is stable, gametophytes are overwhelmingly predominant. Gametophytic predominance is greatest in the lower littoral zone where C. crispus is abundant and space is limited. Under the fucoid canopy where “free-space” exists, the gametophyte to sporophyte ratio is lower. Gametophytic and sporophytic fronds are distributed equally among different size-classes and size-distribution is not considered a competitive factor. Previous studies have shown that sporophytic fronds of C. crispus are more susceptible to infections by endophytic algae and other pathogens, and are more heavily grazed by herbivores than are gametophytic fronds. Thus, mechanistic factors are strongly implied in the selection of gametophytes in the Atlantic population. Key Words: carrageenan; Chondrus; gametophyte; Geni coeficient; life-history ratio; population structure; size hierar- chy; sporophyte INTRODUCTION A regular alternation of independent generations oc- with other life history patterns evolving (e.g., Dixon 1965); curs in the life histories of many algal species, including indeed, there are many species in which sporophytes are the Rhodophyta. Stebbins and Hill (1980) proposed that apparently the dominant generation. Valero et al. (1992) maintenance of a haploid-diploid life history is an adap- pointed out that it is dificult to rationalize the mainte- tation to seasonally variable environments and that dip- nance of a haploid-diploid life history when the ecologi- loidy is more adaptable than haploidy. This implies that cal niches of the two generations are similar, particularly in populations with distinct, alternate haploid and dip- where the two life-history stages are isomorphic. In such loid generations, the trend is towards diploid dominance, instances, the life-history phases are ecologically inde- This is an Open Access article distributed under the terms of the Received 30 January 2011, Accepted 22 February 2011 Creative Commons Attribution Non-Commercial License (http://cre- Corresponding Author ativecommons.org/licenses/by-nc/3.0/) which permits unrestricted * non-commercial use, distribution, and reproduction in any medium, E-mail: [email protected] provided the original work is properly cited. Tel: +1-902-426-8273 Fax: +1-902-426-9413 Copyright © The Korean Society of Phycology 51 http://e-algae.kr pISSN: 1226-2617 eISSN: 2093-0860 Algae 2011, 26(1): 51-60 pendent and, especially if the two generations are peren- McLachlan 1991). The turnover rate for fronds is relative- nial, they will be ecologically competitive, each genera- ly high (Bhattacharya 1985), and largely relates to size, tion obtaining its resources in a similar manner. In such although individual fronds may persist for several years, intraspeciic competitive situations, it can be queried depending upon physical conditions (Taylor et al. 1981). whether one generation can become predominant in a There is no evidence of seasonality of fronds as may occur population and whether this is predictable. May (1986) in other gigartinacean species (cf. May 1986, De Wreede argued that if the two generations are demographically and Green 1990). Perennation occurs through the basal comparable, over time, the proportion of gametophytes holdfast, which can persist for indeinite periods (Taylor and sporophytes should eventually reach equivalency et al. 1981). Once detached, holdfasts will not reattach, because of stochastic events. However, if the two gen- and detached fronds will not generate holdfasts. Growth erations are not equivalent, one of the generations will of plants occurs through expansion of the holdfast with predominate according to May’s mechanistic hypothesis. a continuous production of new fronds (McLachlan et In those species where one life-history generation al. 1989), whereas recruitment into new space occurs dominates in a population, does this extend to all pop- through spore reproduction. The frequency of reproduc- ulations of the species? Reasons for dominance would tive fronds within a population has been reported to be therefore be inherent rather than the result of unique cir- relatively low (Prince and Kingsbury 1973, Chen and Tay- cumstances. This question has been addressed a number lor 1980, Lazo et al. 1989, McLachlan et al. 1989, Taylor of times with respect to the Gigartinaceae, resulting in and Chen 1994). ambiguous conclusions. Gametophytic and / or sporo- Species with isomorphic alternate generations, such phytic dominance have been reported, both for species as C. crispus, should be ideally suited to study intraspe- of Mazzaella (as Iridaea) and Chondrus (e.g., Mathieson ciic competition. Species of Gigartinaceae are particu- and Burns 1975, Hansen and Doyle 1976, Craigie and larly useful in this regard as sporophytic and gameto- Pringle 1978, Bhattacharya 1985, Dyck et al. 1985, Hann- phytic phases have different carrageenans, lambda type ach and Santelices 1985, May 1986, Luxoro and Santelices and kappa type, respectively (McCandless 1981), which 1989, De Wreede and Green 1990, Fernández and Mené- can be distinguished by a simple, rapid colorimetric test ndez 1991, Piriz 1996), whereas Lazo et al. (1989) noted involving resorcinol. Accordingly, the life history phase of a similar proportion of gametophytes and sporophytes fronds can be identiied in their vegetative state, allow- in populations in their study of Chondrus crispus Stackh. ing large numbers of samples to be assessed easily and Chondrus crispus (Irish moss) is the sole species of quickly (Craigie and Pringle 1978, Dyck et al. 1985, Lazo its genus present along shores of the Atlantic Ocean, et al. 1989, De Wreede and Green 1990). where it is widely distributed and abundant throughout In a previous investigation of C. crispus in the south- northern temperate waters, especially in Maritime Can- eastern Gulf of St. Lawrence along the coast of Prince Ed- ada (McLachlan et al. 1987, McLachlan 1991, Taylor and ward Island, where Irish moss is a commercial resource Chen 1994). In some areas, C. crispus occurs essentially (McLachlan et al. 1987), the gametophytic / (tetra)sporo- as ‘monospeciic’ stands, dominating space in the lower phytic (G/S) ratio of the overall population was found to littoral and upper sublittoral zones, in suficient abun- be essentially
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