Genetic Variation in the Epiphytes Tillandsia Ionantha and T. Recurvata (Bromeliaceae)
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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/237979359 Genetic Variation in the Epiphytes Tillandsia ionantha and T. recurvata (Bromeliaceae) Article in American Journal of Botany · April 1987 DOI: 10.2307/2443832 CITATIONS READS 34 75 4 authors, including: Douglas E Soltis University of Florida 995 PUBLICATIONS 55,423 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Systematics of the Saxifragaceae View project Systematics and evolution of Amaryllidaceae tribe Hippeastreae View project All content following this page was uploaded by Douglas E Soltis on 30 August 2016. The user has requested enhancement of the downloaded file. Genetic Variation in the Epiphytes Tillandsia ionantha and T. recurvata (Bromeliaceae) Author(s): Douglas E. Soltis, Amy Jean Gilmartin, Loren Rieseberg and Sue Gardner Source: American Journal of Botany, Vol. 74, No. 4 (Apr., 1987), pp. 531-537 Published by: Botanical Society of America, Inc. Stable URL: http://www.jstor.org/stable/2443832 Accessed: 30-08-2016 12:53 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/2443832?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms Botanical Society of America, Inc. is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany This content downloaded from 128.227.189.58 on Tue, 30 Aug 2016 12:53:31 UTC All use subject to http://about.jstor.org/terms Amer. J. Bot. 74(4): 531-537. 1987. GENETIC VARIATION IN THE EPIPHYTES TILLANDSIA IONANTHA AND T. RECURVATA (BROMELIACEAE)1 DOUGLAS E. SoLTIs, AMY JEAN GILMARTIN, LOREN RIESEBERG, AND SUE GARDNER2 Department of Botany, Washington State University, Pullman, Washington 99164, and 2Corpus Christi Botanic Garden, PO Box 81183, Corpus Christi, Texas 78412 ABSTRACT The genetic structure of two epiphytic species, Tillandsia ionantha and T. recurvata, was investigated using enzyme electrophoresis. Electrophoretic data suggest that T. ionantha and T. recurvata differ in breeding system, in agreement with predictions based on their strikingly different floral morphologies. Electrophoretic data suggest extremely high levels of inbreeding for T. recurvata, whereas Tillandsia ionantha exhibits characteristics of an outcrossing species. Values of P, H, and mean number of alleles per locus are much higher in T. ionantha than in T. recurvata. The mean value of FIS for T. ionantha is low (0.056), closely approaching expec- tations at Hardy-Weinberg equilibrium. In contrast, the mean value ofFIS in T. recurvata (1.000) indicates a complete absence of heterozygotes. The two species also differ in genetic structure. Low FST values for T. ionantha indicate little variation in allele frequencies among populations. In contrast, FST values are high for T. recurvata, suggesting substantial genetic heterogeneity among populations. In addition, the mean value of I is higher in T. ionantha (0.995) than in T. recurvata (0.931). Population genetic data are in agreement with the suggestion of Benzing (1978), who proposed that extreme epiphytes such as T. recurvata, would be characterized by increased autogamy ensuring high seed set. Due to high chromosome numbers in Bromeliaceae (most taxa have x = 25), the family has been considered polyploid. However, with the exception of an additional isozyme for PGM in T. recurvata, the two species are isozymically diploid. IN RECENT REVIEWS (e.g., Hamrick, Linhart and Tillandsia (Bromeliaceae) is a predomi- Mitton, 1979; Loveless and Hamrick, 1984) nantly epiphytic genus confined to tropical and population genetic data are provided for plants subtropical regions of North and South Amer- representing a number of life history strategies. ica. Most species of Tillandsia are at least fa- Noticeably absent from these reviews, how- cultative epiphytes, but different species ex- ever, are plants representing the epiphytic hab- ploit habitats ranging from mesic to xeric it. Epiphytes are well-represented in the world's (Gilmartin and Brown, 1986). As reviewed by flora; approximately 13% of higher vascular Benzing (1978, 1980) xeric and mesic epiphyt- plant species and 27% of fern species are epi- ism are considered to be related ecological phytes (Kress, 1986). Among angiosperms, at strategies located at opposite extremes along a least 2% of Magnoliopsida (dicots) and 44% of single adaptive continuum. As aridity in- Liliopsida (monocot) species are epiphytes creases, rates of plant growth, maturation, and (Benzing, 1986; Kress, 1986). Despite the flower number decline (Benzing, 1978). Benz- prevalence of the epiphytic habit, population ing (1978) suggested that because of the ex- genetic data apparently have not been provid- treme environmental and developmental con- ed for a single epiphyte. In addition, several straints that characterize xeric (or extreme) ecological features make epiphytes intriguing epiphytes belonging to Tillandsia, as well as organisms for population genetic studies. For the unstable and patchy nature of their sub- example, epiphytes usually exist in short-lived strates, these taxa would be characterized by habitats such as the bark of trees. By necessity, mechanisms that ensure high seed set. The epiphytes are exceptionally successful coloniz- mechanisms proposed are increased autogamy ers, but their geographic distribution is patchy. and cleistogamy. In support of this is the pres- ence of cleistogamy in some ofthe most xerical- ly adapted species of Tillandsia (Gilmartin and I Received for publication 10 March 1986; revision ac- Brown, 1985). cepted 2 September 1986. If xeric and mesic epiphytes are character- The assistance and helpful comments of Pam Soltis are ized by different breeding systems, this should gratefully acknowledged. This research was supported in part by NSF grants BSR 84-07573 to AJG and BSR- be reflected in the genetic structure of the 8315098 to DES. species. To begin testing this hypothesis, as well 531 This content downloaded from 128.227.189.58 on Tue, 30 Aug 2016 12:53:31 UTC All use subject to http://about.jstor.org/terms 532 AMERICAN JOURNAL OF BOTANY [Vol. 74 pected to promote at least a tendency toward outcrossing, whereas the latter would be ex- pected to promote selfing. Greenhouse grown plants of T. ionantha rarely set seed, whereas plants of T. recurvata regularly set seed. MATERIALS AND METHODS-A total of 243 plants, each from a distinct clump or clone, representing three populations of Tillandsia ionantha and seven populations of T. recurvata were collected in the field (Table 1) and main- tained in greenhouse culture at Washington State University. All populations sampled were separated geographically by at least 50 km, icm[L with the exception of populations 6 and 7 of A B C T. recurvata. (These were distinct populations from opposite sides of a single host tree.) Young leaves from mature plants were used for elec- trophoretic analysis because they provided the best resolution of banding patterns. The following enzymes were analyzed: al- cohol dehydrogenase (ADH), fluorescent es- terase (FE), glutamate dehydrogenase (GDH), glyceraldehyde-3-phosphate dehydrogenase l]mm ([NAD]G3PDH and [NADP]G3PDH), isoci- trate dehydrogenase (IDH), malate dehydro- genase (MDH), phosphoglucoisomerase (PGI), phosphoglucomutase (PGM), 6-phosphoglu- A B conate dehydrogenase (6PGD), and triose- Fig. 1, 2. 1. Tillandsia ionantha, habit and longitudinal phosphate isomerase (TPI). Samples were pre- section of flower showing exserted genitalia (stamens and pared and electrophoresis conducted following pistil). (Modified from Smith and Downs, 1977.) 2. Til- the general methods of Soltis, Haufler, Darrow, landsia recurvata, habit and longitudinal section of flower and Gastony (1983). The tris-HCl grinding showing the deeply included genitalia (stamens and pistil) buffer-PVP solution of these authors was em- (modified from Smith and Downs, 1977). ployed. For all enzymes a 12.5% starch gel was used. ADH, FE, GDH, PGI, and TPI were as to supply initial insights into the population resolved on a modification of gel and electrode genetic structure of epiphytes, samples of Til- buffer system 8 of Soltis et al. (1983). System landsia ionantha Planchon var. ionantha and 9 was used for MDH, PGM, and 6PGD; sys- T. recurvata (L.) L. were examined using en- tem 1 was used for [NAD]G3PDH and zyme electrophoresis. [NADP]G3PDH. For IDH, systems 1 and 2 Tillandsia ionantha and T. recurvata are ob- were employed. ligate epiphytes or occasionally saxicoles. Til- When more than one locus was observed for landsia recurvata is considered to be an ex- an enzyme, loci were numbered sequentially, treme epiphyte; T. ionantha is semi-xeric in with the most anodally migrating locus des- habit, and not found in as drought-stressed ignated 1. Enzyme variants at individual loci areas as T. recurvata. Tillandsia ionantha is were given letters, with the fastest migrating found