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Morphological and Genetic Variation in the Endemic Seagrass Halophila Hawaiiana (Hydrocharitaceae) in the Hawaiian Archipelago L

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Morphological and Genetic Variation in the Endemic Seagrass Halophila Hawaiiana (Hydrocharitaceae) in the Hawaiian Archipelago L Morphological and Genetic Variation in the Endemic Seagrass Halophila hawaiiana (Hydrocharitaceae) in the Hawaiian Archipelago l Karla J. McDermid,2,3 Monica C. Gregoritza, 3 Jason W Reeves, 4 and D. Wilson Freshwater 4 Abstract: The endemic seagrass Halophila hawaiiana Doty & Stone is fOWld in discrete populations throughout the Hawaiian Archipelago. Morphological characteristics of plants from Midway Atoll, Pearl and Hermes Reef, Kaua'i, O'ahu, Moloka'i, and Maui were measured and compared. Striking variation in leaf length, leaf width, leaf length to width ratio, and internode length was evi­ dent among the 18 collection sites sampled at depths ranging from 0.32 to 18 m. DNA sequence analyses of a chloroplast-genome, single-base repeat locus in ramets from nine different collections found only two repeat haplotypes. Repeat haplotypes were fixed at all collection sites and for all islands except O'ahu. THERE ARE TWO species of seagrass fOWld in this Halophila species (Herbert 1986). A in the Hawaiian Islands. Halophila hawaiiana study by Michael-Taxis (1993) focused on the Doty & Stone was described as a new, en­ surface of H. hawaiiana leaves as a primary demic species in 1966. A second species, site for recruitment of epiphytes. She ex­ Halophila decipiens Ostenfeld, was recently re­ tensively described ecological contributions ported from Midway Atoll, O'ahu, and the of H. hawaiiana meadows on O'ahu, chemical island of Hawai'i (McDermid et al. 2002). content of the leaves, leaf ultrastructure, and Research on Halophila began with Doty and the anatomy of H. hawaiiana. Halophila ha­ Stone (1966), who recorded collections of H. waiiana plants support a native grazing snail, hawaiiana from the islands of Kaua'i, O'ahu, Smaragdia bryanae Pilsbry (Unabia 1984). Moloka'i, and Maui. Herbert (1984, 1986) The leaves of H. hawaiiana also serve as a studied the growth dynamics of one popula­ food source for other grazers such as the tion of H. hawaiiana; estimated biomass, threatened Hawaiian green sea turtle, Chelo­ productivity, and turnover; and developed a nia mydas (Linnaeus) Schweigger (Balazs growth model. High productivity levels were 1980, Russell and Balazs 2000, Balazs et al. observed and applied to a hypothesis about in press). However, knowledge of many other the pioneer-type successional characteristics aspects of H. hawaiiana biology and ecology is limited. The goals of this research were (1) to 1 D.W.F. and ].W.R. were supported by a contract systematically document the morphological from NOAA-NOS and a NSF REU supplement, and M.C.G. was supported by the UH Hilo Marine Science variation among Halophila hawaiiana popula­ Department Senior Thesis Program. Manuscript ac­ tions, both within and among islands in the cepted 6 August 2002. Hawaiian Archipelago, and (2) to conduct 2 Corresponding author. DNA analyses on specimens from different 3 Marine Science Department, University of Ha­ islands to assess genetic diversity among pop­ wai'i at Hila, 200 West Kawili Street, Hilo, Hawai'i 96720 (phone: 808-933-3906; fax: 808-974-7693; E-mail: ulations. Sequences of the intergenic spacers [email protected]). and intron of the trnL region of the chlo­ 4 Center for Marine Science, 5600 Marvin Moss roplast genome in Halophila species have a Lane, Wilmington, North Carolina 28409. number of single-base nucleotide repeats (D.W.F., Robert A. York, and ].W.R., Pacific Science (2003), vol. 57, no. 2:199-209 Wlpubl. data). In this study, variation at a © 2003 by University of Hawai'i Press repeat within the trnL intron was determined All rights reserved for separate ramets of H. hawaiiana. 199 30°.......------t-----+------i1------1-----i-T 500 km •G ~Midway Atoll .t;J" o-Pearl and Hermes Reef I~ I ~ .~ e (~(? 25° ~ • "" "A> \.!J'4' ~ '" Kaua'i .~. O'ahu ~Maui Molo~i~ 20° "" H~ai'iJ,.~ ~ / I , L-.L----------- -.,..J,r I,' ',I Northwestern Hawaiian Islands ,'Main ,Nawaiian Islands ,,' I 1 , I \ Johnston Atoll .,,' " i, lSoN.........-.----t-------t-----..,...:.,-'t-----+,'-t------+'-/---1. I I I 1~55 175"W I ° , I , , , I , , I , I , , , I , , / \ , I , , I Midway , I I , , \ , \ I / , / I I I I / I / \ / Olan 10 I / I! II \ I I / \ / I / \ Okm 10 -r:=::c::=' \ =-, I I I I, I, I \ I I "- (j) ® ® FIGURE 1. Map of the Hawaiian Archipelago. Locations of collection sites are numbered in order from north to south and correspond to site numbers in Table L Variation in Halophila hawaiiana . McDermid et al. 201 TABLE 1 List of 19 Collection Sites Throughout the Hawaiian Archipelago by Island from North to South Collection Site Collection No." Collection Date Depth (m) 1 Midway, cargo pier KM4317 20 September 1996 10 2 Midway, harbor breakwall KM4666 11 June 1999 1.5-3 M Midway, inner harbor KM 5411 27 July 2001 Drift 3 Pearl & Hermes Reef R074 9 October 2000 8 4 Kaua'i, 'Anini Beach Road Bishop 8464 21 February 2000 1-1.5 5 O'ahu, Mark's Reef, Kane'ohe MG35 29 January 2001 3 6 O'ahu, Coconut Island MG31 10 November 2000 0.6 7 O'ahu, Kahala Bishop 14171 21 May 1964 8 O'ahu, Kahala KM 16 28 September 1983 <1 9 O'ahu, Kahala KM 5355 8 June 2001 2 10 O'ahu, WaikIkI Natatorium D&S 3208 1 11 Moloka'i, Keawa Nui Pond KM915 29 November 1985 <1 12 Moloka'i, Keawa Nui Pond KM916 29 November 1985 <1 13 Moloka'i, 'Ohi'a Bridge Pond KM 5347 23 March 2001 0.32 14 Moloka'i, Hotel Moloka'i KM 5349 24 March 2001 1.5 15 Moloka'i, Hotel Moloka'i KM 5348 24 March 2001 1.5 16 Maui, Kihei MG32 15 January 2001 1 17 Maui, Lipoa Street MG33 15 January 2001 1 18 Maui, Makena Beach KM 5357 9 July 2001 18 n KM, K. McDermid; RO, R. Okano; Bishop, Bishop Museum; MG, M. Gregoritza; D&S, Dory and Stone. MATERIALS AND METHODS the outer edge of the blade tip), leaf width (at the widest point), and the internode length Morphometrics of plants from 18 collection between leaf pairs. Means of these measure­ sites on six islands, including Midway Atoll, ments were calculated. In addition, a mean Pearl and Hermes Reef, Kaua'i, O'ahu, Mo­ leaf length: width (L:W) ratio was calculated loka'i, and Maui (Figure 1, Table 1) were for each collection site or meadow. One-way analyzed using techniques based on Tomlin­ analysis ofvariance (ANOVA) was conducted son (1982), Williams and Dennison (1990), on the morphometric data. Larkum (1995), and Procaccini et al. (1999). Seagrass samples for DNA analysis were From November 2000 to July 2001, collec­ taken from nine collection sites between No­ tion sites were visited by wading, snorkeling, vember 2000 and July 2001. Individual sec­ or with SCUBA, and live voucher specimens tions of plants including a growing tip and were uprooted by hand or with a trowel, three leaf pairs along the rhizome were hand­ rinsed in seawater to remove sediment and collected at least 5 m apart, rinsed in sea­ debris, placed in seawater in a watertight water, cleaned of any epiphytes, and then container, and transported in a cooler to the placed in a plastic bag containing silica gel laboratory. Plants were photographed, pre­ desiccant (28-200 mesh) (Chase and Hills served as herbarium specimens that will be 1991). Dried samples in silica were mailed to donated to the Bishop Museum (BISH), and on two of the authors (J.W.R. and D.W.F.) at some occasions preserved in 4% formalin­ the Center for Marine Science, Wilmington, seawater solution. Individual plants (n = 20 North Carolina, for testing. Total genomic whenever possible) were chosen haphazardly DNA was extracted from two to three leaf from the collections, as well as from herbar­ pairs using the DNeasy Plant Mini Kit and ium sheets of collections made before No­ protocol (Qiagen, Inc., Valencia, California). vember 2000. The plants were measured An approximate 400-base pair (bp) section of using calipers to obtain values for leaf length the trnL intron was amplified using the for­ (extending from the node on the rhizome to ward primer 5'-GCA AAT CGG TAG ACG 202 PACIFIC SCIENCE· April 2003 CTA CG-3' and reverse primer 5'-GCA as demonstrated by the small standard error ATT TCG ATA GAA AGA TCC-3'. Am­ values. The leaf L:W ratios showed the plification reactions were set up in 50-J..tl vol­ greatest intermeadow variation. Plants in a umes containing 40-100 ng DNA template, Midway Atoll meadow (site 2) had the 1.25 mM MgCI2, 80 J..LM each dNTP, 0.2 J..LM shortest leaves with the smallest L:W ratio each flanking primer and HotStarTaq DNA (5.21), and plants from Maui (site 18) had polymerase and reaction buffer as suggested the longest, narrow leaves resulting in the by the manufacturer (Qiagen, Inc., Valencia, largest L:W ratio (41.23). The greatest var­ California). The thermocycling protocol fol­ iety of leaf forms was observed among col­ lowed that of Freshwater et al. (2000) with lection sites on Moloka'i. Plants from site 11 the addition of an initial enzyme activation on Moloka'i displayed the narrowest mean step of 15 min at 95°C. Reaction products leaf width, and plants collected at site 13 (less were cleaned of excess dNTPs, enzymes, than 3 km away) the widest. and primers using a GeneClean II Kit (Q­ No correlation was found between latitude biogene, Carlsbad, California) and used as and any of the morphological characteristics. templates in sequencing reactions set up us­ Yet ANOVA showed that for each of the ing the Big Dye Ready Reaction Kit and traits (leaf length, leaf width, L:W ratio, and protocol (Applied Biosystems, Foster City, internode length) there were significant dif­ California).
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