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Limbella Fryei (Williams) Ochyra Distinct from L Journ. Hattori Bot. Lab. No. 63: 395-410 (Dec. 1987) LIMBELLA FRYEI (WILLIAMS) OCHYRA DISTINCT FROM L. TRICOSTATA (SULL.) CM. (MUSCI: AMBLYSTEGIACEAE) JOHN A. CHRISTyl ABSTRACT. Investigation of branching patterns, morphology, karyotype, isozymes and shoot growth confirmed specific differences between Limbella tricostata (Sull .) C. M. and L. /ryei (Williams) Ochyra. Significant differences were observed in branch bud frequency, leaf insertion angle, leaf areola­ tion, chromosome length, electrophoretic mobility of superoxide dismutase, and shoot regeneration. Chromosome numbers of both taxa are n = l1. A key and descriptions segregate the taxa. INTRODUCTION In Moss Flora of the Pacific Northwest, Lawton (1971) reduced the pleurocarpous North American moss Limbellafryei - then known as Sciaromiumfryei - to a synonym of the Hawaiian L. tricostata, because she could find no significant morphological dif­ ferences between plants from the two areas. Morphology of plants from the two regions is superficially similar, but closer inspection indicated to me that differences might exist in branching pattern and leaf insertion angle. The Hawaiian L. tricostata exhibits branching at more or less regular intervals up to I cm apart, giving the plants a pinnate or frondose habit, especially in aquatic forms (Fig. I). In contrast, branches of the North American L. fryei arise in more closely-spaced groups, giving the plants a penicillate, brush-like appearance. Plants are therefore nearly always dendroid (Fig. I). In both dried and living material, leaves of Hawaiian plants are appressed, whereas those of North American plants are erect to wide-spreading (Fig. 3). Also, habitat conditions for Limbella in Hawaii and Oregon are in most respects strikingly dissimilar (Christy 1980). These differences in morphology and habitat, coupled with a disjunctive distri­ bution of 4,600 km of open ocean and 25 ° of latitude, indicated that the Hawaiian and North American populations might not be conspecific, as had been proposed by Lawton (1971). Christy (1985) investigated the differences in order to determine whether or not plants from the two areas were conspecific. MATERIALS AND METHODS 1. Quantification of Branching Patterns. Branching pattern in Limbella was quantified by calculating branch bud frequency per merophyte interval. Bud frequencies were deter­ mined by (1) counting the number of merophytes along the stem, following the spiral growth 1 Botany Section, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233, U.S.A. 396 Joum. Hattori Bot. Lab. No. 63 198 7 FIG. 1. Limbella fricosfata (left of scale) and L. fryei (right of scale). Scale = 5 cm. pattern from the lowermost bud found, toward the distal end of the shoot, until a bud or branch was encountered, (2) recording that merophyte number, (3) resuming the count at one with the next merophyte, following the spiral pattern again until the next bud or branch was encountered, (4) recording that merophyte number and (5) repeating the sequence until the shoot apex was reached. This count gave a measure of the number of times a bud or branch was encountered per merophyte interval. The numbers of buds occurring at each merophyte interval were totaled, the percent occurrence at each merophyte interval (= bud frequency) calculated, and the frequencies plotted against the corresponding merophyte in­ tervals. Twenty shoots (1, J 34 merophytes) of L. tricOSfafa and 17 shoots (689 merophytes) of L. fryei were counted. Differences in frequency were analyzed by an R x C test of inde­ pendence (Sokal & Rohlf 1981). 2. Phenetic Analysis. Twelve specimens of L. tricostafa and 12 of L. fryei, including types, were scored for 28 morphological characters (Table 1). The data were analyzed by one­ way analysis of variance (ANOY A) using the Statistical Analysis System package (SAS Institute 1985). The data for two variables (shortest basal leaf cell length, leaf tooth length) were log transformed prior to analysis to normalize their distributions. Sporophytes were not scored because those of L. fryei are unknown. Leaves and stem sections were measured 1.0- 1.5 cm below the tips of main shoots or largest branches with intact apices. Leaf apical angle and marginal tooth angle were measured using an ocular protractor (Wagner 1951; Christy 1987). J. A. CHRISTY: Limbellafryei distinct from L. tricostata 397 TABLE 1. Morphological characters measured in Limbella tricostata and L. fryei. 1 . Shoot length (cm) 2. Branch length, longest (cm) 3. Leaf insertion angle (degrees) 4. Leaf length, shortest (mm) 5. Leaf length, longest (mm) 6. Leaf width, at widest part (mm) 7. Leaf apical angle, excluding mucro (degrees) 8. Leaf tooth length, longest (jtm) 9. Upper leaf cell length , shortest (jtm) 10. Upper leaf cell length, longest (,Im) 11 . Upper leaf cell width, shortest (jtm) 12 . Upper leaf cell width, longest (jtm) 13. Upper leaf cell length: width 14. Basal leaf cell length, shortest (jtm) 15. Basal leaf cell length, longest (jtm) 16 . Basal leaf cell width, shortest (,lm) 17 . Basal leaf cell width, longest (jtm) 18. Basal leaf cell length : width 19. Costa width at insertion (jtm) 20 . Limbidial width at insertion (,Im) 21. Number of cell layers in costa 22 . Costa thickness (Ilm) 23 . Number of cell layers in limbidia 24. Limbidial thickness (pm) 25 . Stem diameter, longest dimension (Ilm) 26 . Number of cells in stem diameter 27 . Width of stem cortical layer (1/m ) 28 . Number of cells in stem cortical layer 3. Growth Chamber Studies. Sods of living L. tricostata and L. fryei were maintained 2 1 for nine months in a growth chamber under fluorescent lighting of 600--800 pE m- s- , 16 hr light: 8 hr dark at 18°C and 100 % RH, conditions thought to approximate those in the fie ld. Specimens of L. tricostata were grown immersed in tap water and subjected to circula­ tion and aeration. Specimens of L. fryei were placed in closed, transparent plastic boxes and misted with tap water 3- 5 times weekly. Portions of collections of one taxon were also grown in the same container with the other taxon for ten weeks, simultaneously exposing each to the environmental conditions experienced by the other. Growth responses of L. tricostata and L. fryei were observed by examining new shoots produced by Limbella fragments in vitro. Shoots of both taxa were stripped of existing branches, cut into 1 cm segments, and equal numbers from each taxon placed together on moist filter paper in petri plates. The segments were firmly attached to the filter paper with monofilament fishing line. The petri plates were kept in a growth chamber for ten weeks, during which time new shoots developed from buds. Twelve replicate plates were prepared in this manner, containing 120 shoot segments, 60 of each taxon. 4. Isozyme Banding Patterns. Isozyme banding patterns of Limbella were obtained by means of horizontal starch gel electrophoresis, following the methods of Helenurm (1983) and 398 J oum. Hattori Bot. Lab. No. 63 1 9 8 7 Lay ton and Ganders (1984). Living plants were procured from field-collected material that had been maintained in a growth chamber for 1-4 months. Whole plants of 0.2- 3.0 g wet weight were immersed in liquid nitrogen and ground to a powder in a chilled mortar (Fahselt 1980). A few drops of distilled water (Szweykowski et al. 1981 ) were added to the thawing powder to make a paste which was transferred to chilled 1.5 ml polypropylene micro test tubes and centrifuged under refrigeration for 15 min at 20,000 g. The extracts were obtained from (1) field-collected plants that had been maintained in a growth chamber (53 individual Hawaiian plants from four different populations on one island, and 49 Oregon plants from one population), (2) sporelings grown from spores from two different populations on two different islands and (3) reciprocally-cultivated growth chamber material. Extracts were run simultane­ ously on two gels, one with morpholine citrate buffer (Clayton & Tretiak 1972) and one with Ridgway buffer (Ridgway et al. 1970). The buffer system used in the trays was the same as that used in the particular gel being run. Gels were run at ca. 50 mAmps for 3-4 hours, and were stained for 12 enzymes. After initial trials, five enzyme assays were selected because they stained most clearly : malic dehydrogenase (MDH), phosphoglucose isomerase (PG I), phos­ phoglucomutase (PG M), 6-phosphogluconate dehydrogenase (6PGDH) and superoxide dis­ mutase (SOD). 5. Karyotype Morphology. About 150 squash proparations were made from shoot and branch apices, using 14 plants from four different Hawaiian populations and I1 plants from one North American population. Attempts to obtain chromosome figures from L. tricostata sporelings (sensu Nehira 1984) were unsuccessful. Material was obtained from field-collected plants that had been maintained in a growth chamber 4- 5 months. Steel's (1978) procedures were followed with some modifications adopted from Beeks (1955) and Przywara and Kuta (1983). Differences in relative lengths (relative length = absolute length : total length of chro­ mosome complement) were analyzed by one-way analysis of variance (Sokal & Rohlf 1981). 6. Phenolic Spot Patterns. Dried plants were treated following the methods of Gornall and Bohm (1980). Two samples of L. tricosfafa were tested (37.3 g and 69.6 g, respectively) and one of L. fryei (46.0 g) . One-gram quantities of the taxa were also tested separately. Both two-dimensional and sequentially-spotted one-dimensional chromatograms were run . RESULTS 1. Quantification of Branching Patterns. The results indicated that the frequency of branch bud patterns was dependent on moss species (P < .000 I, G = 206; R x C test of independence). In Hawaiian plants, buds occurred most often every fourth mer­ ophyte (26.4 %), with lesser peaks every first, ninth and eleventh merophtye (Fig. 2). A few buds were separated by up to 20 merophytes.
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